THE HECKMAN BINDERY, INC. N. Sidecreeren. INDIANA

AMERICAN Volume 68- FERN 1978—|97 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

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CONTENTS

Volume 68, Number 1, Pages 1-32, Issued April 12, 1978

Rediscovery, Distribution and Phytogeographic Affinities of Leptogramma pilosa in Alabama JOHN W. SHORT and JOHN D. FREEMAN

Chromosome Numbers in the Fern Genus Anogramma UDITH G. Poe isis and GERALD J. GASTONY

Curtis Eugene Delchamps (1925-1977) Geographical Distribution of Isoetes butleri in the

Southeastern United States JERRY M. BASKIN and CAROL C. BASKIN Thelypteris oroniensis, a New Species from Costa Rica LUIS D. GOMEZ P. Additions and Corrections to the Pteridophyte Flora

of Chihuahua, Mexico IRVING W. KNOBLOCH and DONOVAN S. CORRELL The Distribution and Chemical Constituents of the

Farinose Exudates in Gymnogrammoid Ferns ECKHARD WOLLENWEBER

Shorter Notes: The Selaginella apoda Complex in Iowa; Polystichum lonchitis Found in the Black Hills; On the Distribution of Lycopodium flabelliforme in Illinois

Review

Volume 68, Number 2, Pages 33-64, Issued July 11, 1978 The Taxonomic Status of Selaginella eatonii WILLIAM R. BUCK

Microreplicas as a Technique for Rapid Evaluation of Surface Silica Micromorphology in Equisetum RICHARD L. HAUKE

The Establishment of Bracken Following Fire in Tropical Habitats STEPHEN R. GLIESSMAN The Distribution and Ecology of Dryopteris in Southeastern Virginia and Adjacent North Carolina DANIEL L. NICKRENT, LYTTON J. MUSSELMAN, LAURA A. PITCHFORD, and DAVID W. SAMPSON The Fine Structure of the Newly Formed S$ of Onoclea sensibilis NORMAN P. MARENGO and MARIE A. BADALAMENTE

The Anatomy of Equisetum diffusum Tubers S.S. BIR A New Species of Asplenium from Guatemala ROBERT G. STOLZE

Shorter Notes: A New Location for Pellaea glabella in Minnesota; Some Insect Interactions with Azolla mexicana; Notes on North American Lower Vascular Plants; The Ferns of San Salvador Island, II; Cheilanthes microphylla, a Genus and Species New to the Bahama Archipelago; The Chromosome Number of Notholaena cochisensis

Review

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Volume 68, Number 3, Pages 65-96, Issued October 2, 1978

Spread of the Exotic Fern Lygodium microphyllum in Florida CLIFTON E. NAUMAN and DANIEL F. AUSTIN

Chlorophyll and Lipid Changes on Germination in the Non-green Spores of Thelypteris dentata ALLEN V. SEILHEIMER

Gametophytes of Botrychium multifidum as Grown Axenic Culture ERNEST M. GIFFORD, JR. and DOROTHY D. BRANDON

Revision of the Genus Cochlidium (Grammitidaceae) L. EARL BISHOP

Shorter Notes: A Deletion from the ee Flora of Nebraska; dm Lerosiaeimgon in Alabam litorale Recorded for Sesisastae some cernuum in Louisiana

Volume 68, Number 4, Pages 97-124, Issued December 29, 1978

Trichomanes Gametophytes in Massachusetts BRUCE MCcALPIN and DONALD R. FARRAR

A New and Unique, Mat-forming Merlin’s-grass (Isoétes) from Georgia PHILLIP M. RURY

State and Local Fern Floras of the United States,

Supplement II MERYL A. MIASEK

Shorter Notes: Athyrium filix-femina New to Saskatchewan; New Combinations in the Fern Flora of Venezuela; Trismeria. . .trifoliata?

Reviews American Fern Journal Index to Volume 68

Erratum

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AMERICAN FERN JOURNAL

Volume 68 Number 1

January-March, 1978

QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY

Rediscovery, Distribution and Phytogeographic Affinities of JOHN W. SHORT and JOHN D. FREEMAN!

Leptogramma pilosa in Alabama

Chromosome Numbers in the Fern Genus Anogramm

UDITH G. opie and GERALD J. GASTONY

Curtis Eugene Delchamps (1925-1977)

Geographical Distribution abe psa butleri in the Southeastern United Sta’ J

Thelypteris oroniensis, a New Species from Costa Rica Additions and Corrections to the Pteridophyte Flora of Chihuahua, Mexico

The Distribution and Chemical Contituents of the Farinose Exudates in Gymnogrammoid Ferns Shorter Notes: The Selaginella apoda Complex in Iowa; Polystichum lonchitis Found in the Black Hills;

On the Distribution of Lycopodium flabelliforme in Illinois

Review

Missoup; Boranican,

APR 95 1978

GS ; ARDEN LIBRARY

ERRY M. BASKIN and CAROL C. BASKIN

IRVING W. KNOBLOCH and DONOVAN S. CORRELL 1

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LUIS D. GOMEZ P.

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ECKHARD WOLLENWEBER |!

Missouri FPotaNnical

APR 5 1908

GARDEN Li RARE

The American Fern Society Council for 1978 RICHARD L. HAUKE, Dept. of Botany, University of Rhode Island, Kingston, R.I. 02881. President ROBERT M. LLOYD, Dept. of Botany, Ohio University, Athens, Ohio 45701. Vice-President TERRY R. WEBSTER, Dept. of Botany, University of Connecticut, Storrs, Conn. : Secretary JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916. Treasurer JUDITH E. SKOG, Dept. of Biology, George Mason University, Fairfax, Va. 22030. ecords Treasurer

DAVID B. LELLINGER, Smithsonian Institution, Washington, D.C. 20560. Editor-in-Chief JOHN T. MICKEL, New York Botanical Garden, Bronx, N.Y. 10458. Newsletter Editor American Fern Journal EDITOR-IN-CHIEF DAVID B. LELLINGER Smithsonian Institution, Washington, D. C. 20560

ASSOCIATE EDITORS DAVID W. BIERHORST ..Dept. of Botany, University of Massachusetts, Amherst, Mass. 01002 GERALD J. GASTONY ............. Dept. of Biology, Indiana University, Bloomington, Ind. 47401 JOHN T. MICKEL New York Botanical Garden, Bronx, New York 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 Institu- tion, Washington, DC 20560. Second-class postage paid at Washington.

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Library New York Botanical Garden, Bronx, New York 10458, is Librarian, Members any time, the borrower paying all shipping costs. ms Newsletter ar — ~— k Botanical Garden, Bronx, New York 10458, is editor of the SNES eare® ad Forum.” The editor welcomes contributions from members and non- icul : ” mecemnenes notes, offers to exchange or purchase materials personalia, hor- ucultural notes, and reviews of non-technical books on ferns. , a aca Spore Exchange F. Neil D. Hall, 1230 Northeast 88th Street, Seattle. Washi es exchanged and collection lists sent on request. ; ee ee ee

Gifts and Bequests

Dr. John T. Mickel, may borrow books at

Gifts wo 4.

in ferns. Botanical books back iaiee of the ts and ta athers interested fe Journal, and cash or other gifts are al Icomed, and are tax-deductible. Inquiries should be addressed to the Secretary. gifts are always welco

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 1 (1978) l

Rediscovery, Distribution and Phytogeographic Affinities of Leptogramma pilosa in Alabama JOHN W. SHORT and JOHN D. FREEMAN*

Leptogramma pilosa var. alabamensis (Crawford) Wherry was originally de- scribed as Thelypteris pilosa var. alabamensis, based on material collected in Winston County, Alabama (Crawford, 1951). The type locality was stated to be a sandstone cliff on the ‘‘West Fork of the Sipsey River’’ five miles east of Double Springs (presumably where U.S. Highway 278 crosses Sipsey Fork'), at 400 m

FIG. 1. Plants nip: aS pilosa var. alabamensis.

(1300 ft) elevation. Crawford identified two additional collections from the Mexi- can states of Chihuahua and Sonora as var. alabamensis, whereas the typical variety previously had been known from Chihuahua and central and southern Mexico and Guatemala. The species has not been reported until now from any locality in the United States other than the one cited by Crawford.

*Department of Botany and Microbiology, School of Agriculture and Agricultural Experiment Sta- tion, Auburn University, Auburn, AL 36830.

'The nomenclatur es fa! at the collecting site is confused. On recent highway and tie maps, it is called ‘Sines Fork.’’ Co lloquially it is 9 the ‘‘Sipsey River” or . A Sips

The full and correct ae perhaps should be ‘*Sipsey Fork of the ray Warrior Riv

Volume 67, number 4, of the JOURNAL was bite December 31,

2 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

In 1960, when the L. M. Smith Dam was completed several miles downstream from the type locality, the higher water level of Smith Lake in the Sipsey gorge necessitated construction of a new bridge for U.S. Highway 278. This construc- tion leveled the cliff, and with the impoundment of Smith Lake completely de- stroyed the habitat in which L. pilosa var. alabamensis occurred. The species was assumed to have been lost from the Alabama (and, indeed, the U.S.) flora (Dean, 1969).

At a conference on rare and endangered species of Alabama held at Tuscaloosa in the spring of 1975, it was disclosed by Mrs. L. C. Smith (pers. comm. to J.D.F.) that L. pilosa had been observed somewhere along the Sipsey near the reported locality. Topographic maps revealed that a likely location would be the massive sandstone cliffs five miles north of Double Springs where Alabama Highway 33 crosses the river. Since no point in Winston County is more than 1,000 feet in elevation, it was clear that the elevation indicated by Crawford for the type locality had been in error.

In September 1975 and April 1976, several colonies of L. pilosa var. alabamen- sis were found near the Alabama Highway 33 bridge over Sipsey Fork, at the upper reaches of Smith Lake. This site is marked by an overhanging cliff rising more than 60 feet above the river. Leptogramma pilosa occurs in crevices in the north-facing cliff and on a smaller, west-facing cliff do am (Fig. 1). The thin, damp soil has been described as moderately acid (Wherry, 1964). The elevation of Smith Lake is about 500 feet; all colonies of L. pilosa observed were about 10 feet above water level and grew in close association with Trichomanes boschianum and various bryophytes. The type locality downstream was probably several feet lower. Other pteridophytes found nearby included: Osmunda regalis, Athyrium asplenioides, A. thelypteroides, Woodwardia areolata and Selaginella apoda.

Leptogramma pilosa still has not been found in the United States outside the gorge of Sipsey Fork. Besides the localities cited above, it has been seen in the Sipsey River Picnic Area near the Lawrence County line (R. Kral, pers. comm. to J.W.S., 1976). This gorge is over 1,200 miles from the nearest Mexican localities for the species. Located in the Cumberland Plateau just above the Fall Line, all major streams in the Sipsey Fork area have eroded narrow gorges with steep sides and many cliffs. These damp, cool gorges harbor a peculiar and unique as- semblage of plant species including several near endemics as well as disjunct populations of species with principal ranges elsewhere. The Hemlock-Hardwood Forest Association is well developed in the gorges, some 150 miles from the

else. Whether the disjunctions i tween floras or long range dispersals has not been determined. LITERATURE CITED CRAWFORD, L. C. 1951. A new fern for i ; : , the Unit 2 L- DEAN el nited States. Amer. Fern. J. 41: 15-20.

f Alabama, 2nd ed. Southern University Press, Birmingham, AL. xxiv +

222 pp. WHERRY, E. T. 1964. The Southern Fern Guide. Doubleday, Garden City, NY. 348 pp.

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 1 (1978) 3

Chromosome Numbers in the Fern Genus Anogramma, II. JUDITH G. BAROUTSIS* and GERALD J. GASTON Y**:!

In an earlier report on chromosome numbers in the genus Anogramma (Gas- tony & Baroutsis, 1975), three new counts were established and all known previ- ous counts were summarized. At that time, a count of n=26 for A. leptophylla from Europe (Kurita, 1971) was overlooked. We wish now to acknowledge this count, to report new counts made for populations of A. guatemalensis and A. leptophylla, and to suggest an explanation for some of the variant counts previ- ously reported for A. leptophylla.

The general techniques for chromosome preparations were those previously discussed (Gastony & Baroutsis, 1975). To maximize chromosome staining, how- ever, a propionic-iron-haemotoxylin stain (Henderson and Lu, 1968; Rigby, 1973) was applied to fixed mitotic cells of Anogramma gametophytes. The stained chromosomes were visually enhanced by use of phase microscopy in analysis and photographic work.

To promote spreading and separation of mitotic cells during squashing, material was treated with one of two preparations: Glusulase (Endo Laboratories Inc., Garden City, NY), a commercially available enzyme mixture from the intestinal juice of the snail Helix pomatia, was applied full strength to gametophyte tissue for four hours (Fabergé, 1945); Driselase (Kyowa Hakka Kogyo Co., Tokyo, Japan), a fungal-produced enzyme mixture, was applied as a 10% (w/v) aqueous solution according to the Glusulase schedule. Both preparations were equally satisfactory for softening cell walls. The potential of this enzyme technique in working with gametophyte chromosomes has been more fully discussed by Gas- tony (1977).

Chromosome counts for A. guatemalensis, published here for the first time, are based on three unequivocal counts. The counts reported for A. leptophylla, how- ever, are based only on the material illustrated in Figs. 3, 4, 8, and 9.

Sources of spores cultured to provide living material of the taxa herein reported are: A. guatemalensis, Gastony 1037, Depto. Chimaltenango, Guatemala; A. leptophylla, 7 Oct 1972, Mickel, Edo. Oaxaca, Mexico; A. leptophylla, 20 Oct 1972, Esterhuysen, Cape Province, South Africa. Voucher specimens of the plants raised from spores are deposited at IND.

RESULTS AND DISCUSSION

Mitotic squashes from gametophytes of A. guatemalensis and A. leptophylla

show a chromosome number of n=29 (Figs. 1-7). Chromosomes of both these

species have a strong tendency to stick together, particularly at their ends. This stickiness, in conjunction with the form of two of the chromosomes as discussed

*Department of Biology, Augusta College, Augusta, GA 30904. **Department of Biology, Indiana University, Bloomington, IN 47401. _ im ‘We are most grateful to E. A. Schelpe, E. Esterhuysen, and J. T. Mickel for providing us with materials used in this study.

AMERICAN FERN JOURNAL: VOLUME 68 (1978)

BAROUTSIS & GASTONY: CHROMOSOME NUMBERS IN ANOGRAMMA, II 5

below, may explain the variability in the counts that have been reported for Mexican A. leptophylla (Mickel et al., 1966) and in at least some of the other previously reported counts which are at variance with a base number of 29.

An analysis of chromosome morphology in the course of this work has proven useful in ascertaining the actual chromosome numbers present and in determining the source of variability often encountered in Anogramma squashes. From ob- serving numerous cells, it is known that both A. /eptophylla and A. guatemalensis have one very thin chromosome (Figs. /, 3, 5, and 6) and a short chromosome that frequently appears as a knob at the end of another chromosome (Figs. /, 3, and 6). Prior to squashing, and often in squashed preparations, the short chromosome looks like a satellite. After squashing, however, it often lies at right angles to the chromosome with which it is associated (Figs. 3 and 6) or across this chromosome (Fig. 1). The thin chromosome also seems always to be associated with another chromosome, but in several cases it has been found completely free. The reason for these chromosomal associations is unknown. In at least half (ca. 10) of the cells examined, one or both of these chromosomes is not evident, and when counted, these cells appear to have 27 or 28 chromosomes.

