AMERICAN aa FERN 0 JOURNAL PUBLISHED BY THE AMERICAN FERN SOCIETY EDITORS David W. Bierhorst Gerald J. Gastony David B. Lellinger John T. Mickel MERCURY PRESS, ROCKVILLE, MARYLAND 20852 CONTENTS Volume 72, Number 1, Pages 1-32, Issued March 29, 1982 A Brief Report on the Progress of Pteridological Research in China CHING REN-CHANG and WANG ZHU-HAO The Asplenium trichomanes Complex in the United States and Adjacent Canada ROBBIN C. MORAN The First Record of Platycerium ridleyi in Sumatera N. A. P. FRANKEN and M. C. ROOS Comparison of Chromatographic Spot Patterns of Some North American Isoétes Species L. S. KOTT and D. M. BRITTON Anatomical Studies of the Neotropical Cyatheaceae. II. Metaxya and Lophosoria TERRY W. LUCANSKY Aloysio Sehnem (1912-1981) Shorter Notes: An Alternative Simplified Medium oe lanosa and Cystopteris rotund in Flori Reviews Volume 72, Number 2, Pages 33-64, Issued June 28, 1982 The Chromosomes of Lycopodium lucidulum JOSEPH M. BEITEL and FLORENCE S. WAGNER Axenic Culture and Induction of Callus on Sporophytes of the Appalachian Vittaria Gametophy JAMES D. ge MARK WHITTEN, and MICHAEL J, BECK Cystopteris x illinoens a New Natural Hybrid Fern ROBBIN C. MORAN New Records and Distributional Notes on Maryland Pteridophytes STEVEN R. HILL and RICHARD E. RIEFNER, JR. Grammitis succinea, the First New World Fern Found in Amber LUIS DIEGO GOMEZ P. A New Tropical American Species of the Genus Lycopodium BENJAMIN @LLGAARD Specimens from Fée’s Pteridological Collection at the Botanical Garden of Rio de Janeiro PAULO GUNTER WINDISCH Shorter Notes: A Germination Method for Isoétes; A New Station for Lygodium palmatum; The Name of a Hybrid x Asplenosorus; The Deletion of Nephrolepis pectinata from the Flora of Florida; A Filmy Danaea Reviews 40, 44, 48, > pony oo tw fue) a -_ an pee Oo wn w Volume 72, Number 3, Pages 65-96, Issued October 27, 1982 prs Biosystematic, and Nomenclatural Notes on Scott’s Spleenwort, x Asplenosorus ebenoides KERRY S. WALTER, WARREN H. WAGNER, JR. and FLORENCE S. WAGNER Cheilanthes feei New to Virginia THOMAS F. WIEBOLDT and STAN BENTLEY A New Woodsia Hybrid from Kansas RALPH E. BROOKS Polystichum lonchitis in Central Quebec—Labrador MARCIA J. WATERWAY and THOMAS T. LEI A Unique Type of Microsporangium in Selaginella Series Articulatae PAUL SOMERS Cystopteris tennesseensis in Illinois ROBBIN C. MORAN Dieter E. Meyer (July 21, 1926—February 1982) Shorter Note: Flavonoid Chemistry of the North American Lycopodium obscurum Complex Review Volume 72, Number 4, Pages 97-124, Issued December 31, 1982 Pteris x delchampsii, a eenertne Fern meter from Southern Florid . WAGNER, JR. and CLIFTON E. NAUMAN C-glycosylxanthones in the Asplenium adiantum-nigrum Complex P. MICK RICHARDSON and EUGENIA LORENZ-LIBURNAU Taxonomic Notes on Jamaican Ferns-IV GEORGE R. PROCTOR pein amelguita, a New Species of Fossil Fern the Potomac Group (Lower Cretaceous) JUDITH E. SKOG American Fern Journal Index to Volume 72 Errata AMERICAN es FERN | peas JOURNAL ~ APR 96 1982 @ARDEN us RARY QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY A Brief Report on the Progress of Pteridological Research in China CHING REN-CHANG and WANG ZHU-HAO | The Asplenium trichomanes Complex in the United States and Adjacent Canada ROBBIN C. MORAN 5 The First Record of Platycerium ridleyi in Sumatera N.A.P. FRANKEN and M.C. ROOS 12 Comparison of Chromatographic Spot Patterns of Some North American Isoétes Species L.S. KOTT and D.M. BRITTON 15 Anatomical Studies of the Neotropical Cyatheaceae. II. Metaxya and Lophosoria TERRY W. LUCANSKY 19 Aloysio Sehnem (1912-1981) Shorter Notes: An Alternative Simplified Medium for Growing Prothallia; Polystichum munitum on Baranof Island, Southeastern Ajaska; Cheilanthes lanosa and Cystopteris protrusa in Florida Reviews 11, 14, 18 The American Fern Society Council for 1982 DEAN P. WHITTIER, Dept. of Biology, Vanderbilt University, Nashville, TN 37235. hee TERRY R. WEBSTER, Biological Sciences Group, University of Connecticut, Storrs, CT 0626 Vic ee MICHAEL I. COUSENS, Faculty of Biology, University of West Florida, Pensacola, FL 32504 Sec retary JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, TN 37916. Treasurer JUDITH E. SKOG, Dept. of Biology, George Mason University, Fairfax, VA 22030. Records Treasurer DAVID B. LELLINGER, Smithsonian Institution, Washington, DC 20560. Journal Editor ALAN R. SMITH, Dept. of Botany, University of California, Berkeley, CA 94720. Memoir Editor JOHN T. MICKEL, New York Botanical Garden, Bronx, NY 10458. Newsletter Editor American Fern Journal DAVID B. LELLINGER U.S. Nat'l Herbarium NHB-166, Smithsonian Institution, Washington, DC 20560 ASSOCIATE EDITORS DAVID. W. BIERHORST Rt. 3, Box 188, Picayune, MS 39466. GERALD J. GASTONY Dept. of Biology, Indiana University, Bloomington, IN 47401. JOHN T. MICKEL New York Botanical Garden, Bronx, NY 10458. The “American Fern Journal” (ISSN 0002-8444) is an illustrated quarterly devoted to the general study of ferns. It is owned by the American Fern Society, and agent at the Smithsonian Institution, Washington, DC 20560. Second-class postage paid at Washin Claims for missing issues, made 6 months (domestic) to sf aise (foreign) after the date of issue, and the matters for publication should be addressed to the Editor. Changes of address, dues, and applications for membership should be sent to Dr. Judith E. Skog, Dept. of Biology, George Mason University, Fairfax, VA 22030. Orders for back issues should be addressed to Dr. James D. Montgomery, Ichthyological Associates, R.D. 1, Berwick, PA 18603. General inquiries concerning ferns should be addressed to the Secretary. Subscriptions $9.00 gross, $8.50 net if paid through an agency (agency fee $0.50); sent free to members of the American Fern Society (annual dues, $8.00; life membership, $160.00). Back volumes 1910-1978 $5.00 to $6.25 each; single back numbers of 64 pages or sees $1.25: 65-80 pages, $2.00 each; over 80 pages, $2.50 each, plus shipping. Back volumes 1979 et seq. $8.00 each: single back numbers $2.00 each, plus shipping. Ten percent discount on orders of six ike or more. Library Dr. John T. Mickel, New York Botanical Garden, Bronx, NY 10458, is Librarian. Members may borrow books at any time, the borrower paying all shipping costs. Newsletter : John T. Mickel, New York Botanical Garden, Bronx, NY 10458, is editor of the newsletter “Fiddlehead Forum.” The editor welcomes contributions from members and non-members, i including miscellaneous notes, offers to Septet or purchase materials, personalia, horticultural notes, and reviews of non-technical books on fern Spore Exchange Mr. Neill D. Hall, 1230 Northeast 88th Street, Seattle, WA 98115, is Director. Spores exchanged and collection lists sent on request Gifts and Bequests Gifts and bequests to the Society enable it to expand its services to members and to others interested in ferns. Botanical books. back issues of the Journal, and cash or other gifts are always welcomed, and are tax-deductible. Inquiries should be addressed to the Secretary. AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 1 (1982) ] A Brief Report on the Progress of Pteridological Research in China CHING REN-CHANG* and WANG ZHU-HAO** Historical sketch.—Like other branches of botany, serious study of modern pteridology in China was not launched until 1927, when the senior writer decided to switch his interest from Chinese dendrology to pteridophytes. There was then no one in China who knew ferns or how to study ferns. There was no literature available except Engler and Prantl’s “Die Natiirlichen Pflanzenfamilien’” (1898-1902), and there was not a single fern specimen correctly identified in the small herbarium that had just been initiated in Beijing. It seemed unimaginable to start studying ferns under such hopeless circumstances. It was his strong will that surmounted all the difficulties he met with at the beginning. For literature, for example, he wrote to the then world authorities on ferns, such as H. Christ, C. Christensen, W. R. Maxon, and E. B. Copeland, asking for reprints of their papers and monographs on ferns. Thanks to their generous help, he obtained within a few years practically all the available literature chiefly concerning Asiatic ferns of that time to start his work with. After a few years of preparation, including extensive fern collecting particularly in the fern-rich provinces south of the Yangtze River and making preliminary identifi- cations by consulting the literature then available, he soon realized that type specimens are the key to plant taxonomy. The status of any taxon must be decided by its type specimen, not by its description. Therefore, he decided to visit Europe in the early spring of 1930. He went first to Copenhagen where, under the direction of Dr. Carl Christensen, with whom he had had contact by previous correspondence, he began seriously to carry out a study of Chinese ferns. Both Christensen and he participated in the Fifth International Botanical Congress at Cambridge, England in the fall. At the Congress, he met in person with great joy all of the then leading pteridologists from the United States, Germany, England, and other countries. After the Congress, he stayed at Kew for over a year, mostly working on Chinese and Himalayan ferns and taking photographs of type specimens, as well as making critical notes on the types. Afterwards he went back to Copenhagen and stayed there for some months to discuss problems with Christensen concerning the classification of ferns in general and the guiding principles of classification in particular. Leaving Copenhagen in the early spring of 1932, he proceeded to visit most other leading European herbaria as far away as Vienna and Prague, looking up type specimens of ferns and taking photographs. After he returned from Europe in the fall of 1932, he joined the Fan Memorial Institute of Biology in Peiping and devoted himself with great zeal to the cause of Chinese fern study. During the period of the Japanese invasion of China, he evacuated to Yunnan, collecting and studying the rich fern *Institute of Botany, Academia Sinica, Beijing, China. **South China Institute of Botany, Academia Sinica, Guangdong, China. Volume 71, number 4, of the JOURNAL, was issued December 30, 1981. 2 AMERICAN FERN JOURNAL: VOLUME 72 (1982) flora there. After liberation in 1949, he was transferred to the Institute of Botany, Academia Sinica, Beijing, as head of the Taxonomic Section, where he has continued his fern studies until now. Fern hunting in China.—In 1911, the British fleet surgeon Dr. C. G. Matthew published in the Journal of the Linnaean Society of London an “Enumeration of Chinese ferns” totalling a little over 1,000 species then known in China, of which a considerable number have subsequently been reduced as synonyms of earlier- described taxa. Today, after seventy years have elapsed, the species of ferns and fern-allies known in China have been estimated at roughly 2,200 species, about doubling Matthew’s enumeration. Yet this is far from the final figure for the Chinese fern flora. In the spring of last year, for example, the senior writer had a chance to identify four fern collections from different parts of China and found the number of new species is still mounting. In a collection made by the Sino-American Joint Botanical Expedi- tion to Western Hupeh, in central China, 13 new species of ferns were found in five weeks time, including one extraordinary Matteuccia. From another collection made in southwestern Yunnan, nine new species have been identified, one a species of Platycerium, a genus first recorded from China in the evergreen rain forest in the border of Upper Burma. In a third, small collection from northeastern China (formerly known as Manchuria), at least two species are new, one of them a Phyllitis, a genus never known in China before. Five new species were recognized from a collection made in Chekiang province along the eastern coast of China. From these instances, one will certainly be amazed at the extraordinary richness of the Chinese fern flora. In our estimate, by the end of the third decade of the twenty-first century, the number of Chinese ferns will most likely have increased by at least three to four hundred more species. This is simply because of the vast expanse of China’s extremely rugged topography and diversified ecology. The richness of her flora in many provinces never can be exhaustively explored by only a few visits. The famous sacred Mount Omei in Szechuan, for example, has been visited by at least 60 botanists, both foreign and Chinese, since 1883; yet just last year three more new ferns were discovered by the cytotaxonomical team of the Institute of Botany, Academia Sinica. Consequently, fern hunting in China is still a most rewarding adventure. One can easily collect 20-40 different specimens in a day, depending upon the locality one may visit. An exactly similar situation exists with China’s phanerogam flora. Fern herbaria.—As pointed out above, prior to 1927 there were no fern herbaria in China, and in fact only very few specimens had been collected. Strange to say, before 1927 the senior writer had collected thousands of herbarium specimens of flowering plants in different provinces, but very few ferns, in which he then had no interest. Through the effort of many botanists and collectors all over China during the last 50 years or so, the fern herbarium at the Institute of Botany, Beijing, has grown quickly and now has in excess of 150,000 numbers, of which about 95% are of Chinese origin. It is already in a position to serve as a working herbarium for the study of Chinese ferns. In addition to the herbarium specimens, there are kept over 1,000 photographs of types and isotypes of ferns taken during the senior writer’s CHING & WANG: PTERIDOLOGICAL RESEARCH IN CHINA 3 tour in Europe. There is now little difficulty in identifying Chinese fern collections in the herbarium. All other affiliated botanical institutions of the Academia Sinica and most universities in different provinces or regions also have a herbarium of local ferns and fern-allies for compiling local Floras and for research use. Nevertheless, local herbaria still have a long way to go as far as the sufficiency of their collecting activities. At present, even many botanically rich areas within their reach still remain little explored. As a matter of fact, their first and foremost task should be to launch extensive botanical explorations in their respective provinces or regions for many years to come. To meet the urgent need of housing a steady influx of specimens from all parts of the country, a new, modern, 6-storied National Herbarium is under construction and will be completed by the end of 1982. The herbarium of the Institute of Botany will be removed from the Beijing Zoo to its new site in the Botanical Garden, Beijing. Fern research.—In the past 55 years, a considerable amount of research on pteridophytes has been carried out in China, with fern taxonomy the chief activity. Many short papers, monographs, iconographies, and some fern Floras have been published. With over 2,000 species of ferns and fern-allies, the study of Chinese pteridophytes is no mean task, and taxonomic study will continue to be an important aspect of pteridological research in China for many years to come. However, for the purpose of enhancing the scientific level of taxonomic studies, other aspects of the biology of ferns should also be studied. In the past 23 years, a great amount of work on fern spores has been conducted. As a result, a monograph on spore morphology of Chinese ferns, “Sporae Pteridophytorum Sinicorum,” was published in 1976. It is a 414-page volume dealing with 52 families, 174 genera, and over 1,000 species of Chinese pteridophytes. It includes numerous drawings and photographs, a chapter on the geography of Chinese ferns, and another on fossil spores in different geological times. Preparation was made as early as 1964 for the cytotaxonomical study of Chinese ferns, when about 300 species of ferns had been introduced into cultivation in the Beijing Botanical Garden. In the following years, work on chromosome counts was launched successfully. Unfortunately, the entire work was frustrated by the so-called “cultural revolution,” and 300 well grown potted ferns were thrown out of the greenhouse in the winter of 1966. However, with the visit of Mr. A. C. Jermy to China in the fall of 1978 and the lectures he gave on the biosystematic study of Dryopteris, the interest in cytotaxonomy in China was promptly renewed. Studies on chromosomes are being carried out at several institutions at present. Fern Flora of China.—There will be five volumes of the “Fern Flora of China” as part of the 80-volume “General Flora of China.” The first volume was published in October, 1959. Two other volumes were ready for printing, but, alas, this, too, was frustrated by the “cultural revolution” for a time. The remaining two volumes are now in preparation. It is hoped that the first edition of the “Fern Flora of China” will all be published shortly after 1985. Other fern Floras published up to now are the “Fern Flora of Hainan, vol. 1” (1964), “Flora Tsinlingensis, Pteridophyta, vol. 2” (1974), “Herbaceous Flora of Northeastern China, vol. 1” (1959), and the “Flora of 4 AMERICAN FERN JOURNAL: VOLUME 72 (1982) Jiangsu Province, vol. 1 (Pteridophyta).”” The “Fern Flora of Tibert” was scheduled for publication about the end of 1981 as part of the “Flora of Tibet.” Several local fern Floras are in preparation, including those for the provinces of Szechuan, Fukien, and Yunnan, each with 400—1,000 species. It is a strenuous task for local botanists to complete their respective fern Floras. It has been planned that each Chinese province or region will eventually have its own Flora with the pteridophyta as a component. To coordinate and promote fern studies all over the country, the Botanical Society of China in the fall of 1979 set up a section of Chinese Pteridology as an affiliated organ. At the inaugural meeting held in October, 1979 at Tian Mo Shan, Chekiang province, the senior writer was elected president of the section. A plan for developing and popularizing fern study in China was formulated at this meeting. Fern systematics.—Since 1940, the classification scheme pteridophytes has attracted the attention of many fern students and has undergone great changes. Up to now, there have evolved at least 11 different systems by different authors, of which the first breakthrough was “On natural classification of the family ‘Polypodiaceae’” by the senior writer (Sunyatsenia 5:201—270. 1940), which was soon followed by Copeland’s and Holttum’s classifications. In 1954, the senior writer put forth a paper titled “Systematic arrangements of families and genera of Chinese pterido- phytes with corresponding names in Chinese” (Acta Phytotax. Sinica 3:93-99). In 1978, he published a treatise on “The Chinese fern families and genera: Systematic arrangement and historical origin” (Acta Phytotax. Sinica 16(3):1—19, 16(4):16~37. 1978). In spite of all the defects of one kind or another in these systems, we ma well say that the phylogenetic study of ferns has made decidedly marked progress in the last 40 years over the Hookerian and Dielsian systems with which we are all familiar. It may be presumed that to most fern students the interrelationships of fern families and genera as a whole are now self-evident, although a few taxa still remain uncertain as to their proper systematic position, as indicated in the lastest classifica- tion scheme by the senior writer. It may be that many of these isolated ferns are survivors from ancient times whose ancestors or relatives are no longer in existence today, nor are their fossil records available so far. In systematic treatments, we prefer to keep these taxa isolated as such, rather than to combine them with apparently not closely related families. This is of course entirely a matter of personal taste. Regarding this and other problems of fern systematics, the late, revered Carl Christensen had liberal views, but his rather early death prevented him from putting all these down in his writings. AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 1 (1982) 5 The Asplenium trichomanes Complex in the United States and Adjacent Canada ROBBIN C. MORAN* The Maidenhair Spleenwort, Asplenium trichomanes L., is a delicate, evergreen fern that grows in rock crevices or, less frequently, on talus slopes. The species is wide ranging, occurring in North America, Europe, Australia, New Zealand, and Asia. Since Manton (1950) reported the existence of diploid and tetraploid cytotypes in European populations of A. trichomanes, both cytotypes and their sterile triploid hybrids have also been found in North America (Britton, 1953; Wagner & Wagner, 1966). The diploid and tetraploid cytotypes that occur in North America have been named, the diploid as subsp. trichomanes and the tetraploid as subsp. quadrivalens D. E. Meyer emend. Lovis. This paper examines for the first time the occurrence and relative abundance of all three A. trichomanes cytotypes in the United States and Canada, and compares this information to what is known about this species complex in Europe. METHODS Spore size was used to determine cytotype. To determine the validity of this approach, a pilot study was undertaken that involved an examination of spore length from 30 herbarium specimens that were cytologically determined by Drs. F. S. and W. H. Wagner. Tetraploids consistently had much larger exospore diameters than the diploids (Fig. /). In fact, differences in spore size were so obvious that laborious measurements of spore size were unnecessary to distinguish diploids from tetra- ploids. The total number of plants examined for spore size was 115 (1725 spores) for the diploids, and 52 (810 spores) for the tetraploids. Triploid plants were recognized by the presence of aborted spores. Thus, for this study, the presumed cytotype was extrapolated from spore size or spore abortion rather than direct chromosome counts. Over 700 herbarium specimens were examined from the following institutions: EIU, F, ILL, ILLS, ISM, MICH, MO, NCU, NHA, RM, SIU, TENN, VDB, and WTU. For each specimen examined, a permanent spore slide was made using Hoyer’s Solution as a mounting medium and sealing the cover slip with clear nail polish enamel. After the slides had been examined, they were placed in a paper envelope and glued to the herbarium sheet for permanent reference; the specimens were annotated indicating the presumed cytotype. Locality data of individual collections are available from the author. RESULTS AND DISCUSSION The mean spore size for diploid plants was 29.4 ym (s=2.3) and for tetraploid plants 41.0 pm (s=2.6). These results support those of Lovis (1964), who found similar differences in mean spore size of A. trichomanes cytotypes in Europe. *Illinois Natural History Survey, Natural Resources Bldg., 607 E. Peabody Dr., Champaign, IL 61820. 6 AMERICAN FERN JOURNAL: VOLUME 72 (1982) DIPLOID TETRAPLOID mean = 29.4 um mean = 41.0 um €ev. = 2.5 dey. = L146 ym “100 oO O) Q gD S >. UY g e FIG. 1. Outlines of random spore samples mounted in Hoyer’s Solution. A. Virginia, Giles County, limestone along Sinking Creek, | mile north of Newport (specimen cytologically det.), W. H. Wagner 64113 (MICH). B. Illinois, Hardin County, sandstone outcrop in Blind Hollow, southeast of Lamb, R. A. Evers 20091 (ILLS). C. Kansas, Chatauqua County, 3.5 miles east and 4 miles north of Sedan, wooded sandstone canyon, L. Hauser 3134 (NCU). D. Same location as A, specimen cytologically det., W. H. Wagner 64115 (MICH); E. Virginia, Loudoun County along Potomac River, north-northeast of Leesburg, H. E. Ahles 61196 (NCU). F. Canada, British Columbia, limestone cliffs at head of Cumshewa Inlet near Moresby logging camp, Moresby Island, part of Queen Charlotte Islands, J. A. Calder & R. L. Taylor 36254 (NCU). European workers (Jermy et al., 1978; Jermy & Page, 1980; Lovis, 1964) have found that plants of diploid and tetraploid A. trichomanes can usually be distin- guished without recourse to micro-characters. However, this is extremely difficult, if not impossible, to do satisfactorily from herbarium specimens; thus the reliance in this study on the more constant character of spore size. The morphological characters listed in the key given below are best observed in living plants. Lovis (1964) has published shadow diagrams that illustrate morphological differences in well developed fronds of diploid and tetraploid plants. North American pteridolo- gists would do well to become familiar with the characters listed in the key below R. C. MORAN: ASPLENIUM TRICHOMANES COMPLEX 7 and to pay close attention as to their applicability in living specimens of North American plants. Plants mostly of non-calcareous rocks; rhizome scales up to 3 mm long; stipe relatively thin, shiny ery or bronze red; fronds arching upwards and outwards, away from rock faces; pinnae mostly alternate, suborbicular, more distant, thinner, set more oblique to the rachis, the basiscopic margin often auriculate, pinnae usually shed from the rachillae during the first winter, the rachillae retained during first winter, except in exposed habitats; sori 4-6(9), up to 2 mm long; indusia delicate, not conspicuous, spores generally 29-36 jm long; guard cells generally 38-43 ym long; plants diploi subsp. trichomanes Plants of calcareous rocks; rhizome scales up to 5 mm long; stipe relatively thicker, blackish- brown; rarely auriculate; pinnae usually retained during the first winter, the rachillae shed with, or soon after, the pinnae; sori 4-9(12), up to 3 mm long; indusia thicker, conspicuous; spores generally 34-43 jm long; guard cells generally 41-49 ym long; plants tetraploid ........ subsp. quadrivalens The diploid is the most abundant and widespread cytotype, constituting 87% of the specimens examined. Specimens of this cytotype from the southwestern United States (Fig. 2) are from high altitudes in isolated mountain ranges, such as the Rincon and Santa Catalina mountains in southern Arizona, where several collections were made between 2500 and 3000 meters elevation. Diploids are found primarily on acid rock types such as sandstone, basalt, olivine, etc. However, 12 specimens were collected from limestone. There was no discernable geographic pattern to the diploid plants found on limestone. Tetraploid plants are primarily northern in distribution, being common in the middle Great Lakes and New England regions (Fig. 3). Approximately 12% of the 700 specimens “age were tetraploid. This cytotype appears to be an obligate calciphile, as eve ecimen examined was collected from limestone habitats. In Europe and Austaite Lovis (1964) also found the ons rear cytotype an ee to calcareous habitats. Wagner and Wagner (1966) studied a population of A. trichomanes on limestone cliffs along Sinking Creek, about | mile cats of Newport, in Giles County, Virginia. From a random sample of 89 plants, 85 proved to be tetraploid, 3 triploid, and 1 diploid. The abundance of tetraploids at this site emphasizes the calciphilous nature of this cytotype. One may wonder why the tetraploid is not more widespread, as suitable limestone habitats certainly exist in many localities outside its main range, such as the Ozarks of Missouri. The distribution of diploid and tetraploid plants in the Great Lakes region also graphically demonstrates their substrate preferences. The tetraploid is restricted to the Niagara escarpment of Silurian limestone, which forms an arc running from Door County, Wisconsin, across the lower northern peninsula of Michigan to Manitoulin Island, the Bruce Peninsula, and thence southeastward to Niagara Falls in New York. Southwest of Door County, Wisconsin, and southeast of Niagara Falls, the Niagara limestone escarpment becomes gradually covered by glacial tills and is not exposed. The great number of herbarium specimens from locations on the Niagara escarpment suggests that the tetraploid is fairly common there. In contrast, the diploid in the Great Lakes region occurs primarily on Precambrian igneous and metamorphic rocks, such as those that form the Keweenaw peninsula and the escarpments along the north shore of Lake Superior. 8 AMERICAN FERN JOURNAL: VOLUME 72 (1982) FIG. 2. The distribution of diploid Asp/enium trichomanes in the United States and southern Canada. FIG. 3. The distribution of tetraploid (dot) and triploid (triangle) Asplenium trichomanes in the United States and southern Canada. R. C. MORAN: ASPLENIUM TRICHOMANES COMPLEX 9 The triploid is the rarest cytotype; only seven collections (1.0%) were seen from the eastern United States (Fig. 3). The triploid occurs sporadically where the diploid and tetraploid populations overlap in the eastern Appalachians. Limestone was given as the rock type in all cases where herbarium labels provided such information. It is important to note, since many triploid plants are apogamous, that there was no indication of large, globose, unreduced spores in any of the triploids examined. Furthermore, in the above-mentioned population of A. trichomanes studied b Wagner and Wagner (1966), the triploids appeared to be the result of in situ hybridization from diploid and tetraploid plants occurring at the site. Figures 2 and 3 show only the main range of A. trichomanes in the United States and southern Canada, but A. trichomanes also occurs sporadically eastward in Canada to Newfoundland (Cody, 1968) and northward as isolated occurrences in the western cordillera of Alberta and British Columbia to southeastern Alaska at ca. 58° north latitude (Hultén, 1968; Scoggan, 1978). I have not seen specimens from these extremes of the species range. Asplenium trichomanes also occurs on limestone rocks on the Queen Charlotte Islands of British Columbia. All collections examined from this locality were tetraploid, which agrees with the cytological determinations of Taylor and Mulligan (1968). The disjunct A. trichomanes individual reported by Cody (1968) from Newfoundland is probably a diploid since it occurred on serpentine rocks. It is important to note that hexaploid plants are known from Madeira, Australia, and New Zealand. However, only in New Zealand is the hexaploid a prevalent cytotype (Lovis, 1977). No evidence of hexaploid plants was found in this study, based on spore size, although their occurrence in North America could be expected from chromosome doubling of a triploid plant. No apparent morphological differences were found between European and North American A. trichomanes. Indeed, the fact that the same rock habitat differences exist between the diploid and tetraploid cytotypes in both Europe and North America suggests that they are the same taxa. Furthermore, no specimens of A. trichomanes subsp. inexpectans Lovis, a diploid subspecies known only from central and southeastern Europe, were found among diploids in North America which grew on limestone. This subspecies is similar to subsp. trichomanes, but occurs only on calcareous rocks and has a slightly different frond morphology (Lovis, 1964, fig. 2). Unlike other Asplenium species in the eastern United States, A. trichomanes rarely hybridizes and its genome is not implicated in any widespread allopolyploid taxa. In Europe, however, a diploid form of A. trichomanes has hybridized with diploid A. viride Huds. giving rise to the fertile allotetraploid known as A. adulterinum Milde., which is widely dispersed in northern and central Europe (Lovis, 1977). Four sterile. interspecific hybrids involving A. trichomanes have been found in the eastern United States. these are: A. Xclermontae Sim (A. ruta-muraria X 4x A. trichomanes), A. X virginicum Maxon (supposedly A. platvneuron X 2x A. trichomanes), X Asplenosorus herb-wagneri (Taylor & Mohlenbrock) Mickel (A. pinnatifidum X 2x A. trichomanes) and XA. shawneensis R. C. Moran (2x A. trichomanes X Camptosorus rhizophyllus). Each of these hybrids is exceedingly rare. being known from only one or two localities. It 10 AMERICAN FERN JOURNAL: VOLUME 72 (1982) is extraordinary that x A. shawneensis has been found only once, especially since its two parents often grow side-by-side throughout much of the well botanized eastern a States (Moran, 198 Certain aspects of the origin of tetraploid A. trichomanes remain enigmatic. The Shae cytotype appears to be of autopolyploid constitution, as shown by the high levels of autosyndetic chromosome pairing seen oe various hybrids (Lovis et al., 1966; Lovis & Reichstein, 1969; Vida, 1970) and an experimentally produced polyhaploid sporophyte (Bouharmont, 1972). pea Lovis (1977) postulates that the origin of tetraploid A. trichomanes is polyphyletic, in that tetraploid forms may have arisen by different means at different times and in different parts of the world. he North American tetraploid cytotype may have originated by direct autopolyploidy from diploid forms or by hybridization between diploid subspecies, followed by chromosome doubling, producing intersubspecific autopolyploidy (Lovis, 1977, p. ). Whatever the case may be, the calciphilous nature of the tetraploid must be taken into account when postulating its polyploid origin from a diploid ancestor. The tetraploid’s remarkable habitat preference for limestone is not what would be expected from direct autopolyploidy of a diploid taxon that predominantly occurs on acid rocks. Perhaps the North American tetraploid arose via autopolyploidy from a calciphilous diploid; or perhaps this diploid, calciphilous race hybridized with the more common acidic rock race, with subsequent allopolyploidy, and in so doing imparted a dominant adaptive gene complex enabling the tetraploid to thrive in calcareous habitats. Although many techniques seem inadequate to resolve such a tentially complex evolutionary history, an analysis of isozyme variation from populations throughout the entire range of the complex might yield interesting results. I would like to thank the following for their help with various aspects of the study: Drs. Almut Jones, James H. Peck, W. Carl Taylor, Charles Werth, Florence S. Wagner, and Warren H. Wagner, Jr. Special thanks is given to Dr. Kenneth R. Robertson of the Illinois Natural History Survey for providing the writer with research time and encouragement for this project. LITERATURE CITED BOUHARMONT, J. 1972, Meiosis and fertility in apogamously produced diploid plants of Asplenium trichomanes. Chromosomes Today 3:253—258 BRITTON, D. M. 1953. Chromosome studies on ferns. Amer. J. Bot. 40:575-58 CODY, W. J. 1968. Asplenium trichomanes new to Newfoundland. Amer. Fern J. — 179-180. HULTEN, E. 1968. Flora of Alaska and Reeenly Territories. Stanford Univ. Press, Stanford, CA. JERMY, A. C., H. R. ARNOLD, L. FARRELL, and F. H. PERRING. 1978. Atlas of Ferns of the British Isles. Botanical Society of the British Isles and British Pteridological Society. London. , and C. N. PAGE. 1980. Additional field characters separating the subspecies of Asplenium trichomanes in Britain. Brit. Fern Gaz. 12:112-113. LOVIS, * D. 1964. The taxonomy of Asplenium trichomanes in Europe. Brit. Fern Gaz. 9:147-160. . 1977. Evolutionary patterns and processes in ferns. Pp. 229-415 in R. D. Preston and H. W. Woolhouse (eds.). Advances in Botanical Research, vol. 4. Aaa Press, New York. ,H. MELZER, and T. REICHSTEIN. 1966. Asplenium stiriacum D. E. Meyer emend. und A. * aprutianum hybr. nov., die zwei Asplenium lepidium x trichomanes-Bastarde. Bauhinia 3:87-101. R. C. MORAN: ASPLENIUM TRICHOMANES COMPLEX 11 MANTON., I. 1950. Problems of Cytology and Evolution in the Pteridophyta. Cambridge Univ. Press. ridge. ee R. C. 1981. x Asplenosorus shawneensis: a new es. fern a between Asplenium ichomanes and Camptosorus rhizophyllus. Amer. Fern J. 71:85— scoGGAN H. J. 1978. The Flora of Canada, ee bina gett enue Monocotyledoneae. nad. Natl. Mus. Sci. Publ. Bot. 7:9 TAYLOR, R: LL. and GcA. MULLIGAN. ine tas of the Queen Charlotte oye Part: 2. Cytological Aspects of the Vascular Plants. Canad. Dept. Agr. Monogr. 4. Par VIDA. G. 1970. The nature of polyploidy in Asplenium ruta-muraria L. and A. ee C. Presl. ret ee 23:525-5 a WAGNER, W. H.. Jr. and F. S. WAGNER. 1966. Pteridophytes of the yporiees Lake Area. Giles Co.. Virginia: biaeenan studies. 1964-1965. Castanea 31:121— REVIEW “ILLUSTRATION OF THE PTERIDOPHYTES OF JAPAN, VOLUME I,” by Satoru Kurata and Toshiyuki Nakaike. University of Tokyo Press. 628pp. 1979. Yen 50,000.—This book represents the first volume of a series of works devoted to illustrating and mapping the more than 600 species of Japanese (including the Ryu-Kyu and Bonin Islands) pteridophytes. This volume treats 100 species from a variety of families and includes large segments, though regrettably not all. of many genera such as Dryopteris, Polystichum, and Pteris. The treatment of individual species includes photographs depicting habit, line drawings of whole plants as well as details of sori, venation, cuticle, scales, and rhizome anatomy. a short description of the species, and an enumeration of specimens examined with a full page grid-map of Japan showing their location. The entire volume is beautifully produced with excellent paper and binding, something of which more American publishers ought to be enviou Although the descriptive material is entirely in Japanese ( Latin names are included). the illustrations and detailed distributions will be quite useful to Ameri- can botanists, not only pteridologists. but phytogeographers and ecologists as well. It is hard to ignore, for instance, the close resemblance of our Dryopteris ludoviciana with D. tokvoensis after examining the photograph and illustration of the latter. Although it is unlikely that many individuals will be able to purchase this book, they should certainly encourage their institutional libraries to do so.— R. Cranfill, Department of Botany, University of California, Berkeley, CA 94720. 12 AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 1 (1982) The First Record of Platycerium ridleyi in Sumatera N. A. P. FRANKEN* and M. C. ROOS** Platycerium ridleyi Christ, one of the well-known staghorn ferns, was thought to be restricted to a few localities in Malaya, Singapore, northeastern Borneo, and the Lingga Archipelago. It was not known to occur on Sumatera (formerly Sumatra). In general, this species is regarded as rare (Ridley, 1926, p. 110); few collections have been made during the last 20 years, and a to Holttum (pers. comm.), this species must be regarded as extinct in Singapore. During some field trips in Jambi Pesos Central Sumatera, in 1980, the authors visited two places where P. ridleyi appeared to be rather common. One locality is the Nature Reserve of Berbak along the river Air Hitam Laut (+ 1°25’S, 104°30’E) about 15-25 km upstream from the east coast. The second locality is in the center of the island near Pasir Mayang, close to the head of the Batang Hari River (+ 1°0’S, 102°0’E), in a timber concession to P.T.I.F.A. (Asiatic Forest Industry, Ltd.), where the species may become extinct. These two places are rather far from each other, but fit well in the previously known distribution pattern. Platycerium coronarium (Konig ex Miiller) Desv., the sister species of P. ridlevi, can be found in both P. ridleyi localities. The latter is a common and widely distributed species in the ever-wet lowlands of Malaya and the Greater Sunda Islands. Although the two species are closely related, they can be distinguished easily by their different habits. Platvcerium ridlevi has erect foliage fronds, an oval soral lobe, and entire base fronds (with very prominent veins) forming a large bulb. Platyvcerium coronarium has elongate, pendulous foliage fronds, a kidney-shaped soral lobe, and lobed base fronds forming a basket. There is another striking difference between the two species: P. ridleyi grows solitarily, but with many specimens in one tree (at least 10, mostly 15), whereas P. coronarium is present in one or two clusters, which are formed by vegetative reproduction. Ridley (1908, p. 56), who was the first author to distinguish P. ridlevi (as P. biforme var. erectum), mentioned its occurrence on lofty branches of trees like Shorea, at least 30 m above the ground. He never saw P. ridleyi and P. coronarium growing on the same tree. Furthermore, he cited Motley for specimens from Borneo, which were growing on the highest branches of trees in very exposed places. Holttum (1954, p. 140) remarked that P. ridleyi grows cn tall trees, but sometimes also on low branches along rivers. It never grows in open country like P. coronarium. Corner (1978, p. 162) noted P. ridlevi to be frequent on high branches in swamp forests. Our own observations more or less agree with the above findings. In the Berbak Nature Reserve, P. ridleyi occurred along the rivers and smaller streams in the swamp forest and grew exposed to the sun in the crowns of high emergent Shorea trees from the first trunk fork upwards. Platvcerium coronarium, on the other hand, *Rijksherbarium, Schelpenkade 6. Leiden. The Netherlands. *“Instituut voor Systematische Plantkunde. Rijksuniversiteit. Heidelberglaan 2. 