As in mitotic preparations, meiotic squashes also reveal a tendency for chromo- somes to stick to one another. The resultant difficulties were noted earlier (Gas- tony & Baroutsis, 1975) and were experienced again in attempts to count A. leptophylla from South Africa. Only one clear meiotic count of n=29 has been obtained from this South African material thus far (Figs. 8 and 9). The conditions causing chromosomal clumping thus appear to be present in both meiotic and mitotic cells.

The similarity in chromosomal morphology and behavior in gametophyte cells of A. guatemalensis and A. leptophylla parallels other shared features, such as identical spore morphology, similar gametophyte development, and similar physiological response to growth conditions (Baroutsis, 1976). Altogether, this evidence supports Tryon’s (1962, p. 75) suggestion that 4. guatemalensis may be an infra-specific variant of A. leptophylla. Final taxonomic disposition, of course, will require comparative morphological studies of populations throughout the North and Central American ranges of these two species.

LITERATURE CITED

BAROUTSIS, J. G. 1976. Cytology, morphology, and developmental biology of the fern genus Anogramma. Ph.D. Thesis, Indiana University, Bloomington, IN. FABERGE, A. C. 1945. Snail stomach cytase, a new reagent for plant cytology. Stain Tech. 20: 1-4.

FIGS. 1-9. Photographs and camera lucida clarifications of Anogramma chromosomes of specimens cited in the text. FIGS. 1-7. Mitotic figures from gametophyte cells. FIGS. 8-9. Meiotic figure from spore mother cell. T = thin chromosome, S = short chromosome. FIGS. 1-2. A. guatemalensis, n=29, FIGS. 3-4. A. leptophylla, Mexico, n=29. FIG. 5. A. guatemalensis, portion of a squas included to show thin chromosome, T. FIGS. 6-7. A. guatemalensis, n=29.° FIGS. 8-9. A. lep- tophylla, South Africa, n=29.

6 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

GASTONY, G. J. 1977. Chromosomes of the independently reproducing Appalachian gametophyte: A new source of taxonomic evidence. Syst. Bot. 2: 43-48.

, and J.G. BAROUTSIS. 1975. Chromosome numbers in the genus Anogramma. Amer.

Fern J. 65: 71-75.

HENDERSON, S. A. and B. C. LU. 1968. The use of haematoxylin for squash preparations of chromosomes. Stain Tech. 43: 233-236.

KURITA, S. 1971. Chromosome study of four species of leptosporangiate ferns. Ann. Rep. Foreign Students’ Coll. Chiba Univ. 6: 41-43.

MICKEL, J. T., W. H. WAGNER, and K. L. CHEN. 1966. Chromosome observations on the ferns of Mexico. Carylogia 19: 95-102.

RIGBY, S. J. 1973. Induction of apogamy in Pellaea glabella var. occidentalis. Amer. Fern. J. 63:

8-1

158-164. TRYON, R. M. 1962. Taxonomic fern notes. II. Pityrogramma (including Trismeria) and Ano- gramma. Contr. Gray Herb. Harvard Univ. 189: 52-76.

Curtis Eugene Delchamps (1925- 1977)

Curtis Eugene Delchamps was born March 3, 1925, in New Orleans, La., but

grew up in Mobile, Ala. He studied chemistry at the University of Alabama, Pennsylvania State University, and West Virginia University where he received the Ph.D. degree. While still a student at the University of Alabama, he married Earsie Ward, who was also a chemistry major. He started teaching chemistry at the University of Miami, Miami, Florida, in 1955, and continued there until his death September 12, 1977, from a heart attack. Gene had a life-long interest in nature, especially wild flowers, and became interested in photography as a means of studying plants. When he first moved to Florida, he Started to learn about the plant life there, both native and cultivated. He soon discovered that the local plant experts knew very little about the native ferns. Learning about them was a challenge to him, and he worked very hard on the group and soon became an authority in the field. He combined his plant expertise with his photographic skills, and became a popular lecturer on ferns.

He served for two terms as president of the Miami Men’s Garden Club, and helped to bring national recognition to the group by sponsoring a successful can- didate for the Johnny Appleseed Conservation award. He was first president of the South Florida Fern Society in Miami, and continued to serve on its Board of Directors until his death,

j se Gene’s greatest contribution to the world of ferns was his enthusiasm or Carnie and his eagerness to share his knowledge with others. This he did by

ters.

He is survived by his wife, Earsie ad , , aughter, Barbara, and a son, Charles- Mrs. C. E. Delchamps, 18240 $.W. 248th St. Homestead, FL 33031.

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 1 (1978) 7

Geographical Distribution of Isoétes butleri in the Southeastern United States JERRY M. BASKIN and CAROL C. BASKIN*

According to Pfeiffer (1922), Isoétes butleri Engelm. occurs in Tennessee, Mis- souri, eastern Kansas and south into Arkansas and Oklahoma. In Tennessee, /. butleri is restricted to cedar (limestone) glades of the Central Basin, where the soil is water-logged during winter and early spring but may be extremely dry during late spring, summer, and autumn. The University of Tennessee and Vanderbilt herbaria have specimens of /. butleri from Bedford, Davidson, Maury, Ruther- ford, Williamson, and Wilson Counties. In addition, we have found the species in Marshall County on 7 May 1977, along State Road 99, 6.5 miles west of U.S. highway 31A VU. & C. Baskin 1895, V DB). All of the above collections were from cedar glades.

FIG. 1. A county dot distribution map of /soétes butleri in the southeastern United States.

Cedar glades also occur in northern Alabama and in Kentucky, but there has been only one previous report of J. butleri in Alabama, and the species has not been reported from Kentucky. The only report of /. butleri from Alabama is by Kral (1973), who collected the species on a limestone glade near Isbel in Franklin County. We have collected the species on a second cedar glade in Franklin County on 27 May 1977, east of Russellville along County Road 83, 0.6 miles north of State Road 24 VU. & C. Baskin 1324, VDB), and on a cedar glade in Morgan County on 6 Apr 1972, near McKendry, along Cedar Plains Church Road, 0.2 miles north of County Road 55 J. & C. Baskin 1193, V DB). In Ken-

*School of Biological Sciences, University of Kentucky, Lexington, KY 40506.

8 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

tucky we have collected /. butleri on a cedar glade in Warren County on 12 Apr 1973, east of U.S. Road 31W, 0.2 miles north of the Warren-Simpson County line VU. & C. Baskin 1648, MIL, TENN, VDB). The presently known geographical distribution of /. butleri in southeastern United States is shown in Fig. /.

In the southeastern United States, /. butleri has been collected only from cal- careous glades. In a recent study of Jsoétes specimens from Arkansas, Missouri and Illinois, Taylor, Mohlenbrock, and Murphy (1975) concluded that /. butleri ‘*. . . Shows a definite affinity for drier upland sites, but more often calcareous, rather than sandstone ones.”’ In his paper entitled ‘‘Some Features of the Flora of the Ozark Region in Missouri,’’ Steyermark (1934) lists J. butleri as a typical calciphile. However, in his ‘‘Flora of Missouri,’’ Steyermark (1963, p. 11) says that the species ‘*. . . occurs both on sandstone and chert as well as on limestone glades.’’ While discussing the occurrence of /. butleri on sandstone with the senior author, Dr. W. Carl Taylor said that some of the sandstones in the Ozark Region of Missouri are cemented together with calcareous material and that /. butleri growing on them may, in fact, be growing on a calcareous substrate.

We thank Dr. W. Carl Taylor of the Milwaukee Public Museum for verifying the identification of our specimens from Warren County, Kentucky.

LITERATURE CITED

KRAL, R. 1973. Some notes on the flora of the southern states, particularly Alabama and middle Tennessee. Rhodora 75: 366-410.

PFEIFFER, N. E. 1922. Monograph of Isoétaceae. Ann. Mo. Bot. Gard. 9: 79-233, t. 12-19.

itl ones J. A. 1934. Some features of the flora of the Ozark Region in Missouri. Rhodora 36:

————. 1963. Flora of Missouri. Iowa State University Press, Ames.

TAYLOR, W.C., R. H. MOHLENBROCK, and J. A. MURPHY. 1975. The spores and taxonomy of Isoétes butleri and I. melanopoda. Amer. Fern. J. 65: 33-38.

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AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 1 (1978) 9

Thelypteris oroniensis, a New Species from Costa Rica LUIS D. GOMEZ P.*

The genus Thelypteris is one of the largest and most homogeneous of tropical ferns. Morphologically, it forms a very natural group, such as that of Elaphoglos- sum. Over 80 species of Thelypteris are known from Costa Rica, and yet recently a new species has been discovered on the isolated summits of the coastal hills facing the Caribbean, which is here described as:

Thelypteris oroniensis L. D. Gomez, sp. nov. Fig. 1.

Herba terrestris, rhizomate suberecto, lignoso, paucissime squamoso vel quasi glabro, squamis ample-latissimis, ambarinis, clathratis, stipitibus fasciculatis, stramineis, canaliculatis, pilis 1-cellulatis et stellatis dispersis. Frondes pinnatae, pinnis retroflexis, 7 paribus alternibus, rachibus flexuosis, pinnis basalibus ses- ls vel subsessilibus, venis utrinque 7-9 paribus per segmento. Indusium nul- um,

Herbaceous, terrestrial plants with suberect rhizome 1.5-2 cm thick, woody, with a few amber brown, wide, clathrate scales. Stipes fasciculate, stramineous, 21-25 cm long, 2.5 mm in diameter, below with a few paleae, above glabrous, canaliculate, above with 1-celled and stellate hairs mixed. Fronds pinnate, with 7 pairs of alternate, retroflexed pinnae, the apical pinna conform, often with a long, adnate basal segment; fronds lanceolate in outline, truncate at the base, 40-45 cm long, 25-27 cm wide, papyraceous, green above, glaucous-greenish underneath, both surfaces glabrous. Rachis flexuose, helicoidal in growth, with mixed 1I-celled and stellate hairs. Basal pinnae subsessile; middle pinnae and upper pinnae ses- sile. Pinnae 11-15 cm long, 2.5-3.7 cm wide, elongate-lanceolate, the apex nar- rowly acuminate, with 21 pairs of segments, some often lacking and reduced to a narrow wing, the basal pinnules unequal, the upper ones reduced or even absent, the lower ones auriculate, broad, and overlapping the rachis. Veins 7-9 pairs per segment. Sori medial; indusia absent. Sporangia with acicular hairs on the walls and stalks. Many spores aborted.

HOLOTYPE: Monte Oroni (Chase 27), lower Talamanca, Pcia. Limon, Costa Rica, Ocampo 1635 (CR 64453; isotypes UC, US).

Thelypteris oroniensis resembles T. nicaraguensis (Fourn.) Morton in habit and pinna shape, but differs from it and other species of subg. Goniopteris by its flexuose rachises and helicoidal growth of the plants. Among the New World species of Thelypteris, only one Ecuadorian species of subg. Amauropelta has flexuose rachises.

*Herbario Nacional, Museo Nacional de Costa Rica, Apartado 749, San José, Costa Rica.

AMERICAN FERN JOURNAL: VOLUME 68 (1978)

iy A Nae GE ¢ RAM NACHNA SAN SE

x Oronetsis fybr nev. : ki ' | Pasa & Th. (Cyclase.

.

ee

> Orome, Faia Tale- -

» cas) helicwidal, Espo. © Ove 27 jumde 1977, ae

FIG; 1. Holotype of Thelypteris oroniensis,

Ocampo 1635 (CR).

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 1 (1978) 1]

Additions and Corrections to the Pteridophyte Flora of Chihuahua, Mexico IRVING W. KNOBLOCH* and DONOVAN S. CORRELL**

Since the publication in 1962 of our “‘Ferns and Fern Allies of Chihuahua, Mexico,” several additional species have been found in the State. Also, some changes and corrections have been noted that should be made available to others.

ADDITIONS AND DELETIONS

1. Selaginella mutica D. C. Eaton var. mutica

Vicinity of Cerro Bola, Sierra del Paso del Norte (Juarez Mts.), on edge of the city of Juarez, Knobloch 2110 (MSC). This specimen was verified by Rolla M. Tryon, Jr. This addition, which is also new to Mexico, was reported by Knobloch (Amer. Fern J. 56: 36. 1966). In the same publication, new stations in Chihuahua were reported for S. pallescens (Presl) Spring and S. rupincola Underw. 2. Anemia tomentosa var. mexicana (Presl) Mickel

As a result of John Mickel’s work on the genus, we now add this taxon to our flora and eliminate A. anthriscifolia Schrad. The latter, according to a personal communication from Mickel, does not occur in Mexico. All specimens cited in our manual are to be referred to A. tomentosa var. mexicana. 3. Asplenium sessilifolium Desv.

Reported from canyon below Basaseachic Falls by Timothy Reeves (Amer. Fern. 67: 62. 1977). 4. Bommeria subpaleacea Maxon

According to C. Haufler (pers. comm.), B. knoblochii Maxon, which we cited as a valid species, is a synonym of B. subpaleacea Maxon (Contr. U.S. Natl. Herb. 17: 169. 1913). 5. Plagiogyria pectinata (Liebm.) Lellinger

All of the specimens cited in our work as P. semicordata (Pres!) Christ should be referred to this species. According to Lellinger (Amer. Fern J. 61: 110-118. 1971), P. semicordata does not occur in northern Mexico. 6. Pteris cretica L.

Reported from the canyon below Basaseachic Falls by Timothy Reeves ( Amer. Fern J. 67: 62. 1977). 7. Thelypteris puberula var. sonorensis A. Reid Smith

Since var. puberula is not considered to occur in Chihuahua, all specimens in our manual should be referred to the var. sonorensis. 8. Thelypteris ovata var. lindheimeri (C. Chr.) A. Reid Smith

Santa Elena Canyon region in the extreme northeastern part of Chihuahua, Brenckle & Koch 51032 (US), cited by Smith (Univ. Calif. Publ. Bot. 59: 118. 1971).

*Department of Botany, Michigan State University, East Lansing, MI 48824. **Fairchild Tropical Garden, Miami, FL 33156.

12 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

9. Thelypteris resinifera (Desv.) Proctor

La Bufa, southeast of Creel, Knobloch 579 (MSC). This previously unidentified specimen is referrable to this species, according to a personal communication from A. R. Smith.

CORRECTIONS

Page 5, line 21: or Short-thorn Forest.

Page 17: Psilotum nudum (L.) Beauv.; Anemia tomentosa var. mexicana.

Page 34, line 5: omit Pl. 6.

Page 35: Equisetum laevigatum=E. x ferrissii Clute.

Page 36, line 34: after ‘‘ferns’’ add ‘‘and allies.’’

Page 46, line 26: Anemia anthriscifolia=A. tomentosa var. mexicana (Presl) Mickel, Iowa State J. Sci. 36: 427. 1962.

Page 78, line 15: add Baja California.

Page 78, line 34: add Baja California.

Page 85, line 24: for “‘Knobloch 584’ read ‘‘Knobloch 449.”’

Page 85, line 41: for ‘‘bipinnate-pinnatifid’’ read ‘*tripinnatifid.’’

Page 87, line 6: add Arkansas.

Page 93, line 32: add Missouri.

Page 97, line 33: add Sonora.

Page 100, last line: add Hidalgo.

Page 102, line 1: Cheilanthes meifolia is now referred toAspidotis meifolia (D. C. Eaton) Pic.-Ser., Webbia 7: 327. 1950.

Page 104, line 7: add Guanajuato, Jalisco, and Durango.

Page 104, line 26: delete ‘‘“Knobloch 449.”’

Page 105, line 16: after ‘‘glabrous”’ add “‘occasionally hairy.”’

Page 151, line 2 from bottom: add North Carolina.

Page 180, line 4 from bottom: *“*Chihuahua,’’ not ‘‘Chuhuahua.”’

Page 184, line 5: ‘‘Segorachi”’ not ‘‘Sugorachi.”’

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AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 1 (1978) 13

The Distribution and Chemical Constituents of the Farinose Exudates in Gymnogrammoid Ferns ECKHARD WOLLENWEBER*

Many members of the Polypodiaceae sensu lato show a conspicuous, yellow or white deposit on the lower surface of their fronds. Therefore, they are often called gold ferns, gold-back ferns, silver ferns, or silver-back ferns. This is especially true for members of the genus Pityrogramma Link. Such deposits also occur in Cheilanthes Swartz and Notholaena R. Br., although they are less known be- cause species of these genera are not easily cultivated in greenhouses. The culture of the decorative species of Pityrogramma was especially in fashion at the begin- ning of the 19th century. Both P. calomelanos (L.) Link and P. chrysophylla (Swartz) Link were grown at Kew beginning in 1790. Fanciers were attracted by the great variability of species and strains and by the formation of various forms and hybrids that occurred during the culture of ‘‘ Gymnogramma.’’ However, the proliferation of hybrids and horticultural forms led to serious taxonomic confu- sion. Domin’s (1929) statement is still valid: one can easily find completely differ- ent species or hybrids grown in greenhouses under the same name. Even recently in botanical gardens the name P. sulphurea (Swartz) Maxon has been applied to varieties of P. chrysophylla and P. austroamericana Domin.

ANATOMICAL OBSERVATIONS

The special anatomical features of the gymnogrammoid ferns were described rather early. Schkuhr (1804, p. 4, t. 4) reported that the lower surface of the fronds of the Schwefelgelber Vollfarn, now Adiantum poiretii var. sulphureum (Kaulf.) Tryon, were covered with an amorphous, yellow web. We owe to Schkuhr charm- ing drawings of entire plants. As to my knowledge, the first detailed figures showing stalked glands themselves were published by Link (1842, t. III, figs. 7-9). De Bary (1877, p. 105) gave a good description of farina-dusted capitate hairs, or ‘‘pili pulverulenti’’ as he called them. The farinose coating of these plants is not excreted by the entire epidermis, like a true wax coating, but is formed exclusively by the globose terminal cell of small hairs which have a short, unicellular stalk. The wax is exuded on the whole surface of the terminal cells in the shape of rod- or needle-like crystals. Weatherby (1920) also has clearly described the glands and farina of Pityrogramma triangularis (Kaulf.) Maxon. A description similar to that of De Bary (1877, p. 105) was given by Nayar (1962) for Cheilanthes: ‘‘This large terminal cell secretes the waxy substance which appears like minute rods which are radially placed around the cell. On older stipes the hairs wither and the rods break up to form a powdery mass.” De Bary (1877, Pp. 105) also published a drawing of such a capitate gland which is so far unsurpassed, showing the eXx- creted material (Fig. 1). This figure has been copied by many authors, including Blasdale (1893). Comparatively less clear is the drawing by Hohlke (1902), which

*Institut fiir Botanik, Technische Hochschule Darmstadt, Schnittspahnstrasse 3, D-6100 Darmstadt. Federal Republic of Germany.

14 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

was copied even by Ogura (1972, p. 130). Hohlke stated that the cuticle was not lifted by the formation of secretions. He asserted that the exudate was formed in the ‘cell membrane”’ (i.e., cell wall) and he believed he saw pores in which the cuticle had been penetrated by the rods of exudate. Further drawings were pub- lished by Molisch (1923, p. 128), Bower (1923, p. 186), Dous (1927), and Nayar (1964). A small microphotograph was published by Smith et al. (1971). It should be noted here that epidermal waxes may form very similar filaments, as shown for example by Gunning and Steer (1977, ft. 9a, b).

WZ A ZZ

FIG. 1. Pili pulverulenti of Pityrogramma tartarea (B rinsed with alcohol), after De Bary (1877).

There has been only one investigation of the ultrastructure of exudate-secreting glands. Schnepf and Klasova (1973) studied the glands of Pityrogramma chrysoconia (Desv.) Maxon by transmission electron microscopy. They belong to the group of glands with a tubular, smooth endoplasmic reticulum as the dominant cell component; this means they are similar to those excreting volatile oils. The plastids seem to be involved in the formation of the flavonoids, but there is no proof of their accumulation in the vacuole. The flavonoids penetrate the cuticle and crystallize on its surface. These results resemble those obtained from the

glands of Primula by Wollenweber and Sch gee nepf (197 ; garded as merocrinic glands by pt (1770), which are likewise re

gramma sulphurea Desy.) is dimini the structure of a ““cupula,’ the capitate cell during fixation.

The location of the

£. WOLLENWEBER: FARINOSE EXUDATES OF GYMNOGRAMMOID FERNS 15

served on young fronds of Pityrogramma austroamericana (Fig. 2). Occasionally farina is produced abundantly on the rachis, mainly at its base, as in Cheilanthes farinosa (Forsk.) Kaulf., P. calomelanos, and P. chrysophylla. Blasdale (1877) stated, ‘“Though normally occuring on the lower surface only they may appear on the upper, and in all cases they are distributed quite uniformly, that is, without reference to the sori, veinlets or other organs.’’ However, in some species only the fertile pinnules have farinose coatings, as in Onychium siliculosum (Desv.) C. Chr. In Pityrogramma trifoliata (L.) Tryon, very young fronds bear sporadic patches of farina that soon disappear, and the fertile pinnae also have a farinose coating. In P. calomelanos | observed glands on the primary fronds, and accord- ing to Bower (1923, p. 199), they appear on the prothallia of Notholaena trichomanoides (L.) Desv.

Commonly either hairs and scales or a farinose coating—but not both—occur on the laminae of a single species, but exceptions are found in Notholaena.

In N. aschenborniana Klotzsch and N. galeottii Fée, the wax-like indument can be completely covered by scales. On the other hand, in Cheilanthes the separation into genera of the two sections Cheilanthes and Aleuritopteris Fée, as supported by some authors, is possible only because the two indument types exclude each other.

FIG, 2. Farina glands on a young frond of Pityrogramma austroamericana.

CHEMICAL NATURE OF EXUDATES

De Bary (1877, p. 105) was probably the first to distinguish farinose coatings (‘‘mehlige Uberziige’’) from wax coatings. However, the material excreted by gymnogrammoid ferns was usually regarded as a wax-like substance up to the middle of the last century. In 1844, Géppert called it a resin, according to Wiesner (1876, p. 236). Klotzsch (1851) introduced the term ‘‘pseudo-stearoptenes for a chemically undefined group of compounds he characterized as ‘‘parts of volatile

16 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

oils and resins condensed by heat withdrawal.’’ He assumed that these com- pounds contain oxygen and counted among them coumarin from Melilotus of- ficinalis and ‘‘Primula camphor”’ from Primula auricula. Wollenweber (1974) has shown the latter to be a mixture of various flavones, and therefore Klotzsch came rather close to the truth. On the other hand, he erroneously stated that excretion of the dry, farinose mass of fern fronds occurs without glands. Wiesner (1876) called the coatings of ferns ‘‘crystalline efflorescences,’’ which he distinguished from plant waxes. The chemical classification of the substances, which were not precisely defined at that time, followed their solubility in water, alcohol, ether, and other solvents. From such investigations, Géppert derived his opinion that the yellow coatings of what is now called Pityrogramma chrysophylla are related to resins. This opinion was also expressed by Hoéhlke (1902). Christ (1897) used the terms ‘wax flour,”’ ‘‘wax powder,” or ‘‘farina’’ (‘‘Mehl’’), whereas Stras- burger (1905, p. 87) called the exudate a “greasy substance.’’ Haberlandt (1918, p. 477) mentioned the capitate hairs of Gymnogramma in a chapter dealing with oil, resin, slime, and gum glands. Mobius (1927, p. 152) compared the farinose coat- ings with wax exudates, Wetzel in Verdoorn (1938, p. 360) classified them with Volatile oils as ‘resinous substances,”’ and even Ogura (1972, p. 126) mentioned ‘‘fatty or resinous excretions.”

Under these circumstances, it is understandable that taxonomists even at the Present time speak of “‘ceraceous ferns” and of ‘‘wax”’ in keys and descriptions. Apparently the results of recent chemical investigations published in chemical journals have not reached the taxonomists! Even in a phytochemical review by Berti and Bottari (1968, p. 643), the term “‘wax coatings’’ was used, despite the more detailed chemical characterization available. This discrepancy has been mentioned several times in the literature (e.g., Smith et al., 1971).

According to strict chemical definition, waxes are esters of long-chain fatty acids with long-chain primary monovalent alcohols, in contrast to fats and oils, which are composed of glycerol esters. According to botanical terminology,

in describing these coatings, both in ferns and in t interfere with chemical definitions. These terms me authors (e.g., Knobloch, 1976).

primulas, for these terms do no already have been used by so

CHEMICAL INVESTIGATIONS OF FLAVONOIDS Pityrogramma.—The first carried out by Blasdale lated an ether-soluble, y he recognized as a ben Structure. Using mode

chemical analysis of the farina in Pityrogramma was (1893, 1903) on P. triangularis (Kaulf.) Maxon. He iso- ellow substance which he called “‘ceroptene,”’ and which zene derivative, although he did not know its molecular m methods like ultraviolet, infrared, and mass spectros-

£. WOLLENWEBER: FARINOSE EXUDATES OF GYMNOGRAMMOID FERNS 17

copy, Nilsson (1959) established the structural formula for ceroptin as that of a chalcone-like substance (Fig. 3, compound /).' Zopf (1906) isolated a red sub- stance from P. chrysophylla and ‘‘P sulphurea’’* which he called ‘*gymnogram- mene’”’ and from P. calomelanos a white substance he called ‘‘calomelanene.’’ He observed that the color of gymnogrammene depended on the shape and size of the

HCO R,O H3C

OH O OH O 1 Ceroptin Chalcones 2 R,2CH,, R=H 3 R,=R,=CH3 OH ae OH H,CO 0 YO See H,C OH Os: 0 OH 0 Dihydrochalcones § Pityrogrammin

4 R,=CH3, R2=H 5 R,=R2=CH;

HCO OH O 7 Combretol 8 Onychium-chalcone FIG. 3. Structures of some fern flavonoids.

crystals. Actually, the farinose coating of the fern appears yellow due to the small crystals. Gymnogrammene is orange-yellow in solution, and the dry, coarsely crystalline product can be deep red. Nilsson (1961b) obtained a similar material from the Gold Fern P. chrysophylla var. heyderi (Lauche) Domin, and he recog- nized that it was a mixture of two chalcones, compounds 2 and 3. He supposed that the second chalcone was identical with gymnogrammene. However, repeat- ing Zopf’s isolation procedure, I found that gymnogrammene must have been a mixture of 2 and 3 in about equal parts (Wollenweber, 1976a). From the Silver ‘Here and below the compounds mentioned in the text are given numbers in italics corresponding with those used in Figs. 3 and 4 and in Table I.

*Presumably some other species, as P. sulphurea is known to have a unique that of P. chrysophylla.

flavonoid pattern unlike

18 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

Fern P. chrysophylla var. marginata Domin, Nilsson (1961la) isolated and iden- tified a mixture of the corresponding dihydrochalcones 4 and 5. Compound 5 is presumably identical with Zopf’s calomelanene (Wollenweber, 1976a). The yel- low form of P. calomelanos was investigated by Bohm (1968), who by indirect evidence recognized chalcone 3 as the main component of the farina. Star and Mabry (1971) found the dihydrochalcone 5 as the major component in the white form of the same species, but dihydrochalcone 4 in P. tartarea (Cav.) Maxon, together with flavone 27 and flavonol /4. In the P. triangularis species complex, Smith et al. (1971) found chemotypes which mainly produce ceroptin J and the new flavone pityrogrammin 6, whereas others produce methyl ethers of kaempferol (/5, 18). It should be stressed that no other species has been found in the investigations of Star (1977) and Dietz (1978) to form ceroptin. From P. chrysoconia | was able to identify flavonols 9 and // (Wollenweber, 1972), and later I found that the light yellow farina of P. austroamericana is composed of chalcone 3 and dihydrochalcone 5. The thick, white coating on the under surface of the fronds of P. lehmannii Hieron. from Colombia consists mainly of dihy- drochalcone 5 (Wollenweber, 1976a).

FIG. 4. General structure of flavonols and flavones. See Table I for the chemical structures of groups Ri through Ra.

Cheilanthes.—Until recently only two species of Cheilanthes had been investi- gated for farina composition. Erdtman et. al (1966), in C. farinosa from Taiwan, found methyl ethers of kaempferol (/8 and 19) and one other substance which, om our own observations, must be apigenin hese three flavonoids, Rangaswamy and lyer

r substances of the basic pattern may be ed the kaempferol derivatives /6 and 19 and sima (apparently an unpublished name) from the Himalayas. Recently Wollenweber (1976b) found these and other substances : C. bullosa Kunze, C. grisea Blanf., and C.

enweber (1977) to produce a farina very similar xudate of C. viscida Davenp. contains apigenin ~29), probably together with terpenoid materials. nvestigation. Figure 4 shows the general structure nd Table I enumerates them.

E. WOLLENWEBER: FARINOSE EXUDATES OF GYMNOGRAMMOID FERNS 19

Notholaena.—Prior to the report of Wollenweber (1976b), no flavonoid data were published for Notholaena. This report contained results from individual specimens of N. candida (Mart. & Gal.) Hooker var. candida, N. schaffneri (Fourn.) Underw. var. nealleyi (Seaton) Weath., and N. standleyi Maxon. These species also produced some compounds which occur in Cheilanthes, and N. candida var. candida excretes a very rare pentamethyl ether of myricetin, com- bretol 7. Results for many more species, mostly from several specimens each, recently were published by Wollenweber (1977a, b).

TABLE 1. FLAVONOLS AND FLAVONES FOUND AS COMPONENTS OF FERN FARINA.

Flavonols (Ri = OH) Flavones (Ri = OH) 9 Galangin (R2 = OH, R3 = Ra = H) 26 Apigenin (R2 = Rs = OH; R3 = H) 10 Galangin-3-methyl ether 27 Apigenin-7-methyl ether 11 Galangin-7-methyl ether 28 Apigenin-4’-methyl ether 12 Kaempferol (R2 = Ra = OH, R3 =H) 29 Apigenin-7,4’-methyl ether 13 Kaempferol-3-methyl ether 30 Luteolin (R2 = Rs = Rs = OH) 14 Kaempferol-7-methyl ether 31 Luteolin-7-methyl ether 15 Kaempferol4’-methyl ether 32 Luteolin-3’-methyl ether

16 Kaempferol-3,7-methyl ether

17 Kaempferol-3,4’-methyl ether

18 Kaempferol-7,4’-methyl ether

19 Kaempferol-3,7,4’-methy] ether 20 Quercetin (R2 = R3 = Ra = OH)

21 Quercetin-3-methy! ether

22 Quercetin-3,7-methyl ether

23 Quercetin-7,3’-methyl ether

24 Quercetin-3,7,4’-methyl ether

25 Quercetin-3,7,3’,4’-methyl ether

Other genera.—There are a few species of other genera which exhibit farinose excretions. One is Onychium siliculosum, which produces a mixture of chalcone . and the new chalcone 8 (Ramakrishnan et al., 1974, as O. auratum). A second is Adiantum poiretii var. sulphureum (A. sulphureum Kaulf.) which, in addition to chalcone 2, exudes dihydrochalcone 4 and traces of flavonols 9 and 1 1 (Wollen- weber, 1976b). Negripteris scioana (Chiov.) Pic. Ser. and Sinopteris albofusca (Baker) Ching also show a farinose deposit on the lower surface of their fronds; they are under investigation now. Lellinger (1967) reported that some species 0 Pterozonium have a yellow, orange, or reddish farina. This seems to consist of chalcones, for I have identified chalcone 3 in one specimen of P. brevifrons (A. C. Smith) Lellinger.