3508 TC Utrecht. The Nether- lands. FRANKEN & ROOS: PLATYCERIUM RIDLEY! IN SUMATERA 13 occurred distinctly lower in the trees, always in a more shady situation, and apparently was less frequent than P. ridleyi. However, we may have overlooked stands of P. coronarium because of their less exposed habitat. As seen from the river, the branches of emergents and their load of P. ridleyi stand out much more clearly. Where the Pandanus zone starts, P. ridleyi seems to be more common than P. coronarium. Closer to the coast in the Nipa zone, the former is absent, whereas the latter can still be found. Also, outside the reserve near the coast, in open country and anthropogenic situations, only P. coronarium was seen. Pasir Mayang, although drier than Berbak and forming a mosaic of swampy and well-drained places, also produced several localities where P. ridleyi and P. coronarium grew in profusion. But near Muara Bulian, a much drier, non-swampy area, both staghorn ferns were absent. This could indicate that at least P. ridleyi prefers—or may be even entirely restricted to—swamp forest, as Corner indicated. Growing high up in the trees, P. ridleyi can be rather easily seen from a logging road or river. From inside the primary forest, detection is much more difficult. However, when dead, the mass of base fronds falls down intact. Finding these on the forest floor indicates that there is (or was) a tree with P. ridleyi in the neighborhood. The bulging mass of base fronds is very characteristic. It is formed by many dead, old base fronds enclosed in two new green ones. Mostly these fronds are not rounded, but have an elliptical upper part. When the tree branch is not too thick, it is totally enclosed by the base fronds. As a matter of interest, the illustration given by K6nig (Miiller, 1785), who was the first to describe Acrostichum coronarium in his letter to Miiller, shows a fertile frond of P. coronarium and a characteristic base frond of P. ridleyi. This see-siomee is the type of P. coronarium, but the base frond must be excluded from the protologue. We observed the frequent association of P. ridleyi with Lecanopteris, also reported by Ridley (1908). We collected P. ridleyi growing together with L. crustacaea Copel. The Staghorn Fern has colonies of ants in its base fronds, whereas Lecanopteris has them in its thickened rhizome Platycerium coronarium also is a host for ants, but it is never associated with Lecanopteris. Other epiphytes were not found growing with Platycerium. Ideman (in Hallé et al, 1978, p. 332) proposed a method for describing the structure of forests by means of a profile diagram. Of great importance is the distinction between trees of the future, present and past. One of the characters used is the presence of epiphytes. Trees of the present have some epiphytes, whereas trees of the past are overgrown by numerous epiphytes. We do not think that Plarycerium species are an indicator of trees of the past, although, as said before, in the case of P. ridleyi they may occur in great quantity on a single vigorous tree and are almost the only epiphytes present, except for Lecanopteris. We are grateful to Dr. Soetrisno Soewoko of P. P. A. Jambi for giving us the Opportunity to do research in Berbak Nature Reserve. We want to thank the Rijksherbarium, Leiden, where we wrote this paper. Our special thanks to Mr. G. J. de Joncheere for his advice. Finally we would like to thank W. W. F. Holland, Stichting W. S. O. and Studiefonds Van Oosterom for their financial support. 14 AMERICAN FERN JOURNAL: VOLUME 72 (1982) LITERATURE CITED CORNER, E. J. H. 1978. The freshwater swamp-forest of South Johore and Singapore. Gard. Bull., eet pie 1:1-266. HALLE, F., R. . OLDEMAN and P. B. TOMLINSON. 1978. Tropical Trees and Forests: an schizta Analysis. Springer, Berlin. HOLTTUM, R. E. . Flora of Malaya, vol. II. Ferns of Malaya. Gov't. Printing Office, Singapore. MULLER, O. F. a es eines Schreibens des Herrn D. K6nig. Naturforscher (Halle) 21:107-112, 13: RIDLEY.) i A list of the ferns of the Malay Peninsula. J. Straits Branch Roy. Asiat. Soc. 50: pad The ferns of the Malay Peninsula. J. Malay. Branch Roy. Asiat. Soc. 4(1):1-121. REVIEW “FERNS AND FERN ALLIES OF GUATEMALA, PART II. POLY- PODIACEAE,” by Robert G. Stolze, Fieldiana, Botany n.s. 6: 1-x, 1-522. 1981.—This is the second of three parts of Stolze’s monumental and indispensable work on Central American ferns. (The first part, containing the Ophioglossaceae, Marattiaceae, Osmundaceae, Schizaeaceae, Gleicheniaceae, aap 9 eect Plagiogyriaceae, Dicksoniaceae, and Cyatheaceae, was reviewed in Amer. Fern J. 67:94. 1977.) The present part matches the format and quality in eal text and illustrations (all genera and some species are illustrated) of the earlier part and covers the remainder of the ferns, which are all placed in the Polypodiaceae sensu latissimo, alphabetically from Acrostichum to Woodwardia. Although this arrange- ment is handy for finding the 67 genera and over 500 species and infraspecific taxa treated in the volume, it does not group related species or genera together, which can be a disadvantage when browsing through the volume while making identifications. However, the keys are ample and well constructed, and so will surely be used for this purpose. The genera are construed broadly, especially Polypodium, but that is no disadvantage to the user, and the correct names in the segregate genera are given in the synonymies. The species treatments each include a brief synonymy, habitat and range statements, an original description, and often copious notes. Of special value are Stolze’s comments on’ unresolved taxonomic difficulties which point students of ferns toward areas of fruitful research. Dubious and excluded species are accounted for. Presumably an index will conclude the third part, and that will add greatly to the utility of all the parts —D.B AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 1 (1982) 15 Comparison of Chromatographic Spot Patterns of Some North American Isoétes Species L. S. KOTT and D. M. BRITTON* Few comparative phytochemical studies have been done among the species of Isoétes. A comparison of the phenolic compounds in some species in the orders Isoétales, Lycopodiales, and Selaginellales (Vorin, 1972) showed only biflavones, whereas other members of the Isoétales and Lycopodiales had other types of flavones. Vorin (1972) also isolated the flavones apigenin and luteolin from European /. delilei Rothm. and J. durieui Bory. Shortly thereafter, Vorin, Jay, and Hautville (1975) isolated the unique flavone, isoetin, in these same two species, and this clearly separated the quillworts from the lycopods chemically. In further work on four European species (J. durieui, I. delilei, I. lacustris L. and I. velata A. Braun), seven compounds were isolated: apigenin, luteolin, isoetin, selagin, chrysoeriol, tricin, and orientin (or isoorientin) (Vorin & Jay, 1978). Each species showed a specific grouping of these compounds. In other chemotaxonomic studies of /soétes species from the southeastern United States, it was discovered that, very little taxonomic information could be gleaned from chromatographs (Matthews & Murdy, 1969, Boom, 1979). They considered that there was more variation between populations within a species than between species. The present phytochemical study was initiated to survey the patterns of the phenolic compounds as part of a larger taxonomic study of the genus /soétes in northeastern North America (Kott, 1980). No attempt was made to isolate or identify the compounds. MATERIALS AND METHODS Populations of /soétes species used in the chromatographic survey were deter- mined morphologically (Kott, 1980) and cytologically (Kott & Britton, 1980). The species examined were: /. macrospora Dur., I. tuckermanii A. Braun, 1. riparia Engelm., /. acadiensis Kott (Kott, 1981), /. eatonii Dodge, I. echinospora Dur., and one population of /. engelmannii A. Braun. Isoétes hieroglyphica A. A. Eaton was not analyzed chemically because too few collections were available of this rare species. In all, 35 populations were sampled, and several duplicate chromatographs were prepared for each population. Extracts were prepared by powdering dried leaves from several plants in a population and soaking 0.1 gm of material in 0.1 cc of absolute methanol for 48 hours. Paper chromatographs were prepared by applying 200 xml of each sample to standard Whatman #1 paper using the spot method. The chromatographs were run ascendingly in butanol:acetic acid:water (BAW) (12:3:5) in one direction (30 hours) and in a 2% formic acid solution in the other direction (6 hours). Dried chromatographs were examined in the presence of long wave ultra- violet light and ammonia vapor. Average Rf values were determined for all obvious and constantly recurring spots (major spots) (Table /). *Dept. of Botany and Genetics, University of Guelph, Guelph, Ont. NIG 2W1, Canada. AMERICAN FERN JOURNAL: VOLUME 72 (1982) RESULTS AND DISCUSSION The spot patterns were not strikingly different overall from species to species, but some differences existed in the presence or absence of certain major spots. The chromatographs of the seven /soétes species examined fell into one of two spot patterns, the /. echinospora type and the J. macrospora type. Isoétes echinospora, I. eatonii and I. riparia all generally shared a common pattern with a maximum of 8 major spots (Fig. /a). Although only one population of J. engelmannii was sampled, it perhaps should also be included in this group. It differs by the absence of spots 2, 3 and 7 (Table 1). Isoétes eatonii and I. riparia have almost identical patterns to /. echinospora, except that the two former species have one spot each that may be weak or absent at times (Table /). TABLE 1. ANALYSES OF SPOT PATTERNS FOR SEVEN SPECIES OF /soétes. I 2 3 4a 4b 5 6 7 8 color of spot: UV DPF P 1B Se ge B | Saeey 5 P E P UV+tNHe YY P Fi Yoo ¥Geuy OY BP Pp. Rt : BAW BAe Sen oe ea a eae Rf : 2% Formic acid 0 Bs teh re WS sa i oa a ei ae I. echinospora ee 1% a ap — ee + I. eatonii + nue aE a5 - a5 + + I. engelmannii + Ba ae ae !. riparia ae + ne - +: i cs - I, macrospora aR a a am ae -|- I. acadiensis ne ae ak of + I. tuckermanii Sig at + a4 ae Color code: DP = dark purple: P= purple: Y = yellow: YG = yellow-green: OY = orange-yellow The remaining three species, /. macrospora, I. tuckermanii, and 1. acadiensis, generally share the other spot pattern. Basically their pattern resembles the first but with fewer major spots (Fig. /b). In these three species, the purple spots 7 and 8 present in the /. echinospora type are absent. /soétes tuckermanii lacks spot 4b and I. acadiensis lacks spot 2, both of which are present in /. macrospora (Table 1). The large yellow-green spot (4a) at Rf .49/.05 in the J. echinospora type of pattern may be a different compound from the yellow-green spot (4b) at Rf .47/.13 in the J. macrospora type pattern. Until the compound is analyzed chemically, it cannot be determined whether these spots are similar, but it appears that this yellow-green spot is consistently in a slightly different position in each of the two patterns. In conclusion, it appears that gross chromatography can add some taxonomic information to well established morphological and cytological data. Spore mor- phology of J. tuckermanii and I. macrospora share some similarities, and it is not unlikely that they share a common ancestor. This is reflected to some extent in the spot pattern that they share. /soétes tuckermanii has a spot pattern different from that of /. riparia, a species with which it is often confused using spore morphology KOTT & BRITTON: CHROMATOGRAPHIC PATTERNS IN ISOETES 17 i ALAS ” DAVY eo) R$ ae CS Qe () Rt oO > o o a 2 4 oe ‘ie 17 O + ey a eee eee —> FORMIC aa. ¢ se Se ee. R¢# Rf FIG. 1. Comparative spot patterns produced by two dimensional chromatography on /soctes species. a. Composite spot pattern of /. echinospora type. b. Composite spot pattern of /. macrospora type. and habitat preference. /soétes riparia, however, has great affinities with /. echinospora in spot pattern, and perhaps one of the parents of this tetraploid species is the diploid J. echinospora. This is supported by spore morphology of /. ripar id, which is intermediate between the spiny and the ridged spore type. The three diploid species /. eatonii, 1. echinospora and I. engelmannii, although unique in spore pattern, have similar chromatographic spot patterns. The phytochem- ical spot pattern appears to yield no further taxonomic information about these species at this level of comparison. Identification of the compounds that produce the spots may lead to a better understanding of the relationships among the groups, since the unique spore morphology of each of the three diploid species indicates that they are not closely related. AS gle CRU We wish to thank Natural Science and Engineering Research Council for financial Support for this project. 18 AMERICAN FERN JOURNAL: VOLUME 72 (1982) LITERATURE CITED BOOM, B. M. 1979. Systematic studies of the genus Isoétes in the southeastern United States. M.Sc. Thesis, University of Tennessee, Knoxville. KOTT, L. S. 1980. The taxonomy and biology of the ale Isoétes L. in northeastern North America. Ph.D. Thesis. University of Guelph, Guelph. Ontario. . 1981. Isoétes acadiensis, a new species from ee North America. Canad. J. Bot. 59: 259 92-2594. , and D. M. BRITTON. 1980. Chromosome numbers for Isoétes in northeastern North erica. Canad. J. Bot. 58:980—984. Ga J. F. and W. H. MURDY. 1969. A study of Isoétes common to the granite outcrops of the southeastern piedmont, United States. Bot. Gaz. 130:53-61. VORIN, B. 1972. Distribution des composés polyphénoliques chez les Lycopodinées. Phytochemistry 1#:257-262. , and M. JAY. 1978. Etude chimosystématique des Lycopodiales, Isoétales, Selaginellales et Psilotales. Biochem. Syst. Ecol. 6:99-102 , and M. HAUTVILLE. 1975. Isoétine, nouvelle flavone isolée de Isoétes delilei et Isoétes duricat. Phytochemistry 14:247-259. REVIEW “LOUISIANA FERNS AND FERN ALLIES,” by J. W. Thieret. vi + 123 pp., 75 plates. Lafayette Nat. Hist. Museum. 1980. $17.95.—This is a semitechnical floristic treatment, and so is useful to botanists and is at the same time understanda- ble to non-specialists. Each species native to or naturalized in Louisiana is treated. The scientific name, sometimes one or more common synonyms, the vernacular name, a very brief description that is supplemented by a plate of several figures, notes, and a general statement of range are given. The keys to the species of each genus are ample and usable. The introductory portion of the book will acquaint non-specialists with some of the intricacies of pteridology. Unfortunately, there is no conventional key to the families or genera. Instead, there is a “guide” consisting of a series of sequential, non-contrasting statements each leading to one or more genera. By reading through the statements, according to the author, “Eventually you will come upon a statement descriptive of the fern or fern ally you wish to identify.” This procedure is roughly twice as cumbersome as a conventional key. The plates, which are reproduced from other publications with permission, are diagnostic, but any grace or delicacy they had as originals has been lost in reproduction. A dot map by parishes (the equivalent of counties) is included on each plate. The plates are grouped near the center of the book and are unpaged, but are numbered. The work concludes with a checklist of Louisiana pteridophytes, a glossary, literature cited, and an index.. The book is paper-backed and perfect bound, and so will not withstand extensive rough handling. Presumably the book can be obtained only from the publisher, in Lafayette, Louisiana 70501.—D.B.L. AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 1 (1982) 19 Anatomical Studies of the Neotropical Cyatheaceae. II. Metaxya and Lophosoria TERRY W. LUCANSKY* The classification of the monotypic tree fern genera Metaxya and Lophosoria has undergone several revisions. Early taxonomists placed these genera, with the dicksonioid group, in the family Cyatheaceae (Diels, 1902; Christensen, 1906; Maxon, 1912). Bower (1926) stressed the importance of soral position and placed these primitive genera in the family Protocyatheaceae. Later workers (Christensen, 1938; Copeland, 1947) included the two genera in the Cyatheaceae, but excluded the dicksonioid group from that family. Based upon morphological, developmental and anatomical data, Holttum and Sen (1961) again placed Metaxya and Lophosoria and the dicksonioid ferns into the single family Cyatheaceae. More recently, Tryon (1970) revised the classification of the Cyatheaceae, excluding members of the Dicksoniaceae. He recognized Metaxya and Lophosoria as distinctive genera and placed them in an independent position at the base of his phyletic scheme for the family. In this study, the anatomy of Metaxya rostrata (H.B.K.) Presl and Lophosoria quadripinnata (Gmel.) C. Chr. is elucidated and an attempt is made to determine whether the proposed phyletic relationship between these two taxa and the squamate genera of tree ferns is supported by anatomical data Developing shoot tips of M. rostrata and L. quadripinnata were collected in montane and lowland rainforests in Costa Rica. The plant materials were killed and fixed in formalin—acetic acid—alcohol (FAA) and sectioned on a macrotome (Lucansky, 1976a). The slices were partitioned into manageable sizes, dehydrated in a tertiary-butyl alcohol series and embedded in paraffin (Johansen, 1940). Sections (10 xm) were made and stained in a safranin—fast green series. Stained sections were photographed with a Nikon M-35S camera, whereas entire slices were photographed with a Pentax 35 mm single lens reflex camera. Voucher specimens are deposited in the herbarium of Duke University. RESULTS AND DISCUSSION Based upon stem and petiole indument, nodal patterns, and vascular anatomy, Metaxya and Lophosoria are similar and represent distinctive and primitive genera in the neotropical Cyatheaceae (Lucansky, 1974a; Lucansky & White, 1974). Both genera are characterized by unbranched, multicellular trichomes on both the rhizome (stem) and petioles. Metaxya rostrata possesses a prostrate, dorsiventral rhizome, whereas a short, upright radial stem occurs in Lophosoria quadripinnata (Figs. | and 2). The presence of trichomes and a procumbent habit are considered primitive characters in the Cyatheaceae (Bower, 1913). The squamate genera of the New World tree ferns have an arborescent habit, except for an acaulescent habit in Cnemidaria (Tryon, 1970). Metaxya has once-pinnate leaves inserted laterally and dorsally on the rhizome, whereas Lophosoria has decompound leaves inserted over “Department of Botany, University of Florida, Gainesville, FL 32611. AMERICAN FERN JOURNAL: VOLUME 72 (1982) > oS Cw FIGS. 1-6. Habit, transections. and anatomical details of tree fern stems. FIG. 1. Prostrate rhizome of Metaxya rostrata, X0.5. Note adventitic uds; (FRG, 2... Snort, aie stem of eS x 0.6. FIG. 3. Transection oP HDS ac (amphiphloic ae vi M. rostre mT: x 0.4. FIG. Cr eines of M. quadr sap Fl Transection of rhizome (dictyostele) of L. quadripinnata, FIG qc teri tha x 57. tis abbreviations are: m= mucilage-sac cell, sclerenchyma rostrata, X54. . 6. Two-zoned hypodermis of L. b=adventitious bud. c =cortex. e= epidermis. h = hypodermis. iz = inner zone. 0z= outer zone. p=petiole. r=adventitious roots, rt=root trace. s=stelar sheath. st tissue. and vt = vascular tissue. T. W. LUCANSKY: ANATOMICAL STUDIES OF CYATHEACEAE. II. 21 the entire surface of the stem. Adventitious roots arise from the leaf bases, buds and stems of both species (Figs. / and 2). While much is known morphologically about these two genera, comparative anatomical data are almost totally lacking for these neotropical species. Both Metaxya and Lophosoria have an amphiphloic siphonostele, although a dictyostelic pattern with overlapping leaf gaps frequently occurs in L. guadripinnata (Figs. 3 and 4). Bower (1912) found no such stelar pattern in this species, but did indicate an approach to dictyostely in some specimens. No dictyostelic pattern is observed in Metaxya. The siphonostelic axis devoid of overlapping leaf gaps is a relatively primitive feature in the Cyatheaceae, and according to Holttum (1965), a siphonostelic, dorsiventral rhizome is an evolutionary development related to the growth habit of the plant. Stem transections of Metaxya and Lophosoria are anatomically similar in some respects, but differ in certain anatomical features. In both genera, a single-layered epidermis composed of thick-walled, variously shaped cells filled with tannins is generally persistent in the mature sporophyte (Figs. 5 and 6). In Culcita and Cystodium, the outer epidermal walls may be either thickened (Sen & Mittra, 1966) or cutinized (Sen, 1968). In the squamate genera, the epidermal layer is typically sloughed off in the mature plant (Lucansky, 1976b, 1977), although remnants of this layer may persist. Beneath the epidermis in Metaxya is a hypodermis composed of sclerified parenchyma cells filled with tannins and infrequently starch grains (Fig. 5), whereas a two-zoned hypodermis is found in Lophosoria (Fig. 6). The outer zone (8-10 cells thick) is composed of thick-walled parenchyma cells filled with tannin droplets; the inner zone (20-25 cells) consists of smaller, sclerified parenchyma cells filled with tannins. Groups of large mucilage-sac cells are randomly scattered within this latter zone (Fig. 6). Previous workers had reported that the hypodermis of certain tree fern species consists solely of sclerenchymatous fibers (Ogura, 1938; Mehra & Singh, 1955); however, the squamate genera typically possess a two-zoned hypodermis composed of thick-walled parenchyma cells and sclerified parenchyma cells (Lucansky, 1976b, 1977). Except for Metaxya, a two-zoned hypodermis is characteristic of the neotropical Cyatheaceae. In Metaxya, the cortex is composed of large, irregularly shaped, thick-walled parenchyma cells filled with tannins and starch grains (Fig. 5). In older rhizomes, the cortical cells become sclerified and resemble fibers. In Lophosoria, the cortex consists of large, thin-walled parenchyma cells filled with tannins and starch grains, although the outermost layers may be thick-walled. Within the cortex, large mucilage-sac cells are randomly distributed, either singly or in groups of 3-15 (Fig. 7). Distinctive cells called cubical cells occur between the hypodermal and cortical regions in Lophosoria (Fig. 8), whereas these cells are lacking in these regions in Metaxya. Holttum and Sen (1961) reported only solitary cubical cells in the cortex of Lophosoria, but they may form a discontinuous layer. Cubical cells also occur in the cortical region of the squamate genera of the neotropical Cyatheaceae (Lucansky, 1976b, 1977) and are found in the cortex of certain dicksonioid species (Sen, 1964). These distinctive cells usually contain a single, large, irregularly shaped crystal that is thought to be composed of silica (Sen, 1968). Although Ogura (1938) thought that AMERICAN FERN JOURNAL: VOLUME 72 (1982) - a wth ace wean: 2s & a eS OT a 2 tea <= “= ces St ”, ams wi. Se fp. Pad PS se 284e*, Mens cab aay to ack on Ae gr 2g 0 @ Soot eI Se evi Mi: . , ' =~ ‘ ar: ils of tree fern stems. FIG. 7. Cortex of L. guadripinnata, X 127. Note mucilage-sac cells. FIG Cubical cells with crystals in L. quadripinnata, * 429. FIG. 9. Localized sclerenchymatous areas (dissected external stelar sheath) in cortex of L. guadripinnata, x 50. FIG. 10. Meristele of L. quadripinnata, X 124. Note tangential cells. FIG. 11. Tangential cells in stele of M. rostrata, * 220. FIG. 12. Tangential cells with lateral sieve areas in primary phloem of L. guadripinnata., « 464. The abbreviations are: ¢ = cortex, cc =cubical cell. cr=crystal. e=endodermis. m= mucilage- e =pericycle. ph=phloem. sa=sieve area. st= c = tangential ry FIGS. 7-12. Anatomical deta - sac cell. sclerified tissue. t=tannins. cell. and x = xylem. T. W. LUCANSKY: ANATOMICAL STUDIES OF CYATHEACEAE. II. 23 cubical cells were sclerenchyma cells, their living protoplast, wall morphology, position, and resemblance to parenchyma cells in young stems shows that they are thick-walled parenchyma. Sen’s (1964) report that they were not sclerenchyma cells is based upon their rate of cell division and cellular inclusions. Also scattered in the cortex of Lophosoria are localized areas of sclerified tissue (transitional parenchyma-to-sclerenchyma cells) that vary in shape and size (Fig. 9). The stelar sheaths in certain squamate genera arise from the fusion of such localized areas of sclerenchyma cells (Lucansky & White, 1976), and these areas in Lophosoria occasionally fuse to form a dissected stelar sheath and may represent an early evolutionary stage in the development of an external stelar sheath. Localized patches of sclerenchyma tissue have also been noted in the cortical region of Trichipteris microphylla and Cyathea suprastrigosa (Lucansky, 1977) and in Cystodium sorbifolium (Sen & Mittra, 1966). Cubical cells filled with solitary crystals (Fig. 8) frequently occur between these sclerenchymatous areas and the thin-walled paren- chyma cells of the cortex in both Lophosoria and certain squamate species (Lucansky, 1977). No sclerenchyma tissue and cubical cells are found in the cortical region of Metaxya. Cortical bundles are a characteristic feature of certain genera and species in the Cyatheaceae (Lucansky, 1974b, 1976b, 1977), but these vascular bundles do not occur in either Metaxya or Lophosoria. A distinct, single-layered endodermis filled with tanniniferous substances delimits the stelar tissue in both Metaxya and Lophosoria (Figs. 10 and 11). A pericycle composed of 1—5 rows of thin-walled parenchyma cells filled with tanniniferous substances encircles the primary phloem. Although Ogura (1927, 1938) reported anatomical differences between the proto- and metaphloem in certain squamate species, these two tissues are indistinguishable in the present study. Earlier workers (Schiitze, 1906; Ogura, 1927) found that the primary phloem in certain species is composed of distinct rows of sieve cells and phloem parenchyma, but these two cell types are randomly interspersed in M. rostrata and L. quadripinnata. A similar random arrangement has been noted for certain squamate species (Lucansky, 1976b, 1977). The outermost 1-3 layers of primary phloem in both Metaxya and Lophosoria are composed of distinctive cells called tangential cells (Figs /0—/2), whereas these cells typically occur between the proto- and metaphloem in certain squamate species (Lucansky, 1976b, 1977). These cells represent specialized sieve cells that are devoid of nuclei, possess sieve areas on their lateral walls, and accumulate callose (Sen, 1964), although they have been referred to as false sieve tubes (Schiitze, 1906) or as elongated mucilage cells (Ogura, 1927). Although tannin droplets occasionally occur in these cells, they are typically devoid of contents. Previous workers had reported that they contained oil and mucilage (Schiitze, 1906; Ogura, 1927). Tangential cells are a characteristic feature of the Cyatheaceae, and have previously been found in certain neotropical squamate species (Lucansky, 1976b, 1977). The primary xylem in both Metaxya and Lophosoria consists primarily of tracheids with scalariform pitting, with xylem parenchyma interspersed among these xylary elements. Xylem maturation is mesarch in both genera, with metaxylem tissue comprising the bulk of the xylary mass (Figs. /0 and //). The cellular 24 AMERICAN FERN JOURNAL: VOLUME 72 (1982) 3% . ey we ' 26. PAY ee ) * a4, of Sneath s * we ~ - - o% Bed 4 i . $y Ba. ae : * 15 Baer 2 Sc ee emseae | ha he par enone meas inners een maaan met eben A ‘a * . mt Ee - ip thy ‘00. aoe yy pe 18.4. SRE “PSN a OR TA 6 3-18. Anatomical details of tree fern stems. FIG. 13. Pith region of L. quadripinnata, * 58, Note mucilage-sac cells. FIG. 14. Pith region of M. rostrata, x 125, FIG. 15. Internal stelar sheath in — x pith region of L. quadripinnata, x 50. FIG. 16. Transection of adventitious bud of M. rostrata. * Note sclerified inner cortex. FIG. 18. Transection of adaxial petiole strand of L. quadripinnata, * 50, Note mucilage-sac cells in ground tissue. The abbreviations are: ¢ = cortex. e = endodermis. gt = ground = t; ~ ~ tissue. h=hypodermis. ic = inner cortex. m=mucilage-sac cell. mx = metaxylem. oc pi=pith. ps=petiole strand. px =protoxylem. pz= parenchymatous zone. s= internal stelar sheath. sg = starch grain. st=sclerenchyma tissue. t= tannins. and vt = vascular tissue. T. W. LUCANSKY: ANATOMICAL STUDIES OF CYATHEACEAE. II. 25 composition and arrangement of the primary xylem is similar in the squamate genera (Lucansky, 1976b, 1977). The pith in Lophosoria is composed of large, irregularly shaped, thin-walled cells filled with tannins and/or starch grains. Within the pith region of Lophosoria, large mucilage-sac cells are randomly distributed, either singly or in groups of 2-10 (Fig. 13), whereas no such cells occur in the pith of Metaxya (Fig. 14). Mucilage-sac cells, however, have been reported in the pith region of certain squamate genera of the Cyatheaceae (Lucansky, 1976b, 1977). The pith comprises the bulk of the rhizome in Metaxya and consists of thick-walled, variously-shaped parenchyma cells filled with tannins and/or starch grains (Fig. /4). In older rhizomes, these cells become sclerified and closely resemble fibers. An extensive band of sclerified cells in the peripheral region of the pith of Lophosoria occurs and constitutes an internal stelar sheath (Fig. 15) which possibly helps to support both the stem and leaves. The thin-walled parenchymatous zone that separates this stelar sheath from the stele may function in the conduction and storage of carbohydrates (Schiitze, 1906). Stelar sheaths are a characteristic feature of the Cyatheaceae and have been noted for all advanced neotropical genera (Lucansky, 1974b, 1976b, 1977), yet no sclerified stelar sheaths are found in Metaxya. Although the stelar sheaths of the advanced Squamate genera are delimited externally and internally by cubical cells (Lucansky, 1976b, 1977), no cubical cells are found between the stelar sheath and pith region in Lophosoria. The lack of these distinctive cells may be due to the gradual transition, rather than an abrupt one, from the sclerified cells to the thin-walled cells of the pith. In addition to the stelar sheath, one or more large, localized areas of sclerenchyma tissue frequently occur in the central region of the pith in Lophosoria, whereas no sclerenchyma cells occur in the pith region of Metaxya. Although no large, sclerified areas occur in the pith region of the squamate genera, numerous small areas do occur (Lucansky, 1974b) and may or may not be associated with medullary bundles (Lucansky, 1976b, 1977). These latter accessory bundles repre- sent another characteristic feature of the Cyatheaceae (Lucansky, 1974b, 1976b, 1977), but they do not occur in either Metaxya or Lophosoria. In both Metaxya and Lophosoria, adventitious buds arise from the rhizome (stem) and occur on the abaxial surfaces of the leaf bases (Figs. ] and 2). Anatomically these reproductive structures are similar to the stem in each genus, although some differences are noted. The outer boundary of the buds in both genera is a thick-walled epidermis bearing multicellular trichomes. Beneath the epidermis in Metaxya is a hypodermis composed of sclerified parenchyma cells (Fig. /6), whereas-the hypodermal layer in Lophosoria is composed of one (thin-walled or sclerified parenchyma) or two (thin-walled and sclerified parenchyma) layers. The cortex of Metaxya consists of thick-walled parenchyma cells filled with tannins, whereas the cortical region in Lophosoria is composed of thin-walled parenchyma cells. Mucilage-sac cells are randomly distributed singly or in groups of 2—10 in the hypodermal and cortical regions of Lophosoria, but no such idioblasts are found in these tissues in the buds of Metaxya. The stelar pattern in the buds of both genera is an amphiphloic siphonostele with no leaf gaps (Fig. 16). The stele is delimited on either side by distinct endodermal layers filled with tanniniferous substances. Cellular 26 AMERICAN FERN JOURNAL: VOLUME 72 (1982) composition and arrangement of tissues of the bud stele are similar to those of the stem stele. The pith region of a bud in Metaxya consists of either thick- or thin-walled parenchyma cells filled with tannins and starch grains. In Lophosoria, the pith consists primarily of thin-walled parenchyma cells, although the central and peripheral zones of this tissue may become sclerified in older buds. Large mucilage- sac cells are also present in the pith of older buds of Lophosoria. Transections of the adventitious roots of both genera show similar anatomical features (Fig. 17). The epidermis is typically sloughed off in mature roots, and the outer cortex, composed of thick-walled parenchyma cells, forms the outer boundary of the root. The inner cortex is composed of isodiametric sclerenchymatous fibers and forms the bulk of the cortical zone (Fig. 1/7). A two-zoned cortex is also characteristic of the squamate genera (Lucansky, 1976b, 1977), although other workers found the position of these cell layers reversed in certain species (Schiitze, 1906; Sen, 1968). A distinct endodermal layer composed of cells filled with tanniniferous substances delimits the stele. A pericycle (1—2 cells thick) composed of large, thin-walled parenchyma cells filled with tannins surrounds the vascular tissue. The primary phloem consists of sieve cells and phloem parenchyma, whereas the primary xylem is composed primarily of scalariform-pitted metaxylem, with some transitional reticulate-to-scalariform and spiral protoxylem noted. The xylem is typically diarch with exarch maturation (Fig. /7), although triarch and tetrarch xylem occasionally occur in larger roots of both species. A single, undivided leaf trace departs from the stem axis in orderly fashion in Metaxya, whereas a three-parted trace separates from the stem stele in Lophosoria (Lucansky, 1974a). Although Bower (1912) found the leaf trace of Lophosoria to be an undivided, single strand that only became divided in larger leaves, the leaf trace is typically three-parted in both small and large leaves. The petiole strand in Metaxya is an undivided, corrugated, horseshoe-shaped structure with distinct lateral folds and inflexed ends, whereas in Lophosoria the petiole strand is a three-parted structure consisting of an abaxial arc and an adaxial arc composed of two series (Lucansky, 1974a). The horseshoe-shaped configuration of the petiole strands of both genera show an affinity with both the cyatheoid (Ogura, 1927, 1938; Lucansky & White, 1974) and dicksonioid (Bower, 1913; Holttum & Sen, 1961; Sen, 1968) tree ferns, although the division of the petiole strand of Lophosoria anticipates the greater dissection of the strand in the more advanced genera of the Cyatheaceae (Lucansky & White, 1974). The petiole bases of Metaxya and Lophosoria are anatomically similar (Fig. 18). A single-layered, thick-walled epidermis is persistent in the mature leaves, whereas this epidermal layer is sloughed off in certain squamate genera (Lucansky, 1976b). Beneath the epidermis is a hypodermis composed of sclerified, thick-walled paren- chyma cells. In Lophosoria, mucilage-sac cells in groups of 2-6 are randomly distributed in this tissue, whereas no such idioblasts are found in the hypodermis of Metaxya. The bulk of the petiole in both genera is ground tissue composed of thin-walled parenchyma cells filled with tanniferous substances, although thick- walled parenchyma cells may occur in both genera. Mucilage-sac cells are scattered in this tissue in Lophosoria (Fig. 18), but are lacking in the ground tissue of T. W. LUCANSKY: ANATOMICAL STUDIES OF CYATHEACEAE. Il. 27 Metaxya. Numerous small, intercellular spaces also occur in the ground tissue of both genera. In certain squamate genera, parenchyma cells with tannins or mucilage comprise the ground tissue (Lucansky, 1976b, 1977). The vascular tissue of the petiole in both M. rostrata and L. quadripin nata is delimited by a distinct endodermis filled with tannins. Cellular composition and arrangement of each petiole strand is similar to the stem stele, except no ht cells are found in the primary phloem of a petiole. In both species, transitional parenchyma-to- sclerenchyma cells form a sheath on either side of the petiole strands, with the more extensive sheaths towards the center of the petiole base (Fig. /8). TABLE 1. COMPARATIVE ANATOMICAL AND MORPHOLOGICAL DATA ON Metaxya, Lophosoria AND THE SQUAMATE GENERA IN THE CYATHEACEAE. Character Metaxya Lophosoria Squamate Genera Habit prostrate rhizome short, erect stem arborescent single zone two zones two zones Sclerenchyma tissue no in cortex and pith Mucilage-sac cells no yes yes in cortex and pith Cubical cells no yes yes Tangential cells yes yes yes Stelar pattern amphiphloic amphiphloic dicytostele siphonostele siphonostele, dictyostele Stelar sheath no yes = Accessory bundles no no Petiole strand undivided, divided, div ided horseshoe-shaped horseshoe-shaped horseshoe -shaped Previous anatomical and morphological data indicate that Metaxya and Lophosoria are distinctive and primitive genera, and, in some respects, show an affinity with the dicksonioid tree ferns (Lucansky, 1974a). Gastony and Tryon (1976) also found the spore morphology of these two genera to be distinctive. Gastony (1981, 1982) later reported that the spore morphology of Metaxya and Lophosoria is similar to certain genera in the Dicksoniaceae, and on the basis of palynological characters. Although similarities exist between Metaxya and Lophosoria, any relationship between them remains unclear. The present study indicates that significant differences do occur in these two monotypic genera (Table /), and their affinity with the cyatheoid tree ferns is questionable. Based upon anatomical data, Lophosoria is more closely related than Metaxya to the squamate genera in the Cyatheaceae, although some similarities to the Dicksoniaceae are noted. A chromosome number n=65 (Walker, 1966) for the genus also shows a relationship to both the cyatheoid and dicksonioid tree ferns. Metaxya, on the other hand, it quite distinctive from both Lophosoria and the cyatheoid tree ferns and may be better placed in another family or its own family. A chromosome number of n=96 in Metaxya (Roy & Holttum, 1965) also supports its isolated position and the segregation of this genus from the family Cyatheaceae. 28 AMERICAN FERN JOURNAL: VOLUME 72 (1982) LITERATURE CITED BOWER, F. O. 1912. Studies in the phylogeny of the Filicales. II. Lophosoria, and its relationship to the Cyatheoideae and other ferns. Ann. Bot. 26:269-323. . 1913. Studies in the are a si Filicales. II]. On Metaxya and certain other relatively primitive ferns. Ann. Bot O26. The Fems;-vol: II. 2 ages University Zi London. inenansr C. 1906. Index Filicum. Hagerup, Cope —_——: 8. Filicinae. /n F. Verdoorn (ed.), Manual % Pees Martinus Nijhoff, The Hague. anne E. B. 1947. Genera Filicum. Chronica Botanica, New Yor DIELS, = 1902. Cyatheaceae. Jn A. Engler and K. Prantl (eds.), Die Natiiilichen Pflanzenfamilien, eil 1, Abt. 4. W. Engelmann, Leipzi GASTON. G. oh 1981. Spore morphology in the Dicksoniaceae. I. The genera Cystodium, Thyrsopteris, Culcita. Amer. J. Bot. 68:808-819. ———. "1982. Spore morphology in the Dicksoniaceae. I]. The genus Cibotium. Canad. J. Bot. 60: In press. and R. M. TRYON. 1976. Spore Morphology in the Cyatheaceae. II. es genera Ldpliosoria; Metaxya, Sphaeropteris, Alsophila and Nephelea. Amer. J. Bot. 63:738-758. HOLTTUM, R. E. The evolution of the vascular system in ferns with special reference to oelioetane apargis Phytomorphology 14:477-480. d U. SEN. 1961. Morphology and classification of the tree ferns. Phytomorphology : "406-420. JOHANSEN, D. A. 1940. Plant Microtechnique. McGraw-Hill, New York. LUCANSKY, T. W. 1974a. Comparative studies of the nodal and vascular anatomy in the neotropical Cyatheaceae. I. Metaxya and Lophosoria. Amer. J. Bot. 61:464-471. , 1974b. Comparative studies of the nodal “ vascular anatoniy in the neotropical Cyatheascae. II. The squamate genera. Amer. J. Bot. 6 —480. . 1976a. The macrotome: a new Sak for the sectioning of large plant specimens. Stain Techno 51:199-201. 1976b. Anatomical studies of the neotropical Cyatheaceae. I. Alsophila and Nephelea. Aites’ Fern J. 66:93-101. . 1977. Anatomical studies of Cyathea and Trichipteris (Cyatheaceae). Amer. J. Bot. 64:253-259. , and R. A. WHITE. 1974. Comparative studies of the nodal and vascular anatomy in the neotropical Cyatheaceae. III. Nodal and petiole patterns; summary and conclusions. Amer. J. Bot. 61:818-828. : a R. A. WHITE. 1976. — — studies in = sporophytes of tree ferns. I. A primitive and an advance r. J. Bot. 63:463-4 ee W. R. 1912. The tree ferns of North Rea on Panes Inst. eg Rep. 1911:463-491, .1- MEHRA, > N. and J. P. SINGH. 1955. Observations of the anatomy of Alsophila glabra Hook. Sci. Cult: 2273: OGURA, Y. 1927. Comparative anatomy of the Japanese Cyatheaceae. J. Fac. Imp. Univ. Tokyo (Bot.) 1:41-350 . 1938. Handbuch der Pflanzenanatomie. Anatomie der Vegetationsorgane der Pteridophyten. Bead VII. Teil 2. Gerbriider pice Berlin ROY, S. K. and R. E. HOLTTUM. 1965. Cytological . morphological observations on Metaxya rostrata (H.B.K.) Presl. pe Fern J. 55:158-1 SCHUTZE. W. 1906. Zur pr eeniocap Anatomie einiger oe. Farne, besonders der Baumfarne. Beitr. Wiss. Bot. (Stuttgart) 5:329-376. SEN, U. 1964. Importance of anatomy in ages phylogeny of tree ferns and their allies. Bull. Bot. Soc. Bengal 18:26-34. . 1968. Anatomy of Culcita macrocarpa. Canad. J. Bot. 46:43-46. , and D. MITTRA. 1966. The anatomy of Cystodium. Amer. Fern J. 56:97—101. T. W. LUCANSKY: ANATOMICAL STUDIES OF CYATHEACEAE. II. 29 TRYON, R. M. 1970. The classification of the Cyatheaceae. Contr. Gray Herb. 200:1—5S3. WALKER, T. G. 1966. A cytotaxonomic survey of the pteridophytes of Jamaica. Trans. Roy. Soc. Edinb. 