It is striking that the components of fern farina are almost exclusively methyl derivatives of flavonoids, and so are rather non-polar compounds, for oe usually occur as glycosides dissolved in cell sap. Excretions on the winter buds 0 certain trees, e.g., Populus (Wollenweber, 1975a) and Betulaceae Lpimanpa 1975b), also contain methylated flavonoid aglycones. There the “siti * lipophilic properties is more evident because these excretions are often eer ; : remarkable amounts of lipids (volatile oils, terpenoids, phytosterols, and fats),

20 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

and the flavonoids are dissolved in this material. The farina of Primula also consists almost exclusively of a pure mixture of non-polar flavonoids (Wollen- weber, 1974). Excretions of secondary plant products usually are lipophilic (Liittge & Schnepf, 1976, p. 266). It may be assumed that this peculiarity corre- lates with the excretion mechanism, which is almost completely unknown. Considerable amounts of material can be excreted by farinose ferns. In P. austroamericana, we obtained 480 mg of flavonoids from 54 g of air-dried fronds; in P. calomelanos, 32 g from 1 kg; in N. candida var. copelandii (C. C. Hall) Tryon, 0.78 g from 26.2 g; and in P. lehmannii, which produces farina abundantly, 9.1 g from 165 g. These are amounts of 0.9-5.% of the dry weight of the fronds.

CHEMICAL INVESTIGATIONS OF OTHER COMPOUNDS

In a few exceptional cases, white farina on ferns can be due to quite different lipophilic materials. In Cheilanthes argentea (Gmel.) Kunze, we isolated a major component that possibly is a phytosterol; the analysis has not yet been done. Lophosoria quadripinnata (Gmel.) C. Chr. (Cyatheaceae), which appears glau- cous rather than farinose, has a weak deposit probably consisting of a mixture of triterpenes. This material also is under investigation. The n-alkanes, which lead to the glaucous appearance of such polypodiaceous ferns as Phlebodium aureum (L.) Be Smith, have also been found as minor components in Pityrogramma aus- troamericana and P. lehmannii.

Some publications have reported the occurrence of hydrocarbons, long-chain aliphatic alcohols, fatty acids, and terpenoids in ferns. Long-chain alkanes are widely distributed; among them those with odd-numbered carbon chains (C2s- C3s) predominate. Long-chain aliphatic alcohols are found in wax esters; pen- tacyclic triterpenes of the hopan series are abundant in the group of isoprenoids; among the phytosterols, sitosterol is dominant (Bottari et al., 1972; Seigler et al., 1975; Jamieson & Reid, 1975; Lyttle et al., 1976). Unfortunately, it is not obvious from the cited papers whether the substances are internal components of the plants or whether they are deposited externally. The same is true for the sester- penes (Kahn et al., 1969; Iyer et al., 1972, 1973) and the ecdyson analogues (Imai et al., 1969; Faux et al., 1970). When an extract of ground material is made, it can not be seen where the extracted substances were located. Even in some clearly bein species like Cheilanthes farinosa and Onychium siliculosum, where we ome have been ground and extracted only people familiar with the plants i a oft : the flavonoids described are of external origin. When chemical work a ; ay eS iy more attention should be paid to the location of the chemicals

FLAVONOID STRUCTURES AND FARINA COLOR

i. nsible for intense yellow coloration in most cases. The color to chalcones = - a P. chry sophylla, for example, owes its bright yellow ik OR Sched an : The less intense yellow of P. austroamericana is due to sulphureum j cone » with dihydrochalcone 5. The same hue in Adiantum

is produced by chalcone 2 and dihydrochalcone 4. The strong colora-

E. WOLLENWEBER: FARINOSE EXUDATES OF GYMNOGRAMMOID FERNS 21

tion of Onychium siliculosum is due to the presence of chalcones 2 and 6. We observed that the ratio of chalcone 2 to 6 influences the deepness of the hue. Such relationships of colors with ratios of compounds is found especially with mixtures of chalcones and dihydrochalcones, but also occurs to a lesser extent in mixtures of flavones and flavonols. The intense orange of C. mossambicensis Schelpe and C. welwitschii Hooker ex Baker is accounted for by chalcone 2, as shown by Wollenweber (1977b).

In Cheilanthes and Notholaena, white, whitish, and weakly yellow farina dominate (except for some chalcone-colored species and varieties cited below), due to the presence of flavones and flavonols. But it is more difficult to recognize relationships between composition and farina color. In species which produce derivatives of apigenin, like flavones 26-28 in Notholaena grayi and 26-29 in N. greggii (Kuhn) Maxon, the farina is white. Pure white farinas are also encoun- tered in N. candida var. candida (caused by two methyl ethers of myricetin, 7 and an as yet unknown tetramethyl ether) and in N. candida var. copelandii (caused by galangin (9) and the 3-monomethy] ethers of galangin, kaempferol and querce- tin (10, 13, and 21). The white form of N. californica D. C. Eaton produces derivatives of apigenin (27) and luteolin (30 and 3/), kaempferol (/6), and querce- tin (21, 22, and 24). The light yellow farina of N. standleyi consists of derivatives of kaempferol only (12-17). Cheilanthes farinosa is mostly pure white or whitish, the farina being composed of methyl ethers of apigenin (29), kaempferol (14, 18, and 19), and sometimes quercetin (23) as a basic pattern. A faintly yellowish hue may depend on quantitative differences which have not yet been analyzed, but perhaps kaempferol derivative /9 prevails. The white farina of C. albomarginata is due to genkwanin (27) and to two kaempferol methyl ethers (/4 and 16); C. grisea, also white, in addition produces two kaempferol derivatives (18 and 19).

Besides the variation in composition, the density of the deposit and the size of the particles also may play a role in color expression.

THE FUNCTION OF FARINAS

The term secretion, according to the definition of Schnepf (1969), is to be used for exudates produced by organisms or cells as a result of their interaction with the environment, or which are produced as an immediate consequence of such in- teraction. The term excretion, on the other hand, refers to waste matter, the production of which is not directly related to the environment. Schnepf em- phasized that a sharp demarcation of both terms is neither possible nor necessary. Thus the word excretion may well be used when talking about exudates of fari- nose ferns, although the term secretory glands may also be used as a cee

Many years ago, Blasdale (1893) considered the possible function of glandular cells on Pityrogramma fronds and Primula leaves, and he remarked that their existence gave rise to speculation. In his opinion, one could not help recognizing glandular cells as a mechanism for some definite purpose, as he could not — the material excreted as mere waste products. He found one plausible function: protection of young spores as well as the lower epidermis against excessive

22 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

moisture,’ since the position and chemical nature of the farinose material keep water off the lower surface of the fronds. He thought a second role was to protect these natives of arid regions against excessive transpiration, since many allied species without glands had a thick growth of hairs or scales. Hohlke (1902) attrib- uted to the ‘‘resin’’ of Gymnogramma (i.e., Pityrogramma) the function of an insect deterrent ‘‘because the plants in the greenhouse are free of destructive insects even in summer.”’ He felt confirmed in his opinion by the observation that the coating was thick on young fronds and diminished or even vanished on old fronds, which no longer needed this protection. This misinterpretation was proba- bly based on incomplete observation; secretory glands stand much further from each other on adult fronds than on juvenile ones because new glands do not develop during the later stages of frond growth and the glands no longer are active. Hence, the excretion material is dispersed. Partly it crumbles as a dry mass, and partly it is rinsed off by rain. Nayar (1964) states, ‘‘On mature stipes the glandular

hairs shrivel, leaving powdery covering which is often lost on old stipes.”’ Haberlandt (1918, p. 478) stated that the physiological and ecological impor- tance of the epidermal glands in general depends on the nature of the exudate. Apart from the possibility that in some cases useless end products of metabolism may be secreted, secretions usually have some significance, like protection against strong transpiration or against animal attack. These are the same possible functions that Blasdale and Hohlke attributed to fern glands. Bower (1923, p. 198) was convinced of the role of fern excretions as water repellents. Linsbauer (1930, p. 123) accepted primarily their role in preventing excessive transpiration. How- ever, he regarded speculations on the ecological significance as idle. Nilsson (1959) found it tempting to speculate on the possible physiological significance of the ceroptin coating of Pityrogramma triangularis, and he mentioned that the B-triketones (in which he then included ceroptin) are known to exhibit antibacte- rial and sometimes insecticidal activity. I think some antibacterial effect can cer- pone Sena most eee and this could apply to fern farina in different function. He Hae —- ke sired ol 3) bpemiiae? et ae intahee antes a ne py that the chalcones in the sori” of PP. chrysophylla itchiness an a Phe : yzed reaction. However, this is quite unlikely since poksble fineeeea 4 : Rie by glands of the epidermis. All considerations on the arina may be summarized by the comment of Harborne

sa P. X) on flavones and flavonols: ‘The raison-d’étre . . . still remains as

DISTRIBUTION OF FLAVONOIDS IN THE GENERA

All the ferns so far found t Polypodiaceae subfam. G (Onychium), Gymnogram (Adiantum), and Cheilant Pteris), according to the mann, 1954). Hooker an

0 excrete flavonoid aglycones belong to the ymnogrammoideae, and to sects. Cryptogrammeae meae (Pityrogramma and Pterozonium), Adianteae heae ( Cheilanthes, Negripteris, Notholaena, and Sino- System in Engler’s ‘‘Syllabus’? (Melchior & Werder- d Baker (1868, p. 384) proposed Gymnogramma sect.

E. WOLLENWEBER: FARINOSE EXUDATES OF GYMNOGRAMMOID FERNS 23

Ceropteris (Link) Hooker & Baker for farinose species of Pityrogramma and Notholaena sect. Cincinalis (Desv.) Hooker & Baker (1868, p. 373) for farinose species of Notholaena. Farinose Cheilanthes species are called Aleuritopteris by some authors, or are at least separated as Cheilanthes sect. Aleuritpteris (Fee) Hooker & Baker, as in Nayar (1962).

The various species of Pityrogramma in general produce chalcones and/or di- hydrochalcones (2-5, 8); flavones and flavonols can occur as minor components. Cheilanthes and Notholaena, on the other hand, produce flavones and flavonols (7 and 10-32). Known exceptions in Pityrogramma are P. triangularis (Star et al. 1975b) and P. chrysoconia (Wollenweber, 1977a). In both cases, only some forms or chemotypes differ from the chalcone-dihydrochalcone scheme. Exceptions in Cheilanthes are C. aurea Baker, C. aurantiaca (Cav.) Moore, C. chryosophylla Hooker, C. mossambicensis, and C. welwitschii. These species excrete chal- cones. There is also one form of C. welwitschii which produces a dihydrochal- cone. In Notholaena, I know three species which exude chalcones: N. aurantiaca D. C. Eaton, N. nivea var. flava Hooker, and the yellow form of N. sulphurea (Cav.) J. Smith. Dihydrochalcones also may occur, as in N. lemmonii D. C. Eaton and the white form of N. sulphurea. Thus Bohm’s (1975) statement that ‘‘Pityrogramma is the only fern genus known to accumulate chalcones and dihy- drochalcones’’ is no longer true.

CHEMOTAXONOMIC EVALUATION

Alt and Grant (1960) showed that the varieties of Pityrogramma triangular Is constitute a polyploid complex which includes diploids, triploids, and tetraploids. Smith et al. (1971) showed that correlations for these taxa exist between Spore morphology, cytology, and pigment chemistry. According to the composition of farina, they distinguished four chemotypes: ceroptin type, kaempferol-methy! ether types A and B, and a type with kaempferol derivatives and ceroptin. Ac- cording to their data, different ploidy levels can not be distinguished by farina analysis. The complexity of chemical and cytological variation allowed two alter- native interpretations: either P. triangularis is one species existing as an autoploid complex and consisting of genetic variants with the same basic genome, or It 1s a segmental allopolyploid group with several genomes (cf. Mabry, 1973). Later, more detailed chemical investigations included the analysis of internal flavonoid glycosides (Star et al., 1975a). Diploids and tetraploids can be distinguished by this method within the ceroptin type as well as within one kaempferol methyl ether type. Thus, in P. triangularis var. triangularis four taxa can be outlined by means of chemical data. The tetraploid kaempferol methyl ether chemotype 8 7 glycoside pattern composed of those of two diploids, and so may gs her origin. The tetraploid ceroptin chemotype, on the other hand, may be of set . origin. Finally, n-alkanes occuring in these excretions have been spbaserer ; 4 Seigler et al. (1975). As expected, this class of compounds is not peste : chemotaxonomic investigation. However, the average percensade ow wadiias support to the previous suggestions concerning the origin of tetraploids.

24 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

My own investigations as yet have been less far-reaching. Difficulties in obtain- ing plant materials and the small size of most samples received from herbaria has limited analysis to external flavonoid aglycones. Furthermore, the number of samples received of individual species is still very small. Nevertheless, from the high number of species analyzed, certain trends can be observed.

As stated above, excretions of chalcones and dihydrochalcones dominates in Pityrogramma. Apart from the exceptions cited, the occurrence of these sub- stances at present appears to be a genus-specific character. When so far uniden- tified minor components are included, this suggestion is strongly supported. Thus, an Indian P. calomelanos can not be distinguished from a South American P. dealbata. On the other hand, in some species, small differences in external flavonoid patterns are noted, the meaning of which is under investigation. The differences probably are not sufficient to consider the plants as different chemotypes. The presumed specificity of farina composition in a few species (Dietz, 1978) still has to be verified.

As far as Cheilanthes and Notholaena are concerned, it is disappointing that no sharp delimitation of these genera, which are controversial in the taxonomic litera- ture (cf. Knobloch, 1976), is possible by farina analysis. In both genera, methyl ethers of kaempferol, quercetin, and apigenin are synthesized above all others (see Fig. 4 and Table 1). Nevertheless, there are some interesting peculiarities within the genera, even though only a few specimens per species have been analyzed as yet. Notholaena bryopoda Maxon (compounds /3, 16, 19), N. grayi (26-28), and N. greggii (26-29) have species-specific flavonoid patterns. Other species show unique patterns, too, but their substances have not yet been iden- tified. Cheilanthes also has species/specific flavonoid patterns (Wollenweber, 1976c). The basic patterns may vary slightly by the addition of inconstant com- pounds. In this infraspecific variation I am inclined to see an expression of vari- ability in biosynthetic capacity, just as we are used to seeing variability in mor- phological characteristics. Certainly interpretation becomes more difficult when ite complicated flavonoid patterns in species like N. incana Presl or N. stand- a nee SaaS Infraspecific as well as infra- and inter-populational variation

€ studied from single specimens; extensive field collecting is necessary to accomplish this.