66:169—237. ALOYSIO SEHNEM (1912—1981) Aloysio Sehnem was born in Sitio, near Vila Progresso, Brazil, on 24 November 1912, the second son of Joao Sehnem and Rosa Jungblut. His primary schooling was in Vila Progresso. From 1925 to 1930 he studied at a provincial seminary in Sao Leopoldo. He joined the Jesuit order as a novice in 1930 and continued his studies in classical languages and philosophy. In 1940 he became a teacher at the Colégio Catarinense de Floriandpolis, taught courses in Portuguese and in English, and began his theological studies. He was ordained in 1943, and became a lecturer at the Colégio Santo Inacio de Salvador do Sul, where he rose to the post of Rector (1954-57). Following this, he was a member of the Faculty of Philosophy, Sciences, and Letters at Cristo Rei, which became the Universidade do Vale do Rio dos Sinos (Unisinos), where he taught graduate courses in botany. He was awarded the degree of Livre Docéncia in November 1974; his thesis was “The ferns of southern Brazil: their ecology, geographical distribution, and routes of migration.” Padre Sehnem’s interest in mosses and ferns dated from 1935. During his studies of philosophy and theology, he often collected plants with his colleague Padre Balduino Rambo. His specimens in early years were deposited in the herbarium ASSL at Sao Leopoldo. Later Padre Sehnem did much to develop the Herbarium Anchieta (PACA), which was founded by Padre Rambo at Porto Alegre but eventually was moved to Sao Leopoldo. This herbarium concentrates on the fungi and higher plants of southern Brazil, especially of Edo.Rio Grande do Sul. In addition to his work on ferns, which culminated in his publishing almost all the pteridophyte fascicles in Padre Raulino Reitz’s “Flora Ilustrada Catarinense,” Padre Sehnem was interested in growing prize-winning orchids, and he promoted a number of amateur orchid study groups in southern Brazil. Padre Sehnem died-on 19 March 1981; with his death Brazilian pteridology has suffered a great loss.—From material provided through the kindness of Dr. Raulino eitz. 30 AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 1 (1982) SHORTER NOTES AN ALTERNATIVE SIMPLIFIED MEDIUM FOR GROWING PROTHAL- LIA.—It has been brought to my attention that the company which was the source of the nutrient plant tablets specified in my article on nutrient media (Amer. Fern J. 69:122. 1979) is no longer in business. A greenhouse and garden supply store has provided a suitable substitute. Seven drops per liter of agar of “Schultz-Instant Liquid Plant Food” provides a suitable nutrient medium. The wholesale source is. the A. Y. Schultz Company, St. Louis, MO 63043. If this specific nutrient is not available, any commercial liquid plant food would be suitable, using the concentra- tion specified by the manufacturer.—Norman P. Marengo, Department of Biology, C. W. Post College, Long Island University, Greenvale, NY 11548. POLYSTICHUM MUNITUM ON BARANOF ISLAND, SOUTHEASTERN ALASKA.—During August of 1981, while conducting plant surveys from the M/V SITKA RANGER, I found Western Sword-fern (Polystichum munitum’ (Kaulf.) Presl) at two localities on southeastern Baranof Island. These are the first reports of P. munitum from this island and extend the plant’s range westward about 18 km. The first site was on the steep north shore of Port Walter at the western end of the narrows 5 km from the fjord’s entrance (56°23'N, 134°41’W) at an elevation of about 10 m. The plants (Muller 4630) were common on the western edge of an old rock-slide in a well drained, wet area carpeted by mosses, with herbaceous vegetation and an open canopy of deciduous shrubs. I also collected P. munitum 10 km to the south of Port Walter, in Port Armstrong (Muller 4623, 4625, 4626), where it grew on the north shore around the ruins of an abandoned whaling station (56°18'N, 134°40'W). The plants were abundant on rocky, sloping, upper beach meadows and in the adjacent forest of Picea sitchensis (Bong.) Carr. and Tsuga heterophylla (Raf.) Sarg. It is surprising that the fern has not previously been collected here, since the whaling station once was a significant settlement and port. Additional colonies of P. munitum undoubtedly will be found further north and west as Baranof Island is more fully explored. The plant is known to occur across Chatham Strait on Kuiu Island, and its range extends down the Pacific Coast to Idaho, Montana and California. Voucher specimens are at the U.S.D.A. Forest Service herbarium, Sitka, Alaska (TNFS), the University of Washington herbarium, Seattle (WTU), and the herbari- um at the University of Alaska Museum, Fairbanks (ALA).—Mary Clay Muller, Chatham Area, Tongass National Forest, P.O. Box 1980, Sitka, AK 99835. AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 1 (1982) 31 CHEILANTHES LANOSA AND CYSTOPTERIS PROTRUSA IN FLORI- DA. —Recently, Dr. Evans reported (Amer. Fern J. 65:1-2. 1975) a specimen of Cheilanthes lanosa (Michx.) D. C. Eaton, presently in the FLAS herbarium, from Florida. This was the first report of this fern in the state. During a brief visit to the Missouri Botanical Garden, St. Louis in January, 1981, we located a second Florida specimen of C. lanosa. The MO specimen was collected almost one hundred years earlier than the FLAS specimen, but was correctly identified. The herbarium sheet consists of four fronds without rhizomes. The label reads “Cheilanthes lanosa, det. JMG, Jan. 29, 1919, F. Rugel, 1842-1849, Florida, Ex Herbario Musei Britannici.” Although the locality in Florida is not given on the label, we think that this second record should not go unnoticed and that it further substantiates the existence of C. lanosa in Florida, at least in an earlier period. There should also be no doubt that this fern was collected in Florida, as Dr. Rugel is known to have collected plants in Florida during the 1840’s (Bull. Torrey Bot. Club 6:311—312. 1879). In addition to the above specimen at MO, we also found a sheet of Cystopteris protrusa (Weath.) Blasdell consisting of one frond without rhizome, labeled “Cystopteris fragilis, Chipley, Florida, Aug. 1896, E. Pleas, From the Sadie F. Price Herbarium, Bought 1904.” Although the characteristic horizontal, creeping rhizome was absent, there was no difficulty in recognizing this species by its petiolulate basal pinnae and stramineous stipe. A search of pertinent Florida fern literature published by D. S. Correll (Amer. Fern J. 28:11-16, 46-54, 91-100, 1938), J. K. Small (Ferns of Florida, 1931), O. Lakela and R. W. Long (Ferns of Florida, 1976), and R. F. Blasdell (A Monograph of the Fern Genus Cystopteris, Mem. Torrey Bot. Club 21(4):1-102. 1963) did not indicate the presence of any Cystopteris in Florida. To the best of our knowledge, the present paper represents the first report of this genus attributed to Florida. The area about Chipley, Florida (Washington County) is characterized by lime- stone outcrops and springs. It is not unusual that C. protrusa could have been found amongst the more shaded, damp outcrops, although it usually does prefer more circumneutral situations, such as rich humus. Certainly, this area would provide a habitat suitable to C. protrusa. The site in Washington County, Florida represents a range extension of approximately 280 miles south from its closest known site in Walker County, Georgia, according to R. McVaugh and J. H. Pyron (Ferns of Georgia, 1951) and approximately 230 miles south of its closest known sites in Tuscaloosa and Jefferson Counties, Alabama (B. E. Dean, 1969, Ferns of Alabama) _—Aleta Jo Petrik-Ott and Franklyn D. Ott, Department of Biology, Memphis State University, Memphis, TN 38152. AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 2 (1982) ‘US. POSTAL SERVICE STATEMENT OF OWNERSHIP, MANAGEMENT AND CIRCULATION (Required by 39 U.S.C. 3685) ‘A PUBLICATION NO 2 DATE OF Fi AMERICAN FERN JOURNAL ‘2. FREQUENCY OF ISSUE TUNG 0}0j}01218|}4]141]4 | 30 Sept 1981 8. ANNUAL SUBSCRIPTION to PRICE quarterly 4 $8 mem.; $9 subscr. U. S. Nat'l. Herbarium, 10th & Constitution Ave. N.W., Washington, DC 20560 U. S&S. Nat'l. ae 10th & Constitution Ave. N.W., ei ae pe 20560 SAMA Ihc arava Maing aes AMERICAN FERN SOCIETY, INC. (see 5.) Dr. David B. Lellinger (see 5.) ne 7. OWNER (f owned by » corporation, its and address must be stated and also Ihereunder the of stock- frre hacbras ade any pater pal senocepedoysetint yelp sutton andi 2 corporation, the ans snd seielas stan Mdnsaedinac petit eatnia dod govorsmosen snl other unincorporated firm, its ee et ee ee Withe publication la published by 2 nonprofit organization, bddress mus! De stated.) (item must be completed) FULL NAME it AMERICAN FERN SOCIETY, INC. | Gee 5.) | | it & KNOWN BONOHOLDERS. MORTGAGEES, AND OTHER SECURITY HOLDERS OWNING OA HOLDING | PERCENT OR MORF OF TOTAL AMOUNT OF BONDS, MORTGAGES OR OTHER SECURITIES (If there are none, so siate) FULL MAME ‘COMPLETE | if | | I fe) MAIL AT SPECIAL (i) changed, Range with Ide siateenent ) AVERAGE NO COPIES EACH ISSUE DURING PRECEDING 12 MONTHS ACTUAL NO COPIES OF SINGLE ISSUE PUBLISHEO NEAREST TO FILING DATE Co ee ee 1450 1500 8 Pa CIRCULATION 1. SALES THROUGH DEALERS AMD CARRIERS. STREET VEMDORS AD COUNTER SALES URC PT Oe ) 0 2 1193 1128 1193 1128 ‘ FREE OISTIBUT AE Ome 1194 1129 ‘F COMES MOT DISTRIBUTED SL Seeee See rr oven eamnccomrren, sronan 256 371 ‘2 RETURN FROGS MEWS AGENTS ©: TOTAL (un of 8, Prend? prem spe 111 Certify that the statements made by tamnen, O8 c go eon a js neat — ' David B. tel lings! , Editor ito (See Instruction on BRITISH PTERIDOLOGICAL SOCIETY Open to all who are interested in growing and studying ferns and fern-allies. Full members receive THE FERN GAZETTE and the BULLETIN OF THE BRITISH PTERIDOLOGICAL SOCIETY. Membership subscriptions are £ 7 for full members, £ 5 for ordinary members (not receiving the GAZETTE), and £5 for student members (under 25 years of age). For particulars, U. S. residents should apply to Dr. J. Skog, Biology Department, George Mason University, Fairfax, VA 22030. Non-U. S. residents should apply to Lt. Col. P. G. Coke, Robin Hill, Stinchcombe, Dursley, Gloucestershire, England. TRIARCH Over 5©O Wears of slide manufacture and service to botanists. We welcome samples of your preserved research material for slide-making purposes, and we invite your suggestions for new slides that would be use- ful in your teaching. Your purchases have made our 50 years of existence possible. To satisfy your con- tinued need for quality prepared slides, address your requests for catalogs or custom preparations to: TRIARCH INCORPORATED P.O. Box 98 Ripon, Wisconsin 54971 AMERICAN a. FERN a JOURNAL | QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY The Chromosomes of Lycopodium lucidum JOSEPH M. BEITEL and FLORENCE S. WAGNER 33 Axenic Culture and Induction of Callus and Sporophytes of the Appalachian Vittaria Gametophyte JAMES D. CAPONETTI, MARK WHITTEN, and MICHAEL J. BECK 36 Cystopteris x illinoensis: a New Natural Hybrid Fern ROBBIN C. MORAN 41 New Records and Distributional Notes on Maryland Pteridophytes STEVEN R. HILL and RICHARD E. RIEFNER, JR. 45 Grammitis succinea, the First New World Fern : Found in Amber LUIS DIEGO GOMEZ P. 49 A New Tropical American Species of the Genus Lycopodium BENJAMIN @LLGAARD 53 Specimens from Fée’s Pteridological Collection ‘ at the Botanical Garden of Rio de Janeiro PAULO GUNTER WINDISCH = 56 Shorter Notes: A Germination Method for Isoétes; New Station for Lygodium palmatum; The Name of a Hybrid x Asplenosorus; The Deletion of Nephrolepis pectinata from the Flora of Florida; A Filmy Danaea . Reviews 40, 44, 48, 52 The American Fern Society Council for 1982 DEAN P. WHITTIER, Dept. of Biology, Vanderbilt University, Nashville, TN 37235. President TERRY R. WEBSTER, Biological Sciences Group, University of Connecticut, Storrs, CT 06268. Vice-President MICHAEL I. COUSENS, Faculty of Biology, University of West Florida, Pensacola, FL 32504. Secretary JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, TN 37916. Treasurer JUDITH E. SKOG, Dept. of Biology, George Mason University, Fairfax, VA 22030. Records Treasurer DAVID B. LELLINGER, Smithsonian Institution, Washington, DC 20560. Journal Editor ALAN R. SMITH, Dept. of Botany, University of California, Berkeley, CA 94720. Memoir Editor JOHN T. MICKEL, New York Botanical Garden, Bronx, NY 10458. Newsletter Editor American Fern Journal EDITOR DAVID B. LELLINGER U.S. Nat'l Herbarium NHB-166, Smithsonian Institution, Washington, DC 20560 ASSOCIATE EDITORS DAVID W. BIERHORST Rt. 3, Box 188, Picayune. MS 39466. GERALD J. GASTONY Dept. of Biology, Indiana University, Bloomington, IN 47401. JOHN T. MICKEL New York Botanical Garden. Bronx, NY 10458. The “American Fern Journal” (ISSN 0002-8444) is an illustrated quarterly devoted to the general study of ferns. It is Sos by the American Fern Society, and were at the Smithsonian Institution, Washington, DC 20560. Second-class postage paid at Washin Claims for missing issues, made 6 months (domestic) to 2 oak. (foreign) after the date of issue, and the matters for publication should be addressed to the E Changes of address, dues. and applications for neni ould be sent to Dr. Judith E. Skog, Dept. of ef onto George heice University, Fairfax, 22030. Or r back issues should be addressed to Dr. ae D. Montgomery, Ichthyological Associates, R.D. | Lea PA 18603. Geseral inquiries concerning ferns should be addressed to the Secretary. Subscriptions $9.00 gross, $8.50 net if paid through an agency (agency fee $0.50): sent free to members of the American Fern Society (annual dues, $8.00; life membership. $160.00). Back volumes 1910-1978 $5.00 to $6.25 each: single back numbers of 64 pages or less. $1.25; 65-80 pages, $2.00 each: over 80 pages. $2.50 each. plus shipping. Back volumes 1979 et seq. $8.00 each: single back numbers $2.00 each, plus shipping. Ten percent discount on orders of six volumes or more. Library Dr. John T. Mickel, New York Botanical Garden, Bronx, NY 10458, is Librarian. Members may borrow books at any time. the borrower paying all shipping costs. Newsletter Dr. John T. Mickel. New York Botanical Garden, Bronx. NY 10458, is editor of the newsletter “Fiddlehead Forum.” The editor welcomes contributions from members and non-members, including miscellaneous notes. offers to exchange or purchase materials. personalia, horticultural notes. and reviews of non-technical books on ferns. Spore Exchange Mr. Neill D. Hall. 1230 Northeast 88th Street. Seattle. WA 98115. is Director. Spores exchanged and collection lists sent on request. Gifts and Bequests Gifts and bequests to the Society enable it to expand its services to members and to others interested in ferns. Botanical books. back issues of the Journal, and cash or other gifts are always welcomed. and are tax-deductible. Inquiries should be addressed to the Secretary. AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 2 (1982) 33 The Chromosomes of Lycopodium lucidulum JOSEPH M. BEITEL* and FLORENCE S. WAGNER* The Shining Fir-moss, Lycopodium lucidulum Michx. (Huperzia lucidula (Michx.) Trevisan), is an eastern North American endemic which forms colonies in moist coniferous and deciduous woods as well as swamps. The upright, isodichotomous stems root along their length as they become decumbent with age. The evergreen leaves, which are broadest above the middle, have stomates only on the undersides, and are usually coarsely serrate. They are produced in zones of longer and shorter leaves roughly corresponding to sterile and fertile leaves. Kidney-shaped sporangia are produced adaxially on the leaf base, and propagation organs (gemmae) develop at the end of each year’s growth. In areas of overlap between L. Jucidulum and the related northern species L. selago L. (Huperzia selago (L.) Bernh. ex Schrank & Mart.), intermediate sterile hybrids are occasionally found with either or both parents. These hybrids can be distinguished from L. lucidulum by their abortive spores and the presence of stomates on both surfaces of the leaves. The chromosome number of Lycopodium lucidulum Michx. has been reported several times from North America and India. The counts from India (Mehra & Verma, 1957; Ninan, 1958) based on Lycopodium lucidulum sensu Clark (1880), however, are actually from a distinct species, Lycopodium herterianum Kiimmerle (Huperzia herteriana (Kiimm.) Sen & Sen, L. sikkimense Herter 1909 non K. Muell. 1861), and not the nearctic Lycopodium lucidulum Michx. Love and Love (1958) first reported 2n = 264 for Lycopodium lucidulum from Quebec, but without an accompanying photograph, a drawing, locality data, or a voucher specimen citation. Later (1965), they reported chromosome counts from Mt. Washington, New Hampshire (without citation of voucher) of Lycopodium lucidulum and L. selago (which they made into subspecies of Huperzia selago) remarking **. . . it has been at last possible to make exact counts of the chromosomes of material of ssp. selago with appressed and patent leaves from Mt. Washington and also ssp. lucidula from lower levels of that mountain. The chromosome number arrived at is 2n = 272.” The first indication that a lower ploidal level existed in L. lucidulum came from fertile material collected in Minnesota (Clearwater Co., Twin Lakes, 16 June 1977, Wagner 77303, MICH) and examined by F. S. Wagner. It appeared to have either 66, 67, or 68 pairs of chromosomes at meiosis, but the exact number of pairs could not be determined. A similar low count was obtained by J. M. Beitel in fertile material from Michigan (Livingston Co., Schwark Woods, W side of Merrill Road just S of Sheldon Road; TIN, RS5E, SW sec. 35, 20 June 1979, Beitel 79041, MICH), which yielded approximately 66-69 pairs at meiosis. Material of L. lucidulum was collected in an acidic conifer swamp in Gray Co., Ontario (Osprey Township, 0.8 miles W of Rt. 24 on the S side of road dividing Concession 2 and 3) on 25 July 1981 by J. Beitel, W. H. Wagner and F. S. Wagner (Beitel 81024, MICH). The possibility of confusion with L. selago or its hybrids was ruled out by the presence of well formed spores and the lack of stomates on the adaxial leaf *Division of Biological Sciences, University of Michigan, Ann Arbor, MI 48109. Volume 72, number 1, of the JOURNAL, was issued March 29, 1982. 34 AMERICAN FERN JOURNAL: VOLUME 72 (1982) FIG. 1. Phot h of meiotic cl f 1 li idulum Michx. (Beitel 81024), n= 67. FIG..2, Interpretation of chromosome figure (dotted circle = nucleolus). surfaces. The material was kept in a coldroom for two days, then submerged in distilled water in a clear plastic box kept at room temperature in the light for three days. Shoot tips bearing young, developing sporangia were placed in a saturated aqueous solution of Saas ae i and refrigerated at ca. 4°C for 24 hours. This use of a PDB treatment for condensing the meiotic chromosomes prior to fixation by Mehra and Verma (1957), Wilce (1965) and F. Wagner (1980) has reduced some of the eae involved in obtaining interpretable squashes. Excess water was blotted from the tips which were then placed in Newcomer’s solution (Newcomer & Brant, 1954), left at room temperature for two hours, and placed in the freezer. Acetocarmine was used as a stain and squashing medium, Hoyer’s solution was used for permanent mounting, and the slides were ringed with Eukitt plastic mounting medium. Diakinesis was found to be the best stage for counting as the Metaphase chromosomes, although darkly stained and well outlined, have the unfortunate tendency to clump. During diakinesis it was possible to discern the homologous chromosomes of each pair. Those undergoing early disjunction were coupled by a thin chromatin strand (as reported by Wilce, 1965). Figures 1 and 2 clearly show a complement of n=67. Two large chromosome pairs were visible in all the figures examined. They can be seen slightly above the center of the photograph; the lower pair has a characteristic satellite. The nucleolus is seen slightly below and to the left of center. In over fifty figures observed and drawn, none exhibited a higher ploidal level than this figure. BEITEL & WAGNER: CHROMOSOMES OF LYCOPODIUM LUCIDULUM 35 The count of n=67 represents a new chromosome number for Lycopodium lucidulum and a new base number for the L. selago group of the segregate genus Huperzia. It is almost one-half the numbers previously reported from North America (2n= 264, 272), but without voucher specimens or photographs of the chromosomes those records cannot be confirmed as to taxon or interpretation of the figures obtained. The previous low number of n=ca. 68 reported by Léve and Love (1961) for Huperzia selago ssp. appressa was discounted by them (1965) as is - most likely counted on an admixture of roots of a species of Lycopodium s. ‘ The base number of n=67 fits into the gametophytic denominator scheme of Wagner and Wagner (1980) for Lycopodium s. 1. based on multiples of 11 plus one aneuploid addition. Thus, n=78 of Lycopodiella equals (7 11)+1, n=34 of Lycopodium s. s. equals (3X11)+1, and n=23 of Diphasiastrum equals (2x 11)+1. This, however, does not explain the numbers reported for L. cernuum and the various sae hs gee (Léve, Love & Pichi-Sermolli, 1977), nor for L. carolinianum (Bruce, 1976). The field of chromosome numbers in Lycopodium s. /. remains an open one with “ity "ies yet to be counted and many existing counts in need of confirmation. We would like to thank W. H. Wagner, Jr. for his invaluable assistance in the field and lab, and D. M. Johnson, K. S. Walter, N. A. Murray and W. W. Thomas for reading and commenting on the manuscript. LITERATURE CITED BRUCE, J. G. 1976. Comparative studies in the biology of Lycopodium carolinianum. Amer. Fern J. 66:125-137. LOVE, A. and D. LOVE. ag Cytotaxonomy and classification of lycopods. Nucleus 1:1—10. ——_—___—.,, and D. LOVE. 1961. Some nomenclatural changes in the European Flora I. Species and ssieaabactes odie Botaniska Notiser 1 14:33-47. , and D. LOVE. 1965. Taxonomical remarks on some American alpine plants. Univ. Colorado Stud., Biol. Ser. 17:1—43. D. LOVE, and R. E. G. PICHI-SERMOLLI. 1977. Cytotaxonomical Atlas of the Pteridophyta. Cramer Vaduz. MEHRA, P. N. and S. C. VERMA. 1957. Cytology of Lycopodium. Curr. Sci. 26:55—56. NEWCOMER, E. H. ae J. W. BRANT. 1954. Spermatogenesis in the domestic fowl. J. Hered. 45:79-8 NINAN, C. A. 1958. Studies on the cytology ge phylogeny of the pteridophytes. II. Observations on the genus edie ape Proc. Nat. Inst. Sci. India 24,B(2):54—66 WAGNER, F. S. 1980. Chromosome Sais in three interspecific hybrids of Diphasiastrum iycoptaen Bot. Soc. Amer., Misc. Ser. 158:121. WAGNER, W. r. and F. S. WAGNER. 1980. Polyploidy in sarees In W. H. Lewis, (ed.). “seen Biological Relevance. Plenum Publishing, New WILCE, 17. H. nie Section Complanata of the Genus Lycopodium. Now Hedw. Beih. 19. J. Cramer, Weinhei 36 AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 2 (1982) Axenic Culture and Induction of Callus and Sporophytes of the Appalachian Vittaria Gametophyte JAMES D. CAPONETTI,* MARK WHITTEN, ** and MICHAEL J. BECK*** Asexually-reproducing gametophytes of the four tropical fern genera Hymeno- phyllum, Trichomanes, Grammitis, and Vittaria occur in the uplands of the south- eastern United States (Farrar, 1967). The most common and abundant of these is the Appalachian Vittaria gametophyte, which Farrar (1978) has described in detail. Dr. A. J. Sharp of the University of Tennessee was the first to find and collect the Appalachian Vittaria gametophyte in the area of the Mountain Lake Biological Station in Virginia. Since that time, Dr. W. H. Wagner, Jr. of the University of Michigan collected it in the same area and catalogued it into a pteridophyte flora (Wagner, 1963). Other pteridologists have collected it in the area of the Highlands Biological Station in North Carolina (Wagner et al., 1970; Pittillo et al., 1975) and in Tennessee (Wofford & Evans, 1979). Other collections have located these gametophytes in several eastern states in and around the southern Appalachians (Farrar, 1978; Gastony, 1977). Appalachian Vittaria gametophytes are long-lived and produce extensive colonies in deep, protected crevices of non-calcareous rock. Although antheridia and arche- gonia are produced on the gametophytes, reduced, juvenile sporophytes have been found only once in the field by Farrar (1978). They were recognizable as Vittaria, but lacked sufficient characters for identification to species. Since most taxonomic characters for determining fern species are based on sporophyte morphology, the origin and affinities of these gametophytes are uncertain. They were first assigned to the genus Vittaria by Wagner and Sharp (1963) based on finding similar morpholog- ical characters between the gametophytes of the Appalachian Vittaria and those of Vittaria lineata (L.) J. E. Smith from Florida. The chromosome counts reported by Gastony (1977) have provided further evidence of their relationship. The experimental production of sporophytes might provide additional taxonomic characters for comparison with the Vittaria species. Farrar (1978) reported that Appalachian Vittaria gametophytes were difficult to maintain in axenic culture and ceased to grow on defined media. We report here successful axenic culture of the Appalachian Vittaria gametophyte and the spontaneous formation of callus and sporophytes. MATERIALS AND METHODS Gametophytes were collected from shaded, moist, sandstone conglomerate ledges along a cliff in Conkles Hollow State Park, Hocking County, Ohio. They were maintained under non-sterile conditions in culture dishes on washed perlite moist- *Department of Botany, The University of Tennessee, Knoxville, TN 37996-1100. **Department of Biological Science, Florida State University, Tallahassee, FL 32306. ***Carter and Holmes Inc., Newberry, SC 29108. Contribution from the Botanical Laboratory, The University of Tennessee, N. Ser. No. 541. J. D. CAPONETTI ET AL.: VITTARIA GAMETOPHYTE CULTURE 37 ened with quarter-strength Knudson’s medium (Steeves et al., 1955) without sugar. The dishes were maintained in a culture room at 25°C under a table where they received 25 foot-candles of diffuse white light consisting of a combination of fluorescent and incandescent lamps on an 18/6 hour light-dark photoperiod. Dis- tilled water was added to the dishes as required. When the growing gametophytes half filled the dishes, fresh subcultures were initiated from small masses of gametophytes in freshly prepared culture dishes. At the moment, non-sterile cultures are being maintained in this manner. Aseptic gametophyte cultures were obtained by placing a number of small gametophyte pieces into a 25 xX 150 mm screw cap culture tube. About 25 ml of a 1% Alconox solution was added to wash and wet the gametophyte pieces, and the tube was agitated for two minutes. The Alconox solution was decanted, 35 ml of a 10% (V/V) Clorox solution was added, and the tube was agitated gently for one minute. In a UV-sterilized room, the gametophyte pieces were transferred from the Clorox solution to a 100 x 15 mm sterile, plastic Petri dish containing 30 ml of sterile, double-distilled water. Under continued sterile conditions, the gametophyte pieces were then transferred to 25 x 150 mm culture tubes which were plugged with cotton and a plastic cap (Kaput). Each tube contained 20 ml of Knudson’s medium (Steeves et al., 1955) supplemented with 2% sucrose. One gametophyte piece was placed in each tube, with some in liquid media and others on media solidified with 0.8% agar. Some gametophyte pieces in liquid media were placed on filter paper supports above the medium. In an attempt to induce apogamy, gameto- phyte pieces were also cultured on media containing 0 to 4% sucrose in a series of cultures. Aseptic gametophyte cultures can also be obtained from non-sterile stock cultures heavily contaminated with bacterial and fungal spores by forcing such spores to germinate before surface sterilization. Gametophyte pieces were floated on non- sterile, half strength Knudson’s medium without sucrose and agar for one day. The gametophyte pieces were then surface-sterilized and placed into culture as described previously. RESULTS AND DISCUSSION Gametophytes in non-sterile cultures grew slowly in a concentric A sigs They produced gemmae, but did not produce either antheridia or archegon Gametophytes in sterile culture showed signs of growth sit two weeks of culture initiation, and slowly formed a near-spherical mass of ribbon-like prothallial lobes in each tube, whether in liquid or on agar-solidified media, over a period of several months. Based on visual inspection, it appeared that the gametophytes developed more rapidly on media containing 2% sucrose. Gametophytes on media containing sucrose concentrations above and below 2% developed at a reduced growth rate and appeared to be thinner and smaller than those exposed to 2% sucrose. Over a period of seven months, no gemmae or sex organs were observed in any cultures. Although the prothalli appeared normal in overall size and shape on media containing 2% sucrose, they produced an abundance of short, knob-like rhizoids (Fig. /) in addition to the more normal, elongated rhizoids. 38 AMERICAN FERN JOURNAL: VOLUME 72 (1982) FIG, I. hs orbs Vittaria gametophyte from sterile culture, X6.5. FIG. 2. Callus seg ef gametophyte, x 1.3. FIG. 3. Enlarged view of rhizome showing clathrate scales, x FIG, Appalachian Vitraria sporophyte from sterile culture, x 5.2. FIGS. 5 and 6. Enlarged view ae aes scales, x 2 J. D. CAPONETTI ET AL.: VITTARIA GAMETOPHYTE CULTURE 39 Spontaneous callus formation was observed in the cultures kept by one of us (Whitten) in three out of nine tubes maintained for seven months on media containing 2% sucrose without transfer to fresh media. The callus (Fig. 2) formed a friable mass varying in color from yellow to dark brown, and appeared to originate from the center of the near-spherical mass of gametophytes. Adjacent to the callus tissue toward the center of the prothallial mass, occasional aberrant gametophytes with enlarged, irregular cells occurred. A total of six sporophytes were produced in two of the three callus-containing cultures after about nine months. The sporophytes arose from the edge or center of a mass of callus tissue, and did not appear to be associated with normal gametophyte tissue. The largest sporophyte had produced fronds up to 7 mm in length when all cultures died during an air conditioning failure in the laboratory. Before death, the young sporophytes (Figs. 3 and 4) had produced a distinct rhizome which bore clathrate scales (Figs. 5 and 6), which are typical of the genus Vittaria. No roots were observed. The fronds were linear or unequally dichotomous- ly branched, and each contained a distinct vascular bundle. Since the sporophytes appeared to arise from callus tissue and no sex organs were observed, it is presumed that they arose apogamously. Further work on living material is needed to determine their exact origin and ploidy. The production of presumably apogamous sporophytes on Appalachian Vittaria gametophytes has been noted in both natural populations (Farrar, 1978) and in material cultured under non-sterile conditions by Alma S. Stokey, as cited by Farrar (1978). In both instances, the sporophytes aborted after producing a few small fronds. The sporo- phytes in our cultures were also small, but appeared healthy until their untimely death. In all instances, the sporophytes were too immature to provide taxonomic data useful at the species level. Attempts to obtain viable, mature sporophytes are in progress. The appearance of sporophytes in our old, crowded, and tightly closed His suggests that ethylene may play a role in callus formation and apogamy. Ethylene is produced by a wide range of plants and causes many different responses (Burg, 1962). Moreover, Elmor and Whittier (1973) have demonstrated that ethylene can induce apogamy in gametophytes of Preridium aquilinum (L.) Kuhn. They further demonstrated this effect in nine of its strains (Elmore & Whittier, 1975). Future experiments involving ethylene are needed not only on Appalachian Vittaria gametophytes but also on other gametophytes of the southern Appalachians in the genera Hymenophyllum,-.Trichomones, and Grammitis (Farrar, 1967). We thank Mr. Kenneth McFarland for the gametophyte collections and Dr. A. Murray Evans for reading the manuscript and offering suggestions. LITERATURE CITED BURG, S. P. eee Bes Ege canes of ethylene formation. Ann. Rev. Pl. Physiol. 13:265—3 ELMORE, H. . P. WHITTIER. 1973. The role of eihwiene in the induction - apogamous buds in ope gametophytes. Planta 111:85—9 , and D. P. WHITTIER. 1975. Ethylene dkdliackion and ethylene-induced ba bud formation in nine gametophytic strains of Pteridium aquilinum (L.) Kuhn. Ann. Bot. 39:965-971. 40 AMERICAN FERN JOURNAL: VOLUME 72 (1982) FARRAR, D. R. 1967. Gametophytes of four tropical fern genera a independently of their sporophytes in the southern Appalachians. Science 155:1266—1267 1978. Problems in the identity and origin of the Appalachian Vittaria gametophyte, a sporophyteless fern of the eastern United States. Amer. J. Bot. 65:1—-12. erage G. J. 1977. Chromosomes of the ae ceuereny Reproducing Appalachian Gametophyte: New Source of Taxonomic Evidence. Syst. Bot —48. PITTILLO, Fo We WAGER, Jr D:: RR: FARRAR, and S. W. LEONARD. 1975. New pteridophyte records in the Highlands Biological Station area, Southern Appalachians. Castanea 2. 1263-27 STEEVES, T. A., I. M. SUSSEX, and C. R. PARTANEN. oe : vitro studies on abnormal growth of prothalli of the bracken fern. Amer. J. Bot. 42:232-2 WAGNER, W. H., Jr. 1963. Pteridophytes of the Mountain Lake area, 1, Gites County, Virginia, including notes from Whitetop Mountain. Castanea 28:113-150. , and A. J. SHARP. 1963. A remarkably reduced vascular plant in the United States. Science M2 1483-1484. . R. FARRAR, and B. W. McALPIN. 1970. Pteridology uM ay pe Biological e tion Ses Southern Appalachians. J. Elisha Mitchell Sci. Soc WOFFORD, B. E. and A. M. EVANS. 1979. Atlas of the vascular plants of Nie I. Pteridophytes and madabtocantit J. Tenn. Acad. Sci. 54:32-38. REVIEW “THE FERN GENUS DAVALLIA IN CULTIVATION [DAVALLIACEAE],” “DAVALLIA RELATIVES IN CULTIVATION: ARAIOSTEGIA, DA- VALLODES, HUMATA, AND SCYPHULARIA [DAVALLIACEAE],” and “THE GENUS PYRROSIA IN CULTIVATION [POLYPODIACEAE],” by Barbara Joe Hoshizaki, Baileya 21:1-42, 43-50, and 53-76. 1981.—These papers continue the author’s long series of useful treatments of cultivated ferns. Each paper contains an introduction, key to the species, and a brief synonymy and notes on the mo hology, distribution, and cultivation of each species and its cultivars. The cultivars, although not included in the key, are distinguished in the text and, like the wild species, are illustrated with silhouettes. Anyone needing to identify cultivated Davalliaceae and Pyrrosia will find these papers invaluable. —D.B.L AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 2 (1982) 4] Cystopteris x illinoensis: a New Natural Hybrid Fern ROBBIN C. MORAN* Cystopteris is a relatively small, cosmopolitan genus of ferns comprised of about ten species and several hybrids. In eastern North America, the genus as currently interpreted consists of six taxa, two of which are known to be of hybrid origin (Blasdell, 1963). The hybrids are C. tennesseensis (C. bulbifera X protrusa) and C. laurentiana (C. bulbifera x fragilis var. fragilis). Both are fertile allopolyploids which reproduce themselves sexually throughout their respective ranges. This paper concerns a new Cystopteris hybrid having aborted spores and sporangia and presents evidence showing that C. bulbifera and C. fragilis var. mackayi are the parents. 00 OO 00 Q O 9 3 6 C. bulbifera C. X illinoensis C. fragilis var. mackayi FIG. 1. Comparison of patterns of phenolic compounds separated by two dimensional paper chromato- graphy in Cystopteris X illinoensis and its parents. The writer first came across this unusual new hybrid fern when, as a graduate student at Southern Illinois University/Carbondale, he was asked to verify identifica- tions of a box of fern specimens sent to the herbarium. The specimens were donated ten years earlier by Dr. Ralph C. Benedict, of Rockford, Illinois, who was a fern taxonomist and a long time active member of the American Fern Society. In the box of pressed unmounted specimens was a collection identified by Dr. Benedict as “C. bulbifera = C. fragilis var. mackayi’. Location data stated that the hybrid was, “Very rare, found only once in Winnebago County, Ill. Old quarry with colony of var. mackayi—Cystopteris bulbifera a few yards away.” The exact location of the quarry (almost certainly a dolomite quarry, since this is the only rock type in the area) was not given. However, Dr. Benedict removed the hybrid from the quarry and cultivated it in his garden in Rockford. The material sent to the Southern Illinois University herbarium (SIU) consisted of fronds gathered from the hybrid growing in cultivation. Since C. bulbifera X fragilis var. mackayi represented an undescribed hybrid, the writer decided to study the fern further. Unfortunately, Dr. Benedict’s house in Rockford was demolished years ago. A search of the vacant, weedy lot where the house once stood revealed no evidence of any cultivated ferns. Therefore, the current existence of the hybrid is unknown. *Illinois Natural History Survey, Natural Resources Bldg., 607 E. Peabody Dr., Champaign, IL 61820. AMERICAN FERN JOURNAL: VOLUME 72 (1982) FIG. 2. Photograph of the holotype of Cystopteris x illinoensis. R. C. MORAN: CYSTOPTERIS X ILLINOENSIS 43 Morphological evidence indicates that the hybrid is intermediate between C. bulbifera and C. fragilis var. mackayi. The small, scaly bulblets, typical of other taxa of hybrid origin involving C. bulbifera, were found with the specimen material, but not attached to the fronds. Examination of the spores showed them to be aborted; in addition, many sporangia were also aborted. No known Cystopteris hybrid in eastern North America has aborted spores. Presumably this plant represents a sterile triploid hybrid, since C. bulbifera is a diploid and C. fragilis var. mackayi is a tetraploid (Blasdell, 1963). The large size of the fronds also suggests a triploid condition, for many triploid hybrid ferns are large in comparison to their parents (W. H. Wagner, pers. comm., 1980). The fronds have somewhat elongate apices with many pinna pairs and with occasional glandular hairs. These latter characters are typical of C. bulbifera. Dr. Benedict indicated on his handwritten label that the plant was, “a very beautiful large black stemmed hybrid.” However, the pressed specimens have rich, red-brown, rather than deep black stipes. The dark, sclerotized stipe would appear to be a character acquired from C. fragilis var. mackayi. Unfortunately, the rhizome was not preserved; however, a short note found with the specimen stated, “Rhizome not creeping.” Certainly the circumstantial evidence that the hybrid was growing with C. bulbifera and C. fragilis var. mackayi suggests the likelihood that these are the parents involved. It was decided to gain further evidence of the plant’s hybrid nature by using two- dimensional paper chromatography to study patterns of phenolic compounds. It was hypothesized that the hybrid’s chromatogram might be additive with respect to different phenolic compounds observable in the two parental chromatograms. The phenolics were extracted in 80% methy! alcohol and examined by standard methods (Mabry et al., 1970). Plants of C. bulbifera and C. fragilis var. mackayi used in the chromatographic analysis were collected in Winnebago County, Illinois. The results showed that all observable phenolic compounds present in the chromatogram of hybrid material were also present in one or both chromatograms of the parental material, i.e. the hybrid did not contain any unique compounds that were not present in either parent (Fig. /). Most importantly, the hybrid material contained one phenolic compound present in C. bulbifera which was not present in C. fragilis var. mackayi, and one other phenolic compound present in material of C. fragilis var. mackayi but absent in material of C. bulbifera. Thus, the hybrid’s chromatogram is, in large part, additive and supports the hypothesis that C. fragilis var. mackayi and C. bulbifera are the two parent species involved. The following is given as a formal diagnosis: Cystopteris x illinoensis R. C. Moran Fig. 2. Taxon originis hybridae ex Cystopteride bulbifera et C. fragili var. mackayi; parentibus ambobus sporis abortivis et vel bulbilis parvis squamosis statim distinctum. Frondes usque ad 50 cm longi; ae et rhachis inferna atrorubella- brunnea; lamina morphologiae intermediae inter ambas parentes plerumque latissima basi; planta saxatilis saxis calcareis. TYPE: Cultivated in the garden of Ralph C. Benedict at Rockford, Illinois, from wild material found once in an old quarry in Winnebago County, Illinois, date uncertain (probably 1960's), R. C. Benedict (ILL: isotypes ILLS, SIU). 44 AMERICAN FERN JOURNAL: VOLUME 72 (1982) I would like to thank Drs. P. Mick Richardson and David A. Young for their help with the chromatographic analysis and Drs. Warren H. Wagner, Jr. and James H. Peck for many interesting discussions and encouragement. LITERATURE CITED BLASDELL, R. F. 1963. A monographic study of the fern genus Cystopteris. Mem. Torrey Bot. Club. :1-102. MABRY, T. J., K. R. MARKHAM, and M. B. THOMAS 1970. The Systematic Identification of Flavonoids. Springer-Verlag, New York. REVIEW “FLORA OF CHIAPAS, PART 2. PTERIDOPHYTES,” by Alan R. Smith. 370 pp. California Academy of Sciences. 1981. $30.00 + postage.—This is the first pteridophyte Flora of a state of Mexico to be published in twenty years, and the first modern pteridophyte Flora for any of the more tropical Mexican states. The number of pteridophytes is much greater than in the drier states of northern Mexico; Smith treats 46 fern allies in five genera and 563 ferns in 99 genera. The volume begins with an introduction, followed by a key to the genera of ferns: Smith notes that tropical ferns can be keyed more reliably to genus than to family because of the technical characters which must be used to distinguish ferns at the family level. The genera and species are treated alphabetically, rather than in systematic order. The synonymies pertain mostly to species described from Mexico and Central America, and include valuable data concerning types. Many species are handsomely illustrat- ed, and literature references are given to illustrations in other works, which is useful to those who have access to good pteridological libraries. Original descriptions, habitat notes, and phytogeographical notes are included for each taxon. Some specimens, especially those collected by Dennis Breedlove, are cited. The fern-allies are similarly treated following the ferns. An appendix of abbreviations, an adden- um, and an index to accepted names and synonyms concludes the volume. Because of the affinities of the Chiapas flora to that of Guatemala, neighboring countries, and to the adjacent Mexican states, Smith’s Flora will be useful beyond the boundaries of Chiapas. Everyone concerned with Central American pteridophytes should have a copy of this book. Readers interested in a full discussion of the vegetation of Chiapas are directed to “Flora of Chiapas, Part |. Introduction to the Flora of Chiapas,” by Dennis E. Breedlove, who is the general editor of the entire “Flora of Chiapas,” which is projected to appear in many volumes over several years. The California Academy of Sciences is to be congratulated for publishing these well prepared works at an affordable price.—D. B. L. AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 2 (1982) 45 New Records and Distributional Notes on Maryland Pteridophytes STEVEN R. HILL* and RICHARD E. RIEFNER, JR.** During the course of a critical appraisal of the state list of rare and endangered vascular plants of Maryland (Broome et al., 1979), we discovered several interesting range extensions for the pteridophytes. A full treatment of the ferns and fern-allies of Maryland, Delaware, and the District of Columbia was prepared by Reed (1953), who provided dot distribution maps for all of the taxa. These two reports have enabled us to determine gaps in distribution, and our extensive field and herbarium studies have turned up county records for twelve Maryland species. We hope that this information will be of use to those interested in the ferns of the eastern United States in general. Lycopodium clavatum L.—In our experience, the Running Club-moss in Mary- land is most frequent in moist areas of the mountains of Garrett County. If is far less frequent in the piedmont and coastal plain regions of the state. This new locality is at the border between the latter two phytogeographic provinces. Anne Arundel Co.: 2.9 mi SW of Odenton post office, floodplain of the Little Patuxent River. 25 Oct 1970, D. Bystrak s. n. (MARY). Lycopodium tristachyum Pursh—The infrequent Ground Cedar ts historically known from Cecil, Garrett, Prince George’s, Wicomico, and Worcester Counties in Maryland (Reed, 1953), but few current localities are known. Our new record is from an old collection, and the current status of the species in the state should be examined. Montgomery Co.: Cropley Falls, 9 Dec 1950, J. H. Penson s. n. (MARY). Selaginella rupestris (L.) Spring—The Rock Spike-moss in Maryland is known from Allegany, Baltimore, Frederick, and Montgomery Counties, with a possible site in Garrett County (Reed, 1953). It seems to be locally frequent on the slightly calcareous shale barrens of Allegany and Washington Counties, but is quite infrequent overall in the state. The species is declining in Massachusetts (Coddington & Field, 1978), is vulnerable and declining in New Jersey (Snyder & Vivian, 1981), and has apparently become extinct in Delaware (Tucker et al., 1979). It should probably be considered for inclusion in the Maryland list of threatened and endangered species. Washington Co.: Shale barren ledges, High Germany Road ca. 0.3 mi N of Sideling Hill Creek, 2 May 1981, S. R. Hill & R. E. Riefner 9931 (MARY, VT); Shale barren at E side of Sideling Hill Creek at Rte. 40, 2 May 1981, S. R. Hill & R. E. Riefner 9949 (MARY, VT). Ophioglossum vulgatum L.—This new locality for the Adder’s-tongue is a western range extension of ca. 50 miles within Maryland. It also seems significant in that most known populations of the species occur on the coastal plain, rather than in the ridge and valley province of the northwestern portion of the state (Reed, 1953) *Department of Botany, University of Maryland, College Park, MD 20742. **8716 Avondale Rd., Baltimore. MD 21234 = 46 AMERICAN FERN JOURNAL: VOLUME 72 (1982) Washington Co.: Along Rabble Run, Catholic Church Road near Forsythe, 13 June 1981, S. R. Hill & R. E. Riefner 10325 (MARY, NY, TAES, VT). Osmunda claytoniana L.—The Interrupted Fern is the least frequent of the three Osmunda species in Maryland and is nearly always restricted to the piedmont and mountain regions of the state (Reed, 1953). This new record places the species at the edge of the coastal plain, a region where it is rare. Anne Arundel Co.: 1.1 mi S of Odenton post office, hillside, 7 June 1970, D. Bystrak s. n. (MARY). Adiantum pedatum L.—The Northern Maidenhair is frequent in the piedmont and mountain areas of Maryland, but is rare on the coastal plain. If the label data on the St. Mary’s County specimens are accurate (the locality is imprecise), this population would be the southernmost known in the state. Prince George’s Co.: Vicinity of College Park, 14 July 1949, W. S. Glidden 22 (MARY); Paint Branch Creek, College Park, Oct 1953, R. E. Zarza s. n. (MARY). St. Mary’s Co.: Shaded woods near Patuxent River, 21 May 1964, S. A. Riley s. n. (MARY). Asplenium ruta-muraria var. cryptolepis (Fern.) Wherry—The American Wall-rue Spleenwort was included on the Maryland list of threatened vascular plants (Broome et al., 1979) and was given undetermined (UNDT) status. We have located ten healthy populations on the calcareous bluffs along the Potomac River from Allegany to Frederick Counties and feel that while it is local, it is not at present endangered in the state. Reed (1953) recorded it from Washington and Frederick Counties only. Allegany Co.: Limestone crevices along Western Maryland Railroad at Pinto, 10 May 1981, R. E. Riefner 81-78 (MARY). Cheilanthes lanosa D. C. Eaton—The Hairy Lip Fern is occasional on rock outcrops on the piedmont and towards the mountains of western Maryland. The new county record is adjacent to known localities in Baltimore County. The fern has not yet been found in Garrett County, but is to be expected there. Howard Co.: Rock ledges of overlook at Patapsco State Park along Patapsco River near Rte. 40 bridge, 19 Nov 1981, R. E. Riefner 81-850 (MARY). Dryopteris goldiana (Hook.) Gray—Goldie’s Fern has been frequently confused with Dryopteris celsa (Palmer) Small in several fern treatments (e. g., Reed, 1953), and these have combined records for the two. Others recognize D. celsa as D. goldiana subsp. celsa Palmer. Studies on the cytology of the species complex involved (Montgomery, 1975; Walker, 1962) demonstrate that the two species are distinct. In addition, D. celsa is a coastal plain species, whereas D. goldiana is a mountain species. The specimen cited was growing in a rich, moist, beech woodland in the mountains. We have not examined all of the voucher specimens involved in Reed's treatment, but those from Montgomery, Garrett, and Baltimore Counties that we have seen (at US) seem to be D. goldiana, although the specimen from Worcester Co. cited by Broome et al. (1979) is probably D. celsa, which is less frequent in the state. Frederick Co.: Slopes near Owens Creek, Foxville Road. Catoctin Mountain National Park near Thurmont, 4 Oct 1981, S. R. Hill & A. D. Cress 10820 (MARY. NY, VT) HILL & RIEFNER: MARYLAND PTERIDOPHYTE RECORDS 47 Matteuccia struthiopteris (L.) Todaro—The Ostrich Fern is an endangered species in Virginia known from few locations (Porter, 1979), but its status in aryland was DSi ee to be undetermined by Broome et al. (1979). We have located five populations extant in the state in Allegany, Baltimore, Harford, and Washington Counties, and the plant is historically known from Carroll, Howard, Montgomery, and Prince George’s Counties as well (Reed, 1953). We are convinced that the Washington County population is indigenous despite Reed’s (1953) sugges- tion that the western and southwestern populations are escaped from cultivation. The Roundtop population was associated with several other infrequent plants of alkaline alluvial soils, such as Gleditsia triacanthos, Ptelea trifoliata, and Chasmanthium latifolium. Washington Co.: Rich woods and bottomland along the Potomac River just W of Roundtop Mountain, Fae 1981, R. E. Riefner 81-138 (MARY). Woodsia ilvensis (L.) R. Br.—The Rusty Cliff Fern is very rare in Maryland, having a preference for exposed shale cliffs associated with shale barrens. At the Washington County locality the shale is calcareous, although Reed (1953) remarks on the non-calcareous preference of the species. It was growing in full sun with Phlox subulata, Arenaria stricta, Cystopteris fragilis, Comandra umbellata, and Tephrosia virginiana. The only other population that we have seen is at the shale barren at Town Creek (also calcareous shale) in Allegany County, where only a few plants survive (Riefner, 1981). We have discovered one other old collection from Allegany County (Flintstone, 3 June 1928, N. C. Knappa s. n., Patuxent Res. Refuge Herb.). Washington Co.: Shale cliff i Sideling Hill Creek, High Germany Road, 2 May 1981, R. E. Riefner & S. R. Hill 81-55 (MARY Azolla caroliniana Willd.—The Mosquito Fern is said to be very frequent in watercress beds near the Monocacy River by Reed (1953). We have seen only the specimen from Prince George’s County and a specimen from Kenilworth Gardens, Washington, DC (W. Preston, Jr. s. n., MARY). These seem to have been chance introductions and probably did not persist. The Mosquito Fern is undoubtedly a frequent “hitchhiker” on waterfowl migrating north in the spring from warmer regions, where the plant is frequent in swamps and on still-water ponds. The plant may also be introduced from aquaria. Prince George’s ne Golf course pond, 14000 Old Marlboro Pike, Upper Marlboro, 15 July 1976, K. Hummel s. n. (MARY). This is Scientific Article No. A3071, Contribution No. 6136 of the Maryland Agricultural Experiment Station. LITERATURE CITED BROOME, C. R. et al. 1979. Rare and Endangered Vascular Plant Species in Maryland. U. S. Fish and Wildlife Service, Newton Corner, CODDINGTON, J. and K. G. FIELD. 1978. Rare and Endangered Vascular Plant Species in Massachusetts. U. S. Fish and Wildlife Service, Newton Corner. MONTGOMERY, J. D. 1975. Dryopteris celsa and D. clintoniana in New Jersey. Amer. Fern J. 65-69 48 AMERICAN FERN JOURNAL: VOLUME 72 (1982) PORTER, D. M. 1979. Rare and Endangered Vascular Plant Species in Virginia. U. S. Fish and Wildlife Service, Newton Corner, MA. REED, C. F. 1953. The Ferns and Fern-allies of Maryland and Delaware including the District of olumbia. Reed Herbarium, Baltimore, MD. RIEFNER, R. E. 1981. Notes on some proposed rare and endangered vascular plant species in Maryland. Phytologia 47:397—403. SNYDER, D. B. and V. E. VIVIAN. 1981. Rare and aga Vascular Plant Species in New Jersey. “Fish and Wildlife Service, Newton Corner, MA. TUCKER, A O. et al. 1979. Rare and Re ory Seed Plant Species in Delaware. U. S. Fish and Wildlife Service, Newton Corne WALKER, S. 1962. Further studies in Ke pon Dryopteris: The origin of D. clintoniana, D. celsa, and related taxa. Amer. J. Bot. 49:497—-503. REVIEW “ILLUSTRATIONS OF PTERIDOPHYTES OF JAPAN,” volume 2, S. Kurata and T. Nakaike, eds. x + 648 pp. + map. 1981. University of Tokyo Press, yen 10,000 (about $47).—This is the latest of a great array of excellent plant books produced in Japan for the large portion of the population seriously interested in botany. On the glossy paper of this lavish and heavy volume 100 Japanese ferns (from a total of nearly 800) are presented, each with a black and white photo of its appearance in the wild, a line drawing with insets of important details, and a dot map showing precise distribution in Japan. A separate section in the back has a photo- micrograph of a spore of each species. Almost everything is in Japanese, except for botanical names and numbers, so the value of the book for the non-reader of Japanese is in the illustrations. The sharp-detail photos put you inside the forests of Japan looking at moss-covered and lichen-covered trunks, and at boulders and embankments, and the ferns are right there looking just as they really do. The line drawings are not cosmeticized or stylized—the occasional small insect bite or other imperfection is faithfully render- ed. What you see is what you get. This volume deals mainly with Asplenium and the genera allied to Polypodium and Grammitis. Two new combinations are made, presumably due to the reinterpre- tation of a type specimen by Dr. Toshiyuki Nakaike: the fern usually known as Neocheiropteris subhastata (Bak.) Tagawa is renamed N. buergeriana (Miq.) Nakaike, and the fern that was Microsorium buergerianum (Miq.) Ching is now M. brachylepis (Bak.) Nakaike. My only criticism is that about half the book is taken up by lists of localities that to a large extent duplicate infomation already available from the fine dot maps.— G. Price, Herbarium, University of Michigan, Ann Arbor, MI 48109 AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 2 (1982) 49 Grammitis succinea, the First New World Fern Found in Amber LUIS DIEGO GOMEZ P.* Although Dominican ambers are known for their arthropod fauna, no systematic examination has ever been attempted of the plants they contain, even though plant remains usually are well preserved in ambers (Czeczott, 1960). Non-vascular cryptogams (mosses and lichens) also are commonly preserved, but apparently only two vascular cryptogams have ever been reported, both ferns from Baltic ambers (Caspary & Klebs, 1907; Magdefrau, 1957). The present report is the first record of a fern from New World amber and is the first species of the genus Grammitis to be found in amber. There are several deposits of amber in Central and South America, but the most famous New World deposits are at Simojovel, Chiapas, Mexico, and in the Dominican Republic on the Caribbean island of Hispaniola. The latter, considered like the Chiapas deposits to be late Oligocene in age, has been known since Columbus reported on it in the account of his second voyage to the West Indies (Langenheim, 1964). Dominican amber contains many well preserved arthropods and some fragmentary, little-studied plant materials. This has led to extensive exploitation of these fossil resins, which are sold as elaborate jewelry or in bulk, but almost always without precise data as to their location on the islan The plants from which the Dominican amber originated was for many years unknown. Sanderson and Farr (1960) suggested Pinus as a possible source because of the strong, turpentine-like odor obtained from freshly sawed or filed amber. Recently, Langenheim (pers. comm., 1980) has indicated that Dominican amber is the fossilized resin of Hymenaea trees (Leguminosae). The fragment of Dominican amber containing the fern described in this paper was donated to the Museo de Entomologia, Universidad de Costa Rica, where it was polished by abrasion with carborundum and water on rotating, leather-covered wooden disks. The amber was photographed on a light table. All microscopic observations and measurements were made under a dissecting microscope using transmitted light, reducing refraction by immersing the amber in a 95% solution of ethanol. Two small samples of amber were chipped from the specimen in an unsuccessful attempt to find spores of the fern. No attempt was made to extract the fern from the amber because of the scarcity of the material, although extraction is possible using Voigt’s (1936) method. The reconstruction was traced from a photographic print, correcting for curvature in the material. Grammitis succinea L. D. Gomez, sp. nov. Figs. 1-2. Rhizome and stipe unknown. Fronds of unknown length, probably ca. 35 cm long, ca. 60-70 mm wide near the apex, pinnate, herbaceous. Rachis dark brown, terete, saineok scales, densely hairy; hairs unicellular, setose, stiff, 1-1.8 mm long, ca. 0.3 mm in diameter at the widest point. Pinnae 30-40 mm long, 4-6 mm w wide, lanceolate, Sede entire except shallowly crenate towards the apex, rounded to *Museo Nacional de Costa Rica, Apartado 749, San José, Costa Rica. 50 AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 2 (1982) FIG. 1. Fragment of amber bearing Grammitis succinea; an embedded insect is circled, x 3.6. obtuse at the apex, inequilateral at the base, the upper base rounded, subauriculate, truncate, the lower base decurrent. Laminae and costae pilose on both surfaces, the hairs similar to those of the rachis. Costae prominent dorsally, straight in the proximal two-thirds, flexuose and tenuous near the apex. Veins free, 1-forked, never reaching the margin, slightly curved towards the pinna apex. Sori discrete, 12-14 per pinna, medial, subalternate to opposite, round, small, exindusiate, terminal on the acropetal branch of the vein, usually | per vein pair, exceptionally 2. Spores unknown. TYPE: Without precise locality, Dominican Republic, Monge 80—/2, deposited in the Museo de Entomologia, Universidad de Costa Rica. Because of the presence of setose, stiff hairs (rather than scales along the rachis) and sporangium stalks of only one column of cells. this fern is readily placed in the Grammitidaceae, rather than the Polypodiaceae, which is the other fern family to which it could belong. (The free, forked veins are found in both families. particular- ly in species of the Polypodium pectinatum-plumula complex, but setose hairs are L. D. GOMEZ: GRAMMITIS SUCCINEA 51 f Bas ly Fi 2A, 1g BREIL _— portion of Grammitis succinea frond, 3.6. FIG. 2B. Pattern of venation and soriation, unknown in Caribbean species of the Polypodiaceae.) Spores, which are monolete in the species of Polypodiaceae but trilete in those of the Grammitidaceae, would be useful to confirm the placement of the species, but unfortunately have not been recovered from the amber, an absence previously reported in some European ambers (Schubert, 1961), but not others (Wetzel, 1953). Among the New World genera of Grammitidaceae, the plant is obviously referable to Grammitis sect. Cryptosorus, which has pinnatisect laminae bearing several sori on each elongate lobe, rather than to any other genus of the family. Other plant remains known from Dominican ambers need systematic study to develop information on the age of the deposits, as well as to provide floristic data for use in comparing them with plant remains found in Chiapas, Haiti, and Cuba. It is likely that such data would be useful in solving biogeographical problems in the Caribbean area. This research was partially funded by a grant from the Consejo Nacional de Investigaciones Cientificas y Tecnolégicas (CONICIT), Costa Rica. 52 AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 2 (1982) LITERATURE CITED CASPARY, R. and R. KLEBS. 1907. Die Flora des Bernsteins und anderer fossiler Harze des ostpreussischen Tertiars. Abh. Konig]. Preuss. Geol. 4:1—182. CZECZOTT., H. 1960. The flora of the Baltic amber and its age. Prace Mus. Ziemi 4:119-145. LANGENHEIM, J. 1964. Present status of botanical studies of ambers. Bot. Mus. Leafl., Harvard Univ. 20:225—287. MAGDEFRAU, K. 1957. Flechten und Moose im Baltischen Bernstein. Ber. Deutsch. Bot. Ges. : 435 433-435. SANDERSON, M. W. and T. H. FARR. 1960. Amber with insect and plant inclusions from the Dominican Republic. Science 131:1313. SCHUBERT, K. 1961. Neue Untersuchungen iiber Bau und Leben der Bernsteinkiefern (Pinus succinifera (Conw.) emend.). Beih. Geol. Jahrb. Niedersachsische Land. Bodenforsch. Han- nover 45:1-143. VOIGT, E. 1936. Die Lackfilmmethode. ihre Bedeutung und Anwendung i. d. Palaeontologie, Sedimenpetrographie und Bodenkunde. Z. Deutsch. Geol. Ges. 88:272—292. WETZEL, W. 1953. Mikropalaéontologische Untersuchung des schleswigholsteinischen Bernsteins. Neues Jahrb. Geol. Palaontol., Mitt. 7:31 1-321. REVIEW ASPECTS OF PLANT SCIENCES, VOLUME 3. PTERIDOPHYTES, edited by S. S. Bir. Today and Tomorrow’s Printers and Publishers, New Delhi. vi + pp. $15.00.—This volume contains several substantial research papers by well known Indian botanists. The focus of the papers is on the anatomy, classification, cytology and morphology of pteridophytes. B. K. Nayar writes on fern classifica- tion. T. N. Bhardwaja reviews the cytology, morphology, and morphogenesis of Marsilea. S. P. Khullar and S. S. Sharma revise the Himalayan species of Onvchium, with emphasis on their cytology and anatomy. S. N. Patnaik and B. L. Narasama report their experimental work on growing fern leaves in vitro. B. D. Sharma surveys the Indian pteridophyte fossils from the Mesozoic Era. S. S. Bir et al. survey the stomatal types found in 33 species of Indian Athyrium, Cystopteris, Diplazium, Drvoathyrium and Hypodematium. Lastly. S. §. Bir and C. K. Trikha report on the stelar anatomy of 19 species of Indian Polypodiaceae. This reasonably priced volume will be of interest to many pteridologists. It is available from the publisher at 24-B/5, Original Road, Karol Bagh, New Delhi 110005, India.—D. B. L. AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 2 (1982) 93 A New Tropical American Species of the Genus Lycopodium BENJAMIN @LLGAARD* During studies of the Lycopodiaceae for the “Ferns and Fern Allies of Guatemala” pe Robert G. Stolze, a total of 24 species was found to be represented in this untry. One of these is an undescribed species which occurs also in northern Aedes South America. bee pn amentaceum B. Qllgaard, sp. no Fig. 1-4. Species Lycopodio Oe Mett. et L. cuneioi Hieron. Steg affinis, caulibig tenuibus pendentibus usque ad 60 cm longis, dichotomis 3mm crassis in sicco foliis exclusis, icem ae eraatim us ne ‘ad 0.5 mm sate entibus, teretibus vel striato- aiid presertim parte inferiore saepe rubris: mi foliis dim morphi s; rami basales foliis expansis ‘aan ad 50 cm longi, (10)12—20 mm crassi foliis inclusis, usque “ quinquies bifurcati; rami apicales foliis valde doe: seu zonatim vel continuiter sporangiferis, usque ad 30 cm longi, ad dec s bifurcati; folia ramorum basalium decussata vel subdecussata inennens a. mm longis, patentia vel eo iads, oblonge lanceolata vel rad apicibus obtusis vel mage mucronulatis, basibus et apicibus saepe e symmetricis, recta vel recurvata, (4)6—10 mm longa, 2—3(3.5) mm in medio lai laminis plerumque torsione basal verticalibus, plana vel venas secus deorsum rodiis m lon laxe vel arc acuta a ie rcronats, intro subhicna es amplectentibus, abaxialiter rotundata usque ad carinata, 1.2-1.7 mm longa, 1.2—1.5 mm lata; sporangia reniformia, ca. | mm la TYPE: Camino al Paramo de Los Monsalves, Edo. Mérida, Venezuela, epifito péndulo, sobre tronco, en la faja superior de la selva montanera, 3000 m alt., 10 June 1952, Vareschi & Pannier 1817 (VEN; isotype M). According to information kindly provided by Dr. Julian A. Steyermark, the location of the Péaramo Los Monsalves is just northwest of the main city of Mérida, ca. 8°42'N, 71°08’ W. PARATY PES: GUATEMALA: Depto. Quiché: José J gnacio Aguilar 828 and 10/9 in 1942 (F). Depto. Guatemala: On tree in dense forest, slopes of Volcan de Pacaya, between San Francisco Sales and the base of the active cone, 1800-2300 m alt.. 20 Dec 1940. Standley 80538 (F). COLOMBIA: Depto. Cundinamarca: Road to east from Guasca, on wet bank, 3200 m alt.. 6 Jun 1947, Haught 5826 (COL). Monserrate, Bogoté, 2900 m alt., Lindig /504 (P). Depto. Norte de Santander: Ocana. paramos, 8000-10000 ft alt., Schlim 468 p. p. (P). VENEZUELA: Edo. Mérida: Selva de Podocarpus, Paéramo Monsalves, 3190 m alt., 10 Nov 1952, Vareschi 2232 and 2272 (VEN) Lycopodium amentaceum has been found in high montane forest and in cloud forest, as an epiphyte, or occasionally hanging from banks. It is a close relative of L. callitrichifolium Mett. and L. cuneifolium Hieron., a group of very delicate pendent species from the northern Andes and Central America. The main differ- ences which separate L. amentaceum from these species are shown by the typical outline of the expanded leaves (Figs. 5-6). In the size of the expanded leaves it also resembles L. phylicifolium Desv. ex Poir., but its leaves are broader and more obtuse, both at the base and apex. *Botanical Institute, University of Aarhus, 68 Nordlandsvej. 8240 Risskov, Denmark. AMERICAN FERN JOURNAL: VOLUME 72 (1982) Y oe Nt Ni Saw een Years : . “aha ey pe | a oa a = y \ ys — a >i = x Sos a LL sal Ss ‘x Tee aS LGPL 5 2YEPEO re ee be ——/ Fes m, Pest ir os 0 ti a PNT Sa iti “S ee APY > ie Ne te a aS eS NH 3 Tond Habit. FIG. 2. FIGS. of basal division with expanded leaves. FIGS. 3 and 4. Portions of apical sporangiate divisions. 1-4. Isotype of Lycopodium amentaceum, Vareschi & Pannier 1817 (M). FIG. Portion B. OLLGAARD: A NEW TROPICAL AMERICAN LYCOPODIUM 55 lcm FIGS. 5-6. Portions of basal divisions with expanded leaves. FIG. 5. L. cuneifolium Hieron., Schlim 881 (P). FIG. 6. L. callitrichifolium Mett., Holm-Nielsen et al. 3988 (AAU). The plants treated and illustrated as L. callitrichifolium by Vareschi in Lasser (Flora de Venezuela 1(1):41, t. 2A. 1969) belong to L. amentaceum, and include the type of the latter. The Guatemalan specimens had been identified as L. skutchii Maxon, a name which I regard as a synonym of L. myrsinites Lam. From the latter species it can be distinguished by its red stems, smaller sporophylls (more than 2 mm long in L. myrsinites), almost uniform leaves (highly variable and often recurrent to expanded in shape in L. myrsinites), and obtuse leaves (acute in L. myrsinites). 56 AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 2 (1982) Specimens from Fée’s Pteridological Collection at the Botanical Garden of Rio de Janeiro PAULO GUNTER WINDISCH* As cited in a recent note (Windisch, 1981), specimens from A. L. A. Fée’s pteridological, lichenological, and phanerogamic collections can be found in the Herbarium of the Botanical Garden of Rio de Janeiro (RB), as the result of the acquisition made by the Brazilian Emperor Dom Pedro II during his visit to Fée in 1871. Although this has been known by some botanists, for many the latest information about this collection was presented by Stearn (1962, p. 220), who held the traditional view that Fée’s private herbarium had not been traced and was believed to have been lost at sea en route to Brazil. In fact, the pteridophytes at RB are represented by 2,940 specimens bearing Fée’s original annotations and labels, which are a clear indication that the material longed to Fée’s private collection. In addition, some specimens have been remounted without the original labels and with incomplete data on the new labels, making it difficult to ascertain with certainty the origin of the material, although it is likely that these specimens do belong to the Fée collection. Of the original material, 381 specimens appear to be types of taxa described by Fée. A few of these specimens have a note indicating that they were used to prepare lithographs (which were used in Fée’s ““Mémoires’’), and some sheets have pencil drawings copied from original plates or have illustrations cut out from lithographs. A great number of specimens bear Fée’s unpublished herbarium names. In addition to the foregoing specimens, many others are classical collections by such men as Cuming, Galeotti, Martius, and Gardner, some of which are isotypes of taxa described by Fée’s colleagues Kunze, Klotzsch, Bory, and John Smith. Among the pteridophytes, some groups are poorly represented, such as the tree ferns. It may be that the acquired collection was not complete, that parts of it are still misplaced, or that some specimens have been excluded from the Herbarium due to excessive damage. It will be difficult to determine precisely what happened to the specimens since their acquisition. The following list of presently available pteridophyte types (or probable types, in a few cases) with relevant label data will certainly be of help to those working on nomenclatural problems related to the taxa represented. In some cases, no collector was indicated on the specimen label. In other cases, there is no collector’s number, but the year of collection was given; these are cited as “in 1865” or whatever year applies. Numerals which are illegible on the specimen labels but which are known from being published in Fée’s “Mémoires” have been placed in brackets in the list. Similarly, missing collectors or herbaria of origin also have been placed in brackets. Abacopteris truncatum (Philippines, — 298). Acrostichum alatum (Guyane Frangaise, Leprieur in 1839), A. angustifolium (Bolivia, Bridges 1850), A. attenuatum (ex horto Lipsteds A. Hestadvode s (Mexique, Galeotti 6297), A. callolepis (Mexique, Galeotti), A. cochleariaefolium (Andes de Quito, tana 21[3]), A. crispatulum (Quito, Jameson), A. * Instituto de Biociéncias, Universidade Estadual Paulista, Caixa Postal 178. 13500 Rio Claro—SP, P. G. WINDISCH: FEE’S PTERIDOLOGICAL COLLECTION 57 oo (Venezuela, Linden), A. oo (Perou, Dombey), A. a tee Schaffner 2b) iA. a rei 1486), A. Herminieri (Guadeloupe, L’Herminier), A. intermedium oem OEE , A. tea naa (Mexique, serappanl 19), A. tines (les Sandwich, rence in 1836 : ot minutum (Pohl in 1817-21), A. obcrenatum (Guadeloupe, L’Herminier “ 186 A. plumosum (Guyane, obi 44[6]), re rhabdolepis (Quito, Jameson in 1845), Sebliniaise (Amerique Trop., Ocafa, Schlim 622), A. stigmatolepis (Neilgherries), A. stramineum (Brésil, See 3322) A. ae (Venezuela, aig Fendler 272). Adiantum Claussenii (Brésil, Nouv. Fribourg, Claussen 124), A. extensum (Mexique, Veracruz, Daa. Shales r 41), A. fuliginosum (Guyane Francaise, Leprieur 256 ex Herb. Moug.), A. gracile (Brésil, Claussen), A. jacobinae (Brésil, Jacobine, Blanchet), A. jamaicense (Jamaica, Wilson), A. ier), A. - pseudocapillus (Cap de Bonna Spéi, Drége), A. subartetatum (Brésil, Bahia, Blanchet 2373), A. tomentellum (Brésil, Minas Gerais, Claussen), A. trapezoides (Mexique, Veracruz, Galeotti 6317), A. tricholepis (Mexique, Galeotti 64 Alsophila Schaffneriana (Mexigue, San Martin, Schaffner 232). aang latifolia (Nouv. Grenade, Ocafia, Schlim 656). Aspidium acrocarpon (St. Domingue, Tussac), A. agatolepis Ream S. Agostin, cathe co A. ameristoneuron (Cuba, orig. inconnue), A. asperulum (Guadeloupe, L’Herminier in 186(1)}), capitainii (Guadeloupe, L’Herminier in 1861), A. chrysolepis a du Cap Vert, Webb), He ie (Nouv. Grenade, Ocafia, Schlim 323), A. frondulosum (Bourbon, Bory), A. steer (Guadeloupe, Germain in 1861), A. jucundum (Mexique, Galeotti and Cuba, Linden 2115), A. microthecium (Philippines, Cuming 13), A. obtusifolium (Mexique, Huatusco, Schaffner 213), : pachychiamys (Guadeloup e, L’Herminier in 1861), A. pauper (Martinique, Mlle. Rivoire), A. trichophorum (“Cuba,” ex Herb. Bory), A. trichotomum (Cochinchine, Tourane, Gaudichaud). Asplenium bifissum (Cuba, Linden 1888), A. cladolepton (Nouv. Grenade, Ocafia, Schlim 324), A. 5), A. distans (Mexique, Galeotti 6579), A. extensum (Nouv. Grenade, Ocafa, Schlim 629 flageliferum (Nouv. Grenade, Ocania, Schlim 63), ‘ cornea (Mexique, Galeotti 6369), A. herbaceum (Nouv. Grenade, Schlim 326), A. inaequalidens (Guadeloupe, L’Herminier), A. integrum (Guadeloupe, L’Herminier), A. lamprocaulon (Mexique, Galeotti 6340), A. leptophyllum (Mexique, Galeotti 6446 and Mexique, Schlim 479), A. macrodon (Quito, Jameson in 1845), A. myapteron se Orizaba, Schaffner 70 and Mexique, Popocatepetl, Schaffner 294), A. oa ardonetnt (Nouv. Grenade, Ocana, Schlim 492), A. progrediens (Mexique, Huatusco, Schaffner 54), A. pumilum var. hones iophylloides (Abyssinia, ee ), A. surinamense (Guyane Hollandaise, elt 183b), A. set (Quito, Jameson in 1848), A. ternatum (Nouv. Grenade, Ocafia, Schlim 327). ce pea (Bourboun, Montbrison and Madagascar, Nos-Beh, Pervillié), B. villosum (“in hort. ludg. Batavo”’) Blechnum extensum (Brésil, Claussen), B. helveolum (Venezuela, Caracas, Moritz 17), B. heterocarpum (Brésil, Claussen), B. Schlimense (Nouv. Grenade, Schlim 7. Campylonearon caudatum (Mexique, Cordoba, Huatusco, Schaffner 176), C. cubense (Cuba, Linden 1912), C. jamesoni (Quito, Jameson), C. minus (Brésil, Rio, Glaziou and Ameriq. Sud, Brésil, Glaziou ex Herb. Gauthier), C. Moritzianum (Caracas, Moritz 3), C. xalapense (Mexique, Galeotti 6273). Cardiochlaena alata (Philippines, Cuming 148), C. ampla (Guadeloupe, L’Herminier), C. confluens (Guadeloupe, L’Herminier), C. laevis (Philippines, Cuming), C. macrophylla var. crenata (Caracas, Moritz 112), C. macrophylla var. distans (Brésil, Martius), C. pilosa (Brésil, Rio de Janeiro, Weddell 656), C. sinuosa (Philippines, Cumin Cassebera paradoxa (Brésil, Serra ds Orgaos, Gardner 59[3]0), C. Se eee .. . Argen.) Cheilanthes aspidioides (Mexique, Galeotti 6557), C. cucullans (Mex , Schaffner 82), C. ieabeepi Ile Ste. Catherine, Desterro), C. griffithiana (Ind. Oriental., Gril th), C. malaccensis (Malacca, Cuming 408). Chrysodiu iG sonics (Panama, Bonpland). Chrysopteris martinicensis (Martinique, Mile. Rivoire), C. microdictya (Mexique, [Schaffner _ Craspedaria borbonica (Bourbon, Bory), C. Gestasiana (Brésil, Rio de Janeiro, de Gestas), C. 58 AMERICAN FERN JOURNAL: VOLUME 72 (1982 5 4 javanensis (Java, Zollinger 1086), C. lanceolata (Surinam, Hostmann 324). Cc 270). Culcita Schlimensis (Nouv. Grenade, Ocana, Schlim 322). Cyathea Commersoniana (Ile de France, ean Cystopteris azorica (Agores, Fayal, Hochstette Dicksonia incisa (Guadeloupe, L’Herminier in 186(4)), D. neglecta (Philippines, ‘Cuming 1087), D. Didymoglossum fructuosum (Guadeloupe, L’Herminier in 1861), D. Krausii. var. subpinnatifida (Mexique, Schaffner). Diplazium acutale (Mexique, Galeotti 6289), D. anthraxacolepis (Mexique, Huatusco, Schaffner 267b and Mexique, Huatusco, Schaffner 627a), D. camptocarpon (Mexique, Cordoba, Schaffner 69), D. dissimile (Brésil, Ste. Catherine, Mors in 18 . Feei (Mexique, Veracruz, Barrancas de S Martin, Schaffner 26 D. firmum (Ceylon, Thwaites 134 grammatoides (Guadeloupe, {ex Herb. T. aoOReH: D. Schlimense (Nouv. Grenade, Ocafia, Schlim 601), D. Tussaci (St. Domingue, Tussac Doryopteris fddions var. patula (Brésil, Rio de Janeiro, Glaziou 1740). rymoglossum abbreviatum (Cochinchine, Tourane, Voy. de la Bonite, Gaudichaud), D. subcordatum se co a di cha au le Catherine, Gauthier), G. intermedium (Guadeloupe, L’Herminier), G. invertens (Mexico, Orizaba, Schaffner 494), G. lepidotrichum (Mexique, Cordoba, Schaffner 198 and Mexique, Crab Schaffner 451), G. longicaule (Nouv. Grenade, Rio H ema Lah 847), G. molestum (Mexique, Cordoba, Schaffner 180 and Mexique, Orizab a, Schaffner 180), G. pectinans (Brésil, Claussen), G. plec tolepis (Mexique, oe eee 187), G. pleopeltis Pu Lobb 263), G. Villeminianum (Nouv. Grenade, Ocana, Schlim 1 Goniopteris ts LHernt. & Fée (Guadeloupe, L'Herminier in 1861), G. heen umieionpe: 861 | 44), “ey tenera iy on inka L'Herminier i in ‘18 Grammitis limbata (Guadeloupe, Perrotet in 1824), G. longa (Java, Lobb 271), G. zevlanica (Ceylon, Gardner 56). Hemicardion crenatum (Cochinchina, —— H. cumingianum (Philippines, Cuming 68), H. macrosorum (Nouv. Grenade, Ocafia, Schlim 658). Hem ae Phage (Indiae Orientale, fare Dathousie), H. elegantissima (Mexique, Linden), H. insignis (Gi ‘Gu oupe, L’Herminier in 1861), H. lucida (Mexique, Oaxaca, Galeotti 6537), H. repanda (Cuba prpees ents (Brésil, Bahia, Blanchet), H. serrata (Brésil, Bahia. Luschnath). Hydroglossum mexicanum (Mexique, Oaxaca, Galeotti 6419). P. G. WINDISCH: FEE’S PTERIDOLOGICAL COLLECTION 59 Hypolepis chilensis (Chile, sees 6 Gaudichaud), H. helenensis (Ste. Héléne, Cuming 4[3]3), H. parviloba (Brésil, Rio Negro, Spruce 2/19). Jamesonia rotundifolia (Nouv. a nade, Ocana, Schlim 363). Leptochilus subquinquefidus (Philippines, Mindora, Cuming 3), L. Thwaitesianus (Ceylon, Thwaites 1316, L. zeylanicus (Ceylon, Thwaites 1317). Lindsaya curvans (Mexique, Galeotti 6489), L. pC none (Guadeloupe, L’Herminier in 1864 and Brésil, Bahia, Luschnath 22), L. crenulata (Ind. Orient., Malacca, Griffith), L. elegans (Colombie, Moritz 238), L. Galeottii (Mexique, Galeotti 6469), L. L’Herminieri (3 specimens, all Guadeloupe, L’'Herminier), L. montana (Guadeloupe, L’Herminier in 1862 and 1864), L. multifrondulosa (Mexique, Galeotti), L. parvula (Trinité, Germain in 1862). Litobrochia affinis (3 specimens, all Guadeloupe, L’Herminier), L. brevinervis (Guadeloupe, L'Herminier), L. Galeottii (Mexique, Galeotti 6485), L. grandis (Mexique, Mirador, Schaffner 144), L. hemipteris (Mexique, Schaffner), L. inaequalis (Trinité, Germain in 1862), L. mexicana (Mexique, Teotalcingo, Galeotti 6376), L. microdictyon (Philippines, Cuming), L. Montbrisonis (Bourbon, Montbrison), L. organensis (Brésil, Orgaos, ee 5329), L. setifera (Mexique, Cerro S. Martin, Galeotti 6571), L. Tussacii (Ste. Domingue, Tus. Lomariopsis elongata (Brésil, Bahia, a 19), L. erythrodes (Brésil, Martius 366), L Prieuriana (Guyane Frangaise, Leprieur), L. — var. caudata (Guadeloupe, L’ Herminier). Lonchitis stechnochlamis (Cap de b. Spéi, Drége), L. tomentosa (Madagascar, Pervillié). Lophosoria acaulis (Brésil, Serra do aa, Glaziou 3164), L. prostrata (Brésil, Serra do Couto, Glaziou 3165). Lycopodium eriostachys (Brésil, Orgaos, — 1788). Mertensia grandis (2 specimens, both xique, Huatusco, Mirador, Schaffner 230). Microlepia caudata (Mexique, Tele aa Galeotti 6257), M. effusa (Pul-Pinang, Lady Dalhousie, ex herb. Graham), M. Galeotti (Mexique, Teotalcingo, Galeotti 6526). Microsorium longissimum (Philippines, Cuming Myriopteris cheiloglyphys (Mexique, [Orizaba, eke 91]), M. gracilis (Ameriq. nord., Riehl 529), M. marsupianthes (Mexique, pasabagh mr rufa (Mexique, Orizaba, [Schaffner 53)). Nephrolepis Schkhurii var. minor (Linden Notholaena pruinosa (Mexique, Toluca, en: Onychium carvifolium (Philippines, Cuming 3[2]), : a, (Cuba, Linden), Oochlamys Rivoirei (Martinique, Mlle. Rivoire). Pellaea mucronata (Mexique, Shaltaes 150), P. palmescens (Philippines, Cuming), P. Weddelliana (Bolivia, Weddell 3778). Phegopteris adnata (Brésil, Rio de Janeiro, Glaziou 2398), P. Blanchetiana (Brésil, ain het 2928), P. blepharodes (Bourbon, Montbrison), P. brevinervis (Brésil, Claussen), P. cordata (Cuba, Linden 1873), P. ctenoides (2 specimens, both Ste. Domingue, Tussac), P. delicatula (Guadeloupe, i seaoniatie 1864), P. dilatata (2 specimens, both Guadeloupe, L’Herminier in 1861), P. elata (Bourbon, Montbrison), P. fluminensis (Brésil, Rio de Janeiro, Glaziou 965), P. germaniana (Guadeloupe, L’Herminier), P. Helliana (Bourbon, Hell), P. yi eng (Mexique, Huatusco, Schaffner 218), P. inaequalis (Mexique, Huatusco, Schaffner 240, , Huatusco, Schaffner 24la, and Mexique, Tututla, Schaffner 241b), P. lanata (Bourbon, bcs P. leptoptera (Ste. Domingue, Tussac), P. macrotheca (Guadeloupe, L’Herminier in 1864), P. melanorhachis (Mexique, Huatusco, Schaffner 238), P. mollivillosa (Brésil, Martius 320), P. dee ates (Bourbon, Monto ae P. nervosa (Philippines, Cuming), P. nitens (Ceylon, Walker), P. oreopteridastrum (B i Glaziou 963), P. pilulosa (Mexique, Linden), P. alg eae (Cuba, a 2 Cuba, Linden 1874 in 1844), P. rustica (Guadeloupe, L’Herminier in 1861), P. scalpturata (Bourbon, Pervillié and Madagascar, Pervillié), P. scrobiculata (Brésil, Rio de it eiro, Glaziou 1780), P. stenolepis (Mexique, prope Huatusco, Schaffner 239), P. straminea (Bourbon, Montbrison 1898), P. villosa (La Guadeloupe, 861). — & i] . = o Phlebiogonium impressum (Ind. Orientales, Griffith). Plecosorus leptocladon (Nouv. Grenade, Ocaia, Schlim 438), P. peruvianus (Perou, Ruiz). Polybotrya cyathifolia (Guadeloupe, L’Herminier), P. scandens (Mexique, Galeotti 187), P. semi- pinnata (2 specimens, both Brésil, Rio de hints Glaziou 2427). 60 AMERICAN FERN JOURNAL: VOLUME 72 (1982) Polypodium arthropodium (Mexique, Cordoba, Schaffner 185 and Mexique, Orizaba, gi! 186b), P. australe (Tenerife, [Bory] and Corse, Requien in 1845), P. blandum (Amerique merid.), callolepis (Mexique, Mecameca, Schaffner 27] and Mexique, Popocatepetl, Schaffner 272), ; camptoneuron (Cuba, Linden 1886), P. cancellatum (Cuba, Linden), P. cheilostictum (Mexique, Orizaba, Schaffner 453), P. cryptocarpon (Mexique, Cordoba, Schaffner 194), P. cubense (Cuba, Linden), P. echinolepis (Mexique, Cordoba, Schaffner 188), P. ellipsoideum (Mexique, prope S. Angel, Schaffner 211), P. filipendulaefolium (Java, Lobb 269), P. funiculum (Cuba, Linden 1885), P. glycirrhiza (Guadeloupe, L’Herminier in 1861), P. inaequale (Guadeloupe, L’Herminier in 1861), P. paneer (Mexique, Huatusco, Schaffner 199), P. leptostomum (Mexique, Orizaba, Schaffner 210), P. sorum (Quito, Jenson in 1845), P. nivosum (Mexique, Huatusco, Schaaf er 193), °P- salons (Mexique, Huatusco, Schaffner 191, Mexique, Orizaba, ein a 191b, and Mexique, Valle de Mexique, Schaffner 192), P. pubescens (Mexique, Huatus Dos Puentes, Schaffner 181), P. rhagadiolepis (Mexique, Linden, and Mexique, prope Orizab Shee 200b), P. sei (Mexique, Schaffner 310), P. Schaffneri var. crispum (Mexique, ms atepetl, Schaffner ), P. serricula (1 sheet: Guadeloupe, L’Herminier and een Perrotet), P. senile seit eset Ocana, Schlim 364), P. tenuiculum (Guadeloupe, L’Herm Polystichum chlaenosticta (Nouv. Hollande, itereb, P. falcatum (Ste. Domingue, Port-au- Prince, L’Epagnier), P. guadalupense (Guadeloupe, L’Herminier), P. heterolepis (Cuba, Santiago de Cuba, Linden 1742(2)), P. ilicifolium pie Santiago de Cuba, Linden 2/93), P. incisum (Philippines, Cuming 146), P. lepidomanes (Java, Lobb 262), P. rachichlaena (Mexique, Popocatepetl, Schaffner r Pteris aspera (Bourbon, Montbrison), P. croesoides (Bourbon, Montbrison), P. gracilis sees Claussen), P. heteromorpha (Malacca, Cuming , P. melanocaulon (Philippines, Cuming), muricella (Mexique, Cordoba, Schaffner 143), P. mysorensis (Indes, Mysore, Hook fil. & Sega P. oppositi-pinnata (Philippines, Cuming), P. paucinervata (Mexique, Mirador, Schaffner 152), P. Philippinensis (Philippines, Cuming 8), P. prionitis (Philippines, Cuming 46), P. lett (Bourbon, Bory) rostrata (Perou, Spruce), P. semidentata (Nouv. Grenade, Ocafia, Schlim ) Sagenia gemmifera (Madagascar, Pervillié), S. mexicana (Mexique, Galeotti as and Mexique, Oaxaca, Galeotti 6542). Schizoloma javae (Java, Zollinger 1504). Synochlamys ambigua (Nouv. Grenade, Rio Hacha, Schlim 877). Vittaria amboinensis (Amboine, a V. filifolia (Mexique, Linden 168), V. Gardneriana (Brésil, Orgaos, Gardner 147), V. latipes, (Mada agascar, Boivin in 1853), V. loricea (Java, Zollinger 1001), V. minor (Philippines, Mes 38 81), V. Owariensis (Oware et Beni, pane i. in 1820 V. remota (Nouv. Grenade, Ocana, Schlim 611), V. Ruiziana (Perou, Huassa-Haa iz & Pavon), V. sarmentosa (Cap de Bonna Spéi, Drége), V. tenera (Guenzius), V. zeylanica ans comebel 210). Woodsia mexicana (Mexique, S. Angel, Schaffner 306). This survey was made possible by the cooperation of the officers of the Botanical Garden of Rio de Janeiro, and by support from the Brazilian Research Council (Conselho Nacional de Desenvolvimento Cientifico Tecnologico, Proc. 30.1339/77) and the Universidade Estadual Paulista Julio de Mesquita Filho (UNESP). LITERATURE CITED STEARN, W. T. 1962. Fée’s “Memoires sur la famille des fougéres.”” Webbia 17:207-221. WINDISCH, P. G. 1981. Specimens from Fée’s herbarium in Rio de Janeiro (RB). Taxon 30:730. AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 2 (1982) 61 SHORTER NOTES A GERMINATION METHOD FOR ISOETES. — Engelmann (Trans. Acad. Sci. St. Louis 4:359-390. 