In some Cases, variety-specific flavonoid patterns may occur. For example, five specimens of N. candida var. copelandii are characterized by the 3-methyl ethers of galangin, kaempferol, and quercetin, /0, 13, 21. Unfortunately, the poly-0- aa pe of myricetin (7 and unknown) are not so constantly encountered in ona oe. of var. candida. Nevertheless, evaluated jointly with the addi- here of th omponents, they are typical for this variety and permit inclusion

ree specimens which I received unnamed to variety. Similar examples

pein in other Notholaena and Cheilanthes species. ePrsrs st —. in N. affinis (Mett.) Moore and in N. californica, disertbies'thn'’ y oes not distinguish varieties. Tryon (1965, pp. 47-48) € lower lamina surface of N. affinis as having ‘‘pale yellow to yellow

E. WOLLENWEBER: FARINOSE EXUDATES OF GYMNOGRAMMOID FERNS 25

(rarely white) indument.”’ At present I have six samples of this species, all with light yellow farina. Four show kaempferol (/2) as the predominant or even sole component; apigenin (26) and isokaempferid (/3) can occur as minor components. Two collections from Costa Rica, however, show unknown compounds instead; the major component of the farina was identified as a flavonol with butyryl side chain (Wollenweber et al., 1978). It is possible to presume the existence of chemotypes, but as yet it is not known whether these are correlated with popula- tions. For N. californica, Tryon (1956, p. 74) wrote, **. . . lower [surface] whitish to usually yellow ceraceous.’”’ The yellow farina consists of a series of unknown substances now under investigation which show an identical pattern in the eight samples available. But the white farina of three other samples consists of distinct and constant methyl ethers of flavonoids. Here, too, one can presume the exis- tence of chemotypes. It must be left to the taxonomists whether it is justifiable to establish varieties or not. But in both cases both color differences and differences in flavonoid pattern exist.

Tryon (1962, 1964) did not take into account ‘‘wax color forms”’ in ferns in which the color is not correlated with any other characteristic or with geography. For P. chrysoconia, he wrote, ‘Plants with white wax on the leaves and those with yellow wax both occur nearly throughout the range of species and there seems to be no reason to recognize these variants.” For P. chrysophylla, he wrote, ‘‘The white and yellow color forms, although especially striking in this species, do not merit recognition.” For P. calomelanos and P. tartarea, however, he wrote, ‘‘In these species the strong correlation of the character with geography seems to provide it with an importance it would otherwise not have.”’ Tyron distinguishes varieties of these species by the farina color (P. calomelanos vat. calomelanos, var. aureoflava (Hooker) Weath. ex Bailey, and var. ochracea (Presl) Tryon; P. tartarea var. tartarea, var. aurata (Moore) Tryon, and var. jJamesonii (Baker) Tryon). _

Species with very variable patterns of external flavonoids, like N. schaffneri in which each sample is different from every other one and no correlation with the established varieties schaffneri and nealleyi can be detected, are still very puzzi- ing and dictate caution in interpreting flavonoid data In addition, special difficul- ties are expected with those species in which different colors are observed on a single plant, possibly depending on the age of the frond or plant (Tryon, 1956). These phenomena require further investigation, as do flavonoid studies in general. I would appreciate contributions of fresh material and herbarium specimen frag- ments (even of widespread species) to support such studies.

My investigations reported here were initiated by Prof. W. Hagemann, of the University of Heidelberg, who first supplied samples of farinose ferns. In view of the rarity of many species and their restriction to tropical regions of the world, the work on this subject would have been impossible without the kind support of many pteridologists. To all of them I am greatly obliged, and I wish to express my gratitude. Thanks are also due to Prof. W. Ullrich, of the Technische Hochschule Darmstadt, for critically revising the manuscript and for his kind help with the English translation.

26 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

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new sesterpene from Cheilanthes farinosa. Indian J. Chem. 10: -484.

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a a and E. H. REID. 1975. The fatty acid composition of fern lipids. Phytochem. 14: KHAN, H., *: ZAMAN, H. L. CHETTY, A. S. GUPTA, and S. DEV. 1971. Cheilanthatriol—a cee sci, undamental type in sesterpenes. Tetrahedron Letters 1971: 4443-4446

a J. abelniaies necting welche auf der Aussenseite der Pflanzen vor- r. Verh Preuss. Akad. Wiss. 1851: 751-752. KNOBLOCH. I. W. 1976, Morphological characters in Cheilanthes together with a key to North and

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KREGER, D. R. 1958. Wax. Jn W.Ruland, ed. Handbuch der Pflanzenphysiologie, vol. 10.

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AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 1 (1978) 29 SHORTER NOTES

THE SELAGINELLA APODA COMPLEX IN IOWA.—In 1958 R. F. Thorne and R. L. Hulbary discovered the only Iowa locality for specimens of the S. apoda (L.) Spring complex (Muscatine Co., Thorne 20171, UI). This population also represents the most northwesterly station of the S. apoda complex in North America. The population occurred in a seepage bog at the base of a sandy bluff. Shortly after the discovery of this Selaginella, the locality was ditched, drained, and placed into intensive agricultural use. The population was presumed to have been exterminated. In 1975, one of us (JHP) visited the locality and discovered that the population was still extant. Drainage and grazing had great impact on the larger vascular plants, but the prostrate Selaginella appeared to have been pro- moted by the disturbance. The population now exists as a dense ground cover and forms a strip 1-2 m wide and over 200 m long. In addition, the population, which was not fertile in 1958, was abundantly fertile in 1975-1977.

The plants originally were identified as S. apoda and have subsquently been so treated by Iowa workers (e.g., Peck., J.H., 1976. The pteridophyte flora of lowa. Proc. Iowa Acad. Sci. 83: 143-160). Recently, however, investigations by - of us (WRB) led to the description of Selanginella eclipes Buck as a species distinct from S. apoda (Buck, W. R., 1977. A new species of Selaginella in the S. apoda complex. Canad. J. Bot. 55: 366-371). Selaginella eclipes is distinguished from S. apoda by features of the megaspore wall and by leaf morphology. Selaginella eclipes is found to the north and west of S. apoda, and occurs in an arc from Oklahoma northeast to the Great Lakes region and then northeastward along the St. Lawrence River. The Iowa population of the S. apoda complex was not examined during the original study of S. eclipes. Subsequently, lowa specimens were examined and found to be S. eclipes. Selaginella apoda (L.) Spring - there- fore excluded from the Iowa pteridophyte flora. The purpose of this note Is to call attention to the importance, persistence, and identity of the lowa population.— James H. Peck, Dept. of Biology, University of Wisconsin-La Crosse, La CG rosse WI 54601 and William R. Buck, Dept. of Botany and Herbarium, University of Michigan, Ann Arbor, MI 48109.

POLYSTICHUM LONCHITIS FOUND IN THE BLACK HILLS.—A single plant of Polystichum lonchitis (L.) Roth was found in the Black Hills of oo on September 4, 1977. The location is in Crook County, in Upper Dugout Gulc about 8 miles south of Beulah and about 1.5 miles west of the South Dakota border, at an elevation of 4800 feet. The plant was growing in a damp ravine eed Paper Birch, Betula papyrifera Marsh., and Beaked Hazelnut, ee Marsh. The plant was vigorous and had numerous fronds, several of whic ne of removed to be deposited at Harvard University (GH) and the iO a Wyoming (RM) (Dorn 3042). The closest known localities are about m

30 AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 1 (1978)

southwest, in Carbon County, Wyoming. The species also occurs in northwest Wyoming, south-central Montana, and further west. This new locality is not unexpected since the species distribution now parallels that of several other vas- cular plants which jump from the Rocky Mountains, to the Black Hills, to the upper Great Lakes, and to the Gaspé area of Québec.—Robert D. Dorn, Box 1471, Cheyenne, WY 82001.

ON THE DISTRIBUTION OF LYCOPODIUM FLABELLIFORME IN ILLINOIS.—The Ground-pine, Lycopodium flabelliforme (Fern.) Blanch., was collected on 11 Oct 1976 from the northwest part of Lake Argyle State Park, McDonough County, Illinois (NE 1/4 of S36, T6N, R4W). This collection is the first report of this species in western Illinois and the fourth report of a native station for the state. It is interesting to note that the three previously reported native stations, in Pope, Ogle and Crawford Counties,! are on the southern, northern, and eastern perimeters of the state, and McDonough County is at the western edge. Lycopodium flabelliforme, therefore, may occur in interior Illinois counties as well. Three adventive stations in two other peripheral northeastern counties (Ogle and Cook) have also been reported.2, The McDonough County population of L. flabelliforme is about 112 miles from the nearest of the two reported Iowa collections and 140 miles from the nearest Illinois location; the species has not been reported from Missouri.

The Mc Donough County plants were scattered in a shaded area of about 30 ft? on a slope several feet above the lake margin. Cones were not present. The canopy trees of the collection site were Acer saccharum Marsh., with Ostrya virginiana (Mill.) K. Koch and Ulmus rubra Muhl. the dominant understory trees. The mesic deciduous woodland is a relatively young, secondary one. Characteris- tic herbaceous vascular plants included C ystopteris fragilis (L.) Bernh., Adiantum pedatum L., Equisetum arvense L., Sanguinaria canadensis L., Hepatica acutiloba DC., and Dicentra cucullaria (L.) Bernh. Mosses present were Mnium oe Hedw., Brachythecium acuminatum (Hedw.) C. F. Austin, and ite ie : pi owes (Brid.) Grout. The soil was a sandy loam over sandstone

A voucher specimen Illinois University Herb Depa 61455

(R. D. Henry 4052) has been deposited in the Western eo _eroarium (MWI) at Macomb.—R. D. Henry and A. R. Scott, ment of Biological Sciences, Western Illinois University, Macomb, IL

Ill. State Acad. Sci. 64( : 9. distribution record for Lycopodium flabelliforme in Illinois. Trans. + 1967. The Illustrated Flora of Illinois: Ferns. Southern Illinois University

Section Complanata of the genus Lycopodium. Nova Hedw. 19: 142,197,200.

REVIEW 31

a REVIEW

‘‘4 MONOGRAPH OF THE FERN GENUS BOLBITIS (LOMARIOPSID- ACEAE)”’, by E. Hennipman, Leiden Botanical Series 2: xii + 331 pp., 12 pl., 87 fig. 1977. Leiden University Press. ISBN 90-6021-405-6. $38.50/Dfl. 92.—This is an impressive and intricate work, unquestionably the most complete monograph that exists for any pantropical fern genus. Hennipman recognizes 44 species (a reduction from the 97 of Copeland) and 13 hybrids. He divides Bolbitis into ten series, based primarily on venation, spore morphology, rhizome anatomy, and rhizome scales. One of these series (Egenolfianae) comprises the species often included in the segregate genus Egenolfia, but Hennipman offers convincing rea- sons for treating it as a part of Bolbitis. Further, he suggests that the free-veined series Egenolfianae may represent a derived condition in a genus that is usually characterized by anastomosing and reticulate venation. Generally, the series seem well characterized; however, no key is provided and only 34 of 44 species are placed in this scheme, so that it is a little difficult to understand series limits. Hennipman postulates that several of the ten unplaced species may have arisen through hybridization between species of different series. One American species (B. bernoullii) is postulated to have arisen through hybridization between B. lin- digii and a species of another genus, possibly Polybotrya, a suggestion that cries out for experimental verification. If Hennipman is right, hybridization has been extremely important in the evolution of Bolbitis and, by extrapolation, in tropical ferns in general.

The taxonomic account includes keys (separate keys for American, African, and Asian-Pacific Bolbitis), descriptions, synonymy, distribution maps, and eXx- cellent illustrations. There is a thorough introductory account of such topics as morphology and anatomy, karyology, habitat, juvenile leaf characters, intra- and inter-generic relationships, and biogeography. One of the most difficult aspects in dealing with Bolbitis is the great variation in leaf morphology within species, many of which show a succession of leaf forms (heteroblastic series). Certain preco- ciously fertile juvenile forms had previously been treated as distinct species, but Hennipman, through the study of herbarium specimens and living plants, has convincingly shown their identity with more developed forms.

Several unusual or novel features of the taxonomic treatment are exemplary. There is an abbreviated list of specimen citations: where species are common Hennipman gives the number of collections seen in a given political unit; where uncommon or rare, collector, number, and herbarium acronym are cited. To compliment this list, there is a complete index of collections organized alphabeti- cally by collector. Following the description of each species, there are extensive notes on habitat, elevation, relationships, variability, aberrant specimens, typifi- cation, and spore morphology. The last-named character Is given more complete coverage than in any fern monograph I have seen, tropical or temperate. ,

One possible source of confusion is the treatment of certain “taxa as hybrids and others as species of hybrid origin. Hennipman attributes abnormal spores . aborted spore mother cells to most of the hybrids. However, Bolbitis x lancea, B.

32 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

x prolifera, and B. x sinuosa nm. foxii are all described as having ‘‘normally shaped spores.”’ It is also a little disconcerting, even frightening, that most of the 44 species described by Hennipman have some specimens (even a majority, e.g., in B. pergamentacea) that show abnormal spores. Curiously, there are many more inter-series hybrids (13) than intra-series hybrids (3).

If the quality of this monograph is to be approached for other fern genera, there is plenty for all pteridologists to do for the foreseeable future.—Alan R. Smith, University Herbarium, Department of Botany, University of California, Ber- keley, CA 94720.

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AMERICAN PS FERN a 2 JOU RN AL April-June, 1978

QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY

The Taxonomic Status of Selaginella eatonii WILLIAM R. BUCK

Microreplicas as a Technique for Rapid Evaluation of Surface Silica Micromorphology in Equisetum RICHARD L. HAUKE

The Establishment of Bracken Following Fire in Tropical Habitats STEPHEN R. GLIESSMAN

The Distribution and Ecology of Dryop in Southeastern Virginia and Titcccst North Carolina DANIEL L. NICKRENT, LYTTON J. MUSSELMAN, LAURA A. PITCHFORD, and DAVID W. SAMPSON

The Fine Structure of the Newly Formed of Onoclea sensibilis fase r. *». MARENGO and MARIE A. BADALAMENTE

The Anatomy of Equisetum diffusum Tubers S. S. BIR A New Species of Asplenium from Guatemala ROBERT G. STOLZE zene Notes: A New Location for Pellaea glabella n Minnesota; Some Insect Interactions with Azolla mexicana; es on North American Lower Vascular Plants; + Ferns of San Salvador Island, II; Cheilanthes microphylla, a Genus and Species New to the Bahama Archipelago; The Chromosome Number of Notholaena cochisensis

Review

MIRA BOTANICN

UL 25 1978

GARDEN LIBRARY

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37

The American Fern Society Council for 1978

RICHARD L. HAUKE, Dept. of Botany, University of Rhode Island, Kingston, R.1. 02881. President ROBERT M. LLOYD, Dept. of Botany, Ohio University, Athens, Ohio 45701. Vice-President TERRY R. WEBSTER, Dept. of Botany, University of Connecticut, Storrs, Conn. 06268. ' Secretary JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916. Treasurer JUDITH E. SKOG, Dept. of Biology, George Mason University, Fairfax, Va. 22030. Records Treasurer

DAVID B. LELLINGER, Smithsonian Institution, Washington, D.C. 20560. Editor-in-Chief JOHN T. MICKEL, New York Botanical Garden, Bronx, N.Y. 10458. Newsletter Editor American Fern Journal EDITOR-IN-CHIEF DAVID B. LELLINGER Smithsonian Institution, Washington, D. C. 20560 ASSOCIATE EDITORS DAVID W. BIERHORST ..Dept. of Botany, University of Massachusetts, Amherst, Mass. 01002 GERALD I GASTONY <2.55.02%: Dept. of Biology, Indiana University, Bloomington, Ind. 47401 JOHN T. MICKEL New York Botanical Garden, Bronx, New York 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 Institu- tion, Washington, DC 20560. Second-class postage paid at Washington.