1886), Campbell (Ann. Bot. 5:231-258, tr. XV-XVII. 1891), La Motte (Amer. J. Bot. 20:217-—233. 1933; Ann. Bot. n.s. 1:695-715. 1937), and Boom (Amer. Fern J. 70:1—4. 1980) all described a similar procedure for germinating /soétes spores in order to obtain endosporic megagametophytes. Their methods, however, allow the introduction of soil on the spore wall, archegonial pad, and among the rhizoids. This makes necessary the tedious separation of all mud and grit from the plant material before embedding and sectioning if torn ribbons and nicks in the microtome knife are to be avoided, according to Johansen (Plant Microtechnique, 1940). Recently I was successful in germinating megaspores of /. engelmannii A. Br. on an inorganic nutrient medium, which totally avoids the problem of soil on the specimens. Material of 7. engelmannii was collected on 5 Sept 1981, at a time when the spores were nearly mature. All plants were from the lake at Hanging Rock State Park, one of the most easterly mountain areas in North Carolina, located 32 miles north of Winston-Salem on roads NC-89 and NC-66. Mature, fertile plants along with adhering mud were removed from the bottom of the lake with as little disturbance as possible and were brought to the Duke University greenhouse. Inact plants were transplanted for future studies. Sporophylls broken off in moving the plants were used in culture studies. The nutrient medium was that of Dr. B. C. Parker, Washington University, St. Louis, Missouri, which was used by Klekowski (J. Linn. Soc. London 62:361-377. 1969) for culturing gametophytes of the Blechnaceae. This medium was slightly modified by adding 0.30 g per liter of Mycostatin (=Nystatin, available from SIGMA) and 15 g (rather than 10 g) of agar per liter of nutrient solution. All but the very base of the sporophylls was removed and discarded. The sporangia were then sterilized in 1.31% aqueous sodium hypochlorite solution and rinsed thoroughly with sterile, charcoal-distilled water in a germ-free transfer case in a P| physical containment laboratory to insure that the culture plates were not contaminated with fungal spores. Both micro- and megaspores were inoculated on each plate using sterilized needles and forceps. The plates were sealed with parafilm to help retain moisture. Initially, the plates were kept in the laboratory where they were exposed to normal ambient light fluctuations. After about a month in which no prothallia were observed, the cultures were placed in darkness at a temperature of 0°C for 2 days. Following removal from these conditions, megagametophytes at various stages of development were observed in 7-10 days. The mature female gametophyte of /. engelmannii exhibits an abundance of rhizoids, as well as a conspicuous protrusion of gametophytic tissue. This technique should be suitable for other species of Jsoétes with little or no modification. Appreciation is extended to Bryan J. Taylor, Chief Park Naturalist of the North Carolina Division of Parks and Recreation Department, for granting a plant collecting permit.—Sterling J. Sam, Department of Botany, Duke University, Durham, NC 27706. 62 AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 2 (1982) A NEW STATION FOR LYGODIUM PALMATUM—The senior author dis- covered L. palmatum (Bernh.) Swartz along the Blue Ridge Parkway in Swain County, North Carolina in 1980. Together we visited the site in August 1981 and collected a voucher specimen (Stupka & Sharp 811, TENN). The stand, which seems to be in very good health, is of interest because of its proximity to the Great Smoky Mountains National Park, where L. pa/matum is rare. Only four stations for the American Climbing Fern have been found in the Park, and at these the species is no longer extant or the colony is diminishing. The population at the Swain County site will be monitored to see if it, too, declines or disappears. Radford, Ahles, and Bell, in their “Manual of the Vascular Flora of the Carolinas,” indicate that L. palmatum is known from only ten counties in North Carolina. The discovery in Swain County adds an eleventh county to the list. Lygodium palmatum does occur in counties near Swain, but none of them are contiguous to the Park.—Arthur Stupka, R. #3, Gatlinburg, TN 37738 and A. J. Sharp, Department of Botany, University of Tennessee, Knoxville, TN 37916. ; Contribution from the Botanical Laboratory, University of Tennessee, n.s. No. 9. THE NAME OF A HYBRID x ASPLENOSORUS.—The backcross hybrid between Asplenium platyneuron (L.) B.S.P. and x Asplenosorus ebenoides (Scott) Wherry has never received a formal epithet. I wish to name it in honor of Kathryn Boydston of Fernwood, Niles, Michigan, in recognition of her many contributions to the culture of native ferns, especially of asplenioids. x Asplenosorus boydstonae K. S. Walter, hybr. nov. Herba inter Asplenium platyneuron et X Asplenosorum ebenoidem intermedia et ex hybridatione harum specierum genita. Frons ad 21 cm longa, ad 3.5 cm lata, lanceolata, ad basim truncata, ad apicem caudata; pinnis irregulariter dispositis, inaequaliter deltoideo-lanceolatis quasi subfalcatis, obscure retroflexis. A. platy- neurone apice minore differt, a x A. ebenoide circumscriptione pinnarum differt. Fertile fronds up to 21 cm long and 3.5 cm wide (in cultivation), lanceolate, truncate at the base. Rachis and midrib very dark brown, more or less glossy. Pinnae up to 27 on each side of the rachis, irregular in size and in placement on the rachis, inequilaterally deltate, lanceolate or subfalcate, obscurely retroflexed, occasionally auricled, the margins crenulate. Sori linear, up to 2 mm long, irregularly placed. Chromosomes 36 pairs and 36 univalents. TYPE: Havana Glen, 1.2 km NNW of the Havana Post Office, Hale County, Alabama, Wagner & Walter 70011 (MICH). x Asplenosorus boydstonae is distinguishable from Asplenium platyneuron by its narrow apex and from x Asplenosorus ebenoides by its greater dissection of the blade. See Walter, Wagner, and Wagner (Amer. Fern J. 72(3). 1982) for illustrations and a full discussion of this hybrid.—Kerry S. Walter, Division of Biological Sciences, University of Michigan, Ann Arbor, MI 48109. AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 2 (1982) 63 THE DELETION OF NEPHROLEPIS PECTINATA FROM THE FLORA OF FLORIDA. — On November 22, 1959, in company with Mr. John Beckner of St. Petersburg and others, I visited the Big Cypress region east of Naples on a botanical expedition into Collier County, Florida. Leaving Bonita Springs before heading for the royal palm section west of Deep Lake, where some of the rarest Florida pteriodphytes were to be seen, we traversed a new road recently cut through the wilderness, in the hope of finding something unusual in a little explored area southeast of Corkscrew Swamp. Along State Route 858 about 20 miles northeast of Naples we made what appeared to be an interesting discovery, a Nephrolepis high in a cypress tree. Since N. exaltata (L.) Schott (Wild Boston Fern) is the most common species in Florida, being both epiphytic and terrestrial, this would have been the most logical identifica- tion; but this fern somehow looked different, and we speculated that it might just possibly be N. pectinata (Willd.) Schott, a species widely distributed in tropical America. I collected and pressed one specimen, while Mr. Beckner obtained a living plant for his fern garden. Unfortunately, his plant died shortly thereafter in an early frost. Since all of his other Nephrolepis plants survived that freeze, this seemed further evidence that the plant we collected might actually be N. pectinata. However, when I showed my material to Conrad Morton, then Curator of Ferns at the Smithsonian Institution, he said that it was beyond question N. exaltata, calling my attention to the round or auricled pinna bases, rather than the acuminate triangles that characterize N. pectinata. I wrote Dr. Edgar Wherry following publication of his “Southern Fern Guide” in 1964, in which N. pectinata is listed among the ferns of Florida (from Collier County), and informed him of Morton's identification. He replied that he had not actually seen any living or pressed material, but had included N. pecinata in his book on the basis of Mr. Beckner’s description and report. In Clifton E. Nauman’s article “The Genus Nephrolepis in Florida” (Amer. Fern. J. 71:35—40. 1981), he noted that he had “seen no specimens of this species in the herbaria or field. It is doubtful that this species exists in Florida.” After reading his article, | wrote Mr. Nauman, enclosing photocopies of my herbarium sheet. He agreed that the fern was indeed N. exaltata. In view of this, N. pectinata should be deleted from the floristic record for Florida and the United States —Thomas Darling, Jr., 5008 Larno Drive, Alexandria, VA 22310. A FILMY DANAEA.— One of the rarest species of Costa Rican Danaeas is D. crispa Endres in Reichenb. f., which was first collected by Endres in the 1870's, probably in the northern Atlantic lowlands of the country. Danaea crispa is related to D. wendlandii and to D. jenmanii (the latter two may not be distinct from each other), but differs from them in having thin, crispate pinnae. A recent collection provided the following information: the rhizome is creeping, tortuous like that of D. carillensis Christ, and bears a fascicle of fronds more or less 64 AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 2 (1982) spirally arranged around the tip. The fronds are pinnate and have up to 17 pairs of subalternate pinnae. The basal pinnae are much reduced, and the apical pair subtends a usually proliferous bud. The pinnae are translucent, with distant, free, forked veins. In cross section, the lamina is made up of three layers of cells, all photosynthetic. The epidermal cells are thin-walled, elongate, measure 33-61 wm in the upper surface, 53-76 ym in the lower surface, and 25-38 jm deep in each layer. The mesophyll is one-layered; its cells are equal to or smaller than the epidermal cells. From upper to lower cuticle, the laminae are 215-219 wm thick. No palisade tissue is differentiated. Chloroplasts are more or less evenly distributed in the cells, and vary from 4.1 to 10.6 wm in diameter. Around the veins, the lamina is multilayered: one row of epidermal cells, 1 or 2 layers of parenchyma, one layer of sclerenchyma, and amphiphloic vascular tissues are present. At the margins and apices of segments, the two epidermal layers join, with or without small parenchyma cells between them. There are no stomates in my specimen. With such thin laminae, D. crispa is the only known filmy species in the genus, a quality which restricts its distribution to areas of extremely high relative humidity. The fronds dehyrate in less than five hours at 50% r.h. or in three hours at 45% rh. Rehydration can take place in 2 or 3 hours by total submersion of the leaves, which regain complete turgidity and erectness.—Luis Diego Gomez P., Museo Nacional de Costa Rica, Apartado 749, San José, Costa Rica. BRITISH PTERIDOLOGICAL SOCIETY Open to all who are interested in growing and studying ferns and fern-allies. Full members receive THE FERN GAZETTE and the BULLETIN OF THE BRITISH PTERIDOLOGICAL SOCIETY. Membership subscriptions are £ 7 for full members, £ 5 for ordinary members (not receiving the GAZETTE), and £5 for student members (under 25 years of age). For particulars, U. S. residents should apply to Dr. J. Skog, Biology Department, George Mason University, Fairfax, VA 22030. Non-U. S. residents should apply to Lt. Col. P. G. Coke, Robin Hill, Stinchcombe, Dursley, Gloucestershire, England. ... from TRIARCH We are grateful for the purchase orders and collec- tions of material received during Triarch’s 56 years of service. We try to say thank you by maintaining quality of product, variety of listings, and by supporting biolog- ical societies with memberships, donations, and advertising. Having completed 25 years in the management of Triarch, | add this written word of thanks for your support, without which Triarch would not exist! Paul L. Conant, President TRIARCH PREPARED MICROSCOPE SLIDES P 8 Ripon, Wisconsin 54971 a Fems and lhed Plants Sept. 30, 1982, With Special Reference to Tropical America and you save 20%. ROLLA M. TRYON and ALICE F. TRYON, Harvard University MORE THAN 850 PAGES This exceptional volume is the first modern, in-depth treatment of ferns and related plants, particularly in tropical America. Written by Rolla M. and Alice F. Tryon, well known for their work in this area, FERNS AND ALLIED PLANTS presents a new classification of the Pteriodophyta based on a wealth of data published during the last few decades as well as the Tryons’ own wide-ranging research. Extensive bibliographies of modern literature on fern biology are provided. In addition to tropical American genera, this volume also has extensive information on ferns of temperate and paleotropical regions, making it of worldwide perspective. Each of the 127 tropical and temperate American genera, represented by 29 families, is organized by accounts of its: © ecology O nomenclature and typification O geography O spores O generic characteristics O cytology MORE THAN 2,000 ILLUSTRATIONS AND PHOTOGRAPHS Carefully compiled, magnificently produced, and superbly illustrated, FERNS AND ALLIED PLANTS has more than 2,000 individual illustrations and photographs prepared especially for this volume, including: © photographs of living plants in their natural habitats O photographs and illustrations of biological and systematic features O distribution maps © more than 700 electron micrographs of fern spores Geologists who use palynological records in hydrocarbon exploration will find the spore micrographs of immense practical value. Unrivaled in its coverage, FERNS AND ALLIED PLANTS will serve as the authoritative reference for years to come. 1982/approx. 896 pp./2022 individual illustrations and photographs in 639 figures/ISBN 0-387-90672-X/cloth $148.00 B SPECIAL 20% DISCOUNT PRICE: $118.00 (Orders must be received by September 30, 1982.) For more information, or to order, please write to: SPRINGER-VERLAG NEW YORK INC., Dept. $5490, 175 Fifth Avenue, New York, New York 10010 ry SPRINGER-VERLAG New York Heidelberg Berlin AMERICAN FERN humo JOURNAL cre QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY ee Biosystematic, and Nomenclatural Notes on Scott’s Spleenwort, x Asplenosorus ebenoides KERRY S. WALTER, WARREN H. WAGNER, JR. and FLORENCE S. WAGNER Cheilanthes feei New to Virginia THOMAS F. WIEBOLDT and STAN BENTLEY A New Woodsia Hybrid from Kansas RALPH E. BROOKS Polystichum lonchitis in Central Quebec—-Labrador MARCIA J. WATERWAY and THOMAS T. LEI A Unique Type of Microsporangium in Selaginella Series Articulatae PAUL SOMERS Cystopteris tennesseensis in Illinois ROBBIN C. MORAN Dieter E. Meyer (July 21, 1926—February 1982) Shorter Note: Flavonoid Chemistry of the North American ycopodium obscurum Complex Review The American Fern Society Council for 1982 DEAN P. WHITTIER, Dept. of Biology, Vanderbilt University, Nashville, TN 37235. President TERRY R. WEBSTER, Biological Sciences Group, University of Connecticut, Storrs, CT 06268. Vice-President MICHAEL I. COUSENS, Faculty of Biology, University of West Florida, Pensacola, FL 32504. Secretary JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, TN 37916. Treasurer JUDITH E. SKOG, Dept. of Biology, George Mason University, Fairfax, VA 22030. Records Treasurer DAVID B. LELLINGER, Smithsonian Institution, Washington, DC 20560. Journal Editor ALAN R. SMITH, Dept. of Botany, University of California, whens CA 94720. Memoir Editor JOHN T. MICKEL, New York Botanical Garden, Bronx, NY 1 Newsletter Editor American Fern Secret EDITOR DAVID B. LELLINGER U.S. Nat'l Herbarium NHB-166, Smithsonian Institution, Washington, DC 20560 ASSOCIATE EDITORS DAVID W. BIERHORST Rt. 3, Box 188, Picayune, MS 39466. GERALD J. GASTONY Dept. of sai Indiana hs Bloomington, IN 47401. JOHN T. MICKEL ew York Botanical Garden, Bronx, NY 10458. The “American Fern Journal” (ISSN 0002-8444) is an illustrated quarterly devoted to the general study of ferns. It is owned by the American Fern Society, and published at the Smithsonian Institution, Washington, DC 20560. Second-class postage paid at Washington. Claims for missing issues, made 6 months (domestic) to 12 months (foreign) after the date of issue, and the matters for publication should be addressed to the Editor. Changes of address, dues. and applications for membership should be sent to Dr. Judith E. Skog, Dept. of Biology, George Mason University, Fairfax, VA 22030. rik for back issues should be addressed to Dr. James D. Montgomery, Ichthyological Associates, 8603. . Berwick, PA 1 Gen - inquiries concerning ferns should be addressed to the Secretary. Subscriptions $9.00 gross, $8.50 net if paid through an agency (agency fee $0.50): sent free to members of the pei Fern Society (annual dues, $8.00; life membership, $160.00). Back volumes 1910-1978 $5.00 to $6.25 each; single back numbers of 64 pages or less, $1.25; 65-80 pages, $2.00 each; over 80 pages, $2.50 each, plus shipping. Back volumes 1979 et seq. $8.00 each: single back numbers $2.00 each, plus shipping. Ten percent discount on orders of six volumes or more. Library Dr. John T. Mickel, New York Botanical Garden, Bronx, NY 10458, is Librarian. Members may borrow books at any time, the borrower paying all shipping costs. Newsletter n T. Mickel, New York Botanical Garden, Bronx, NY 10458, is editor of the newsletter “Fiddlehead Forum.” The editor welcomes contributions from members and non-members, including miscellaneous notes, offers to exchange or purchase materials, personalia, horticultural notes, and reviews of non-technical books on ferns. Spore Exchange Mr. Neill D. Hall, 1230 ela 88th Street. Seattle, WA 98115, is Director. Spores exchanged and collection lists sent on reque Gifts and Bequests Gifts and bequests to the Society enable it to expand its services to members and to others interested in ferns. Botanical books, back issues of the Journal, and cash or other gifts are always welcomed, and are tax-deductible. Inquiries should be addressed to the Secretary. AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 3 (1982) 65 Ecological, Biosystematic, and Nomenclatural Notes on Scott’s Spleenwort, < Asplenosorus ebenoides KERRY S. WALTER, WARREN H. WAGNER, JR., and FLORENCE S. WAGNER* The best known and historically most interesting fern of the Appalachian region is the Scott’s Spleenwort, x Asplenosorus ebenoides (Scott) Wherry (Alston, 1940; Weatherby, 1949), a bigeneric hybrid between Asplenium platyneuron and Camptosorus rhizophyllus. The Ebony Spleenwort, A. platyneuron (L.) Oakes, has tall, pinnately compound leaves with shiny, blackish midribs; its veins are simple and bear parallel sori. The Walking Fern, Camptosorus rhizophyllus (L.) Link, has attenuate, simple leaves with green midribs; its veins are reticulate and bear arching sori. Proliferous leaves of the Walking Fern bear a plantlet at their long and thread-like tip. The hybrid fern SEs the parental characteristics in a spectacular and often highly asymmetric mann The hybrid was first found near " Philadelphia along the Schuylkill River around 1862 by a horticultural writer, Robert Robinson Scott, and has been known as Scott’s Spleenwort ever since (Weatherby, 1949). Scott published a short description in Thomas Meehan’s “Gardener’s Monthly” in 1865 and called it Asplenium ebenoides (i.e., “like ebeneum,” the then-current synonym of platyneuron). Howev- er, Meehan in an editorial note suggested that it might be a hybrid, and the following year the British naturalist Rev. M. J. Berkeley proposed its parents correctly (Weatherby, 1949). A number of problems center around Scott’s Spleenwort. There has been a question about the correctness of the epithet ebenoides. A fern considerd by some to be identical to it had been named Asplenium hendersonii by Houlston a decade and a half earlier than A. ebenoides. It was assumed that Scott’s Spleenwort was a sterile hybrid because in the original locality it occurred singly with the parents. However, around 1874, Julia L. Tutwiler, a teacher, discovered a large population of Scott’s Spleenwort in Rock Hollow, near Havana in Hale Co., Alabama, a ravine later known to botanists as “Havana Glen.” This apparently fertile population grew on a conglomerate rock of “pudding stone.” Although Slosson (1902) proved hybridity experimentally using the gametophytes of the parents, fertility was demonstrated only much later by showing that the Havana Glen plants have doubled chromosomes and so are fertile and sexual (Wagner, 1954; Wagner & Whitmire, 1957). Further studies have involved experimentally recreating fertile forms of XA. ebenoides from sterile ones, backcrossing it, and demonstrating by chromatography that x A. ebenoides contains the combined flavonoid compounds of both parents and that the 2x and 4x forms are alike in this respect (Wagner, 1954, 1956; Wagner & Whitmire, 1957; Wagner & Boydston, 1958; Smith & Levin, 1963). *Division of Biological Sciences, University of Michigan, Ann Arbor, MI 48109. MISSOURI BOTANICA SEP 19 1988 GARDEN LiBRaRy 66 AMERICAN FERN JOURNAL: VOLUME 72 (1982) 10 cm C A ; £ FIG. 1. Silhouettes. A. Type specimen (BM) of Asplenium hendersonii with stipe increased to approximate length of original description. B-E. x Asplenosorus ebenoides. B. Sterile form, Mercer Co., KY, Wagner 9127 & D. M. Smith (MICH). C. Fertile form. Hale Co., AL, Maxon & Pollard 335 (US). D, E. Sterile form, Giles Co., VA, Wagner 10324.5 (MICH). K. S. WALTER ET AL.: NOTES ON SCOTT'S SPLEENWORT 67 NOMENCLATURE For over 75 years there has been some question about the proper epithet for x A. ebenoides. Carl Christensen (1905) first suggested that it might be Asplenium hendersonii Houlston (1851), rather than A. ebenoides R. R. Scott (1866). The late _ V. Morton wrote (in litt., 7 February 1968), “I have really no doubt that hendersonii is an earlier name for ebenoides. It is too bad to have this well-known name abandoned, but I see no alternative.” To support this proposition, Morton sent a copy of his photograph of the type specimen of A. hendersonii from the British Museum. We compared the type photo (silhouette is shown in Fig. /A) with fronds of equivalent size of variations of unquestioned x A. ebenoides (Fig. 1B—E). We found two important differences: in A. hendersonii, the basal pinnae are much broader and rounded apically while in x A. ebenoides, large fronds always have narrowly caudate tips, inherited from Camptosorus rhizophyllus. Careful examination frequently reveals a tiny plantlet or proliferation at the extreme apex (Fig. /D). Asplenium hendersonii has tips ca. 3—5 times broader than those of comparable x A. ebenoides specimens. The type specimen of A. hendersonii is annotated as folows: “. . . this plant was raised by Mr. Henderson (“Gardener to Earl Fitzwilliam at Wentworth’) from spores, but from what country received he was unable to say.” According to the original description (Houlston, 1851), the stipes are “about three inches long, and thinly covered with long brown narrow scales,” the latter character not found in mature specimens of XA. ebenoides. Examination of the type of A. hendersonii revealed the existence of several significant differences. The sori of XA. ebenoides are usually 1-3 mm long and only rarely are longer (Fig. 2A); those of A. hendersonii are mostly 2-5 mm long (Fig. 2B). Many sori and indusia in the upper half of the blades of A. hendersonii are double, and arise from two adjacent veins which meet one another, precisely as is found in the Hart’s-Tongue, Phyllitis. The other distinctions of A. hendersonii can be readily explained if Phyllitis, rather than Camptosorus, is postulated as one of the parents. The “stubby” pinnae tips, the broad leaf apex, and the long sori are, of course, all features of Phyllitis and not Camptosorus. We conclude, therefore, that A. hendersonii is really an X Asplenophyllitis Alston, i.e., a spontaneous hybrid between Phyllitis scolopendrium and some species of Asplenium, perhaps the Sea Spleenwort, A. marinum, which is wide- spread along the coasts of England and Europe. Natural hybrids of Phyllitis with Asplenium are well known (Alston, 1940, listed three) as are artificial ones (Lovis, 1973). It is even possible that spores of Phyllitis, a very popular species in cultivation, germinated together with spores of an Asplenium in cultures sown by Henderson, giving rise to A. hendersonii. It is clear, therefore, that hendersonii is not the correct epithet for the Appalachian hybrid. If Scott’s Spleenwort is kept in the genus Asplenium, the name should appear as Asplenium X ebenoides, using the multiplication sign, as prescribed by the Interna- tional Code of Botanical Nomenclature, to indicate its hybrid nature. As to the rare fertile form, it has been argued elsewhere (Wagner, 1969) that it is irrelevant whether AMERICAN FERN JOURNAL: VOLUME 72 (1982) K. S. WALTER ET AL.: NOTES ON SCOTT'S SPLEENWORT 69 the plant is sterile or fertile. Since it is a hybrid, diploid or tetraploid, the hybrid sign should be used, and the cytological condition be indicated by a descriptive phrase “sterile diploid form” or “fertile tetraploid form.” In the case at hand, the fertile tetraploid form has been produced in culture directly from the sterile form Bees = Whitmire, 1957). Ho r, we believe the name of the plant should be x Asplenosorus ebenoides (Scott) Whend: with the multiplication sign before the generic name, as this plant is, according to most present thinking, an intergeneric hybrid (Mickel, 1974). Camptosorus should be maintained as a genus because its differences from Asplenium are of the same magnitude as those of other genera in the Asplenioideae such as Diellia and Phyllitis. There is no reason to abandon the genus Camptosorus simply because it hybridizes with Asplenium. Lovis (1973) discussed intergeneric fern hybrids and pointed out that not only are there Asplenium * Camptosorus (= x Asplenosorus Wherry) hybrids known, but Asplenium x Phyllitis (= Aspleno- phyllitis Alston), Asplenium x Ceterach (= XAsplenoceterach D. E. Meyer), Camptosorus X Phyllitis, and Asplenium x Pleurosorus as well. STATUS OF THE HAVANA GLEN POPULATION This report is based upon field studies by W. H. Wagner and K. S. Walter in 1971 and by Dr. R. R. Haynes of the University of Alabama in 1978; the most recent previous report is that by Wherry and Trudell (1930). The locality is approximately 40 km south of Tuscaloosa outside of Havana. Winters are relatively mild, with the leaves off the deciduous trees and shrubs, and most annual herbaceous shoots withered. What we call “Havana Glen” is known locally as “Rock Hollow.” It is owned by Mrs. R. B. Lavender, of Mobile, Alabama, who has strict rules governing who can enter the area and whether logging is permitted in or near it. We discussed the past history of the unusual fern populations with her relative, Mrs. T. N. Lavender, who remembered the botanists who had visited, especially H. E. Ransier of Manlius, ew York, who traveled to Havana at least twice Miss Tutwiler originally described the habitat as “about eight miles from the Black Warrior River . . . The soil is either red clay or a mixture of sand and gravel except in the creek and river bottoms. The country is rolling, covered with hills j : at ie x ~ ey ot ith: SOW, yong 1912—G. L. a wa 1918, 1920—E. W. Graves; ant 1933—H. E. Ransier; 1929—E. T. and H. W. a ; 1967, 1974—B. E. Dean; 1971—W. H. Wagner, Jr. & K. S. Walter, 1978 R- R. Hayne . 2. Comparison of x Asplenosorus ebenoides and Asplenium hendersonii. A-B iegnieeteagsey ehensides. A. Abaxial side of frond near tip showing short, unpaired sori, Wagner 76016 (MICH). Same, Wagner s. n., 5 Dec 1955 (MICH). C. Type specimen of Asplenium hendersonii. Abaxial side a frond from a coumerable area showing longer, paired sori, and broader lamina. FIG. 3. Havana Glen, Hale Co., AL, habitat for x Asplenosorus ebenoides, showing moss- and lichen-covered boulders of conglomerate rock. 70 AMERICAN FERN JOURNAL: VOLUME 72 (1982) about 200 feet above sea level” (Dean, 1969). The Hollow lies along a brook below farm fields 1.2 km NNW of the Havana Post Office. The valley side where the rock crops out slopes approximately 20-35° (Fig. 3). The slopes are occupied mainly by deciduous trees: toward the bottom of the valley increasing numbers of mountain laurel, Kalmia latifolia, appear. Haynes (pers. comm.) considers the forest an excellent representative of the deciduous forest of Alabama, “as natural as you can get in the state.” It shows no evidence of logging. Some of the prominent trees include Quercus alba, Q. rubra, Q. prinus, Fagus grandifolia, Ilicium floridanum, Liriodendron tulipifera, and Magnolia macrophylla. Scattered shrubs and woody vines include Vitis spp., Smilax spp., Euonymus americanus, and Hydrangea quercifolia. The forest is somewhat open and park-like. The rock outcrops in the valley are 0.5-8 m tall, and most are north-facing. Wherry and Trudell (1930) described the rock as a brownish conglomerate of siliceous pebbles in a ferruginous and slightly calcareous cement and the soil reaction at the roots of the x Asplenosorus plants as being minimacid’. The more or less crowded pebbles in the sandy substratum are smoothly rounded, indicating stream wear prior to consolidation. The rocks are coated with green bryophytes and/or fine, white, crustose lichens. Those collected in association with X Asplenosorus ebenoides were identified by Howard A. Crum of the University of Michigan Herbarium. The crustose lichen is a species of Parmelia. The most abundant mosses are Anomodon attenuatus (Hedw.) Hub. (Wagner 71167) and Bryoandersonia illicebrum (Hedw.) Robins. (71/68). Together with the liverwort Metzgeria furcata (L.) Dum. (71169) were mixed small quantities of the moss Brachythecium oxycladon (Brid.) Jaeg. & Sauerb. and two liverworts of the genera Radula and Frullania. Two additional mosses occur frequently, viz. Fissidens cristatus Wils. ex Mitt. (71170) and Anomodon rostratus (Hedw.) Schimp. (7//7/). Vouchers are deposited in the University of Michigan Herbarium (MICH) The majority of ferns and herbs occur on and around the rock outcrops rather than on the leaf-covered forest floor. Scattered plants of Polystichum acrostichoides, Dryopteris marginalis, and Hexastylis sp. may grow in the flatter, non-rocky areas, but all three tend to be more common at and around the rock bases. On the boulders themselves are such herbs as Mitchella repens and Saxifraga virginiana. The latter, together with Hepatica americana, may sometimes be found in bloom as early as January or Febru In addition to the Polystichum and Dryopteris, we found nine additional fern species or hybrids at the locality. The most common are Polypodium polypodioides, Asplenium platyneuron, and A. trichomanes. Along the upper slopes are large plants of Cheilanthes lanosa. Masses of tangled green filaments in sheltered, deeply Shaded crevices of the rock are clonal gametophytes of the filmy fern genus Trichomanes. A large patch of sporophytes of T. boschianum occurs on the vertical wall in a recess of one of the larger cliffs. *Donald Farrar reports (pers. comm.) that this substrate is Pottsville sandst with « tightly chic pO © 6) ndstone of Pennsylvanian age, K. S. WALTER ET AL.: NOTES ON SCOTT'S SPLEENWORT 4. Young spleenworts on moss-covered rock at Havana Glen, AL. PPR = Asplenosorus baydstonae (= Asplenium platyneuron X Asplenosorus ebenoides). PP = A. pinnae PPRR = . ebenoides. We did not find Camptosorus rhizophyllus in our explorations. Wherry and Trudell (1930) had the same experience, although Wherry wrote (in litt., 1972) that “The St. John brothers told us that they had visited the ‘glen’ and found . . . the two parents,” and the original discoverer, Miss Tutwiler, had written that she found walking fern there (Dean, 1969). It is possible that all of these people were in some other valley nearby. For example, Wherry, (op. cit.) wrote, After I visited the “Glen” I thought it would be interesting to see what was in other ravines nearby. ssuming that I started north, the road descended down the conglomerate-rock escarpment (on the east side of the Glen) and joined an east-west road running along the valley. I walked eastward along this road, repeatedly ascending small cliffs, and entering small ravines, probably for a distance of a mile. Asplenium platyneuron was conspicuous everywhere; but the ebenoides did occur repeatedly on both moist and dry rock surfaces. Although Wherry did not find Camptosorus, his finding other nearby ravines suggests that a detailed exploration of the whole area in the vicinity of Havana Glen might be profitable. Not only might populations of Camptosorus turn up, but other interesting pteridophytes as well. 72 AMERICAN FERN JOURNAL: VOLUME 72 1982) 4 5 cm FIG 5. Silhouettes of medium- and large-sized fronds of plants collected at Havana Glen, AL, and cultured in greenhouse. Left = x Asplenosorus boydstonae. Right = Asplenium platyneuron. THE TETRAPLOID EBENOIDES POPULATION AND ITS BACKCROSSES In view of Wherry’s comments quoted above, it is possible that A. x ebenoides is considerably more widespread in the Havana area than has been believed. The following is based upon our observations made during the 1970’s. The fertile Scott’s spleenwort is much less common in the Glen than either A. platyneuron or A. trichomanes. Juvenile specimens are abundant intergrown with juveniles of other K. S. WALTER ET AL.: NOTES ON SCOTT'S SPLEENWORT 73 spleenworts. The number of XA. ebenoides plants is estimated to be between 200 and 300. Most are small, the number of fronds ranging from 3 to 7 and the frond size 3 cm or less (Fig. 4). Isolated, single plants are exceptional; most of the individuals are in groups of several to 2-3 dozen. Plants with sori had fronds as short as 4 cm long including the petiole when they occurred in relatively dry, exposed places, and up to more than 14 cm long in damper, shady places. We observed fewer than three dozen soriferous plants, these widely scattered and not confined to a single area. No plants, even the largest, showed frond-tip proliferations visible to the naked eye, although such proliferations have been reported in the past. According to Wherry (pers. comm.), the proliferous tip illustrated by him and Trudell (1930) was not from the Glen, but from outcrops of dry conglomerate half a mile or so east. Functional proliferations must be extremely rare, and vegetative reproduction is of little or no importance in maintaining the population. Virtually all of the plants evidently arise from spores. The small plants grow slowly, especially during dry years (even though in culture they may be vigorous and fast growing, almost becoming greenhouse weeds). Similarly, many of the spleenwort populations in peninsular Florida are noted for waxing and waning year to year, depending upon climate. During drought, popula- tions become reduced to tiny plants, many so small as to simulate sporelings. In moister periods, the spleenworts spring up and form large, soriferous plants of mature form. The conditions at Havana Glen may be similar. In the Appalachian spleenworts, backcross hybrids are rare, the exception being the sterile triploid hybrid x Asplenosorus pinnatifidum x Asplenium montanum, (= x Asplenosorus trudellii (Wherry) Mickel) (W. Wagner, 1954). This is so common in some localities that it must have some method of propagation, although the spores are highly abortive, and meiosis involves both univalents and bivalents. Some backcrosses of %Asplenosorus ebenoides have been produced in the laboratory, including the tetraploid, fertile form with its parents. The backcross to Asplenium platyneuron (= XAsplenosorus boydstonae K. S. Walter) was first synthesized experimentally at the University of Michigan (Wagner, 1956); and this as well as the backcross to Camptosorus rhizophyllus at Fernwood, Niles, Michigan (Wagner & Boydston, 1958). Backcross leaves are intermediate in structure. The A. platyneuron backcross has leaves like A. platyneuron but possesses a caudate tip shorter than that of XA. ebenoides. The leaves of the Camptosorus rhizophyllus backcross look like irregular- ly lobed Walking Fern leaves. In describing the A. platyneuron backcross, we noted that it “has very little likelihood of ever being discovered in the wild, except possibly in one small area in Hale Co., Alabama,” by which we meant Havana Glen. A particularly fine mixed colony was found by K. S. Walter in 1971 on the east-facing side of a boulder. Here we encountered the first wild examples of x Asplenosorus boydstonae. This boulder was approximately 1 m high and 2.5 m wide. Mosses covered roughly one-half of its surface; the remainder was white with crustose lichens. In the mosses were 35 plants of x Asplenosorus ebenoides, 12 large, fertile, up to 15 cm long, and the remainder small, like those in Fig. 4. There were 23 74 AMERICAN FERN JOURNAL: VOLUME 72 (1982) Q & = »* an mw &S B 3890 09 a ¢y Sage 6 0 ? whe <> . iy) an ¢ . is ¢ > e? ae. = a’ xe = ON oO Ov An J oe » , | bad ee Ba 7 ee 3 tan Soe - > es = O go Me ded o® "0.02 mm FIG 6. Chromosomes of a natural x Asplenosorus boydstonae hybrid from Havana Glen, AL, showing 36 pairs (PP) and 36 singles (R). plants of Asplenium platyneuron, of which only four were fertile and up to 15 cm long. One small plant of A. trichomanes was present. Growing with them were three plants of small size but obviously greater blade division than in equal-sized plants of typical x Asplenosorus ebenoides (see Fig. Two specimens were taken alive for culture purposes. We grew them to full size (Fig. 5), and found them to be morphologically identical with the experimentally produced backcrosses. They have 108 chromosomes (i.e., the triploid condition, with 36 pairs of chromosomes from x Asplenosorus ebenoides and 36 singles from Asplenium platyneuron, Fig. 6), as had the laboratory specimens (Wagner & Boydston, 1956, pl. V, figs. A The discovery of the sterile cherie x Asplenosorus boydstonae, at Havana Glen may be the first case in pteridology in which a taxon was produced initially in the laboratory under experimental conditions and only discovered later as a wild plant growing under natural conditions. We wish to express thanks to R. R. Haynes, Clive Jermy, Mrs. T. N. Lavender, Mary R. Rainey, Donald Rainey, James Rainey, the late Edgar T. Wherry and to the late Blanche E. Dean for their assistance in this study. LITERATURE CITED ALSTON, A. H. G. 1940. Notes on the supposed es in the genus Asplenium found in Britain. Linn. Soc. 152 Sess. (1939-40):132-1 CHRISTENSEN, C. 1905-06. Index Filicum. th Co openhagen DEAN, rE . 1969. Ferns of Alabama and Fern Allies, rev. ed. Southern University Press, Northport, sore J. 1851. The genera and species ~ cultivated ferns. Sub-order Polypodiaceae: Tribe Asplenieae (cont.). Gard. Mag., Bot. 3:257-265. K. S. WALTER ET AL.: NOTES ON SCOTT'S SPLEENWORT 75 LOVIS, J. D. 1973. A biosystematic approach to the phylogenetic problem and its application to the Aspleniaceae. In A. C. Jermy et al. (eds.), The Phylogeny and Classification of the Ferns cademic Press, NY. MICKEL, J. T. 1974. The status and composition of Asplenosorus. Amer. Fern SLOSSON, M. 1902. The origin of Asplenium ebenoides. Bull. Torrey Bot. Club 29: SMITH, IN. 1963. A chromotographic study of reticulate pois in the Appalathing ahtlenines. Amer. J. Bot. 50:952-958. WAGNER, W. H., Jr. 1954. Reticulate evolution in the Appalachian aspleniums. Evolution 8:103—109 ——_—. 1956. Asplenium ebenoides x platyneuron, a new triploid hybrid produced under stitial conditions. Amer. Fern. J. 46:75-82. . 1969. The role and taxonomic treatment of hybrids. BioScience 19:785—789. and K. E. Boydston. 1958. A new hybrid spleenwort from artificial a at Fernwood and its relationship to a peculiar plant from West Virginia. Amer. Fern J. 48:146—159. d R. S. Whitmire. 1957. Spontaneous production of a cerca distinct, fertile tlopolypoid by a sterile diploid of Asplenium ebenoides. Bull. aes tte Club 84:79-89. WEATHERBY, C. 1949. One chance in a thousand. Horticulture 27(3):85, WHERRY, -E:.G: P e W. TRUDELL. 1930. The Asplenium ebenoides eis near Havana, Alabama. Amer. Fern J. 20:30—32. 