Matter for publication and claims for missing issues (made within six months of the date of issue) should be addressed to the Editor-in-Chief,

Changes of address, dues, and applications for membership should be sent to Dr. J. E. Skog, Dept. of Biology, George Mason University, Fairfax, Va. 22030.

Orders for back issues should be addressed to the Treasurer.

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, $5.00; sustaining membership, $10.00; life membership, $100.00). Extracted offprints, if ordered in advance, will be furnished to authors at cost, plus postage.

Back volumes $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. Ten percent discount on orders of six volumes or more; postage additional.

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Dr. John T. Mickel, New York Botanical Garden, Bronx, New York 10458, is editor of the newsletter ‘“Fiddlehead Forum.” The editor welcomes contributions from members and non- bers, including miscellaneous notes, offers to exchange or purchase materials, personalia, hor- ticultural notes, and reviews of non-technical books on ferns.

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h nd to others interested

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 2 (1978) 33

The Taxonomic Status of Selanginella eatonii WILLIAM R. BUCK*

Selaginella eatonii Hieron. ex Small has long been relegated to synonymy under S. armata Baker, but actually it is a distinct species. The plants are among the smallest known in this predominantly tropical genus. Selaginella eatonii and the West Indian S$. armata are both members of subg. Stachygynandrum (Pal. Beauv.) Baker, which is characterized by having dorsiventrally flattened shoots comprising two lateral rows of larger leaves plus two medial rows of smaller leaves. According to Baker’s (1883-1885) scheme of classification, S. eatonii is closely related to the other heterophyllous Selaginellae of eastern North America, namely S. apoda (L.) Spring, S. eclipes Buck, and S. ludoviciana A. Braun.

When J. K. Small (1918, p. 67) originally described S. eatonii, he credited Hieronymus with being the author. No specimens of it were cited, although Small reported the plant as growing on limestone in the ‘‘Everglade Keys;”’ he also stated that the plant was first collected in 1903. The species is quite rare in South Florida.

Much of the pertinent literature has either been misleading or in error. Small (1938, p. 422) transferred S. eatonii to the genus Diplostachyum, which was erected by Palisot de Beauvois to accommodate many of the heterophyllous species of Selaginella. Today this segregate genus is mostly unaccepted. Prior to this time, Britton and Millspaugh (1920, p. 477) reported S. eatonii as occurring in the Bahamas. However, Alston (1952, p. 44), a world authority on Selaginella, cited their Bahamian material as S. bracei Schmidt, which also is a plant of diminutive habit; it was described from Andros and Abaco Islands, Bahamas. Alston (1952, p. 43; 1955, p. 246) placed S. eatonii in synonymy under S. armata, which was described from Cuba. Unfortunately, S$. eatonii was not typified by Alston in his publications, nor were descriptions given to any of the species; only keys, specimen citations, and brief annotations were provided. Subsequent au- thors who have dealt with the South Florida species, e.g., Wherry (1964, p. 276), Ward (1968), Long and Lakela (1971, p. 69), Mickel (1974), and Lakela and Long (1976, p. 27), have used the name S. armata.

Recent examination of type material of the three species in question has led to the conclusion that Alston (1952, 1955) was mistaken in equating S. eatonii and S. armata,. Rather, Selaginella eatonii is synonymous with S. bracei. However, as S. eatonii was described in 1918 and S. bracei in 1924, Selaginella eatonii has to be the correct name for the species. The following key, along with descriptions and illustrations of the type material of the three species, as well as my lectotypifi- cation of S. eatonii and S. bracei, should prevent further misunderstanding, espe- cially of South Florida material.

*Department of Botany and University Herbarium, University of Michigan, Ann Arbor, MI 48109. Volume 68, number 1, of the JOURNAL was issued April 12, 1978.

34 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

Leaf margins serrate, light-green; stomata on abaxial side of lateral leaves and adaxial side of medial leaves scattered over the whole laminar surface; megaspores yellow to orange, less than 220 wm in diameter 1. S. eatonii

Leaf margins ciliate, especially at the base, hyaline; stomata on abaxial side of lateral leaves and adaxial side of medial leaves arranged linearly only along the midrib; megaspores white, more than 220 «m in diameter 2. S. armata

4 Ne

agai Leaves of Selaginella. FIGS. 1-2. S. eatonii (Eaton in 1903, NY). FIG. 1. Lateral leaves. ab - Medial leaves. FIGS. 3-4. S. bracei (Brace 1834, NY). FIG. 3. Lateral leaf. FIG. 4. Medial eat. FIGS. 5-6. S. armata (Wright 3908, NY). FIG. 5. Lateral leaves. FIG. 6. Medial leaves.

1. Selaginella eatonii Hieron. ex Small, Ferns Trop. Florida 67, fig. 1918. Figs. 1-4. ‘ Selaginella bracei Hieron. ex O. C. Schmidt, Repert. Sp. Nov. Fedde 20: 156. 1924, syn. nov. pnaite —— Harbour, Great Abaco Island, Bahamas, 21 Dec 1904, L. J. K. Brace 1834 j PES: Marsh Harbour, Great Abaco Island, Bahamas, Brace 1618 (NY); Mastic Point, Andros Island, Brace 7024, 7113 (both NY). Diplostachyum eatonii (Hieron, ex Small) Small, Ferns Southeast. States 422. 1938.

cas ap Se About lime-sinks, border of Everglades, Black Point Creek, oa County, Florida, 13 Nov 1903, A. A. Eaton 265 (NY; isolectotype US not

W. R. BUCK: TAXONOMIC STATUS OF SELAGINELLA EATONI| 35

Plants yellow-green, small, creeping over limestone with mosses; stems 1-4 cm long. Lateral leaves ovate, gradually acute, 1.25-1.4 mm long, 0.75-0.9 mm wide, serrate; midrib ending 75-125 xm below leaf apex; margin light-green, composed of 2-4 rows of linear, slightly to distinctly papillose cells, papillae arranged lon- gitudinally; upper epidermal cells + isodiametric, many-sided, with several small chloroplasts per cell, lower epidermal cells + rectangular, 2-6 times longer than wide, with strongly sinuous walls; stomata of abaxial (aligular) surface scattered over whole lamina, those of the adaxial (ligular) surface confined to the margin. Medial leaves lanceolate, long-acuminate, 1.0-1.2 mm long, 0.3-0.35 mm wide, serrate; midrib ending 300-350 um below leaf apex; margin light-green, composed of 1-3 rows of linear, slightly to distinctly papillose cells, the papillae arranged longitudinally; apex long-acuminate, serrate, composed of linear cells continuous from the leaf margin, not papillose; epidermis as in lateral leaves; stomata on adaxial (ligular) surface only, scattered over the whole lamina. Strobili 2-5 mm long; sporophylls ovate, ca. 1.4 mm long, 0.75 mm wide, acuminate, serrate; margin as in the trophophylls; midrib ending ca. 400 wm below the leaf apex, strongly ridged on the abaxial side from linear, papillose cells, spinose, the spines ending at about the costal apex; stomata on the abaxial surface only, scattered over the whole lamina. Megaspores yellow to orange, Ca. wm diameter, slightly roughened to almost smooth. Microspores orange, ca. 30 wm in diameter, rough.

2. Selaginella armata Baker, J. Bot. Brit. & For. 22: 90. 1884. Figs. 5-6.

Type: Cuba, C. Wright 3908 (K fide Alston, 1952; isotypes BM, NY!).

abaxial surface confined to the midrib region, those of the adaxial surface confined to the margin. Medial leaves elliptic-lanceolate, short-acuminate, 1.0-1.2 mm long, 0.3-0.35 mm wide, ciliate; midrib ending 300-400 zm below the apex; margin hyaline, composed of 2-4 rows of linear, smooth to slightly papillose cells, the cilia single-celled, becoming longer toward the leaf base, to 130 um long; apex short-acuminate, with only a single pair of linear cells confluent at the apex; epidermis as in the lateral leaves; stomata on the adaxial surface only, arranged longitudinally along and over the midrib. Strobili 2-5 mm long; sporophylls ovate, card mm wide, short-acuminate, ciliate; margin as in the trophophylls but more distinctly papillose; midrib ending Ca. 500 4m below the leaf apex, spinose but not ridged at the back, the spines ending above the costal apex. Megaspores white, ca. 230 «m in diameter, reticulate. Microspores orange, ca. 30 wm in diameter, + granular.

Selaginella eatonii is the correct name for the native heterophyllous Selaginella of South Florida, and S. armata is excluded from the North American flora. The two species are most easily separated on the basis of the serrate versus ciliate leaves in S. eatonii and S. armata, respectively. However, numerous additional diagnostic characters are present at the microscopic level. Most conspicuous of these are the differences in stomatal distribution. In S. eatonii the stomata of the aligular surface of the lateral leaves and of the ligular surface of the medial leaves are scattered over the whole laminar surface, whereas for the same laminar re-

36 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

gions in S. armata, the stomata are confined to the midrib area. Stomatal distribu- tion was first suggested by McNab (1887) as of possible taxonomic importance in the genus. More recently, Buck and Lucansky (1976) used it as an aid in the separation of S. apoda and S. ludoviciana. An additional microscopic character useful in separating S. eatonii from S. armata is the distance from the distal end of the costa to the apex of the lateral leaf. In S. armata the midrib ends 250-400 wm from the apex, whereas in S. eatonii the equivalent distance is only 75-125 pm. Megaspore differences also exist between the two species. Although Hellwig (1969) found megaspore color of little use in segregating higher taxonomic groups of Selaginella, it is of value in separating these species. In S. eatonii the mega- spores are yellow to orange; those of S. armata are white. Also the megaspores of S. armata are larger than those of S. eatonii. Further research on the apparent plastid differences between S. eatonii and S. armata would be rewarding.

r. C. E. Delchamps, of the University of Miami, Coral Gables, Florida, planned to contribute ecological data from personal field experience with S. eatonii. Unfortunately Dr. Delchamps died prior to contributing. I am indebted to him for first calling this problem to my attention by sending me living South Florida material.

I thank Dr. John Mickel of the New York Botanical Garden for permission to examine the type material and for the use of facilities while I was visiting there. I am also grateful to Drs. Howard Crum and W. H. Wagner, Jr. for comments and criticisms concerning the manuscript.

LITERATURE CITED

ALSTON, A. H. G. 1952. Hist.) 1(2): 27-47. ——-—. 1955. The heteroph Hist.) 1(8): 221-274, BAKER, J. G. 1883-1885. A synopsis of the genus Selaginella. J. Bot. Brit. & For. 21: 1-5, 42-46, 80-84, 97-100, 142-145, 210-213, 240-244, 332-336: 22: 23-26, 86-90, 110-113, 243-247, 275-

278, 295-300, 373-377; 23: 19-25, 45-48, 116-122, 154-157, 176-180, 248-257, 292-302. BRITTON, N. L.

New York.

BUCK, W. R. and T. W. LUCANSKY. 1976. An anatomical and morphological comparison of Selaginella apoda and Selaginella ludoviciana. Bull. Torrey Bot. Club 103: 9-16.

HELLWIG, R. L. 1969, Spores of the heterophyllous Selaginellae of Mexico and Central America. Ann. Missouri Bot. Gard. 56: 444-464

LAKELA, O. and R. W. LONG. 1976.

LONG, R. W. and O. LAKELA. Coral Gables, Florida.

McNAB, W. R. 1887. On the stomata and ligules of Selaginella. Brit.

MICKEL, J. T. 1974. Checklist of pteridophytes of North America Forum 1(3): 1-4,

SMALL, J. K. 1918. Ferns of Tropical Florida. - 1938. Ferns of Southeastern States. Science Press, Lancaster, Pennsylvania.

WARD, D. B. 1968. Checklist of the Vascular Flora of Florida. I. IFAS, University of Florida, Gainesville.

WHERRY, E. T. 1964. The Southern Fern Guide. Doubleday,

A revision of the West Indian species of Selaginella. Bull. Brit. Mus. (Nat.

yllous Selaginellae of continental North America. Bull. Brit. Mus. (Nat. t 56.

Ferns of Florida. Banyan Books, Miami, Florida. 1971. A Flora of Tropical Florida. University of Miami Press,

Assoc. Rep. 1887: 743, 744. north of Mexico. Fiddlehead

Published by the author, New York.

Garden City, New Jersey.

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 2 (1978) 37

Microreplicas as a Technique for Rapid Evaluation of Surface Silica Micromorphology in Equisetum RICHARD L. HAUKE*

The genus Equisetum has caused problems to field botanists and herbarium workers because the species are often difficult to identify. The general morphol- ogy is so plastic and responsive to environmental influence that the same species can look quite different under different circumstances, and two different species can under Certain conditions closely resemble one another. One source of charac- ters which has recently come to the foreground is the micromorphology of surface Silica.

Equisetum has long been noted for its ability to take up silicon dioxide and deposit it on the surface of the plant as a hard, outer coat. For this reason, certain species have been used to scour pots and pans (scouring rush), or to polish wood (joiner’s rush). The silica deposits have been suspected of varying among species, and in studying subgenus Hippochaete, | used the silica profile of the branch ridges as a character in distinguishing E. giganteum, E. myriochaetum, and their hybrid, E. x schaffneri (Hauke, 1963). I also illustrated the silica rosettes in the valleys of E. ramosissimum subsp. debile. Milde (1867) had illustrated these, as well as some vague differences of surface pattern among species of subg. Equisetum, particularly on the stomata.

With development of the Scanning Electron Microscope (SEM), surface fea- tures of biological entities have become more amenable to study. Various people have used this technique to look at Equisetum (Laroche, 1968, 1969a, 1969b; Laroche et al, 1970; Kaufman et al., 1971; Page, 1972, 1974; Tanowitz, 1975; Dayanandan, 1977; Lawry, unpublished). Page (1972) revised the taxonomy of Equisetum subg. Equisetum largely utilizing silica micromorphology. Since the SEM technique is somewhat laborious and requires special equipment, it does not lend itself to convenient and rapid evaluation of surface micromorphology of a number of specimens from different localities, of plants of different ages, or of different parts of the same plant. Apparently Page looked at only one specimen for each species, and did not appreciate the possibility of silica pattern variation within a plant or between plants of the same species.

The silica micromorphology does show some consistent characters by which species of Equisetum subg. Equisetum can be recognized (Hauke, 1978), and it is useful in helping to identify aberrant specimens. However, it is impractical for the working taxonomist, who must sort stacks of specimens, to stop and prepare for the SEM material from a problematic specimen, assuming he even has ready access to a scanning electron microscope.

I sought a practical alternative to the SEM for rapidly evaluating surface silica micromorphology of herbarium specimens of Equisetum. The technique I adopted is the *‘microreplica’’ method, as published in the Turtox News some years ago. | have used this successfully on dried, pressed herbarium specimens. It works best

*Department of Botany, University of Rhode Island, Kingston, RI 02881.

AMERICAN FERN JOURNAL: VOLUME 68 (1978)

plicas and SEM micrographs (from Page, 1972: bar scales sido arvense stomate with pilules. FIGS. 3-4. E. pratense stomates : aligned, distinct mamillae. FIGS. 5-6. E. palustre stomates scattered, wit

FIGS. 1-6. Photographs of microre imate) of Equisetum. FIGS, 1-2. Bi lines, with longitudinally transversely aligned, confluent mamillae

R. L. HAUKE: MICROREPLICAS IN EQUISETUM 39

on branches that have been flattened well and are lying against the herbarium sheet.