76 AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 3 (1982) Cheilanthes feei New to Virginia THOMAS F. WIEBOLDT* and STAN BENTLEY ** The Slender Lip Fern, Cheilanthes feei Moore, is a widely distributed species of the western United States which occurs eastward to Arkansas, Missouri, Kentucky, and Illinois (Fernald, 1950; Steyermark, 1963; Cranfill, 1980; Mohlenbrock, 1967). The easternmost station for this fern was along Cedar Creek in Bullitt County, Kentucky, in a region known as The Knobs. This small population, discovered originally by Clyde Reed (Reed, 1952), is thought to be disjunct from the next nearest stations along the Ohio River in southern Illinois (Cranfill, 1980). Cheilanthes feei was discovered in 1979 by Bentley during a routine survey of the plants of Claytor Lake State Park, Pulaski County, Virginia. The actual identity of the plant went unknown until the station was visited by the authors on May 8, 1981, when the first voucher collections were made. This discovery is a remarkable disjunction of approximately 450 km eastward from the Bullitt County, Kentucky station and nearly 650 km from the more contiguous portion of its range from Illinois westward. The preferred habitats of this fern are dry, exposed crevices in limestone or dolomite cliffs (Wherry, 1961; Mohlenbrock, 1967). Extensive exposures of this kind may be found along the New River in Virginia. Claytor Lake is a large impoundment of some 4500 acres formed in 1939 when the Appalachian Power Company built its Claytor Dam on the New River just upstream of Radford, Montgomery County, Virginia. In the southeastern sector of Pulaski County, the New River is deeply entrenched where it has cut through thick beds of limestone of the Rome formation and Elbrook dolomite, both of Cambrian origin. The more resistant dolomite forms nearly vertical cliffs, frequently several hundred feet high. The station for C. feei is a rather small but precipitous cliff, perhaps fifty feet high, which forms a nose-like projection along a westward facing portion of the shoreline. Because of the configuration of the rock, the exposures range from northwestern to southern. During a second visit to the outcrop on May 14, 1981, an estimate of 1200 plants was made. Most of these are extremely small and occupy nearly every minute fracture in the rock. Several hundred larger, mature plants crowd the deeper cracks and more protected, overhanging sites. Several smaller, shaded outcrops in oak woods adjacent to the main cliff support a considerable number of additional plants. Adjacent areas of the lake shore, where similar outcrops and exposures occur, were searched for additional populations of C. feei, but only six other plants were found on a very small ledge at the water level several hundred feet to the north. Since the normal pool elevation is approximately eighty-feet above the former river level, this same outcrop was undoubtedly much more extensive prior to flooding. A much larger population may have existed and, of course, other nearby populations may have been inundated by the lake waters. ee Dept. of Biology, Virginia Polytechnic Institute and State University, Blacksburg, VA **1201 MacGill Street, Pulaski, VA 24301. WIEBOLDT & BENTLEY: CHEILANTHES FEE! NEW TO VIRGINIA 77 The flora of this outcrop is typical of many other outcrops in this portion of Virginia. Chestnut Oak (Quercus prinus) and Red Cedar (Juniperus virginiana) occur on a rather broad ledge and in deep crevices of the rock. Other woody species include Rhus aromatica, R. radicans, Rhamnus lanceolata, and Clematis viorna. Where soil accumulation is sufficient, a mat of vegetation has developed which is made up principally of Poa compressa, Carex eburnea, Aquilegia canadensis, Aster oblongifolius, and Solidago sphacelata. The Carex is particularly representative of such sites. Other ferns associated with the Lip Fern site are Pellaea atropurpurea and Asplenium ruta-muraria. A few individuals of Pellaea glabella were also found. Interestingly, the Slender Lip Fern overwhelmingly dominates the available space on the outcrop, despite the presence of other rock ferns characteristic of the habitat and common in this portion of the state. The presence of many individuals of various ages and colonization of such minute cracks is indicative of reproduction by spores. It is interesting that Cheilanthes feei, seemingly prolific at one station, has not colonized other available spaces in its proximity. Other important fern distributions in Virginia are associated with the New River. The Venus Maidenhair, Adiantum capillus-veneris, was known only from a single 1879 collection from the same region of Pulaski County (Stevens, 1973). One of only two former stations for Cheilanthes alabamensis was a dolomite cliff along the New River in Giles County, Virginia, this being the northeasternmost record for the species. The occurrence of C. feei in the Appalachian Valley of Virginia is particularly notable for several reasons. This record adds another species to the list of pterido- phytes distributed primarily in the West or Southwest which occur as long-range disjuncts in the Southern Appalachians. Included in this list are Asplenium septentrionale (Emory, 1970), Cheilanthes castanea (Knobloch & Lellinger, 1969), and Notholaena sinuata (Univ. of Georgia Herbarium, pers. comm.). How these species have come to occur in the eastern states is an interesting question. The newly discovered Lip Fern station could be regarded as a relict from a former, more widespread distribution, or it could be considered an example of a long-distance dispersal. For other kinds of plants, the objections to the latter hypothesis are varied, but for leptosporangiate ferns, these are largely unfounded. Aerial dissemination of the minute spores over great distances does not stretch the imagination. This is pointed out by Shaver (1954) in a discussion of the disjunct occurrence of a coastal plain species, Woodwardia virginica, in Tennessee. Since the possibility of such an event is very small, time is a major constraint on the theory. The relictual theory attibutes a more eastern (and presumably broader) distribution to more favorable climatic conditions sometime in the past. This is a concept familiar to most phytogeographers in the region. The shale barren flora, with its western affinities, has already drawn speculation as to how that endemic flora may have evolved (Keener, 1971). An eastward migration of several of these species (or their ancestral populations) may have occurred in response to a presumed warmer and drier climate, perhaps the Xerothermic Period (Sears, 1942). Even if such a climatic extreme did happen, it is reasonable to assume that the precipitous, xeric cliff habitat preferred by C. feei would still have occurred as isolated habitats and necessitated some rather 78 AMERICAN FERN JOURNAL: VOLUME 72 (1982) long “jumps” to get from one place to another. This kind of migration would have required long periods of time, as well. The fact that C. feei is found along New River is important in this regard. The New River, despite its name, is believed to be one of the oldest rivers in the world, being a remnant of the old Teays River which drained the area since the late Cretaceous Period. Consequently, there has been plenty of time—so much, in fact, that either hypothesis is plausible. The antiquity of the river affords us a dimension unavailable in most studies in plant geography. It is important to consider the many interesting plants which occur along New River, their distribution, and biology. Since there is so much more than a single, bizarre disjunction associated with the river, we prefer to think of this as another example in support of a relictual interpretation of the facts. The occurrence of C. feei in Pulaski County, Virginia substantiates the importance of the New River as an ancient source of specialized habitats and as a corridor for plant migrations through eons. LITERATURE CITED CRANFILL, R. 1980. Ferns and Fern Allies ey Kentucky. Kentucky Nature Preserves Commission Scientific and Technical Series Num EMORY, D. L. 1970. A Major North American sot Extension for the Forked Spleenwort, Asplenium septentrionale. Amer. Fern J. 60:129-134. ssiidiseoes M. cee 1950. Gray’s Manual of Botany, 8th ed., corrected printing. D. Van Nostrand, New KEENER, c “i eh . The natural history of the mid-Appalachian shale barren flora. Jn P. C. Hol .). The Pinerihational History of the soem a the Southern Appalachians, Part II. Haig Virginia Lae sc a Univ. Res. Div ee KNOBLOCH, I. W. and D. B. LELLINGER. 1969. sae castanea and its allies in Virginia and West Virginia. mots 34:59-60. MOHLENBROCK, R. H. 1967. The ets Flora of Illinois, Ferns. Southern Illinois University Press, Carbondale and Edwardsville REED, C. 1952. Notes on a ferns of Kentucky, III. Cheilanthes feei on Silurian limestone in ucky. Amer. Fern J. 42:53—-56. SEARS, P. B. 1942. Xerothermic ie Bot. Rev. 8:708-736. baa J. M. 1954. Ferns of Tennessee. Bureau of Publications, George Peabody College for eachers, Nashville, TN See C. E. 1973. Looking for Virginia’s maidenhair fern. hla Edie STEYERMARK, J. A. 1963. The Flora of Missouri. University Press, Ames, low WHERRY, E. T. 1961. The Fern Guide, Northeastern and Midland United mikes ie Adjacent Canada. Doubleday, Garden City, NY. AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 3 (1982) 79 A New Woodsia Hybrid From Kansas RALPH E. BROOKS* Woodsia R. Br. is a genus of about 25 species distributed worldwide except for Australia and Antarctica (Brown, 1964). Rosendahl (1915) was the first to describe a hybrid Woodsia (alpina X ilvensis) from specimens collected near Stockholm, Sweden. Butters (1941) reported W. cathcartiana (= W. oregana var. cathcartiana) x ilvensis and Tryon (1948) reported both W. glabella x ilvensis and W. cathcartiana X scopulina, all from southern Minnesota or southwestern Ontario. Thus, compared to genera such as Asplenium or the promiscuous Dryopteris, hybridization would appear to be an unusual event among woodsias. In June 1976, while collecting along a sandstone outcrop in an upland prairie in the Smoky Hills of central Kansas, I discovered a mixed colony of W. obtusa (Spreng.) Torr. and W. oregana D.C. Eat. var. oregana. A thorough search of the outcrop revealed an abundance of W. oregana in the drier, exposed sites, a fair number of W. obtusa in more protected, mesic sites, and a few individuals which appeared to be morphological intermediates of the two species. A limited sample of specimens was removed from the colony for further study. Examinations of the hybrid and parental plants revealed morphological intermedi- acy occurring in several characters on the hybrid plants. This is demonstrated by polygonal graphs of mean values for six characters (Fig. 4), including frond length, blade width and texture, stipe base color, stipe vestiture, and the shape of indusium segments. The indusia, which are traditionally used as a primary diagnostic character in Woodsia, were viewed with the aid of a Phillips 501 scanning electron microscope (Figs. 5-10). The indusia of W. obtusa consist of 4-6 wide, plate-like segments while those of W. oregana consist of 5-9 filamentous segments. Hybrid individuals have indusia with 5—10 segments that were each several cells wide and 2-3-furcate or otherwise irregularly lobed apically. Spore characteristics were also examined with the SEM. Brown (1964) stated that Woodsia spores were of no diagnostic value. While this may be true for light microscope studies, it would appear not to be so for SEM work. Surface ornamenta- tion for W. obtusa and W. oregana in Kansas, as well as several other sites in the Great Plains, are consistent with those illustrated herein (Figs. 1]—16). Of particular interest are the walls forming the reticulate surface in each species. Those of W. obtusa are 34m high and form complete areoles. In W. oregana the walls are 1-2y.m high and frequently “dead end,” i.e. they do not always connect to form complete areoles. In addition, there is an obvious difference in the surface texture in each species. Spores from the hybrids were typically abnormal and abortive, although a very few nearly normal spores were observed (Figs. 13-14). The latter spores display some degree of intermediacy, having the disconnected walls of W. oregana and surface texture of W. obtusa. * University of Kansas Herbarium, 2045 Avenue A, Campus West, Lawrence, KS 66044. 80 AMERICAN FERN JOURNAL: VOLUME 72 (1982) FIGS. 1-3. Holotype of Woodsia x kansana, Brooks 12259d (KANU). FIG. |. Habit FIG. 2. Pinna FIG. 3. Sorus. ne R. E. BROOKS: NEW WOODSIA FROM KANSAS 81 W. oregana W. x kansana FIG. 4. Polygonal graphs of mean values of six characters for central Kansas Woodsias. The abbreviations are: A=frond length (cm), B=frond width (cm), C=blade texture (coriaceous to membranaceous), D = stipe base color (dark reddish brown to stramineous), E=stipe vestiture (lacking scales to scaly), and F=indusium segments (filamentous to wide and plate-like). Cytologically, Brown (1964) indicates that W. obtusa is a tetroploid, n= 76, probably derived from W. oregana, n= 38. A firm chromosome count has not been established for the hybrid plants. However, my colleague Dr. Christopher Haufler has observed meiotic irregularities, as one might expect given the differences in chromosome numbers of the parental species. he morphological intermediacy, spore morphology, and meiotic irregularities fulfill the criteria for hybridity as outlined by Wagner and Chen (1965). Therefore recognition of this new hybrid is set forth in the following diagnosis and description. Woodsia X kansana R. E. Brooks, hybr. nov. Figs. 1-3, 7, 8. Laminae inter parentes W. obtusam et W. oreganam var. oreganam. Indusia 5—] segmentis; composita segmentis pluribus cellulis latis, apicibus 2—3-furcatis vel irregulariter lobatis. Sporae abnormales et abortivae. 82 AMERICAN FERN JOURNAL: VOLUME 72 (1982 — FIGS. 5-10. Aisa sage and interpretive drawings . isp indusia. FIGS. ee W. obtusa. FIGS. 7-8. W. x kansana, FIGS. . W. oregana. Plants morphologically intermediate between W. obtusa and W. oregana. Rhi- zomes to 4 cm long, 2—4 mm in diam; scales brownish with a darker, thicker median stripe, lanceolate, 2-5 mm long, entire. Fronds 1-2 dm tall; blade coriaceous to subcoriaceous, lanceolate to narrowly ovate, pinnate-pinnatifid or bipinnate, 6.5—12 cm long, 2.5-4 cm wide, glabrous or sparsely glandular, apex acute; pinnae subopposite to alternate, deltoid to ovate, apex acute to obtuse, short petiolate or subsessile; ultimate segments oblong or ovate, apex obtuse to rounded, margins R. E. BROOKS: NEW WOODSIA FROM KANSAS 83 15 FIGS. 11-16. Ornamentation of eva spores. FIGS. 11-12. W. obtusa. FIGS. 13-14. W. Xkansana. FIGS. 15-16. W. oregana. mostly slightly reflexed and entire to serrate; petiole dark reddish brown at the base and becoming stramineous above or infrequently all stramineous, scales few, pale brownish, lanceolate, and membranaceous. Sori medial, discrete, sometimes appear- ing confluent with age, roundish; indusium basal with 5—10 segments; segments everal cells wide, hs tie 2-3-furcate or otherwise eeu lobed; spores mostly sbricaial and abortive 84 AMERICAN FERN JOURNAL: VOLUME 72 (1982) TYPE: KANSAS. McPherson Co.: 2 mi west and 2 mi south of Marquette, east facing sandstone outcrops along a high prairie ridge, 25 June 1976, Ralph Brooks 12259d (KANU; isotypes MICH, NY). Associated with its parents W. obtusa and W. oregana var. oregana on sandstone outcrops. Known only from central Kansas (McPherson and Ellsworth counties, geese poe, eth Co.. Kansas, 1928, Clement Weber 2 (MO). The habit drawing is the work of Carol Kuhn Teale, a graduate student in botany at the University of Kansas. LITERATURE CITED BROWN, D. F. M. 1964. A monograph of the fern genus Woodsia. Beih. Nova Hedwigia 16:1—-154 + 40 BUTTERS. Py K. 1941. Hybrid woodsias in Minnesota. Amer. Fern J. 31: 15-21. ROSENDAHL, H. V. 1915. Om Woodsia alpina och en Dydlig Inlands form of Fenna Samt Woodsia alpina < ilvensis nov. hybr. Svensk. Bot. Tids. 9:414-420. TRYON. R. M.. Jr. 1948. Some woodsias from the north shore of Lake Superior. Amer. Fern. J. 5 WAGNER. W. H.. Jr. and K. L. CHEN. 1965. Abortion of spores and sporangia as a tool in the detection of Dryopteris hybrids. Amer. Fern. J. 55:9-29 REVIEW The Genus Polypodium in Cultivation [Polypodiaceae], by Barbara Joe Hoshizaki. Baileya 22:1-52, 53-99. June 1982.—This long work continues Barbara Joe’s invaluable series on cultivated ferns. A key to subgenera and keys to the species in each subgenus lead to pertinent synonymy, a brief description, the native range, and horticultural notes for each species. Cultivated varieties are fully discussed under each species and also are illustrated. The taxa are illustrated by a silhouette of a frond or sometimes by a photograph. Occasionally a line drawing or silhouette of some detail is also provided, for instance, of a rhizome scale where that is diagnostic. Addenda and an index conclude the paper. In the addenda, the synonym Pleopeltis revoluta (Spreng. ex Willd.) A. R. Smith should be added to Polypodium astrolepis. Also, Polypodium revolutum C. Chr. appears to be a nomen nudum; the species apparently does not have a valid name. Under subgenus Niphidium, the name Polypodium albopunctatissimum Linden is mentioned as if it were a valid name, but it, too, is a nomen nudum. All who need to know or to identify cultivated polypodiums will make use of this paper constantly.—D.B.L. AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 3 (1982) 85 Polystichum lonchitis in Central Quebec—Labrador MARCIA J. WATERWAY* and THOMAS T. LEI*: ** North American distribution maps for the Northern Holly Fern, Polystichum lonchitis (L.) Roth, show a disjunct range from alpine areas in the western United States and Canada to the northern Great Lakes region, the Gaspé Peninsula, Nova Scotia, Newfoundland, and Greenland (Fernald, 1935; Hultén, 1958, 1968; Wagner, 1979). One isolated station at Richmond Gulf on the eastern shore of Hudson Bay (Rousseau, 1974; Payette & Lepage, 1977) links the northern range of this species in Alaska, British Columbia, and Alberta with its distribution on the coast of Green- land. Rouleau (1956) reported P. lonchitis from Labrador but without locality data or specimen citation. Since a specimen has never been located to verify this report, the occurrence of the Northern Holly Fern in Labrador has been considered doubtful (Rousseau, 1974; Payette & Lepage, 1977) links the northern range of this species in Grenfell-Forbes Northern Labrador Expedition of 1931, noted that Rev. Hettasch had seen P. lonchitis growing on Ogualik Island (Cod Island) on the Labrador coast (57°47'N, 61°47’W) but had not collected it. This brief mention may have been the basis for Rouleau’s including it in his list for Labrador. Our recent collections from the Schefferville region (Lei s. n., 20 July 1980, MTMG; Waterway 2140, 30 July 1981, MTMG, CAN) confirm the presence of P. lonchitis in the central interior of the Quebec-Labrador peninsula. In 1980, a specimen was collected from a population of about ten to fifteen individuals growing on a protected, east-facing slope of Geren Mountain in the saddle between Geren Mountain and Sunny Mountain (55°04'N, 67°14’W). The plants were small ( 10-15 fronds each) and separated from one another by distances of about 05° m. Phegopteris connectilis, Carex scirpoidea, Salix vestita and Anemone parviflora were observed at the same site. An attempt to relocate this population in 1981 in order to gather more information about the habitat and associated species resulted in locating a second site in the same general area. This site is also on an east-facing slope of Geren Mountain, facing Sunny Mountain, at an elevation of approximately 740 m. Treeline occurs at about 585 m on this slope, but a few, very stunted Picea marina trees were found near the P. lonchitis. Both sites are in Quebec but located only 2 km from the provisional border with Labrador. At the site found in 1981, a single clump of P. lonchitis with more than 100 fronds was found growing on a step-shaped dolomitic boulder located in a sheltered draw a short distance below a talus slope which still had a large snowpatch remaining in late July. Both hematite and dolomite occur in the vicinity. Most of the ground surface in the draw is moss-covered, giving the area a distinctly greener appearance than the surrounding dry, alpine tundra. Species diversity is greater in this protected area below the snowpatch than in the surrounding tundra vegetation. *McGill University Herbarium, Dept. of Plant Science, Macdonald College of McGill University, 21,111 Lakeshore Road, Ste. Anne de Bellevue, Quebec H9X 1CO, Canada. **Present address: Dept. of Botany, Erindale College, University of Toronto, Mississauga, Ont. L5L 1C6, Canada. 86 AMERICAN FERN JOURNAL: VOLUME 72 (1982) Co-occurring calciphilic species include Carex sini C. vaginata, Salix vestita, Anemone parviflora, Arabis alpina, Saxifraga aizoon, Dryas integrifolia, Bartsia alpina, Castilleja septentrionalis and Solidago ukinaeay Several addi- tional species usually found in damp, often forested habitats, including Gym- nocarpium dryopteris, Streptopus amplexifolius, Coptis trifolia, Viola adunca, Pyrola minor, Trientalis borealis, Linnaea borealis, and Viburnum edule indicate the moist, protected nature of the site. Other tundra species growing in the immediate vicinity of the P. lonchitis are Lycopodium alpinum, L. clavatum, Carex bigelowii, Poa arctica, Juncus trifidus, Salix herbacea, S. uva-ursi, Oxyria digyna, Cerastium alpinum, Ranunculus allenii, Cardamine bellidifolia, Rubus acaulis, Sibbaldia procumbens, — latifolium, Pyrola grandiflora, Phyllodoce caerulea, an Taraxacum lapponicum. reviously onset localities for P. lonchitis in eastern North America are either in the Great Lakes region, often associated with limestone or dolomite of the Niagara Escarpment (Fernald, 1935; Soper, 1954; Thompson, 1962; Marquis & Voss, 1981) or in coastal or oceanic regions including the Gaspé Peninsula (Rousseau, Wynne-Edwards & Dansereau, 1937; Scoggan, 1950), Cape Breton, Nova Scotia (Roland & Smith, 1969), Newfoundland, the east coast of Hudson Bay (Rousseau, 1974) and possibly the Labrador coast (Abbe, 1936). Habitat data for most of these coastal localities also suggest a calcareous substrate, either of limestone, dolomite, or gypsum. Most eastern habitats are talus slopes or other rocky areas and the majority are shaded. In western North America, P. lonchitis has a continental as well as a coastal distribution, and is usually found in the subalpine forest zone (Wagner, 1979). The stations we have discovered represent the first report of P. lonchitis from a continental situation in northeastern Canada. Their location in the Labrador Hills, just north of the iron-mining district of Schefferville is more than 600 km east from the nearest station on Hudson Bay and more than 600 km north from the localities in the Gaspé Peninsula. The occurrence of two sites in the same area suggests that P. lonchitis can become established there when the spores land on a suitable substrate (dolomite) in a favorable microhabitat. Many other such lush, protected sites with dolomitic substrate occur in the Labrador Hills, and P. lonchitis very likely occurs in some of them. Further exploration in this and other subarctic regions of Canada may well result in the discovery of other such isolated localities, thus eliminating the apparent disjunctions in the North American range of P. lonchitis. We would like to thank the curators of CAN, DAO, and MT for allowing us to examine specimens, Dr. D. E. Swales for identifying some of the associated species, and the McGill Subarctic Research Station for providing accommodation and transportation in Schefferville. LITERATURE CITED ABBE, E. C. 1936. Botanical results of the Grenfell—-Forbes Northern Labrador Expedition, 1931. Rhodora 38:102—161. BOIVIN, B. 1966. Enumeration des plantes du Canada. Naturaliste Canad. 93:265. FERNALD, M. L. 1935. Critical plants of the upper Great Lakes region of Ontario and Michigan. Rhodora 37:197—222. WATERWAY & LEI: POLYSTICHUM LONCHITIS IN EASTERN CANADA 87 HULTEN, E. 1958. The Amphi-Atlantic plants and their Phytogeographical Connections. Almqvist & Wiksell, Stockholm. . 1968. Flora of Alaska and Neighboring Territories. Stanford Univ. Press, Stanford, California. MARQUIS, R. J. and E. G. VOSS. 1981. epnae of some western North American plants disjunct in the Great Lakes region. Michigan Bot. 20:53-82. PAYETTE, S. and E. LEPAGE. 1977. La flore gare du golfe de Richmond, baie d’Hudson, Nouveau-Québec. Provancheria 7:1—68. ROLAND, A. E. and E. C. SMITH. 1969. The Flora of Nova Scotia. Nova Scotia Museum, Halifax, ROULEAU, E. 1956. A checklist of the vascular plants of the province of Newfoundland (including the French Islands of St. Pierre and Miquelon). Contr. Inst. Bot. Univ. Montréal 69:41-98. pt - 1974. Géographie floristique du Québec-Labrador. Les Presses de I’ Université Laval, ROUSSEAU! = oy -C. WYNNE-EDWARDS, and P. M. ee aia 1937. Quelques additions anante a he flore de la région du Bic. Ann. ACFAS SCOGGAN ci = . The flora of Bic and the Gaspé sree ae Natl. Mus. Canada Bull. SOPER, s as a The Hart’s Tongue Fern in Ontario. Amer. Fern J. 44:129-147. Lenin ek P. W. 1962. An unusual fern station on South Manitou Island, Michigan. Amer. Fern J. WAGNER, H. 1979. Systematics of Polystichum in western North America north of Mexico. Pteridologia 1:1—64. 88 AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 3 (1982) A Unique Type of Microsporangium in Selaginella Series Articulatae PAUL SOMERS* One of the most distinct taxonomic groups in the genus Se/aginella is the series Articulatae, comprised of about 40 taxa endemic to Latin America, one species that is in South Africa, the Azores and Latin America, and several that are restricted to Southeast Asia. This heterophyllous group, first described by Spring (1850) and maintained in later classifications by Hieronymus (1901) and Walton and Alston (1938), takes its name from the articulations located just below each bifurcation of the stem. These appear either as dark-colored swellings in living material or as dark constrictions in some dried specimens. Other morphological characteristics that unify the group are: (1) each strobilus has only a single (or rarely two) basal megasporangia; (2) each enlarged fertile megasporophyll is subtended by one or more enlarged sterile leaves; (3) the megaspores are exceptionally large and have high crestoreticulate muri; (4) the microspores are pale (usually buff to tan) and have ornamentation of sharp spines on their exines; and (5) the aerial roots arise dorsally from the stem. Also, about three-fourths of the taxa in the series have bistelar or multistelar stem anatomy, a feature not found elsewhere in the genus. During a taxonomic survey of the Articulatae, a previously undescribed type of microsporangium was found to be another unifying feature. A survey of nearly all of the articulate taxa and many non-articulate ones supported this generalization. An explanation of the survey and a description of the microsporangia follows. MATERIALS AND METHODS Specimens of Selaginella with articulate stems from the following herbaria were examined: ENCB, F, FSU, G, GA, GH, H, K, MO, NY, P, PR, PRC, TENN, UC, UMEX, UPS, US and VT. A comprehensive survey of other Selaginella taxa was not undertaken, but representatives of many non-articulate taxa were examined during the course of the study. The basic structural differences between articulate and non-articulate micro- sporangia were discernable to the naked eye, but in order to see anatomical details, slides were prepared using Hoyer’s solution (Anderson, 1954). For even closer scrutiny, a few microsporangia were examined with a scanning electron microscope. Initially an AMR model 900 SEM was used, but later an ETEC autoscan model U-1 was employed. Samples were mounted on standard aluminum studs (Ladd Research Industries) using air-drying silver conductive paint (GC Electronics) or double-faced cellulose tape. The prepared studs were coated with vaporized carbon and gold in a Denton model 515 vacuum-coater with a randomly rotating head. RESULTS AND DISCUSSION The microsporangia of typical non-articulate Selaginella are simple, bivalved structures. They are hinged at the base and split open along a lateral suture which runs from the apex down each side of the sporangium, forming two equal halves * Tennessee Natural Heritage Program, 701 Broadway, Nashville, TN 37203. P. SOMERS: UNIQUE MICROSPORANGIUM IN SELAGINELLA FIG. . Features of open microsporangia in non-articulate and articulate species of steeper ee A. One aie from microsporangium of non-articulate S. oaxacana (Evans B. & Bowers 3223, TENN). FIG. SEM view of completely opened paleo eee from S. subarborescens (Hermann Fo US). FIG. C. Open meponenan 1 of S. sertata (Allen 1959, GH), with the D. SEM view ¥ ager e folded under. FIG. of open eres of S. fr net (Lindeman 4007, US). FIG SEM view of basal cata of sree abe aen ee of § J (Lindeman 4007, US) s seats tends = annuloid cells on either of sporangial s _ F. SEM view of portion of microsporangium « of S. fragilis ees 4007, US) pea eave bands of cells around sporangial stalk and cells forming suture ine of dehiscence. 90 AMERICAN FERN JOURNAL: VOLUME 72 (1982) upon dehiscence. Each half is nearly hemispherical and consists of uniform cells (Fig. 1A). According to Goebel (1901), microsporangial cell walls in S. flabellata Spring, a non-articulate species, are differentially thickened, being thinner on their outer walls than on their inner tangential and radial walls. He described and illustrated microsporangia from a number of species, none of them articulate. Steinbrinck (1902) considered microsporangia to be much simpler in design than megasporangia; he described the former as “primitive” and “of unartistic origin.” He based his judgement on material labeled S. flabellata sent to him by Goebel. Microsporangia of the articulate species of Selaginella are more complex than those of non-articulate ones. They possess two distinct tissue types. At the base of each sporangium are two broad bands of relatively thick-walled cells. These bands converge on both sides of the short sporangial stalk (Figs. /C, E, F and 2A, B, E) and continue as broad annuloid bands up the curved faces of the sporangium. The adaxial and abaxial sides of the sporangia are not identical. Dehiscence occurs along a distinct suture line that leaves a tongue-shaped segment on the adaxial side and a larger, two-lobed portion, representing the remainder of the sporangium (Fig. /B-D, F), on the abaxial side. The two lobes, which were the radial faces of the sporangium, are composed of flat, thin-walled cells (Figs. 1B, D, F and 2C, D). These lobes remain concave on their inner faces and often carry masses of micro- spores upwards during dehiscence. Goebel (1901) and Steinbrinck (1902) found that both mega- and microsporangia possessed cells with thick walls except on their outer faces. Goebel was uncertain about how the sporangia dehisced, but Steinbrinck strongly advocated the idea that cohesive forces of water played a major role. To support his hypothesis, Steinbrinck placed ripe megasporangia in absolute alcohol for 24 hours, and then dried them out in the air. Dehiscence did not occur. Then he placed the same sporangia in water until the cell lumina were filled with water. When they were allowed to dry the second time, they dehisced normally, demonstrating the importance of water in the dehiscence mechanism. The same principles of cohesion and adhesion probably apply to the dehiscence of the unique microsporangia of the articulate species of Selaginella. When these microsporangia are immersed in water, they remain round and turgid; but when removed and allowed to dry, they gradually open. A combination of cohesive and adhesive forces during desiccation may cause the outer tangential walls of the cells in the thick walled regions to be pulled inward (Fig. /E), thus causing tension along the suture line and the subsequent curling back of both ends of the banded region. My observations of excised microsporangia, when removed from water, show that they may curl completely inside out upon drying. The annuloid banding is somewhat different than in ferns. A cross-sectional view through a microsporangium of S. atirrensis Hieron. (Fig. 2A, B) reveals that the cells are higher in the center of each band and taper in size laterally. Also, in this particular example the radial and inner tangential walls of the cells do not appear noticeably thicker than the outer tangential walls. In these respects they resemble the bulliform or “motor” cells described by Esau (1953, p. 145) as epidermal features of certain grasses that curl or fold their leaf blades. P. SOMERS: UNIQUE MICROSPORANGIUM IN SELAGINELLA 9] FIG. 2. Microsporangial features in articulate species of Selaginella. FIG. A. SEM view of portion of microsporangium with microspores of S. atirre nsis (Pittier 3638, US) showing cross-section of annuloid bands of cells. FIG. B. Close-up SEM view of annuloid band of cells shown in Fig. A. FIG. C. SEM view of microsporangium of S. pub (cult. at Columbia Univ., NY), showing ‘split through region of see! walled cells. FIG. D. SEM view of se Aa from Fig. C. showing split along suture line. FIG. E. SEM view of microsporangium of S. parkeri (Killip 37400, US) showing split along suture line. FIG. F. Strobili from S. atirrensis (Tonduz 14552, US) showing open megasporangium at base of left strobilus and open microsporangium near top of right strobilus. 92 AMERICAN FERN JOURNAL: VOLUME 72 (1982) The opening of the microsporangium is not accompanied by a sudden recoiling of the annulus and catapulting of the spores. Instead, it breaks first at the apex along the suture line and then continues gradually downward along the border between the annuloid bands and the pockets of thin-walled cells. Each microsporangium is oriented so that its longer axis, with its pockets of thin-walled cells, is situated outwardly against the subtending sporophyll. When dehiscence occurs, the force of the uncurling annuloid bands may be exerted against the appressed sporophyll, forcing it downward. As the sporangium continues to curl back, it becomes exserted from the sporophyll (Fig. 2F), exposing the two concave lobes containing micro- spores. At this time the spores can be picked up by the wind, splashed away by rain, or washed down the sides of the strobilus, possibly into an open megasporangium at its base. In summary, the Articulatae possess a type of microsporangium not found elsewhere in the genus and considered to be more advanced evolutionarily because of the presence of annuloid bands of cells that probably facilitate dehiscence. Unlike the explosive dehiscence in sporangia of ferns, the microsporangia of the Articulatae just slowly uncurl, breaking along a defined suture line of border cells and gradually lifting the masses of spores upward. While a hygroscopic mechanism is obviously responsible for dehiscence, the anatomical and physiological details are yet to be described. The discovery of this unique and highly developed type of microsporangium within the Articulatae makes it more apparent than ever that the group is a well defined taxonomic unit within Selaginella. Combined with the other characteristics noted above, it can be argued that the group should be elevated to a subgenus. Existing classifications recognize 2-4 subgenera. Hieronymus (1901) split the genus into the homophyllous and heterophyllous taxa, and Walton and Alston (1938) followed Baker’s (1887) basic classification, where the heterophyllous taxa were subdivided into three groups based mainly on whether the sporophylls were uniform or dimorphic. If one assumes equal weighting of taxonomic characters, the Articulatae should have, at least, as high a rank as any of the traditional subgenera. LITERATURE CITED ANDERSON, L. E. 1954. Hoyer’s solution as a permanent mounting medium for bryophytes. Bryologist 57:242-244 BAKER, J. G. 1887. Handbook of ae Fern bec Peace Bell, London. ESAU, K. 1953. Plant Anatomy. J. GOEBEL, K. 1901. Schnee IX. Space Sporenverbreitung und Blutenbildung bei Selaginella. Flora 88:207—22 HIERONYMUS, G. 1901. Selaginellaceae. In A. Engler & K. Prantl, eds. Natl. Pflanzenfam. 1(4):621--715. W. Engelmann, Leipzig. SPRING, eee F, Heap aie famille des Lycopodiacées, seconde partie. Mém. Acad. Roy. Sci. STEINBRINCK, C. 1902. Ueber den Schleudermechanismus der Selaginella- -Sporangien. Ber. Deutsch. Bot. Ges. 20:117—128. WALTON, J. and A. H. G. ALSTON. rei Lycopodiinae. Jn F. Verdoorn, ed. Manual of Pteridology. Martinus Nijhoff, The Hagu AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 3 (1982) 93 Cystopteris tennesseensis in Illinois ROBBIN C. MORAN* Cystopteris tennesseensis Shaver is a fertile allotetraploid that presumably arose from an ancient cross involving C. bulbifera (L.) Bernh. and C. protrusa (Weath.) Blasdell. Shaver (1950) pointed out that C. tennesseensis is morphologically intermediate between C. bulbifera and C. protrusa and concluded that these were the two parental species. Morphological and cytological work by Blasdell (1963) supports this interpretation. Plants of C. tennesseensis are usually misidentified as either C. bulbifera or C. protrusa. As a result, the range and abundance of this fern are not precisely known. This paper presents the distribution of C. tennesseensis in Illinois and shows that it is much more common and widespread than generally realized. Specimens of all Illinois Cystopteris from the following herbaria were examined: EIU, ILL, ILLS, ISM, MO, and SIU. Those specimens found to be C. tennesseensis were annotated as such and recorded. Locality data of individual collections are available from the author. In addition, numerous areas of suitable rock habitats were visited in the field, especially in southern Illinois and along the lower Mississippi and Illinois rivers. Previously, this fern was reported from only three Illinois counties: Champaign, Jackson, and Will (McGregor, 1950; Mohlenbrock & Ladd, 1978); however, I was unable to locate specimens from Champaign County. Cystopteris tennesseensis occurs most frequently along major watercourses of the state, such as the Illinois, Mississippi, and Wabash rivers (Fig. /). Along these major rivers, habitat for this fern is provided by the rocky escarpments and ravines that run along or into the river floodplains. The writer’s field experience in these areas indicate that C. rennesseensis is commonly present in suitable habitats and is often abundant where found. Cystopteris tennesseensis is also present, although to a more limited extent, in the Driftless Area of northwestern Illinois and in the Shawnee Hills of southern Illinois. In Illinois, C. tennesseensis grows on both limestone and sandstone; however, the great majority of specimens were collected from limestone, indicating a preference for calcareous habitats. Limestone outcrops in Illinois primarily occur along the Mississippi River, the lower Illinois River, and in portions of the Driftless Area in the northwestern corner of Illinois. Major sandstone outcrops where this fern occurs are the Shawnee Hills of southern Illinois and in north-central Illinois at Starved Rock State Park along the Illinois River. Observations in Illinois by the writer support the observations of McGregor (in Wagner & Hagenah, 1956, p. 144) that C. tennesseensis produces bulblets much more abundantly on limestone than on sand- stone. Furthermore, the Illinois populations of this fern on limestone are more extensive and contain a higher density of individuals compared to sandstone populations where similar amounts of rock habitat exist. The preference of this fern for limestone as opposed to sandstone has been noted in other states (Cranfill, 1980; Shaver, 1950, 1954). *Illinois Natural History Survey, Natural Resources Building, 607 East Peabody Drive, Champaign, IL 61820. 94 AMERICAN FERN JOURNAL: VOLUME 72 (1982) FIG. |. The distribution of Cystopteris tennesseensis in Ilinois. All 90 herbarium specimens of C. fennesseensis examined for this study were originally misidentified. This is primarily due to the fact that C. tennesseensis is not keyed to in many commonly used floristic manuals and, in part, because the fronds are highly variable and confusingly intermediate between those of its parents. The qualitative characteristics of frond dissection are difficult to describe and to be appreciated by botanists unfamiliar with this fern. However, there is one quantitative character that can be used to distinguish this fern from both C. bulbifera and C. protrusa: spore size. In the genus Cystopteris polyploidy is correlated with spore size. Thus, C. bulbifera and C. protrusa, both diploids, have spores that are generally 27—32 ym in length, whereas C. tennesseenis, a tetraploid, has spores that are generally 32—42 ym in length (Blasdell, 1963). The writer has found spore size to be a reliable character and recommends its use if one is uncertain about identification based on frond morphology alone. Further discussion concerning the identification of this fern can be found in Cranfill (1980) and Shaver (1950, 1954). Cystopteris tennesseensis is probably more common in many states than is generally realized. A careful search of other herbaria would add greatly to our knowledge of the frequency and occurrence of this fern elsewhere in its range. R. C. MORAN: CYSTOPTERIS TENNESSEENSIS IN ILLINOIS 95 LITERATURE CITED BLASDELL, R. F. 1963. A monographic study of the fern genus Cystopteris. Mem. Torr. Bot. Club 21(4): 1-102. CRANFILL, R. 1980. The Ferns and Fern Allies of Kentucky. Ky. Nat. Pres. Comm. Sci. Tech. Bull. 1:1-284. McGREGOR, R. L. 1950. Two varieties of Cystopteris fragilis. Amer. Fern J. 40:201—207. MOHLENBROCK, R. M. and D. M. LADD. 1978. Distribution of Illinois Vascular Plants. Southern Ill. Univ. Press, Carbondale, IL. SHAVER, J. M. 1950. A new fern, Cystopteris tennesseensis sp. nov., from Tennessee. Tenn. Acad. Sci. 25(2):107-113. —_——.. 1954. Ferns of Tennessee. George Peabody College for Teachers, Nashville, TN. WAGNER, W. H., Jr. and D. J. HAGENAH. 1956. Observations on some bulblet-producing popula- tions of the Cystopteris fragilis complex. Amer. Fern J. 46:137-146. DIETER E. MEYER (July 21, 1926-February 1982) Dieter E. Meyer was born in Gorlitz on 21 July 1926, son of Erich and Elisabeth Meyer. His primary and secondary schooling took place in G6rlitz prior to and during World War II, and his early interest in natural history led to a volunteer position with a local natural history society. From 1946 to 1950 he studied botany, especially of the cryptogams, in Jena. In 1952 he earned a doctorate from the Free University of Berlin with a thesis on hybridization in Asplenium, which became a life-long interest. In 1954 he joined the staff of the Botanical Garden and Museum in Berlin-Dahlem; he was appointed a Keeper in 1964 and a Supervisory Keeper in 1969. His taxonomic interest in ferns was whetted by the excellent living and herbarium collections at Berlin. He himself made collections in Germany, Austria, Switzerland, England, and Canada, often searching for specimens of rare Asplenium hybrids that could be brought into cultivation for further study. He reorganized and restored the fern collections at Berlin, which had lain nearly untouched after Hieronymus’ death and which fortunately had escaped destruction during World War Il. In addition to his broadly based researches on Asplenium, Meyer published a variety of biographical, bibliographical, and floristic papers. Music was among his extra-botanical interests. A full biography and bibliography is to be published in Willdenowia, from which most of the foregoing comments have been taken.