The portion of branch selected is wetted with acetone, and then a plastic cover. slip (22 mm*) is pressed firmly against it. One can apply maximum pressure by using the end of the thumb, pressed vertically with the weight of the body. After about 12 minute, the coverslip is removed. It is placed on a microscope slide impression side down and observed under a standard light microscope. acetone will have softened the plastic and the pressure will have caused it to conform to the surface of the branch. Microscopic physical features of the surface will be impressed into the plastic, hence the name ‘‘microreplica.”’

As with any technique, certain precautions are necessary for it to work well. If too much acetone is used, some will move over the coverslip and the technician’s fingerprint will be impressed into the plastic. If the surface is not reasonably flattened, only the high spots will be replicated. If there are abrupt heights and depths, the plastic will develop fine fracture lines that will obscure any replica. Care must be used in removing the coverslip from the branch, to avoid having the branch break and its surface stick to the plastic coverslip. I have found that a dissecting needle slid between the two helps to separate them. The coverslip, as it dries, often tends to bend. I immediately attach it to a microscope slide with permanent transparent mending tape to minimize this distortion.

The microreplicas produced by this technique show enough micromorphology to reveal the patterns detected with the SEM. It is true that the contrast is not so great and the resolution of finer details is often poor, but one can see the size, arrangement, and distinctness of mamillae and the type and distribution of pilules on the stomata. Figure | is a photograph of a microreplica showing the stomate and pilules of Equisetum arvense, and Fig. 2 is an SEM micrograph from Page (1972) of the same species. Figures 3 and 5 are photographs of microreplicas showing the mamillae and stomatal arrangement of E. pratense and E. palustre, respectively, and Figs. 4 and 6 are SEM micrographs from Page (1972) of the same species.

As can be seen, there are micromorphological characters useful in species identification, and these characters can be detected quickly on questionable her- barium specimens using the microreplica technique.

LITERATURE CITED DAYANANDAN, P. 1977. Stomata in Equisetum: A structural and functional study. Ph.D. Thesis, University of Michigan, Ann Arbor HAUKE, R. L. 1963. A meen monograph of the genus Equisetum subgenus Hippochaete. Nova Hedw. Beih. 8: 1-123 + . 1978. A taxonomic ee of Equisetum subgenus Equisetum. Nova Hedw. 30 (1, 2):

1-72. KAUFMAN, P. B., W. C. BIGELOW, R. SCHMID, and N. S. GHOSHEH. 1971. Electron microprobe analysis of silica in epidermal cells of Equisetum. Amer. J. Bot. 58: 309-316 LAROCHE, J. 1968. Contribution a l'étude de I’ Equisetum arvense L. II. Recherches sur la nature et la localization de la silice chez le sporophyte. Rev. Gén. Bot. 75: 65-116. . 1969a. Etude des concrétions siliceuses de l’épiderme de |’Equisetum arvense L. au micro- scope a balayage. C. R. Acad. Sci. Paris 268: 2417-2418.

40 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

. 1969b. Etat de la silice sur et dans la membrane épidermique des organes aériens stériles d’Equisetum arvense L. Rev. Gén. Bot. 76: 483-489.

———, C. GUERVIN, C. LECOQ, and VO THI DAO. 1970. Intérét taxonomique de I’excrétion siliceuse chez les Equisétacées. C. R. Acad. Sci. Paris 270: 2958-2960.

MILDE, J. 1867. Monographia Equisetorum. Nova Acta Acad. Leop.-Carol. 32(2): i-viii, 1-605

PAGE, C. N. 1972. An assessment of interspecific relationships in Equisetum subgenus Equisetum. New Phytol. 71: 355-369, . 1974. Equisetum subgenus Equisetum in the Sino-Himalayan region—a_ preliminary taxonomic and evolutionary appraisal. Fern Gaz. 11: 25-47.

TANOWITZ, B. D. 1975. Patterns of epidermal silicification in Equisetum. Bot. Soc. Amer. Abstr. 1975: 61.

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AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 2 (1978) 41

The Establishment of Bracken Following Fire in Tropical Habitats STEPHEN R. GLIESSMAN*

The spread of Bracken, Pteridium aquilinum (L.) Kuhn, as a vigorous and dominant weed is well recognized in many vegetation types throughout the world (see Braid, 1959, for a review). It has been demonstrated that this dominance is a consequence of the fern’s rapid extension of underground stems and abundant vegetative reproduction (Watt, 1943, 1947), its strong allelopathic potential (Gliessman & Muller, 1972), and its resistance to fire (Vogl, 1964). Very little attention has been directed to spore germination, gametophyte formation, and sporeling establishment in relation to the dominance of Bracken.

Bracken is capable of producing large numbers of spores on each frond, and the time of spore release can extend through much of the growing season (Conway, 1957). In temperate regions, however, most spore dispersal takes place during the drier part of the year or just before the winter begins, times of the year that do not particularly favor sporeling establishment. This would explain, at least in part, reports of the small number of Bracken sporelings which become established under natural field conditions (Conway, 1953). In the tropics, where conditions of temperature and humidity are much more equable, such climatic control of spore germination and early growth presumably is less important. I have observed con- tinual growth of new Bracken fronds throughout the year in several locations in Costa Rica (Gliessman, 1976). It is possible, then, that spore release is not re- stricted to a certain period, as it is in temperate regions (Conway, 1957), but may be much more haphazard over the entire year. Thus, a constant source of spores could be available for any potentially habitable area.

Observations were made in the field in Costa Rica to determine the conditions under which Bracken sporelings become established, which has possible implica- tions for Bracken control. In Costa Rica, Bracken is encountered frequently from just above sea level on well-drained soils up to more than 3000 m elevation. As in other areas of the tropics (Richards, 1966, pp. 391-399), Bracken most often forms a type of deflected succession in regions formerly covered with dense, tropical forest. These are areas that have an annual rainfall in excess of 2500 mm, the majority of which is concentrated in the wet season that extends from mid-May to late December. Due to frequent cloud cover (especially at higher elevations) and the occurrence of sporadic rainfall even in the dry season, humidity at the soil level is quite favorable for sporeling establishment all year around.

The pattern of land use in this part of the tropics appears to lend itself quite well to the establishment of Bracken. Most forest clearing, using the well known system of ‘“‘slash and burn,’’ takes place towards the end of the wet season (December) until late in the dry season (late April). The felled material is allowed to dry as much as possible. Before the more frequent rainfall begins in early May, the slash is burned. As a consequence, when the wet season rains begin in earnest,

*Departmento de Ecologia, erate hs Biologia, Colegio Superior de Agricultura Tropical, Apartado 24, H. Cardenas, Tabasco, Méxi

42 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

the conditions that are encountered following the fire very closely approximate the ideal conditions described by Conway (1949) for sporeling establishment and young sporophyte growth. She demonstrated that spore germination took place very soon after release from the fronds, and that it was best on soils with an alkaline reaction (pH 7.0-7.7), especially on sterilized soils.

50 7 i4 —~ 40: 2 ; : V oO 3 504 ( \ \ ae i: Zz 205 ? q V | 10 ’ i} — ass : ; yt ey JUN DEC FIG. 1. Biweekly totals of rainfalls at Finca Loma Linda in 1973. 1 = initiation of forest clearing; 2 =

widespread burning of slash; 3 = widespread occurrence of gametophytes: 4 = establishment of young sporophytes.

On the western boundary of Finca Loma Linda (1300 m elevation) approxi- mately 2 km south of Canfas Gordas, Coto Brus, in southern Costa Rica, a section of montane moist tropical forest was cut beginning in the early dry season (Fig. /)- In late April, the slash was burned. Immediately following the first heavy rains two weeks later, I performed soil pH analyses of the upper 5 cm of soil, including the ash layer. Readings ranged from pH 7.0 to 8.0, there being a positive correla- tion between higher pH and greater ash depth. The ashes were compacted into a dense layer on the soil surface by the rains and ranged from 3 to 11 mm deep. Low areas and depressions where rainwater had accumulated had the greatest ash depth. In a part of the deforested area that escaped burning, I found pH readings ranging from 5.2 to 5.8, the soil surface being covered with a mat of organic matter a osed of humus and intact plant detritus up to 3 cm thick. Thus, only on the ih ze hae bir ag of ie net season did conditions combining a high pH a nly n microviotic diversity exist which were ideal for spore- “ey t weeks after the soil pH analyses were completed, on 7 June 1973, | made

ct counts of gametophytes easily visible with the naked eye in ten randomly

S. R. GLIESSMAN: BRACKEN IN TROPICAL HABITATS 43

placed, 10 cm? plots. I found (Table 1) that a considerable number of well-formed Bracken gametophytes already had become established. Coverage of the soil surface by the developing gametophytes was practically complete. Much closer inspection would probably have revealed more recently germinated spores and very young gametophytes. Nevertheless, the quantities observed are much higher than any others that have been reported in the literature for observations made under natural field conditions (Whyte, 1930; Conway, 1953). Careful inspection of those cutover areas that escaped burning failed to reveal any Bracken gametophyte establishment.

On 18 August 1973, I reexamined the same sites for the establishment of young sporophytes. Of the gametophytes originally observed, approximately 20% had formed sporophytes (Table 1). The number may actually be less than 20%, be- cause individual gametophytes were not marked and new gametophytes could have developed during the time following the initial observations. Still, the num- bers of sporophytes in such small areas is impressive.

TABLE 1. NUMBERS OF BRACKEN GAMETOPHYTES IN 10 cm? SAMPLES TAKEN 4 WEEKS AFTER SOIL pH ANALYSIS AND SPOROPHYTE ESTABLISHMENT 10 WEEKS LATER

Sample no. No. gametophytes No. — 1 115 d 2 85 > g 68 3 4 127 6 5 94 23 6 73 4 7 78 7 9 96 > 10 84 0 Average 92.2 16.9

On more favorable sites (e.g., near downed logs or burned-out stumps) several plants had produced as many as six fronds, some up to 25 cm long, but the majority had two or three fronds with an average length of 5-10 cm. If we consider that at the time of these observations there remained at least 3.5 months of addi- tional frequent rainfall and abundant soil humidity, the growth rates of which the young bracken sporophytes are capable (Conway, 1949) would certainly allow the plants to become well established before the more difficult conditions of the following dry season arrived.

The menace posed by the vegetative spread of Bracken in many parts of the world, including the tropics, is well known (Page, 1976). The capability of Brack- en to occupy large tracts of land with former agricultural value has long been observed in Costa Rica (Standley, 1937, p. 29). Once having become established, the fern is very difficult to eradicate manually or mechanically, and only the widespread application of new chemical fernicides offers control (Martin, 1976). Because of the delicate nature of gametophytes in general and the rather narrow tolerance for environmental stress of the germinating spores, I believe it would be easier to prevent the establishment of Bracken than to remove it following its introduction.

44 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

During the early stages of development, especially initial establishment, indi- vidual plants are most susceptible to adverse environmental factors. In the case of Bracken, understanding that the optimum conditions for sporeling establishment are very similar to those encountered following fire, management practices that avoid these conditions would best prevent its introduction. In the tropics, where conditions of temperature and humidity are very favorable for gametophyte estab- lishment and growth all year around, control of soil characteristics such as pH and microbial diversity might be a positive preventive. Fire should not be used in regions especially susceptible to Bracken infestation or close to areas already dominated by the fern after the original vegetation has been cleared. If high labor cost or physical obstruction to planting caused by the downed slash makes the use of fire necessary, it could be applied only if the slash were gathered in mounds, burned, and then the concentrated ash carefully observed and clinically treated or repeatedly disturbed if gametophytes or young sporophytes appear.

Bracken rapidly takes advantage of conditions created after fire in the tropics. Young sporophytes become established in a very short time in areas where Bracken did not exist before. The widespread use of fire in the tropics thus favors an ever-increasing spread of Bracken. Observations on the establishment of sporelings following fire may provide the necessary tools for preventing domi- nance by this fern.

LITERATURE CITED BRAID, K. M. 1959. Bracken: a review of the literature. Hurley: Commonwealth Agricultural ureaux. ae E. — autecology of bracken (Pteridium aquilinum (L.) Kuhn): the germination of € spore and the development of the prothallus and the voun ophyte. Proc. Roy. Soc. Edinburgh. 63: 325-343. young sporophyte. Pr y PGS die and sporeling survival in bracken (Pteridium aquilinum (L.) Kuhn). J. Ecol. 41: i Spore production in bracken (Pteridium aquilinum (L.) Kuhn). J. Ecol. 45: 273-284. AN, S.R. 1976. Allelopathy in a broad spectrum of environments as illustrated by bracken. Bot. J. Linnean Soc. 73: 95-104, »and C. H. MULLER. 1972. The phytotoxic potential of bracken (Pteridium aquilinum (L.) Kuhn). Madrofio 21: 299-304. pool D. J. 1976. Control of bracken. Bot. J. Linn. Soc. 73: 241-246. »C. N. 1976. The taxonomy and phytogeography of bracken—a review. Bot. J. Linn. Soc. 73:

1-34. RICHARDS, P. W. 1966. The Tropical Rain Forest, rev. ed. Cambridge University Press, Cam-

bridge. a 1937. Flora of Costa Rica. Part I. Field Mus. Nat. Hist., Bot. Ser. 18: 1-398. RJ. 2 The effects of fire on the vegetational composition of bracken-grasslands. Wiscon- sin Acad. Sci. Arts Lett. 53: 67-82. TT, A. S. 1943. Contributions to the ecology of bracken (Pteridium aquilinum (L.) Kuhn). I. The frond and the plant. New Phytol. 42: 103-126, . 1947. Contributions to the ecology of bracken (Pteridium aquilinum (L.) Kuhn). IV. The wie Structure of the community. New Phytol. 46: 97-121. E, J. H. 1930. The spread of bracken by spores. Trans. Bot. Soc. Edinburgh 30: 209-211.

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 2 (1978) 45

The Distribution and Ecology of Dryopteris in Southeastern Virginia and Adjacent North Carolina!

DANIEL L. NICKRENT, LYTTON J. MUSSELMAN, LAURA A. PITCHFORD, and DAVID W. SAMPSON*

The pteridophyte flora of southeastern Virginia and northeastern North Carolina has received considerable study during the past century. Such notable botanists as Chickering, Palmer, Kearney, Small, Wherry, and Fernald have botanized in this region (see the literature review in Kirk et al., 1978). More recently, the Dismal Swamp and contiguous areas have been studied, and the results of these efforts have been included in the Floras published for North Carolina (Radford et al., 1968) and Virginia (Harvill et al., 1977). The genus Dryopteris has received special attention, for it was in 1899 that Palmer first collected D. celsa (Palmer) Small in the Dismal Swamp. More recent studies on the genus in the Dismal Swamp through 1973 are summarized in Wagner and Musselman (1978).

The present paper presents new information on the distribution of Dryopteris species and hybrids collected since 1974, not only in the Dismal Swamp but throughout the Virginia counties of Norfolk (now the City of Chesapeake), Nan- semond (now the City of Suffolk), Southampton, Isle of Wight, and Surry, as well as the North Carolina counties of Hertford, Gates, Chowan, Perquimans, Pas- quotank, and Camden. This is an area roughly delimited by the James River to the north, the Atlantic Ocean to the east, the Chowan River to the west, and Al- bemarle Sound to the south (Fig. 1). This work indicates that Dryopteris is more widely distributed in the area than formerly thought.

Our field observations indicate that in this area Dryopteris species always are ecotone plants. Although they are invariably found in areas contiguous with swamps, they seldom grow in inundated sites. A description of the habitat at the study location in Gates County illustrates such a habitat best. The Suffolk es- carpment rises abruptly to the west. Along the slope of the escarpment are Pinus taeda, Liquidambar styraciflua, Fagus grandifolia, Quercus michauxii, Osmunda cinnamomea, and Polystichum acrostichoides.