—D.B.L. 96 AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 3 (1982) SHORTER NOTE FLAVONOID CHEMISTRY OF THE NORTH AMERICAN LYCOPODIUM OBSCURUM COMPLEX.—The Lycopodium obscurum complex is taxonomically difficult because of environmental effects on the subtle characters used to distin- guish the taxa. A major problem has been whether to recognize as distinct the species L. dendroideum Michaux (Hauke, Bioscience 19:705-707. 1969). Using phyllotaxy, Hickey (Amer. Fern J. 67:45—-48. 1977) re-defined L. obscurum L. var. obscurum and the problematic L. dendroideum and described another taxon, L. obscurum var. isophyllum. The current study of flavonoid chemistry was initially undertaken to provide an independent source of evidence regarding the status and relationships of these taxa. Material for flavonoid analysis was collected from natural populations, and voucher specimens are on deposit at TENN. From each taxon, 50 g of vegetative tissue and 5 g of reproductive tissue (spores, sporophylls and strobilus axes) were extracted 2 or 3 times in 85% methanol for at least 24 hrs. Isolation by paper chromatography and identification by UV-spectral analysis for flavonoids followed the general procedures as outlined by Mabry, Markham and Thomas (The Systemat- ic Identification of Flavonoids, 1970). The identity of the flavonoids was confirmed by co-chromatography with authentic compounds on cellulose-TLC. Flavonoids were found to be absent in the vegetative material of all three taxa. In the reproductive tissue, however, chrysoeriol was uniformly present. Chrysoeriol has been found commonly throughout the genus Lycopodium sensu lato, according to Voirin and Jay (Biochem. Syst. Ecol. 6:95—-97. 1978). Localization of flavonoids solely in reproductive tissues has not been previously reported for the fern allies. The function of this localization is unknown. Since all three taxa had identical flavonoid characteristics, flavonoid chemistry does not distinguish among them. Thanks are due to R. James Hickey for kindly confirming the identity of the voucher material.—Frank Fusiak, Department of Botany, University of Tennessee, Knoxville, TN 37996-1100. Contributions from the Botanical Laboratory, The University of Tennessee, n.s. No. 542. BRITISH PTERIDOLOGICAL SOCIETY Open to all who are interested in growing and studying ferns and fern-allies. Full members receive THE FERN GAZETTE and the BULLETIN OF THE BRITISH PTERIDOLOGICAL SOCIETY. Membership subscriptions are £7 for full members, £ 5 for ordinary members (not receiving the GAZETTE), and £5 for student members (under 25 years of age). For particulars, U. S. residents should apply to Dr. J. Skog, Biology Department, George Mason University, Fairfax, VA 22030. Non-U. S. residents should apply to La: Corr. G. Coke, Robin Hill, Stinchcombe, Dursley, Gloucestershire, England. _.. from TRIARCH We are grateful for the purchase orders and collec- tions of material received during Triarch’s 56 years of service. We try to say thank you by maintaining quality of product, variety of listings, and by supporting biolog- ical societies with memberships, donations, and advertising. Having completed 25 years in the management of Triarch, | add this written word of thanks for your support, without which Triarch would not exist! Paul L. Conant, President TRIARCH PREPARED MICROSCOPE SLIDES P.O. Box 98 Ripon, Wisconsin 54971 es Soe eee ry SPRINGER-VERLAG New York Heidelberg With Special Reference to Tropical America ROLLA M. TRYON and ALICE F. TRYON, Harvard University MORE THAN 850 PAGES This exceptional volume is the first modern, in-depth treatment of ferns and related plants, particularly in tropical America. Written by Rolla M. and Alice F. Tryon, well known for their work in this area, FERNS AND ALLIED PLANTS presents a new classification of the Pteriodophyta based on a wealth of data published during the last few decades as well as the Tryons’ own wide-ranging research. Extensive bibliographies of modern literature on fern biology are provided. In addition to tropical American genera, this paleotropical regions, making it of worldwide perspective. Each of the 127 tropical and temperate American genera, represented by 29 families, is organized by accounts of its: O ecology O nomenclature and typification O geography O spores O generic characteristics O cytology MORE THAN 2,000 ILLUSTRATIONS AND PHOTOGRAPHS Carefully compiled, ~~ produced, and superbly illustrated, FERNS AND ALLIED PLANTS has more than 2,000 individual illustrations and photographs prepared especially for this volume, a ng: © photographs of living plants in their natural habita O photographs and illustrations of biological and Sac features O distribution ma O more than 700 electron micrographs of fern spores Geologists who use palynological records in hydrocarbon exploration will find the spore micrographs of immense practical value. Unrivaled in its coverage, FERNS AND ALLIED PLANTS will serve as the authoritative reference for years to come. 1982/approx. 896 pp./2022 individual illustrations and photographs in 639 figures/ISBN 0-387-90672-X/cloth $148.00 For more information, or Se order, please write SPRINGER-VERLAG NEW YORK INC., Dept. $5490, 1 175 Fifth Avenue, New York, New York 10010 Berlin AMERICAN sais FERN ERE JOURNAL QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY Pteris x delchampsii, a Spontaneous Fern aie from Southern Florida W. H. WAGNER, JR. and CLIFTON E. NAUMAN _ 97 C-glycosylxanthones in the Asplenium adiantum-nigrum Complex P. MICK RICHARDSON and EUGENIA LORENZ-LIBURNAU 103 Taxonomic Notes on Jamaican Ferns—IV GEORGE R. PROCTOR 107 Pelletixia amelguita, a New Species of Fossil Fern in the Potomac Group (Lower Cretaceous) JUDITH E. SKOG 115 American Fern Journal Index to Volume 72 Errata The American Fern Society Council for 1 DEAN P. WHITTIER, Dept. of Biology, Vanderbilt University. Nashville, TN 37235 —— TERRY R. WEBSTER, Biological Sciences Group, University of Connecticut. Storrs. CT 06268. Vice- 38 sident MICHAEL I. COUSENS, Faculty of Biology, University of West Florida, Pensacola, FL 32504 Seaeiey JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, TN SLILG. Treasurer JUDITH E. SKOG, Dept. of Biology, George Mason University. Fairfax, VA 22030. Records Treasurer DAVID B. LELLINGER, Smithsonian Institution, Washington, DC 2056 Journal Editor ALAN R. SMITH, Dept. of Botany, University of California, Berkeley, - 94720. Memoir Editor JOHN T. MICKEL. New York Botanical Garden, Bronx, NY 10458. Newsletter Editor American Fern Journal EDITOR DAVID B. LELLINGER U.S. Nat’l Herbarium NHB-166, Smithsonian Institution, Washington, DC 20560. ASSOCIATE EDITORS DAVID W. BIERHORST Rt. 3. Box 188, Picayune, MS 39466. GERALD J. GASTONY Dept. of ae Indiana Pein Bloomington, IN 47401. JOHN T. MICKEL ew York Botanical Garden, Bronx, NY 10458. e “American Fern Journal” (ISSN 0002-8444) is an illustrated quarterly devoted to the general study of ferns. It is owned by the American Fern Society, and —— at the Smithsonian Institution, Washington, DC 20560. Second-class postage paid at Washing Claims for missing issues, made 6 months (domestic) to +s month (foreign) after the date of issue, and the matters for pabieaine shawl be addressed to the Edito Changes of address, dues, and applications for membership iguld be sent to Dr. Judith E. Skog, Dept. of Biology, George Mason University, Fairfax, VA 30. Orders for back issues should be addressed to Dr. James D. Montgomery, Ichthyological Associates, .D. 1, Berwick, PA 18603. General inquiries concerning ferns should be addressed to the Secretary. Subscriptions $9.00 gross, $8.50 net if paid through an agency (agency fee $0.50): sent free to members of the American Fern Society (annual dues, $8.00; life membership, $160.00). Back volumes 1910-1978 $5.00 to $6.25 each: single back numbers of 64 pages or less, $1.25; 65-80 pages, $2.00 each; over 80 pages, $2.50 each, plus shipping. Back volumes 1979 et seq. $8.00 each: single back numbers $2.00 each, plus shipping. Ten percent discount on orders of six volumes or more. Library Dr. John T. Mickel, New York Botanical Garden, Bronx, NY 10458, is Librarian. Members may borrow books at any time, the borrower paying all shipping costs. Newsletter Dr. John T. ae New York Botanical Garden, Bronx, NY 10458, is editor of the newsletter “Fiddlehead Forum.” The editor welcomes contributions from members and non-members, including miscellaneous ae offers to exchange or purchase materials, personalia, horticultural notes, and reviews of non-technical books on ferns Spore Exchange Mr. Neill D. Hall, 1230 Northeast 88th Street. Seattle, WA 98115, is Director. Spores exchanged and collection lists sent on request Gifts and Bequests Gifts and bequests to the Society enable it to expand its services to members and to others interested in ferns. Botanical books, back issues of the Journal, and cash or other gifts are always welcomed, and are tax-deductible. Inquiries should be addressed to the Secretary. AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 4 (1982) 97 Pteris X delchampsii, a Spontaneous Fern Hybrid from Southern Florida W. H. WAGNER, JR.* and CLIFTON E. NAUMAN** The genus Preris comprises over 200 tropical and subtropical species, and hybrid- ization among its species is well known. Many of the species-groups, such as the P. longifolia and quadriaurita groups, include both Old World and New World endemics. The Chinese Ladder Brake, P. vittata L., which is a member of the P. longifolia group, has been introduced into the Americas, where it frequently grows with or near closely related native species. For instance, Maxon (1926, p. 436) wrote that P. vittata “has escaped from cultivation and become well established in St. Thomas, Bermuda, Dominica, Martinique, Barbados, Trinidad, eastern Brasil, Florida, Alabama, and Louisiana.” It is with the Florida occurrences that we are concerned here. This study began when C. E. Delchamps discovered some unusual brakes growing along the Coral Gables Canal in Miami in the late 1950’s. He sent examples to Wagner, who found them to be intermediate between the Chinese Ladder Brake and the Bahama Ladder Brake, P. bahamensis (Agardh) Fée. In April 1966, Delchamps and Wagner visited the locality and found the two species and the intermediate growing togther. The intermediate, which we have found to have abortive spores, as would be expected in an interspecific hybrid, was surprisingly common at this locality. It is of special interest because it is a hybrid between a naturalized exotic species and an endemic species, which were brought together through man’s activities. Recently Nauman made special searches for these plants in southern Florida and discovered several new localities for the hybrid, which is herewith described. Inter P. bahamensem et P. vitta rhizomatis atrogriseae; squamae petioli pallido-brunneae, pinnae mediae frondis fertilis maturae (3)4(5) mm latae, remotae, 1.2-1.8 i pinnae ab imo ad summum g denticulatum; coenosori maturi usque ad 1-2 mm a costa extens!. TYPE: North bank of the Coral Gables Canal at 61st Ave. and Waterway Drive, Miami, Dade County, Florida, 17 April 1966, W. H. Wagner 66013 & C. E. Delchamps (MICH). PARATYPES (All Dade County, Florida): Coral Gables Canal bank, SW 39th St. near 62nd Ave., Delchamps in 1960 (MICH); Ca. 30 ft from N bank of the Mowry Canal (C-103) at junction with Tennessee Road, plants in full sun, ca. 10 individuals, 28 June 1979, C. E. Nauman & D. F. Austin 738 (MICH), 739 (FAU); Nixon-Lewis Hammock, W end of Avocado Road, 27 Feb 1979, Nauman et al. 652 (FAU, MICH); Sweetwater, open field near 122 SW 127th Ave., 8 April 1979, T. M. Thurmond 4b (FAU, MICH); Fairchild Tropical Garden, Limestone Wall on SE side of Glade Lake, 22 Dec 1981, Nauman 1490 A-D (MICH). *Department of Botany, University of Michigan, Ann Arbor, MI 48109. **Department of Botany, University of Tennessee, Knoxville, TN 37916. Volume 72, number 3, of the JOURNAL, was issued October 27, 1982. 98 AMERICAN FERN JOURNAL: VOLUME 72 (1982) yy yY EZ S24 4 Zo ime oe Ne FIG. 1. Specimens from the type collection of Pteris x delc seid along the Coral Gables Canal, heneley FL. Middle: Rhizome with sterile frond. Scale = 30 c WAGNER & NAUMAN: PTERIS x DELCHAMPSII 99 We name this fern for the late Dr. C. E. Delchamps, warm friend and field companion over many years, who took time out from his career as Chemistry Professor at University of Miami to explore southern Florida for rare and interesting pteridophytes. His guidance in the natural areas of Florida aided numerous research- ers in their studies of problematic ferns. (See also the obituary by Mrs. C. Bi; Delchamps, 1978.) At first sight, P. x delchampsii resembles a narrow, somewhat skeletonized P. vittata. However, the rhizome scales are less conspicuous, shorter (2-3 mm long), and dark brown. The scales along the petiole are fewer than in P. vittata, but they do have the latter’s pale tan color. The pinnae are more stiff, remote, and considerably narrower. The apical pinnae are less ascending, and the sterile pinna margins (wholly sterile pinnae, and tips of partially fertile pinnae) are not so coarsely or so sharply dentate as in P. vittata. The coenosori of P. x delchampsii do not expand so much as do those of P. vittata when the sporangia have matured and discharged, probably due in part to abortion of the spores and failure of many sporangia to open, but the coenosori of the hybrid nevertheless come closer to the costa than do those of P. vittata. Hybrids found in open, disturbed areas in full sun more closely resemble P. bahamensis in their stiffly erect habit and slightly revolute pinnae. Under shaded conditions, the fronds are more lax and arching and the pinnae are not revolute, and so closely resemble those of P. vittata. Specimens suspected of being P. x delchampsii can be confirmed as that by observing the extensively aborted spores, which are unlike the normal, uniform spores of the parents. A diagnostic key to the parents is given below as an additional aid in separating the hybrid from its parents and the parents from one another, as well as in detecting backcrosses. The key is based upon our Florida collections. KEY TO THE PARENTS OF PTERIS x DELCHAMPSII pinna widths distant, linear, round at the apex, the apical ones usually spreading, not strongly ascending, the sterile margins subentire to entire, lacking pointed teeth; middle fertile pinnae narrow, usually (2)3(4) mm wide; mature coenosori nearly touching the costa, thus exposing little lamina surface; old fronds persisting as naked rachises, the pinnae having disarticulated. P. bahamensis |. Fronds rather lax and spreading; rachises with more or less numerous, conspicuous, narrow, pale tan scales or fibrils; rhizome scales densely tufted, pale tan, 2-4 mm long; stipes scaly throughout; Bahamas, Cayman Islands and Cuba (Maxon, 1926, p. 433; Proctor, 1977, p. 141). 100 AMERICAN FERN JOURNAL: VOLUME 72 (1982) crevices of oolitic limestone, usually rooted in circumneutral to slightly acid marl and in pockets of humus. The canopy of these habitats is composed of Slash Pine, Pinus elliottii. The shrub layer varies in its dominant species but typically contains Serenoa repens, Coccothrinax argentata, Metopium toxiferum, Guettarda scabra, and a variety of other species. The herb layer is dominated by assorted grasses, of which Schizachyrium rhizomatum is a conspicuous example. Plants of P. bahamensis may attain a height of over one meter in this habitat. Older specimens often have litter a decimeter or more deep at their bases composed of old fronds, pine leaves and other debris. Fires in these areas frequently destroy plants completely, but recolonization evidently occurs quickly, and within a period of a few months new plants appear. The Chinese Ladder Brake, P. vittata, occurs scattered throughout Florida and other parts of the southeastern United States at least as far north as the Carolinas and as far west as Louisiana. The plants are frequent in disturbed locations on calcareous substrates, often on limestone ledges, walls, and sinks. The plants are generally more abundant and luxuriant (up to | m tall) in areas of high humidity, such as canal banks and limestone sinks with standing water. In dry places, the plants are sparser and dwarfed (usually less than 0.5 m tall). The introduced species can be found together with its native relative in pinelands, particularly around small sinkholes. However, P. vittata is not common in these habitats. Conversely, the ledges and walls favored by P. vittata are seldom inhabited by P. bahamensis. Hybrids have not been found in the pineland habitats, but only in adjacent areas. The type locality in southwest Miami is a canal bank in an urban, residential area. The rocky sides of the canal had numerous weeds in addition to the brakes. The other ferns were: Anemia adiantifolia (Wagner 66010, MICH), Sphenomeris clavata (66011), and Thelypteris cf. augescens (66009). The canal bank had been widened and most of the ferns, including the parent and hybrid Ladder Brakes, had been destroyed when Nauman visited the locality in 1977. However, he found four new sites for the hybrid during the past five years. Three plants of P. x delchampsii were found along a fringe of the Nixon-Lewis hammock in a border of Schinus terebinthifolius. The Glade Lake population at Fairchild Tropical Garden consisted of 11 plants. Two were terrestrial in a muddy substrate on the south bank of the lake; the remaining plants were confined to a small colony about 1.5 m in diameter on the westernmost portion of a rock wall. Pteris bahamensis was abundant on the north face of the wall, an atypical habitat for this species. Pteris vittata was also abundant near the site, but there were only three small, sterile plants within 50 m of the hybrid plants. The other sites (see specimen citations) were similar in being disturbed, open areas having loose, calcareous substrates covering oolitic limestone. The presently known range of the hybrid in Florida is in Dade County from south Miami to Homestead. We predict that P. Xdelchampsii may be discovered in Monroe and Collier counties. Because the habitat that the hybrid prefers is becoming more common in all of these counties, we expect that its range will increase in the future, although the available natural habitat of P. bahamensis is disappearing. Unless the hybrids develop some means of reproduction, the fate of P. WAGNER & NAUMAN: PTERIS x DELCHAMPSII 101 x delchampsii in years to come depends upon the success of P. bahamensis in surviving man-made changes. In all likelihood, the weedy P. vittata will be the ultimate survivor. BIOSYSTEMATICS OF PTERIS x DELCHAMPSII It is possible that P. x delchampsii may develop a means of reproducing itself, in spite of its abortive spores. (The hybrid spores display all sorts of irregularities in size, shape, and exine development, and were illustrated by DeBenedictus, 1969, pl. 26.) Commonly the number of spores or spore-like bodies per sporangium in P. x delchampsii is less than 64 because normal meiosis is interrupted and a number of large, presumably unreduced spores are formed, in addition to many tiny, aborted spores. The giant spores are evidently viable, as described below. Both the hybird and its parents are tetraploid with 4x = 116; the base number of Pteris is 29. In her studies of this plant, DeBenedictus was unable to obtain a precise count of P. bahamensis, but her evidence supported the conclusion that it is a tetraploid. It is interesting to note that P. longifolia L., of which P. bahamensis may be only a geographical variety, is reported by Walker (1966) also to have 4x = 116, a number evidently characteristic of the Ladder Brake group. Unlike its sexual parents, P. Xdelchampsii shows peculiar pairing behavior. Counts by W. H. Wagner, Jr., K. L. Chen, and V. M. DeBenedictus (vouchers in MICH) show ca. 32-35 bivalents, with the remaining chromosomes univalents. DeBenedictus grew spores of P. x delchampsii because of the possibility that it possessed a low-level expression of apogamy which enabled it to produce gameto- phytes that could proliferate directly into sporophytes. Such a mechanism would allow it to reproduce in its habitats. She found that gametophyte production from hybrid spores was much lower than in the parents. However, those gametophytes that did form bore many antheridia and archegonia on an especially well developed cushion region. Some gametophytes also produced peculiar growths of tissue on the wings. A few of the hybrid gametophytes became filamentous. After flooding her cultures of P. xdelchampsii with water, DeBenedictus ob- served a few sporophytes developing. These arose from archegonia and appeared to be sexual in origin. Unlike P. x delchampsii, the hybrid progeny gave 6x = 174, which DeBenedictus explained as possibly resulting from backcrossing of P. x delchampsii with one of its parents that was present as a contaminant. According to her, “Sperms produced by gametophytes of the hybrid would have 116 chromo- somes. Fertilization of a parental gametophyte [with 58 chromosomes] would produce a sporophyte with 2n = 174” (DeBenedictus, 1969, p. 70). The phenomenon of apparent backcrossing between “sterile hybrids” and their parents in nature also has been suspected in Asplenosorus X ebenoides x Camptosorus rhizophyllus, Dryopteris (goldiana x intermedia) X intermedia (Wagner, 1971), Asplenium Xcurtissii X abscissum (Morzenti, 1967), and Polystichum (acrostichoides X braunti) X acrostichoides (Morzenti, 1962). It is possible that “sterile hybrid” ferns are especially likely to produce unreduced spores because of their faulty pairing behavior, and that the gametophytes produced by the unreduced spores are especially prone to backcrossing with their parents. We 102 AMERICAN FERN JOURNAL: VOLUME 72 (1982) encourage field workers in southern Florida to make special efforts to discover new populations of P. Xdelchampsii and its parents and to examine in detail those already known in order to find both hybrids and backcrosses. Because of its vigorous growth, P. x delchampsii may become an ideal research tool, both in the wild and in the laboratory. We wish to thank Katherine Lim Chen and Virginia Morzenti DeBenedictus for their contributions to this investigation. The University of Michigan Matthaei Botanical Gardens kindly grew cultures of the hybrid plants for our research. Most of the work described here was carried out under National Science Foundation Grant GB-3366, “Evolutionary Characters and Classification of Ferns.” LITERATURE CITED DeBENEDICTUS, VIRGINIA M. MORZENTI. 1969. Apomixis in ferns with special reference to sterile hybrids. Ph.D. thesis, University of Michigan, Ann Arbor, MI. DELCHAMPS, MRS. C. E. 1978. Curtis Eugene Delchamps (1925-1977). Amer. Fern J. 68:6. MAXON, W. R. 1926. Pteridophyta of Porto Rico and the Virgin Islands. Sci. Surv. Porto Rico Virgin 6:373-571. MORZENTI, V. M. 1962. A first report on pseudomeiotic sporogenesis, a type of spore reproduction by which “sterile” ferns produce gametophytes. Amer. Fern J. 52:69—78 67. Asplenium plenum: a fern which suggests an unusual method of species formation. 068. PROCTOR. G. | oe Preridophiyta’ In R. A. Howard (ed.), Flora of the Lesser Antilles, vol. 2. rnold Arboretum, Jamaica Plain, MA on W. H., Jr. 1971. Evolution of Dryopteris i in relation to the Appalachians. /n P. C. Holt (ed.), The Distributional History of the Southern Appalachians. Virginia Polytechnic Inst. State Univ. Res. Div. Monogr. 2, Blacksburg, VA. WALKER, T C. 1966. A ae ae survey of the pteridophytes of Jamaica. Trans. Royal Soc. Edinburgh 66:169—237, pl. 1-5. AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 4 (1982) 103 C-glycosylxanthones in the Asplenium adiantum-nigrum Complex P. MICK RICHARDSON and EUGENIA LORENZ-LIBURNAU* The polyphenolics of the Appalachian Asplenium complex (Smith & Harborne, 1971; Harborne et al., 1973) confirmed in all respects the concept of reticulate evolution in the species complex which had been proposed by Wagner (1954) on the basis of morphology and hybridization and chromosome studies. For example, A. montanum Willd. contained four C-glycosylxanthone compounds which were pres- ent in all hybrids containing the A. montanum genome. It has now been discovered that a parallel situation occurs in the chemistry of a group of European spleenworts, namely the Asplenium adiantum-nigrum complex. Asplenium adiantum-nigrum L., is an allotetraploid derived from A. cuneifolium Viy. and A. onopteris L. (Shivas, 1969). Asplenium onopteris is also a diploid progenitor of the tetraploid A. balearicum Shivas, the other diploid parent being A. obovatum Viv. (Shivas, 1969; Lovis et al., 1972). Thus, A. onopteris is analogous to A. montanum, the diploid ancestor of the tetraploids A. bradleyi D. C. Eaton and A. pinnatifidum Nutt. The analogy is further supported by the presence of C -glycosylxanthones in A. onopteris, A. adiantum-nigrum, and A. balearicum (Fig. 1). The European spleenworts are well known cytologically and morphologically (Lovis, 1977; Walker, 1979), but chemically they are virtually unknown. Several species have previously been surveyed for the presence of xanthones and proved negative: A. adiantum-nigrum (from Spain), A. ruta-muraria L., A. septentrionale (L.) Hoffm., A. trichomanes L., and the related species Ceterach officinarum DC. and Phyllitis scolopendrium (L.) Newm. (Smith & Harborne, 1971). A_ later reinvestigation of A. adiantum-nigrum indicated the presence of a xanthone-O- glycoside (Imperato, 1980). This particular compound had a 1,3,7,8-hydroxylation pattern in contrast to the 1,3,6,7-hydroxylation pattern of the C-glycosylxanthones found in the Appalachian spleenworts. It also had its sugar molecules attached by an oxygen linkage rather than by a direct carbon-carbon linkage as found in the C-glycosylxanthones. Imperato (1980) suggested that the absence of xanthones in the Spanish sample of Smith & Harborne (1971) may be due to phytogeographical factors. A survey of ferns for the presence of C-glycosylxanthones is currently being undertaken at the New York Botanical Garden (Richardson, in press). The discovery of the C-glycosylxanthones mangiferin and isomangiferin in Asplenium adiantum- nigrum growing in the Enid A. Haupt Conservatory at NYBG led to a more intensive study of this plant and its related species. The possibility of geographic variation in xanthone production was examined by studying herbarium samples from almost the complete range of A. adiantum-nigrum. This led to the sampling of the diploid progenitors and ultimately to A. balearicum. *Harding Laboratory, New York Botanical Garden, Bronx, NY 10458. 104 AMERICAN FERN JOURNAL: VOLUME 72 (1982) MATERIALS AND METHODS Apart from the A. adiantum-nigrum plant (NYBG 2066/76) which initiated the study, all experimental work was performed on dried material. Dr. T. G. Walker (Department of Plant Biology, University of Newcastle upon Tyne) kindly provided the sample of A. balearicum. The other samples were all in the herbarium at NYBG. Asplenium adiantum-nigrum.—Scotland, Nicholson in 1881; England, Taylor et al. 1731; Switzer- land, Morthier in 1879; Yugoslavia, Richter in 1910; Gran Canaria, Cook 308; Tenerife, Kuntze in 1888: Madeira, Wilkes 44; Lebanon, Stutz 3045; Punjab, Stewart 7881; Kashmir, Stewart 17448 and 21784: Caucasus, Radde Ex herb. horti. Petropolitani; South Africa, Ex. herb. Mt. Holyoke Seminary, Colorado, Bethel 270; Hawaii, Degener H186 and Degener & Wiebbe 3889. Asplenium cuneifolium.—Germany, Missbach 5591] and Luerssen 5706; Austria, Werderman & Meyer 232; Hungary, Richter 5592. A. onopteris.—Switzerland, Leroy; Italy, St. Lager in 1894; Dalmatia, Ronniger in 1926. Small amounts of plant material were extracted in 80% methanol using a Polytron Homogenizer (Richardson, 1982). The extracts were initially screened for the presence of C-glycosylxanthones by one-dimensional chromatography on Whatman No. | paper in both water and 15% acetic acid. C-glycosylxanthones are revealed as orange spots under ultraviolet light (360 nm), turning fluorescent yellow with ammonia vapor. Extracts which proved positive were further analysed by two- dimensional chromatography on Whatman No. 3 paper in TBA (t-BuOQH: HOAC: H3O, 3:1:1) and HOAc (15% acetic acid). The C-glycosylxanthone spots were eluted in 80% methanol and co-chromatographed with mangiferin and isomangiferin (isolated from Asplenium montanum, Bozeman & Radford 11552, NYBG) in TBA, BAW, 15% HOAc, and H30. Acid hydrolysis of the extract (2N HCl, | hr, 100°C), followed by two-dimensional chromatography, was performed on the one fresh sample of A. adiantum-nigrum. For further details of the chromatography solvents and methods, see Harborne (1973) and Markham (1982). RESULTS AND DISCUSSION The two-dimensional chromatogram of the A. adiantum-nigrum from the conser- vatory revealed the presence of eleven polyphenolic compounds. Four were dark/yellow spots, typical of flavone and flavonol glycosides, with Ry values in TBA and HOAc as follows: 14,40; 17,51; 23,51; and 27,58. Seven were orange/yellow spots, typical of xanthones, with Ry values: 36,24; 37,49; 46,63; 20,72; 55,57; 55,68 and 70,64. The rutin marker was 39,68. After acid hydrolysis, the latter four xanthone spots disappeared and were replaced by a new xanthone spot at 48,26. This suggests that these latter four compounds were xanthone-O-glycosides, one of which was characterized by Imperato (1980). The other three xanthones were resistant to acid hydrolysis and were suspected to be C-glycosylxanthones. For two of the compounds this was confirmed by co-chromatography with mangiferin and isomangiferin, but the identity of the third C-glycosylxanthone remains unknown until larger quantitities of the compound become available for analysis. Extracts from herbarium specimens of A. adiantum-nigrum produced either two or three xanthone spots on the two-dimensional chromatograms. In every case mangiferin and isomangiferin, and in some cases the unidentified C-glycosyl- xanthone, were readily visible and confirmed by co-chromatography. In no cases RICHARDSON & LORENZ-LIBURNAU: C-GLYCOSYLXANTHONES IN ASPLENIUM 105 were the four dark/yellow or the four xanthone-O-glycoside spots visible. This may be due to the small amounts of herbarium material available for analysis. It was usual to use only ca. 10-20 mg per chromatogram, compared to a whole plant sometimes used by Smith & Levin (1963). It is also possible that some of the missing compounds have been oxidized since some of the herbarium specimens date from the late nineteenth century. The three samples of A. onopteris yielded two-dimensional chromatograms which were identical to those of the ‘three-spotted’ herbarium material of A. adiantum- nigrum. Each sample contained mangiferin, isomangiferin, and the same unknown C-glycosylxanthone. The four samples of A. cuneifolium produced blank two- dimensional chromatograms, even when repeated with comparatively large amounts of material. It can be stated firmly that C-glycosylxanthones were absent from all four samples. Only one specimen of A. balearicum was available, and the two- dimensional chromatogram indicated the presence of mangiferin, isomangiferin, and the same unknown C-glycosylxanthone, as well as a large dark/yellow spot at 32,55. However, there was insufficient material to verify the three xanthones by co- chromatography with the compounds from the other species. It would be interesting to examine larger amounts of A. onopteris and A. balearicum chemically in order to determine if they contain the xanthone-O-glycosides which occur in A. adiantum- nigrum, Samples of A. adiantum-nigrum from throughout its range contained C-glycosyl- xanthones. This contrasts sharply with the earlier negative report by Smith & Harborne (1971). Possibly there is a genuine geographical variation in xanthone occurrence (this could be revealed by more intensive sampling). Alternately, the plant examined by Smith & Harborne (1971) may have been misidentified and may actually be A. cuneifolium. Most of the A. cuneifolium plants at NYBG were found to be misidentified as A. adiantum-nigrum, presumably due to the great similarity between the two species. Asplenium onopteris is the first diploid European Asplenium species which has been found to contain C-glycosylxanthones. The compounds are absent from A. cuneifolium. Asplenium obovatum may or may not contain the compounds, and this cannot be predicted by the occurrence of the compounds in A. balearicum. If A. obovatum actually contains C-glycosylxanthones, then the compounds can be pre- dicted to be present in both A. billotii Schultz, the autotetraploid derivative of A. obovatum, and A. foreziense Heribaud, another allotetraploid derived from A. obovatum. An investigation of the whole European Asplenium complex for C-glycosylxanthones would perhaps confirm the relationships established by cyto- logical studies. The occurrence of C-glycosylxanthones in the European Asplenium complex also may help to relate them to the Appalachian spleenworts. Smith & Harborne (1971) mention that the ocurrence of xanthones in other Asplenium species may possibly suggest a relationship to A. montanum. Experimental hybrids of A. onopteris and A. montanum would reveal any chromosomes held in common. Finally, the occurrence of C-glycosylxanthones in the diploid A. onopteris and its tetraploid derivatives helps to confirm the suggestion by Smith & Levin (1963) that chemical investigations have significant potential for evolutionary studies involving polyploidy. 106 AMERICAN FERN JOURNAL: VOLUME 72 (1982) The authors wish to thank W. H. Wagner, Jr. for assistance with various aspects of this study, T. G. Walker for the Asplenium balearicum material, and Linda L. Oestry for a critical reading of the manuscript. LITERATURE CITED HARBORNE, J. B. 1973. Phytochemical a Chapman & Hall, Lond —— ce A. WILLIAMS and D. M. SMITH. 1973. Species-specific Kacmplerol derivatives in the splenium complex. Biochem. “hi sec. F. 1980. A xanthone-O-glycoside from Asplenium adiantum-nigrum. Phytochemistry 19:2030—2031. LOVIS, J. D. 1977. Evolutionary patterns se processes in ferns. Advances Bot. Res. 4:229-415. ———., P. ROWNSEY, A. SLEEP and M. G. SHIVAS. 1972. The origin of Asplenium balearicum. Brit. Fern Gaz. 10: sm 268. MARKHAM, K. a sag wegen of Flavonoid Identification. Academic Press, London. RICHARDSON, . 1982. A speedy and efficient extraction procedures for flavonoids. Phytochem. Bull. 4 en . In press. C-glycosylxanthones in the fern genera Davallia, Humata and Nephrolepis. Phytoc SHIVAS, M. G. 1969. A cytotaxonomic study of the Asplenium adiantum-nigrum complex. Brit. Fern Gaz. 10:68-80. SMITH, D. M. and J. B. HARBORNE. 1971. Xanthones in the Appalachian Asplenium complex. pare! 10:2117-2119 EVIN. 1963. A ‘cisomtusraphic study of reticulate evolution in the Appalachi- an yer complex. Amer. J. Bot. 50:952-958. WAGNER, W. H., Jr. 1954. Reticulate evolution in the Appalachian Aspleniums. Evolution 8:103-118. WALKER, T. G. 1979. The . of ferns. Jn A. F. Dyer (ed.). The Experimental Biology of Ferns. Academic Press, Lon AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 4 (1982) 107 Taxonomic Notes on Jamaican Ferns-IV GEORGE R. PROCTOR* This paper concludes my series of short papers on Jamaican ferns (Proctor 1965, 1968, 1981) written in conjunction with the preparation of a book about these plants, now essentially completed. For the sake of brevity, the rationale for new combinations will not be discussed at the present time. The reasons for these changes will become evident when the Jamaican fern volume is published. I thank Dr. John Wurdack for his help in preparing the Latin descriptions, and Margi von Montfrans for drawing the two figures. LYCOPODIACEAE Lycopodium hippurideum var. montanum (L. M. Underw. & Lloyd) Proctor, comb. & stat. nov. Lycopodium montanum L. M. Underw. & Lloyd, Bull. Torrey Bot. Club 33:107. 1906. TYPE: Summit of Blue Mt. Peak, Jamaica; Underwood 2537, (NY: = Maxon 1412, isotype US). HYMENOPHYLLACEAE Trichomanes micropubescens Proctor, sp. nov. Fig. 1. ubg. Didymoglossum. A speciebus cetera subg. Didymoglossi laminis fertilibus nguste linearibus vel late oblanceolatis 6-13 mm gis 1-6 mm latis omnino nneis bus, marginibus pilis maioribus stellatis 3—5-ramosis remotis, soris solitariis terminalibus, involucro anguste obconico exserto, labiis abrupte expansis reflexis rotundis zona marginali fuscobrunnea ampla 5-8 cellulis lata differt. Plants imbricate, formin ats: rhizome filiform, clothed with dark brown S tis ‘ always solitary; involucre narrowly obconic, 2-2.5 mm long, half or less immersed, the free distal portion marginate or very narrowly flaring in two broadly rounded, usually reflexed lips, these with a wide, brown- darkened marginal zone 5-8 cells broad. TYPE: Along the Drivers River above Soyo Falls, parish of Portland, Jamaica, on deeply shaded tree-trunks in wet forest near river, 400-500 ft (122-152 m), 23 Jan 1980, Proctor 38554. (IJ; isotype GH). *Jardin Botanico Nacional, Apartado 21-9, Santo Domingo, Dominican Republic. 108 AMERICAN FERN JOURNAL: VOLUME 72 (1982) This species is unique in its combination of minute, simple pubescence along the margins and abaxial side of veins and the very broad, darkened marginal band (up to 8 cells wide) on each lip of the involucre. mm Fig. 1. Holotype of Trichomanes micropubescens Proctor Trichomanes radicans var. cee age) (v. d. Bosch) Proctor, comb. & stat. nov. Trichomanes antillarum vy. d. Bosch, Ned. Kruidk. Arch. 5:164. 1861. TYPE: “Woods above Fort Stewart”, parish of St. Mary, eottts see 1843, Purdie s.n. (K). CYATHEACEAE Cyathea x bernardii Proctor, hybr. nov. x lewisii stipitibus longioribus ae cm vs. 20-35 cm), pinnis side (non fonictamien chet stipitibus 10-12 mm longis, caudicis stipitisque squa omnino albidis solum bicoloribus ad basim apicemque, costulis sbaxiaiiter squamis HRer. minutis albis bullatis armatis differt Caudex to 5 m tall, 6-10 cm in diameter, not spiny, densely clothed at the apex with whitish, Be, concolorous (or faintly bicolorous at base and apex) scales, these a to 4 cm long. Fronds ar poe eee 2.5-3.2 m long; aie 50-70 cm the adaxial side; pinnae lance- pseske 45- 65 m lon ng, up to 22 cm broad, stalked, acuminate; costae densely hirsute on adaxial side, sparsely so beneath; pinnules up to 32 pairs, ayaa -oblong, up to 12 cm long, 1.7-2.5¢ cm broad, sessile, attenuate at the apex, cut nearly to the costule; cine and midveins more or less hirsute on both sides, shaGally bearing numerous, small, whitish bullate scales with hairlike tips; segments mostly 24-27 pairs, narrowly oblong- falcate, 10-15 mm long, 2-2. mm broad, the apex blunt, the margins crenulate-lobulate and more or less revolute; G. R. PROCTOR: NOTES ON JAMAICAN FERNS-IV 109 veins mostly 12-15 pairs, chiefly 1- or 2-forked below the middle. Sori 4-10 er ge inframedial or nearly medial; indusium flabelliform, irregularly lacerate or lobed, few of the lobes often hair-pointed, brownish-hyaline; receptacle bearing scaall, golden-yellow, blunt-tipped paraphyses and much longer, colorless, sharp-pointed, hairlike ones, these exceeding the sporangia in length; spores often eno or abortive. YPE: Corn Puss Gap, parish of St. Thomas, Jamaica, in lower montane rain forest, 2000 ft (610 m), 13 Feb. 1950, Proctor 400]. (US; isotypes IJ, MO). This entity is postulated to represent the hybrid Cyathea arborea x C. armata (R. Tryon, 1976). It was formerly equated with Cyathea x calolepis (D. C. Eaton) Domin of Cuba, which is now thought to be a hybrid of C. arborea with a species other than C. armata, one not so far identified. Because the Jamaican plant has a rather wide local distribution (having been found at six localities in the parishes of St. Mary, St. Andrew, and St. Thomas), it has been provided with a specific epithet as a matter of convenience. It is named for Mr. C. Bernard Lewis, former Director of the Institute of Jamaica. The type specimens of C. x bernardii were collected from a plant growing beside the type plant of C. x J/ewisii (Morton & Proctor) Proctor, named for the same Mr. Lewis, and also having C. arborea as one of the parent species (C. aspera (L.) Swartz is the other). POLYPODIACEAE Arachniodes argillicola (Proctor) Proctor, comb. n Polystichopsis argillicola Proctor, Amer. Fern. J. 51: 147. (sei. ers 2 miles NE of High Peak, above Murdock’s Gap, parish of Portland, Jamaica, on shaded clay banks, 3500-4500 ft (1069-1372 m), Proctor 5842 (IJ; isotype GH). Arachniodes chaerophylloides (Poiret) Proctor, comb. nov. Polypodium chaerophylloides Poiret in Lam., Encycl. Méth. Bot. 5:542, 1804. TYPE: Puerto Rico, Ledru s.n. (P. Herb. Lam., photos GH, US). Polystichopsis chaerophylloides (Poiret) Morton, Amer. Fern J, 50: 155. 1900. Arachniodes lurida (Jenman ex a M. Underw. & Maxon) Proctor, comb. nov oe. rasan Jenman ex L. M. Underw. & Maxon in Slosson, Bull. Torrey Bot. Club 40: 183, Lene tne eg | . TYPE: Jamaica, Jenman, “1874- 79” (NY; isotype US). ph aikeils lurida (Jenman ex L. M. Underw. & Maxon) Morton, Amer. Fern J. 50:155. 1960. Arachniodes macrostegia (Hooker) Proctor, comb. no Sse tere macrostegium Hooker, Sp. Fil. 4:148. 1862. TYPE: Bo Uaupés, Brazil, Spruce 2245 (K; isotype U Nephrodium ones Baker, Ann. Bot. 5:325. 1891. TYPE: Fox's Gap, Jamaica, April 1886, Hart s.n. (K; isotype I). er aety oc een ils (Baker) Morton, Amer. Fern J. 50:155. 1960. Arachniodes pubescens (L.) Proctor, comb. n Polypodium pubescens L. Syst. Nat. ed. 10, 2:1327. nes TYPE: Jamaica, P. Browne s.n. (LINN 1251.4 Polvetichapsis pubescens (L.) Morton, Amer. Fern J. 50:155. 1960. Asplenium erosum var. duale (Jenman) Proctor, comb. & stat. nov. Asplenium duale Jenman (as Adiantum in error), Gard. Chron. III, 13:10. 1893. TYPE: Jamaica, Sherring s.n. (K ?, not seen). 110 AMERICAN FERN JOURNAL: VOLUME 72 (1982) Asplenium gilbertii Proctor, sp. nov. Fig. 2. Ab Asplenio sciadophilo \aminis deltato-oblongis minus quam 10 cm longis basiliter truncatis, divisionum ultimarum venis furcatis differt, ab A. trichomanes- dentato, A. myriophyllo et cetero laminis 2-pinnatis differt; et ab A. clutei laminis 2.5-4 cm latis (vs. minus quam 1.5 cm) et pinnarum plus quam 10 paria ferentibus differt. h e short, erect, 3-4 mm thick, clothed at apex with blackish, fee puen narrowly deltate- Shea or deltate-attenuate scales 1-3 mm long, these sometime atshaedapet Fronds few, erect or spreading, 15-20 cm long; stipes ee or brownish, 8—11 cm long, aie than ib blades, spay ae marginate, beans pairs, short-stalked, slightly inequilateral, lance-deltate, 7-11 mm broad near base, pinnate in the basal half with 3 or 4 pairs of free pinnules, the Tage acroscopic one the largest; distal portion of pinnae ee or Jona to an uae apex, the lobes 1-1.5 mm ong: indus m bro rown a eh the Sats bane sage ee sinks! mostly immature or abcetia TYPE: Mandeville, Jamaica, B. D. Gilbert s.n. in 1895 (GH). The stated type locality is about 2000 ft (610 m) above sea level. Known only from the type collection. Asplenium myriophyllum var. brevisorum ope var. nov. 0.4-0.8 m dilute viridi differt Differs from var. myriophyllum in having shorter sori (0.5—1.3 mm long, vs mostly 1.5-2.5 mm long), these never diplazioid; in having mostly n narrower ane divisions (0.4—0.8 m Sapstrte vs. 0.7—1.4 mm broad); and in the rather stiff, pale to light green tissue of the bla The two Jamaican varieties of AA species appear to be completely allopatric. Variety brevisorum occurs exclusively on limestone rocks and has been found chiefly in the central and western parts of the island; var. myriophyllum, on the other hand, occurs both on calcareous and non-calcareous rocks at somewhat higher elevations and is confined to the region of the Blue and Port Royal Mts. Both forms need to be investigated cytologically. Probably neither is endemic to Jamaica. Asplenium radicans var. tripinnatum (Hieron.) Proctor, comb. nov. Asplenium flabellulatum var. tripinnatum Hieron. Hedwigia 47:229. 1908, LECTOTYPE (chosen here): Ecuador, Stuebel 85] (B, photo US). The other syntypes are: Colombia, Stuebel 3/ and /5/ (both B, photos US) and Ecuador, Stuebel 780 (B). = trichomanes-dentatum var. jamaicense (Jenman) Proctor, comb. & Stat. a sence: Jenman, J. Bot. Brit. For. 24:268. 1886. TYPE: Jamaica, Sherring s.n. (K, not seen G. R. PROCTOR: NOTES ON JAMAICAN FERNS-IV lll Fig. 2. Holotype of Asplenium gilbertii Proctor Diplazium montediabloense Proctor, sp. nov. Diplazium brevisorum John Smith, Cat. Kew Ferns 6. 1856, non John Smith, 1841. TYPE: A cultivated specimen originating from Jamaica (K, not seen). A Diplazio pectinato magnitudine maiore, costis nudis et abaxialiter fere glabris, segmentis in quoque pinna paucioribus (minus quam 23, vs. plus q 5 in quoque latere) et latioribus (8-10 mm latis, vs. 3-7 mm latis), differt. Rhizome unknown, but plant stated to be “arborescent,” i.e., with an erect caudex like a tree fern. Fronds evidently large, more or less oblong, of unknown length but 112 AMERICAN FERN JOURNAL: VOLUME 72 (1982) probably between | and 2 meters, 35-50 cm broad, acuminate at the apex; rhachis and other vascular parts stramineous, glabrate, bearing but a few scattered pluri- cellular hairs chiefly at the base of costae abaxially. Pinnae lance-oblong, short- cm long (the lowermost pair of each pinna somewhat reduced), 8-10 mm broad, obtuse at the sharply serrate apex, the margins crenate- or bicrenate-lobulate, cut to 0.5-1.5 mm of the costa, the sinuses rounded; veins mostly forked below the middle, prominulous especially on adaxial side; tissue light green, thin-membranous. Sori inframedial, ca. 1.5 mm long; indusium thin, light brown, the margin more or less erose. TYPE: “Near the top of Mt. Diablo,” parish of St. Catherine (?), Jamaica, R. C. Alexander s.n. in 1849-50 (US). This specimen is marked “Asplenium brevisorum Hs, Sai. Known only from the type collection. This species seems obviously related to Diplazium pectinatum (Fée) C. Chr., but differs in its much larger size, fewer segments per pinna, and almost entire lack of pubescence. Diplazium stokeyae Proctor, sp. nov. Asplenium radicans var. pallidum Jenman, Bull. Bot. Dept. Jamaica II, 2:88. 1894. TYPE: Second lumina parallelogramoid in outline, averaging 0.2 mm long and 0.1 mm wide, often ronds ca. 1 m long; stipes shorter than the blades, trisulcate adaxially, brown darkening to nearly black at base, the basal part bearing a few Q. oO ie’) m broad; rhachis light brown, glabrescent, slightly viscid-glandular;, pinnae 8-10 pairs, subopposite to alternate, stalked, lance-oblong or oblong, acuminate, -8 cm broad, the lowermost | or 2 pairs slightly inequilateral with basiscopic pinnules a little longer than the acroscopic ones; free pinnules 3-5 pairs, the remainder of the ultimate divisions more or less adnate; costae very narrowly green-marginate; costae and costules brown and glabrescent to very sparingly puberulous with short pluricellular hairs, each costule subtended at the base by a single very small, pale brown, narrowly deltate-attenuate scale; pinnules and larger adnate divisions oblong, obtuse, 2.5-4 cm long, 1.3—-1.6 cm broad, the margins crenate-serrate or crenate-biserrate to very shallowly lobulate; tissue pale green, membranous, glabrous on the adaxial side, sparsely puberulous abaxially with very minute white, erect, apparently unicellular hairs. Sori 5-7 pairs per division, inframedial on distal vein-branches, 4-5 mm long; indusium narrow, brown, with a subentire to slightly erose margin. TYPE: Near the waterfall 0.5 mile N of Hardwar Gap, parish of Portland, Jamaica, ca. 3800 ft (1158 m), Proctor 9849 (IJ). Named for the late Alma G. Stokey in recognition of her valuable studies of fern gametophytes (based in part on Jamaican materials) and also in fond remembrance. Both of the cited localities for Diplazium stokeyae were sources of fern spores that she used in her work. G. R. PROCTOR: NOTES ON JAMAICAN FERNS-IV 113 Diplazium wilsonii var. eae ne (Jenman) Proctor, comb. & stat. nov. Asplenium brunneo-viride Jenman, J. Bot. Brit. For. 24:269. 1886. TYPE: Vicinity of Cinchona, parish of St. Andrew, Jamaica, risen Rue eal dS Grammitis minor (Jenman) ange comb. sieges minor Jenman, Bull. Bot. Dept. ae é a 212. 1897. TYPE: Murray’s Flat, Mt. Moses, parish of St. Andrew, Jamaica, iat s. n. in 1876 (NY; isotype US). Cochlidium minus (Jenman) Maxon, Sci. Surv. Porto Rico Virgin Is. 6:407. 1926. Grammitis seminuda (Willd.) Proctor, comb; ee seminudum Willd. Phytogr. 13, ¢. 8, fig el "TYPE: Martinique, /sert s.n. in 1787 (B-Herb. . £9587). Coc Le seers (Willd.) Maxon, Sci. Surv. Porto Rico Virgin Is. 6:407. 1926. ee fimbriata Maxon ex nage s nov. uebelii magnitudine multo €, pinnis oppositis vel suboppositis, costis cost peanane esecuiniter calanduless pilis paucis et grossioribus, differt. Rhizome 2.5—4 mm in diameter, ing numerous glistening, reddish-brown, eS apiculate are often serial in finely bristle-like tips. Alias laxly ascending, up to | m or more long; stipes 40-45 cm long, lustrous brown, shallowly grooved adaxially, rough especially toward the basal end by the ini scherctie te of broken-off hairs, otherwise bearing a few narrowly conic spines 0.5-1. long, also minutely glandular- ene and wi ith scattered, colorless, aie: articulate hairs up to 1.5 mm long. Blades deltate, up to 60 cm or more broad at base, 2- uate ean or -sect with lobed ultimate segments, acuminate at the apex, the vascular axes and veins on both sides with scattered, colorless, articulate hairs, the ‘cn tissue glabrous; rhachis with a few s cattered spines like those of the stipe; pinnae spreading, opposite or subopposite, “es oblong, with up to | pairs of pinnules, these narrowly deltate-oblong, the larger ones up to "10 c m long, 2.5 cm broad at base , mostly subsessile; ee segments lobed, the femile lobes lobulate at the acroscopic base. Sori located on fertile lobules; nee ery membranous, densely ciliate with a tuft “Of colorless, hyaline acicular T : Lower slopes of Mt. Moses, parish of St. Andrew, Jamaica, 000-2500 ft (610-762 m), 14 Apr 1903, Maxon 1060 (US). Known only from the type collection and an unlocalized specimen gathered by Wiles (BM, fragm. US) during the period 1793-1805 Hypolepis jamaicensis Maxon ex Proctor, sp. nov. A H. bogotensi stipite et rhachidi spiniferis costis costulisque eglandulosis side . nigriscenti laminis minus se (3-pinnato-pinnatifi 4 pacer vl 5-pinn atis), spinis multo minoribus aus quam | mm, VS. oe y a 2 mm) rectis, costis eeenitis isque DObENCEHREALE diff Rhizoie 25 mm i nei mostly oxen bit on one or two sides toward the Owing apex and ne es bearing lustrous, yellowish-brown, articulate hairs, these with nary pointed tips. Fronds sprawling or vinelike, up to m long, sometimes forming tangles; stipes 50-100 cm long, dull yellowish-brown, darker toward the base, grooved rsucoeed finely muriculate or with scattered, small, sharp spines 0.5-1 mm long, glabrescent. Blades narrowly deltate-oblong, att peg up to | m broad at base, 3-pinnate- ~pitinatifid: rhachis and costae muriculate finely spiny like the stipe: pinnae oblique or spreading, sori or subopposite, deltate-oblong, acuminate, with up to 15 pairs of opposite t alternate primary pinnules, these oblong-acuminate, the larger ones up to 15 cm nee and 6 cm broad at base, all but the most distal ones short-stalked; vascular parts on both sides 114 AMERICAN FERN JOURNAL: VOLUME 72 (1982) pubescent with pluricellular, colorless hairs, these somewhat curved or flexuous but mostly not strongly articulate and not glandular; ultimate segments lobed or lightly crenate, pale-veined. Sori solitary at acroscopic bases of segment lobes; indusium le-membranous, the margin (at least when young) coarsely toothed and sparingly septate-ciliate. YPE: Vicinity of New Haven Gap, parish of St. Andrew, Jamaica, ca. 5500 ft (1688 m), 21 June 1904, Maxon 2610 (US). Probably endemic in Jamaica, this species occurs rather widely in the Port Royal, Blue, and John Crow Mountains at elevations of 1200-6500 ft (366-1982 m). It is clearly related to H. bogotensis Karsten (which also occurs in Jamaica) and H. repens (L.) C. Presl (which does not). By implication, A. R. Smith (1981) included this entity within his concept of H. bogotensis, at the same time rightly pointing out that true bogotensis does occur in our area. Hypolepis jamaicensis differs from H. bogotensis in its larger fronds and more rampant habit; its muriculate or finely spiny stipe, rhachis, and costae; the finer character of its pubescence; in its lack of glandularity; and in its pale (vs. dark) ultimate veins. Hypolepis jamaicensis seems most closely related to H. repens (with which it has often been confused), a species found widely in the West Indies and continental American tropics, but not so far authentically identified as occurring in Jamaica. Hypolepis repens is more spiny and less pubescent than H. jamaicensis, and its indusium is subentire and glabrous. Polypodium triseriale var. gladiatum (Kuhn) Proctor, comb. nov. Polypodium brasiliense var. gladiatum Kuhn, Engl. Bot. Jahrb. 24:130. 1897. TYPE: “Interior” of Cuba, Sept 1822, Poeppig s. n. (LZ, destroyed; isotype BR, photo US). Polypodium gladiatum Kunze, Linnaea 9:45. 1834, non Vell., 1827. Goniophlebium ampliatum Maxon, Contr. U.S. Nat. Herb. 10:492. 1908. Polypodium ampliatum (Maxon) Proctor, Bull. Inst. Jamaica Sci. Ser. 5:45.1953. Polytaenium lineatum var. intramarginale (Baker ex Jenman) Proctor, comb. & stat. nov. : Vittaria intramarginalis Baker ex Jenman, J. Bot. Brit. For. 15:266. 1877. TYPE: Jamaica, Jenman 58 in 1877 (K). Polytaenium intramarginale (Baker ex Jenman) Alston, Mutisia 7:9. 1952. Pteris arborea f. regia (Jenman) Proctor, comb. & stat. nov. Pteris regia Jenman, Gard. Chron. III, 17:39. 1895. TYPE: Jamaica, D. Morris s. n., ex herb. J. Hart (K, isotypes IJ, photos US). LITERATURE CITED PROCTOR, G. R. 1965. Taxonomic notes on Jamaican ferns. Brit. Fern Gaz. 9:213-221. _______. 1968. Taxonomic notes on Jamaican ferns-Il. Brit. Fern Gaz. 10:21-25, t. I. . 1981. Taxonomic notes on Jamaican ferns—III]. Amer. Fern J. 71:57-61. SMITH, A. R. 1981. Pteridophytes, Part 2. Jn: D. E. Breedlove (ed.), Flora of Chiapas. California Acad. Sci., San Francisco. TRYON, R. 1976. A revision of the genus Cyathea. Contr. Gray Herb. 206:19-98. AMERICAN FERN JOURNAL: VOLUME 72 NUMBER 4 (1982) 115 Pelletixia amelguita, a New Species of Fossil Fern in the Potomac Group (Lower Cretaceous) JUDITH E. SKOG* A new species of the fossil fern Pelletixia Watson and Hill (1982) has been identified from fragments of pinnules discovered in sediments belonging to the Lower Cretaceous Potomac Group of Maryland. The locality was exposed in a stream bank at high flood stage during a tropical storm in 1972. Subsequently, material was collected from the site for several years until erosion by the stream completely destroyed the locality. The material described here is placed in a genus previously known only from the Wealden beds (Lower Cretaceous) of England. The Potomac Group and Wealden Formation have been shown to have many plants in common (Hueber, 1982), and so it is not surprising to find an additional example of a genus reported only from these two localities. Locality and Stratigraphy.—Pelletixia amelguita was isolated as pinnule frag- ments from material brought to the U. S. National Museum of Natural History by Mr. Howard Hruschka. He had collected the material from a high, newly eroded bank along Paint Branch about one mile north of the University of Maryland campus in College Park, Maryland (Washington East Quadrangle 7.5 minute series topo- graphic sheet, 39°00’ W lat., 76°56’ N long.). The plant-bearing strata are in Zone 1 of the Potomac as defined by Doyle (1969) and are probably of Barremian—Aptian age within the Lower Cretaceous. The stratum whence came the specimens de- scribed herein is probably the same which produced the rhizome of the fern Loxsomopteris anasilla J. Skog (1976). The age of the specimens is approximately 130 million years. Materials and Methods.—The clay matrix was bulk macerated in 48% hydroflu- oric acid. The residue was thoroughly washed with several changes of water until acid-free, sieved, and then sorted under a dissecting microscope for all sizes of material (Hueber, 1982). The specimens were examined under a Wild M-5 dissect- ing microscope, photographed, and then drawn using a camera lucida attachment. One specimen was air dried and mounted on the surface of a 12 mm cover glass that had been thinly coated with white glue. The cover glass was attached to a SEM stub with silver paint, coated with gold-palladium, and observed with a JOEL JSM-35C scanning electron microscope at the Palynology Laboratory, United States Geologi- cal Survey, Reston, Virginia. The specimen was cracked open on the stub and recoated to release some spores after the first viewing indicated the presence of sporangia and spores. Unfortunately, the inclusion of thymol granules in the water used for separating material did not prevent the second specimen from being destroyed by bacteria and fungi. This problem has also been noted by Hueber (1982) for material of the same age from a different locality. Photographs were taken with a Wild M-5 dissecting scope with 35mm camera attachment using Kodak Plus-X Pan film or with the SEM using Polaroid Type 52 positive-negative film. “Department of Biology, George Mason University, Fairfax, VA 22030. 116 AMERICAN FERN JOURNAL: VOLUME 72 (1982) FIG. 1. Pinnules of Pelletixia amelguita, X26. FIG. 2. Pinnule of Pelletixia aap Type specimen care 326733, x52. The abbreviations are: |= ike. m= matrix, mr= midrib, s = stalk. Pelletixia amelguita J. E. Skog, sp. nov. (Schizaeaceae) Figs. 1-12. The fragments isolated from the goles represent two tightly enrolled Reng 13 to 2.5 mm long on short stalks mm or less long (Figs. / and 2). The specimens are fractured, probably due to sirtaenae as the matrix dried out in the jaboratory (Figs. 2 and 3). Ridges and clay-filled grooves can be seen on both specimens anc J. E. SKOG: PELLETIXIA AMELGUITA 117 may represent veins or lobes of the pinnule. Such detail would be better detined were the pinnules unrolled. A definite midrib is seen along the adaxial surface of the pinnules (Fig. 3). Only one specimen (Fig. 2) remained for further study after general handling while studying; the additional characters described here are based on that specimen alone. When first viewed with the scanning electron microscope, the pinnule was clearly seen to be fertile (Figs. 4 and 5). The epidermis shows few cellular details other than it is composed of smooth, even cells. The sporangia were broken when the specimen was dissected, with the result that no details of the annulus can be described. Remnants of what may be an annulus can be seen in Figs. 4 and 8. At least 120 (128) spores can be counted in one area (Fig. 5), probably derived from a single sporangium. The spores are well-preserved and distinctive. They are trilete (Fig. 6), about 28-35 jm in diameter, and have raised lips around the laesura which seem to extend into a ridge that extends to the margin of the spore (Fig. 7). The spores are triangular with the margins at times slightly convex and the apices somewhat rounded. Two or three muri parallel the margins of the spore on both the proximal and distal surfaces (Figs. 6 and 8). These muri are narrow, 0.5—2 wm wide, 1-2 ym high and are separated by spaces 1.5—3.5 xm wide. On the distal surface the muri join at the pole to form a triangle about 5.0 4m in diameter. The shape of the triangular area is variable, ranging from sharply triangular (Figs. 8 and 9) to subdivided (Figs. /0 and //), to circular within a triangle (Fig. /2). The outer muri extend to the apices. TYPE: USNM 326733, Paleobotany Collections, U. S. National Museum of Natural History. TYPE LOCALITY: East bank of Paint Branch, College Park, MD, U.S.A. Washington East Quadrangle 7.5 minute series, 39°00’ Lat. 76°56’ Long. Stratigra- phy: Patuxent Formation, Potomac Group, Barremian—Aptian, Lower Cretaceous. SPECIES NAME: Derived from the Spanish diminuitive of “amelga” meaning a ridge between valleys—to emphasize the smaller muri on the spores of this species in contrast to the spores of Pelletixia valdensis (Seward) Watson & Hill. Seward (1913) described fertile pinnae or fronds from the Wealden (Lower Cretaceous) in England which he named Pelletieria valdensis. Pelletieria Seward, 1913, was shown by Watson and Hill (1982) to be a later homonym of Pelletiera St.-Hilaire, 1822, a member of the Primulaceae. They proposed the new generic name Pelletixia and formed the new combination Pelletixia valdensis for the monotypic species. Watson (1969) restudied Seward’s type specimen for the species and determined that the fronds were quadripinnate, comprising slender axes without laminae. The fertile pinnules were tightly enrolled in this genus and there was little sterile tissue associated with the clusters of sporangia. The specimen discovered in the Potomac Group and described here exhibits the form of pinnule identical to the generitype. It is identified with the genus even though details of the morphology of the frond are not known for the new material. No other ferns of this age have been described with these very tightly revolute fertile pinnules. The spores of both species belong to the dispersed spore genus Cicatricosisporites. Those of Pelletixia valdensis have been well described and identified with C. brevilaesuratus (Hughes & Moody- Stuart, 1966). The spores of P. amelguita are closest in details of size and morphology to the dispersed spore species Cicatricosisporites aralica (Bolkhovitina) Brenner (see Brenner, 1963, Plate 7, fig. 4, 5; Phillips & Felix, 1971, Plate I, fig. 16). Brenner commented that these spores are similar to those in the fossil fern Ruffordia goeppertii Seward and that they occur commonly in Zone I of the Potomac AMERICAN FERN JOURNAL: VOLUME 72 (1982) FIGS, 3- iegaie Vee is details of the holotype of Pelletixia mola itl FIG. 3. Whole pinnule before breaking 3 apart, 54. Spores can be seen in upper center. FI a of spores enlarged. Remnant of sporangial wail and annulus can be seen at top center of sihod oie x 200. FIG. 5. Area of spores after further breaking of specimen. Approximately 120 spores can be counted, x 200. J. E. SKOG: PELLETIXIA AMELGUITA 119 Group and are very common in the Wealden of England. He noted that similar spores are produced by extant species of Anemia. Both Hughes & Moody-Stuart (1966) and Watson (1969) commented that C. brevilaesuratus, the spore of Pelletixia valdensis, is comparable to the spores of Ruffordia goeppertii. The spores of P. amelguita are smaller than those of P. valdensis, actually about half the size, and the ridges are widely spaced, whereas in P. valdensis the ridges are quite close together. The spores of P. ame/guita are certainly more similar to those of some specimens of Ruffordia goeppertii in size and spacing of muri, especially as described by Watson for specimen V 2192 of R. goeppertii. However, no specimen of R. goeppertii has been described with the tightly revolute pinnules of Pelletixia. hus, while the newly described pinnules are placed in the genus Pelletixia, | suggest that the genus may actually represent a stage in the development of the fertile frond. The open (unrolled), dispersing stage of the pinnules has been described and identified as a specimen of Ruffordia goeppertii. There is support for this interpretation within the modern genus Anemia. In modern affinities the fossil plants most closely resemble the genus Anemia, particularly plants found in the subgenus Anemiorhiza. Anemia aurita Swartz (endemic to Jamaica) has fertile pinnules that are at first tightly rolled and then unroll somewhat to release the spores (Mickel, 1981). The spores of this species and others within the subgenus resemble those of Pelletixia and also Ruffordia. Mickel (1981) considers this subgenus to be the most primitive of the genus. The interpretation of the habitat of Pelletixia has been uncertain. Seward (1913) suggested it was possibly aquatic. Hughes and Moody-Stuart distinguished it from Ceratopteris, but suggested exploring the possibility that it was aquatic and not related to the rest of the family Schizaeaceae. Watson (1969) said that “the form of the fertile fronds cannot be matched” within the family Schizaeaceae and compared the structure of them to Onoclea sensibilis L. | suggest that there are comparable fertile pinnules within the Schizaeaceae, especially in tropical species of the genus Anemia, and that following further study, Pelletixia, Ruffordia, and Anemia may all prove to be congeneric. The author acknowledges the aid of Mr. Howard Hruschka of College Park, MD, for collecting the matrix material; Mrs. Effie Shaw, Palynology Laboratory, U.S. Geological Survey, Reston, VA, for SEM assistance; and Mr. Jan Endlich, Depart- ment of Biology, George Mason Univ., for preparing photographs. LITERATURE CITED BRENNER. G. J. 1963. The spores aes hae of the Potomac Group of Maryland. Maryland Dept. eol. Mines Water Res. Bu DOYLE, J. A. 1969. Cretaceous La ee ol of the Atlantic coastal plain and its evolutionary cutee J. Arnold Arbor. 50:1—35 HUEBER, F. 1982. Megaspores and a alynomorh from the Lower Potomac Group in Virginia. See Contr. Paleobiol. 49:1—69. HUGHES, N. F. and J. MOODY-STU ee 1966. Descriptions of Schizaeaceous spores taken from early Cretaceous macrofossils. casa 9:274-289 MICKEL, J. T. 1981. Revision - ie subgenus Anemiorhiza (Schizaeaceae). Brittonia 33:413-429, PHILLIPS, P.. P. and C: J. FE 18th. A study of lower and middle Poh eareg spores and pollen from the southeastern set States. I. Spores. Pollen Spores 13:279-34 AMERICAN FERN JOURNAL: VOLUME 72 (1982) FIGS. 6-9. Spores of the holoty ‘. of Ai lixia seed) Gon FIG. 6. Detail of spores cae ee ha . Proximal face of spore, x 2200. e that lips i ‘Shoe in teeth showing d ak face with may be seen at edges of photograph, x 940. FIG. face with trilete mark and raised | join with muri and extend to the usta: is irregular polar triangle. Possible remnants of cobs 9. Isolated spore with polar triangle and detail of muri on distal hae X 2200. J. £. SKOG: PELLETIXIA AMELGUITA 121 FIGS. 10-12. Spores of the a ae of Pelletixia amelguita. FIG. 10. Cluster of spores showing variability of polar triangle, x 618. . 11. Spore with tripartite polar ogee (one section occluded), * 1854. FIG. 12. Spore with ie oe a in the polar triangle, SEWARD, nee “sg Contributions to our knowledge of Wealden floras. Quart. J Geol. Soc. London —1] SKOG, J. : "ore "Loxsomoptrs anasilla, a new fossil fern rhizome from the Cretaceous of Maryland 14. Amer. Fern J. WATSON, J. 1969. A revision of the English Wealden flora, I. Charales-Ginkgoales. Bull. Brit. Mus owe Hist.) Geol. 17:207—254. es and C. R. HILL. 1982. Pelletixia: a new name for Pelletieria Seward (fossil). Taxon 31:553 122 AMERICAN FERN JOURNAL: VOLUME 72 (1982) AMERICAN FERN JOURNAL Manuscripts submitted to the JOURNAL are reviewed for scientific content by one or more of t editors, and, often, by one or more outside reviewers as well. During the past year we have received ie kind assistance of C. Haufler, H. T. Horner, Jr., F. M. Hueber, E. E. Karrfalt, J. D. Montgomery, D. J. Paolillo, H. E. Robinson, J. E. Skog, R. G. Stolze, R. A. White, and D. P. Whittier, to whom we are deeply indebted. We welcome suggestions of other reviewers.—D.B.L. INDEX TO VOLUME 72 Abacopteris truncatum, 56 Bathmium ree 57; villosum, 57 Acrostichum, 14; alatum, 56; angustifolium, 56; attenuatum, 56; Beck, M. J. of oe ponetti pharodes, 56; callolepis, 56; cochleariaefolium, 56; crispatu- eitel, J of Wagner. The chromosomes of Lycopodium lum, 5 , 13; dom um, 57; erythrolepis, 57 lucidulum, 33 gratum, 57; hartwegii, 57; herminieri, 57; int dium, 57; Bentley, S. (see T. F. Wieboldt) foncophyllum, 57; micradenium, 57; minutum, 57; obcrenatum, Bir, S. S. Aspects of me sciences, vol. 3. pteridophytes (rev.), 52 lu , 57; rhabdolepis, 57; schlimense, 57; stigmato- Blechnum exten 57; helveolum, 57; heterocarpum, 57; s, 57; stramineum, 57; tenuiculu schlimense, 51; Semings udum, Lee Adiantum capillus-veneris, 77; claussenii, 57; nsum, 57; A pterid 1 h in Ch 1 fuligi , 57; gracile, 57; jacobinae, 57; pe icense, 57; Britton, D. M. (see E $ Kott) igrescens, 57; obliquetruncatum, 57; ovalescens vifolium, Brooks, R. E. A new Woodsia hybrid from Kansas, 79 57; pedatum, 46; pseudocapillus, 57; subaristatum, 57; tomentel- Camptosorus, 67, 69, 71; rhizophyllus, 9, 65, 67, 71, 73, 101 bm, oda j rapeao oudes, ot trict seg 57 Campyloneurum caudatum, 57; cubense, 57; jamesoni, 57; morit- ice), 29 zianum, 57; xalapense, 5 , M. Whitten & M. J. Beck. Axenic culture and poe ila schaffnerian Sh An re simp medium for growing prothallia, 30 oo lat Rg A an rer “ournal, 122 induction of callus and sporophytes of the Appalachian Vittaria gametophyte, 36 —— alata, 57; ampla, 57; confluens, 57; ion p 1 Cyatt . Il. Metaxya and crophylla var. crenata. 57, var. distans, 57; pilosa, Pe sinuosa vr Lanta 19 Anemia, 119; adiantifolia, 100; subg. Anemiorhiza, 119; aurita, Cassebeera — 57; petiolata, 57 119 Cormopiers see — ges 109; chaerophylloides, 109; lurida, 109; , 103 bescens, 109 Cessna in the Asplenium adiantum-nigrum complex, pee? of ip a Ba 3: oe (rev.), 5 bag acrocarpo agatolepis, 57; ame ae ron, 57; chattel alabamensis, 77; aspidioides, 57; castanea, rulum, 57; cin, 57; chrysolepis, 57; dissectum, 57; cucullans, 57; feei, 76-78; glaberrima, 57; griffithiana, a fn ndulosum, 57; germani, 57; jucundum, 57; mierothecium, =e _, lanosa, ai, 46, 70; ‘Mialaccensis, 57 peagtorcscs. Mag _postchiays 37: pauper, 57; trichophorum, ja, 31 57; t hing, .& ZH. ang. A brief pate on grt progress of Asweease 48. 68, 103, 105; abscissum, 101; adiantum-nigrum, ica nee research in China, | 103—105; pride 9; balearicum, 103, 105, 106; bifissum, The chromosomes of aa ee lucidulum, 33 57; billotii, 105; bradleyi, 103; brevisorum, 112; brunneo-viride, Chrysodium Bes ROY 113; cladolepton, 57; Xclermontae, 9; clutei, 110; coriaceum, Chrysopteris martinice ye microdictya, 57 57; cuneifolium, 103-105, cress 101; debile, 57; depau- pcanaoiara er es, ori ae 119; Ba ie lvls 119 peratum, 57; 57; duale, var. duale, 109; Cnem extensum, 57; flabellatum var. tripinnatum, 110; flageliferum, Coc hlidium minus, 113; seminudum, 113 57; foreziense, 105; galeottii, 57; gilbertii, 110, 111; hendersonii, omparison of ue ane spot patterns of some North Amer- , 69; herbaceum, 57; inaequalidens, 57; integrum, 57; ican Isoétes species, 15 jamaicense, 110; lamprocaulon, 57; leptophyllum, 57; macrodon, Craspedaria borbonica, 57; gestasiana, 57; javanensis, 58; lanceolata, 57, marinum, 67; montanum, 73, 103-105; myapteron, 57; 38 myriophyllum, 110, var. brevisorum, 110, var. myriophyllum, Cryptosorus dionaea, 58 110; neo-granatense, 57; atum, a ; onopteris, Culcita, 19; schlimensis, 58 103-105; pinnatifidum, 103; platyneuron, 9, 62, 65, 70-74; Cyathea arborea, 109; armata, 109; aspera, 109; bernard dii, progrediens, 57; pumilum var. hy lloides, 57; radicans var. 108, 109; xcalolepis, 109; commersoniana, 58; lewisii, 108, pallidum, 112, var. tripinnatum, 110; rutamuraria, 9, 77, 103, 109; suprastrigosa, 23 var. cryptolepis, 46; sciadophilum, 110; septentrionale, 77, 103; Cyatheaceae, 14, 19, 21, 23, 25-27, 108 Penren 57; tenellum, 57; ternatum, 57; tri es, 5-10, Cystodium, 19; sorbifolium, 23 70, 72, 74, 103, subsp. _ ivalens, 5,7, ” Dep. ag Cystopteris, 31, 41, 43, 52, 93, 94; azorica, ni nee 41, 43, asp. prrouoms, t 3, 94; fragilis, 31, var. fragilis, 41, v ackayii, 41, 43; jamaicense, 110; Xv sree a le, 9 x illinoensis, 41-43; laurentiana, ‘41: aaa “31, 41, 93, 94; he Avec trichomanes pele: in the United States and tennesseensis, 41, 93, 94 adjacent Canada Cystopteris tennesseensis in oe 93 x Asplenocetera Cystopteris illinoensis: a new natural hybrid fern, 4 x Asplenophyllitis, 67, 69 Danaea carillensis, ge a; 25 64; jenmanii, aw wendlandii, 63 Asplenosoru ; boydstonae, 62, 71-74; ebenoides, 62, Darling, T. J., Jr. The deletion of Seaesene pectinata from the 65-67, 69-74, 101: herb-wagneri, 9; fe: 95-73; flora of Florida, He shawneensis, 9, 10; trudellii, 73 Davallia in cultivation [Davalli ] (rev.), 40 Athyrium, 5 Davalliaceae, 40 Axenic ees and induction of er" and sporophytes of the Death Notices: Dieter E. Meyer (July 21, 1926—February 1982). 95; Aloysio Sehnem (1912-1981), 29 palachian Vita gametophyte, The deletion of Nephrolepis pectinata from the flora of Florida, 63 Azolla anni INDEX TO VOLUME 72 Diphasiastrum, 35 sseocbea: incisa, 58; neglecta, 58: stenochlaena, 58 Dicksoniaceae, 14, 19 Didymo ea lossum fructuosum, 58; krausii var. subpinnatifida, 58 Diellia, Dieter E. Meyer (July 21, 1926—February 1982) ew. ental 95 Diplazium, 52. acutale, 58; anthraxacolepis, 58; bre 111: mptocarpon, 58; dissimil ; expansum, 112 ; ee 38; firmum, 58; fuertesii, 112: grammatoides, 58: rm) nopodium, 58; mohillense, 58; mont salut locae: LEE; Giesiehcarerunaies. Hcl: Praelon gum, 58; schlimense, 58: sonii var. brunneoviride, | 13 peoreiars ae var. ie atula, 58 Drymogl 58; subcordatum, 58 Drynaria acuminata, 58: compacta, 58: crassinervata, 58: elastica, 58; mexicana, 58. pinnata. 58; prieurei. 58: stenoloma, 58: rium, 52 Dryopteris, 3, 11, 70, 79; celsa, 46; goldiana, 46, 101, subsp. celsa, 46; intermedia, 101; uae: 11; lurida, 109; marginalis, 70: tokyoensis, Ecological, biosystematic, and nomenclatural notes on Scott's Spleenwort, id Eriosorus ruizianus 58 e fern mm genus T). Wh Iti th [D hh ] 40 Ferns and | — ng of G part II. Polypodi ( 14 A filmy D The first re aot Platycerium ridleyi in Sumatera, 12 Flavonoid phe of the North American Lycopodium obscurum complex Flora of Chiapas, part 2 pags (rev.). 44 Franke M. C. Roos. The first record of Platycerium ridleyi i in Sumatera. a Fusiak, F. F istry of the North American Lycopodium obscurum com . 96 - genus Po ack um in pha ii dongs ogee (rev.). 84 A germination method for Isoétes Gleicheniaceae ay me ilmy Danaea, a Grammitis succinea, the first or und in ambi Goniophlebium acuminatum, 5 otiieie 114; anisomeron, 58; arcuatum alaguala, 58: chrondrocheilon, 58: coriaceum, 58; gauthieri, 58 qguesantabe m, 58: in ns, 58: lepidotrichum, is pectinans, 58: plectolepis, 58: ferax, 58: elon 58; leptocladia, 58: ' macroclada, $ 8; ensis, 58: mollis, ro 58: pyramida' Pose! suadanglars 58: repanda, a, 58; stri pe 8: tenera, 58 Peon ie {o0os Grammitis, 36, 39, * sect. succinea, 49-5 Gramimaitis succinea, the first New emu fern found in amber, 49 , 58: cumingianum, 58; macrosorum. 58 Hemistegia contracta, 58; pei 58: insignis, 58: lucida. ; repanda, 58 Hevardia Pagid me Hi a =e tee t, Jr. New records and distributional means! secon ytes, 45 fanart B. J. Davallia relatives in cultivation (rev.). 8 The fern genus Po ium in ee sie aiareacatt: (rev. Huperzia lucidula, 33; selago, ubsp. lucidula, 33. selago, 33 4 Cryptosorus, 51; seminuda, 113; — 4 subsp. Hymen Il Hypodematium, 5 Hypolepis bogotensis, 113, 114; chilensis, 59: fimbriata, 113: helenensis, 59: " maicensis, 113, 114; parviloba, 59: repens. 114; stuebelii, 1 oe of the re of Japan, vol. 1 (rev.), 11, vol. 2 .4 nv 15-17, 61; acadiensis, 15, 16; delilei, 15: durieui, 15: 123 eatonii, 15-17; echinospora, 15-17: engelmannii, 15-17, 61; Satan - agi me mea 16, 17; riparia, 15, 17; tuckermanii, 15, 16; v Jamesonia rund, tt, L. S. & D. M. Britton. priate of some Sica ace spot patterns ts some No n Isoétes species, Kurata, ‘9 & T. oe last “of the pteridophytes peat vol. | (rev.), v.), 48 i Lei, T. T. (see M. J. Waterway) Leptochilus subquinquefidus, 59; thwaitesianus, 59: a benatiste 59 Lindsaya curvans, 59; consanguinea, 59; crenulata, 59; elegans, 59; galeottii, 59; I"herminieri, 59; montana, 59; gubthioadandne, 59; parvula, 59 Litobrochia affinis, 59; brevin nervis, 59. galeottii, 59; grandis. rh na, it microdi 5 montbrisonis, 59; organensis, 59; setifera, 59: tussacii, 59 scarring decile: 59; erythrodes, 59. Sr iarass, 59: sorbi- folia var. caudata, 59 Lonchitis praca 59; tomentosa, 59 Lophosoria, 19, 21, 23, 25-27; acaulis, = 59; prostrata, 59: "7 Lore a see P. M. Richardson) Louisiana ae and fem allies (rev.), 18 115 crn peri of the neotropical Cyatheaceae. Lucansky, T. W. Ani Il. Metaxya and Lophos: Lycopodiaceae, 53, 107 Lycopodiella, 3 copodium, 35, 96; alpinum, 86; amentaceum, 53-55: ages folium, 53, 55; carolinianum, 35; cernuum, 35; clavatum, 45. cuneifolium 5; dendroideum, 96 ostachys, 59; ‘8 terianum, 33 urideum var. ser , 107; lucidulum at montanum, 10 Si obsc var, obscurum, 96; iim, 33: selago, pontine 33; tristachyum, 45 tum, 62 arsile Mattenccia, sie iit 47 Mertensi dis, 5 etaxya, 19, 21, 2. bagioiele rostrata, 19, or 23527 Microlepi Jat ef "59: mee fus Microsorium rabies 48: ae erianum, oo longissimum, 59 Monogramme a nor, 113 Moran, R. C. eaten cane conipnen in the ses States and Bhai Can natural hybrid fern, 41; Cy. i — M. C. Polystichum munitum on Baranof Island, southeast- ska, naar cheiloglyphys. 59; gracilis, 59: marsupianthes, 59; rufa. Nakaike, T. (see S. Kura The name of a hybrid oe ON 62 W. H. Wagner, Jr.) Nephrolepsis, 63; exaltata, 63: pectinata, 63: schkhuri ~~. apes 59 New 45 A new station for nde east 62 a Woodsia hybrid fro nsas, A new rp American err of the genus Lycopodium, 53 Niphidi Noto oo osia, 59; sinuata 7 Ollga pa A new tropical A sp f the g Lycopo- ium, Onoclea Sw Onychium, 52: ae. 59; multifidum, 59 Oochlamys rivoi = - Ophioglossace Ophigolossum Seta 45 124 46 Osmundaceae 14 Ott, F. D. (see A. J. Petrik-Ott) Pellaea emi ce 77, glabella, 77; mucronata, 59; palmescens, ; we Pellet HT: vans 117, 119 i amelguita, | 15—121; valdensis, 117 Pelletixia nang new species of fossil fern in the Potomac Group (Lowe: Crease), 115 Petrik-Ott, A rah . Ott. Cheilanthes lanosa and Cystopteris vis, 59; connectilis, sn cordat a, 59. cteno ides, 59; delicat dilatata, 59: elata, 59; ‘luminensis, 59; germaniana, 59; helliana, 9, impressa 50: inaequalis, 59; lanata, 59 toptera, 59 cuss — n saute = mollivillosa, 59; montbri- na, 59; 59; oreopteridastrum, 59; sitdeos ratchitornis "59; ‘rustica . 59; sca! _— 59; scrobiculata, a stenolepis, 59; straminea, ‘59: villosa, 59 Phlebiogonium impressum, 59 Phyllitis, 1, 67, 69: sc circle 67, 103 Plagiogyriaceae, 14 et = 13; biforme var. erectum, 12; coronarium, 12, 13; 13 Mbt stl, 59; peruvianus, 59 Pleope sina revoluta, Pleur Polybe ace toedbiboia: 59; scandens, 59; semipinnata, 59 Polypodiaceae, 14, 50-52, 109 Polypodium, 14, 48; albopunctatissimum, 84; ampliatum, 114: arthr i astrolepis, 84; australe, 60; blandum, 60; brasiliense var. gladiatum, 114; callolepis, 60; cam ‘on, 60; cancell chaerophylloides, 109; cheilostictum, 60 crypt n, 60: echinolepis, 60; ellipsoideum, 60: filipendulaefolium, 60; funiculu ; glycirrhiza, 60; i 60; in ides, 60; leptostomum, 60: rosorum 60; nivosum, 60; oulolepis, 60; polypodioides, 70: pubescens 09; re 84; aneerng ns 60; — eri, 60, var. a ‘ulum, 60; triseriale tum, crispum, 60; serricula, 60; senile, 60; tenu r. gl , 114 folyihiconee aigitients, 109; chaerophylloides, 109; lurida, 109; ochropteroides, 109; pubescens, 109 Polystichum, 11,70; acrostichoides, 70, 101; bra 101; chlaenosticta, 60; UE si 60; guadalupense, 60; haem ; ilicifolium, 60; incisum, 60; lepidomanes, 60; lonchitis, 85, 86; munitum, 30; ‘chisel 60; schizolobium, 60; viviparum Polystichum lonchitis in central Quebec—Labrador, 85 Polystichum munitum on Baranof Island, southeastern Alaska, 30 Polytaenium intramarginale, 114; lineatum var. intramarginale, Proctor, G. R. Taxonomic notes on Jamaican ferns-IV, 107 ae sg hamensis, - croesoides, ham, ii, pores gracilis, teromorpha, 60: longifolia, 101; lon, 60; muricella, 60; nsis, 60; oppositi-pinnata, 60; paucinervata 60; philippinensis, 60; prionitis, as i goer 60; regia, 114; ‘ . ttata, 97, 99-10 Pteris x delchampsii, a spontaneous. ss hybrid from southern Reviews: Aspects of plant sciences, vol. 3. Pteridophytes, 52; ER Page 70, line 11: For “>” read “ ER AMERICAN FERN JOURNAL: VOLUME 72 (1982) Davallia relatives in cultivation, 40; The fern genus Davallia in cultivation, 40; Ferns and fern allies of Guatemala, ii rt Il, 14; — of Chiapas, part 2. Pteridophytes, oe The genus Polypo- in cultivation — _ stations of sie oe of Japan, vol. 1, fern allies, 18 eavagpeans P. M. - = Lorenz-Liburnau. C-glycosylxanthones in 103 nienim ompl ex ahaa R. mare & aoe R. Hill) Roos, M. C. (see N. a . Franken) Rufforida, 119; goeppe' ist Sagenia gemmifera, aie mexicana, 60 Sam, S. J. A germination ar ery for Isoétes, 61 Schizeeaceae, 14, 117, 119 Schizoloma javae, 60 92; ser. Articulatai tae, 88, 92: atirrensis, 90, 91; na, 89; parkeri, a new — of fossil fern in the — us), 115 eridophytes (re v.), ha: Fée’s Lae sea collection at the botanical garden of Rio < Janeiro, 56 Sphenomeris clava aren : Stolze, R. G. Fach nt fern allies of Guatemala, part II. dose gener (rev.), Stupka, A. & A. J. sap A new station for Lygodium palmatum, 62 Synochlamys ambigua, 60 Taxon notes on Jamaican ferns—IV, 107 100 Thieret, J. W. ‘Lou uisiana ferns and fern allies (rev.), 18 Thier Sega clgom 23 Trichomanes, 36, 39, 70; owt 70; subg. ee ie 107; mic seats ubescens, 108; radicans var. antil 08 A unique type of ee ciate in Selaginella ser. Tecate: — 36, 38, 39; amboinensis, 60; filifolia, or —s ana, 60; sigalg 114; sh — 60; line: ata, a ee minor, ariensis, 60; , 60; ruiz . sarmentosa, 60; ‘a, 60; zeyl anica, one Peg F. S. (see J. M. Beitel; ates F. Walter) Wa: Hn 7 Gai Nauman, a Preis x delchampsii, a 97 + spontaneous fe hvhrid f; Wagner, W. H., r) Walter, : S. name of a hybrid rd fe piagsise Walter, W. H. Wagner, Jr. & F. S. Wagner. pon bossa aire nomenclatural notes on Scott's Spleenwort, ts Wang, z -H. “re Chi aN Rs ve se Polystichum lonchitis in central 85 Whitten, M. (see J. D. Caponetti) Wieboldt, : - & s. Bentley. ee feei new to Virginia, 76 Windisc . Specimens from Fée’s pteridological fi st at the acid garden of Rio de Janeiro, 5 sia, 79, 82, ~ alpina, 79; ethan, 79; aa 19% ilvensis, 47, 79; 0-83: a, 60; oregana, 79, 81-83, var. co. 79, 81, 84; pre "39, “gi- 84, ean Woodwardia, 14; virginica, 77 ee FOR VOLUME 71 1981 RATUM FOR VOLUME 82 1982 Page 27, line 15 should read: * . genera in the Dicksoniaceae, and concluded that both genera are more dicksonioid than cyatheoid on the basis. BRITISH PTERIDOLOGICAL SOCIETY Open to all who are interested in growing and studying ferns and fern-allies. 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Conant, President TRIARCH PREPARED MICROSCOPE SLIDES .O. Box 98 Ripon, Wisconsin 54971 hed Plants With Special Reference to Tropical America ROLLA M. TRYON and ALICE F. TRYON, Harvard University MORE THAN 850 PAGES This exceptional volume is the first modern, in-depth treatment of ferns and related plants, particularly in tropical America. Written by Rolla M. and Alice F. Tryon, well known for their work in this area, FERNS AND ALLIED PLANTS presents a new classification of the Pteriodophyta based on a wealth of data published during the last few decades as well as the Tryons’ own wide-ranging research. Extensive bibliographies of modern literature on fern biology are provided. In addition to tropical American genera, this volume also has extensive information on ferns of temperate an paleotropical regions, making it of worldwide perspective. 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