The wet, low area which forms the eastern boundary of this Dryopteris site has frequent periods of inundation, especially during the winter months. As in the well drained slopes of the escarpment, no Dryopteris is found in this area. Dominant tree species here are Nyssa aquatica, Taxodium distichum, Acer rubrum, Salix nigra, and Populus heterophylla. The herbaceous vegetation is characterized by Saururus cernuus, Pilea pumila, Arundinaria gigantea, Lorinseria areolata, and Smilax spp.

*Department of Biological Sciences, Old Dominion University, Norfolk, VA 23508. Address reprint request to LJM.

'This work was supported by NSF grant SMI77-01237 and was incidental to research sponsored by a grant from the National Geographic sagt Miss Pitchford was a -Undergraduate Research Participant in 1977. We wish to thank Prof. W. H. Wagner, Jr. for initiating and encouraging this work and for providing some previously unpublished data.

46 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

The ecotone where Dryopteris occurs contains elements from both the escarp- ment and the low areas. Nyssa aquatica, Acer rubrum, and Taxodium distichum usually are the dominant trees. The understory consists of //ex opaca and Asimina triloba. Lonicera japonica (the weedy Japanese Honeysuckle) is common throughout the swamp; however, it serves as a good indicator of a Dryopteris site when present with the woody plants noted above. Other ferns, such as Athyrium asplenioides and Lorinseria areolata, also are abundant here. This area has high leaf litter accumulation throughout the year, usually is damp, and receives very little light.

ttn", a Cty ae an

eh ig Set

4 D. X australi pet: oli: : D.X triploidea D.X separabilis D. celsa X cristata — (Palmer) Small

FIG. 1. County map and th i ue p € study region. FIG. 2A-H. Distribution of Dryopteris taxa in the study

The center of Dryopteris dis site, where all eight taxa found i The sexual species are D. celsa

tribution in our study region is the Gates County

(D. celsa x intermedia), ILG; D. edia x spinulosa), 11S; D. x australis (Wherry)

Small (D. ici all (D. celsa x ludoviciana), GLL; and the unnamed hybrid D. celsa x cris-

tata, GLLS.

D. L. NICKRENT ET AL.: DISTRIBUTION AND ECOLOGY OF DRYOPTERIS 47

DISTRIBUTION OF SEXUAL SPECIES

The following list summarizes the distribution of each sexual species by county (Fig. 2). Voucher specimens cited in this paper are deposited in the herbaria of Old Dominion University (ODU) and the University of Michigan (MICH). Dryopteris celsa (Palmer) Small Figs. 2A and 3A.

Harvill et al. (1977) listed D. celsa as occurring in Norfolk, Nansemond, and Southampton counties in Virginia. Two additional counties may now be added:

Isle of Wight: Two plants at margin of upland forest and small tributary of the Blackwater River, Musselman 5055. Surry: About ten plants at bottom of steep slope along small stream, Musselman et al, 4948.

Dryopteris celsa was described as being ‘‘rare and sporadic”’’ and occurring no farther north than Martin County in North Carolina (Radford et al., 1968). Our field studies have shown, however, that D. celsa is considerably more widespread than once thought, at least in the northeastern section of the state. Recently, Hardin (1977) recorded D. celsa as threatened throughout all floristic provinces of North Carolina. The following list gives new county records for this fern not recorded in Wagner and Musselman (1978) or Musselman et al. (1977). In each county, the habitat is very much like the one described for the Gates County site.

Hertford: A large population of ca. 50-75 plants growing with D. cristata, Nickrent 1271. Chowan: A large population of ca. 75-100 plants growing with D. spinulosa, Nickrent 1295. Perquimans: Two small populations of less than 20 plants, one with D. x separabilis, Nickrent 1298, 1299. Pasquotank: Two small populations of less than 20 plants, Nickrent 1301, 1302. Camden: Many scattered popula- tions along a creek bank; one site with four plants of D. spinulosa, Nickrent 1278. Nansemond: A population of ca. 500 plants near Whaleyville, Musselman 4944.

Dryopteris cristata (L.) Gray Figs. 2B and 3B.

Harvill et al. (1977) recorded D. cristata from Norfolk and Southampton coun- ties. Efforts to find a population in the interposed Nansemond County have so far not been successful. Hardin (1977) considered this species to be threatened in the mountain and piedmont portions of North Carolina. Dryopteris cristata was first collected in Gates County in 1974 (Teulings s. n., NCU, ODU). Since then, the Crested Shield Fern has been discovered in adjacent Hertford County (Mussel- man 5472). Only two plants were found here among a large D. celsa population near Parker’s Ferry. Aside from the presence of Pinus taeda, this habitat was generally like the ecotone described earlier.

Dryopteris spinulosa (Muell.) Watt Figs. 2C and 3C.

The Spinulose Wood Fern previously was known from only Cabarrus and Mecklenburg counties in North Carolina (Radford et al., 1968), until Wagner and Musselman (1978) found it in Gates County. Hardin (1977) considered this fern to be threatened in the southern Piedmont area. The range of this fern is now ex- tended to two more counties within our study area.

Camden: About four plants among a large D. celsa population, Nickrent 1274. Chowan: Only two plants among a large D. celsa population, Nickrent 1296.

The rarity of this species makes the possibility of finding the hybrid between it and D. celsa somewhat remote. If the hybrid is to be found, however, the most probable location would be where the two parents are abundant: in the Dismal Swamp (Wagner & Musselman, 1978).

AMERICAN FERN JOURNAL: VOLUME 68 (1978)

ee) oa 8

LAA Ml

va / ‘4

D. L. NICKRENT ET AL.: DISTRIBUTION AND ECOLOGY OF DRYOPTERIS 49

Dryopteris intermedia (Muell.) A. Gray Figs. 2D and 3D. In our study area, the Fancy Fern is found in the Virginia counties of Norfolk and Nansemond and in Gates County, North Carolina. Apparently this fern has more restricted habitat requirements than the other Dryopteris taxa. DISTRIBUTION OF DRYOPTERIS HYBRIDS Dryopteris < australis (Wherry) Small Figs. 2E and 3E. The postulated parents of the Southern Wood Fern Hybrid are D. celsa (GGLL) and D. ludoviciana (Kunze) Small (LL) (Wagner, 1971). To date, D. ludoviciana has not been collected in the Dismal Swamp area. This would be an important discovery, since only D. celsa is currently known to exist in the vicinity of the hybrid. In this area, D x australis is known only from the Gates County site. The ie alos and cytology of this hybrid will be published separately (Wagner, pers. comm.). Dryopteris < ‘tito Wherry Figs. 2F and 3F. Two sites in Gates County are the only known locations for the Glandular Spinulose Fern Hybrid in our study area. The rarity of the parents, D. intermedia and D. spinulosa, and the difficulty in proper identification probably account for the fact that this hybrid is seldom seen or collected. If the two parents existed together in great enough numbers, hybridization should occur, for D. x triploidea is common and even abundant elsewhere in its range (Wagner, 1971). For exam- ple, Montgomery (1976) mentions that this hybrid may be even more abundant than either parental species in New Jersey. Dryopteris < separabilis (Palmer) Small Figs. 2G and 3G. Previous collections of the Glandular Log Fern Hybrid have been made from Norfolk and Gates counties. A new site in Perquimans County (Nickrent 1297) was especially interesting, for it contained only two plants of D. x separabilis and only about 15 plants of one of the parents, D. celsa. A thorough search revealed no D. intermedia, the other parent. This raises the question of how plants of this ‘*sterile’’ triploid originated at this site. Dryopteris celsa < cristata Figs. 2H and 3H. This hybrid is one of the more interesting fern discoveries made in the Dismal Swamp in recent years. Its genomic formula is GLLS, and the plants yield mainly Sterile spores. This hybrid has been reported from only three other locations in the United States, each of which is many hundreds of miles distant from the others. The first report of D. celsa x cristata was from the East Bergen Swamp in Genesee County, New York (Wagner & Wagner, 1965). The presence of aborted spores and later the recognition of 164 chromosomes confirmed the identification. In 1968, while exploring a swamp in Kalamazoo County, Michigan, W. H Wagner and D. J. Hagenah again discovered this Log Fern hybrid (Wagner et al.,

FIG. 3. Representative pitino of fronds of Dryopteris taxa from the study area. FIG

D. celsa (Nickrent 1271). FIG. 3B. D. cristata ames 1023). FIG. 3C. D. spinulosa aceon 102). FIG. 3D. D. intermedia (Sampson 135). F 3E. D. X australis (Nickrent 1292). FIG. 3F

x triploidea (Pitchford 2004). FIG. 3G. D. x hanes (Pitchford 2031). FIG. 3H. D. celsa x cristata (Pitchford 1005). Bar=5 cm.

50 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

1969). The third report was by Montgomery (1975) from Bergen County, New Jersey.

In 1975, this hybrid was collected from Gates County (Musselman 4911). Fig- ure 3H shows a frond of D. celsa x cristata from this site. As with most Dryo- pteris hybrids, it is intermediate in morphology between it parents.

DISCUSSION

Data presented here indicate that Dryopteris taxa are much more common in southeastern Virginia and adjacent North Carolina than previously was realized. Among the sexual species, one of the more significant findings of this study is the collection of D. spinulosa in Camden and Chowan counties, North Carolina. With the Gates County report of Wagner and Musselman (1978) and the two Piedmont counties recorded by Radford et al. (1968), the Spinulose Wood Fern is now known from five North Carolina counties.

Certainly the rarest taxon included in this study is the unnamed hybrid D. celsa x cristata. This hybrid will be given a specific epithet for consistency in referring to Dryopteris hybrids (Wagner, pers. comm.).The large population (ca. 100 plants) of the hybrid is located in the midst of a huge D. celsa stand (over 1,000 plants) and about 25 plants of D. cristata.

At this same site are a few scattered plants of D. intermedia and abundant D. spinulosa, yet the only D. cristata hybrid found here is the one noted above. We have searched carefully for D. x boottii Underw. (D. cristata x intermedia) and D. xX uliginosa Druce (D. cristata x spinulosa) to no avail.

The Glandular Log Fern Hybrid (D. x separabilis) is a very rare fern, but one which will grow in disturbed habitats. We have been able to grow large numbers of sporophytes of this triploid from its ‘‘giant’’ spores. While we have yet to deter- mine with certainty the chromosome makeup of either the ‘‘giant’’ spores or the sporophytes, it appears that spores may play an important role in the reproduction of this plant. Wagner (1971) has proposed a mechanism which he termed “‘hy- bridization by remote control,’’ which may be an explanation for the hybrid’s occurrence. This mechanism, as opposed to others, is in agreement with field observations, for the habitat of the hybrid can only be described as weedy. The plants of D. x separabilis grew along the roadside among Rubus cuneifolius, Phytolacca americana, and abundant Lonicera japonica. This habitat may be unfavorable for D. intermedia sporophytes, but adequate for the gametophytes to persist and provide the necessary gametes for hybridization. A third alternative. which we consider unlikely, is that both parents once existed at this site but only D. celsa persisted.

The Log Fern (D. celsa) has long been the most intriguing vascular plant of the Dismal Swamp, one of the few areas where it is truly common and found in large populations. Our work indicates, however, that the Log Fern is much more com- mon in the geographical region under consideration than was realized previously. It is to be expected in well drained soil at the border between swamps and upland areas. An exception to this is its absence along any large river. This may be due to the fact that it cannot tolerate inundation. In fact, the Log Fern appears to be an

D. L. NICKRENT ET AL.: DISTRIBUTION AND ECOLOGY OF DRYOPTERIS 51

aggressive invader of disturbed areas in swamp systems. It regularly spreads to

roadbanks within the Dismal Swamp. Large, vigorous plants were abundant on

dredge spoils from a drainage canal at the Camden County site. Activities which impede water movement, thus making a drier habitat, favor the spread of the Log

Fern. The construction of U.S. Highway 158 in Gates County apparently blocked

water movement to the south, with resultant drying and the spread of D. celsa.

Similar examples could be given from the sites noted earlier in this paper. It may

be significant that of the eight county records reported in this paper, we consider

only two sites (Isle of Wight and Chowan counties) to be relatively undisturbed.

In these areas, D. celsa does not form large stands, but rather occurs scattered

among plants of Woodwardia areolata, Athyrium asplenioides, and Onoclea sen-

sibilis. Thus, we consider D. celsa to be more abundant now than at any other time in recent history.

Dryopteris celsa is morphologically quite variable within our study area. One of us (DWS) has begun a study of variability in D. celsa using such characters as the pinna angle in relation to the rachis and the shape of the pinnules closest to the rachis. Of particular interest is the presence of aborted and ‘‘giant’’ spores in morphologically typical D. celsa plants.

LITERATURE CITED

HARDIN, J. W. 1977. Vascular Plants. Jn J. E. Cooper, S. S. Robinson, and J. B. Funderburg, eds. Endangered and Threatened Plants and Animals of North Carolina. N. C. State Museum of Natural History, Raleigh.

HARVILL, A. M., Jr., C. E. STEVENS, and D. M. E. WARE. 1977. Atlas of the Virginia Flora, part |. Virginia Botanical Associates, Farmville, VA.

KIRK, P. W., Jr., H.G. MARSHALL, and P. STEWART. 1978. Scientific and Technical Literature Concerning the Dismal Swamp Area. Jn P. W. Kirk, Jr., ed. The Great Dismal Swamp. Univ. Press of Virginia, Charlottesville, V

MONTGOMERY, J. D. 1976. The dicirtoution and abundance of Dryopteris in New Jersey. Amer. Fern J. 66: 53-59.

MUSSELMAN, L. J., D. L. NICKRENT, and G. F. LEVY. 1977. A contribution towards a vascular flora of the Great Dismal Swamp. ange 79: 240-268.

RADFORD, A. E., H. E. AHLES, and C. R. BELL. 1969. Manual of the Vascular Flora of the Carolinas. Univ. of North Carolina Press, cee Hill, NC.

WAGNER, W. H., Jr. 1971. Evolution of Dryopteris in Relation to the Appalachians. Jn P. C. H ed. The Distributional History of the Biota of the Southern Appalachians, part II, Flora. ginia Polytech. Inst. and State Univ. Res. Div. Monogr. 2: 147-

, and F. S. WAGNER. 1965. Rochester area log ferns fPrvopiers celsa) and their hybrids. Proc. Rochester Acad. Sci. 11: 57-71. ,and D. J. HAGENAH. 1969. The oe re (Dryopteris celsa) and its hybrids in Michigan—a preliminary report. Michigan Bot. 8: 137- and L. J. MUSSELMAN. 1978. a acs (Dryopteris celsa) and their relatives in the ae Swamp. Jn P. W. Kirk, Jr., ed. The Great Dismal Swamp. Univ. Press of Virginia, Charlottesville, VA.

52 AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 2 (1978)

The Fine Structure of the Newly Formed Spore of Onoclea sensibilis NORMAN P. MARENGO and MARIE A. BADALAMENTE*

The fine structure of the dividing meiocyte of Onoclea sensibilis L. was de- scribed by Marengo (1977), and that of the mature spore of the closely related Matteuccia struthiopteris (L.) Tod. by Marengo (1973). It is of interest to estab- lish the ultrastructure of the cells formed at the termination of the second meiotic division. Marengo (1949) reported that cytoplasmic inclusions resolvable by light microscopy apparently disappeared during spore enlargement and that proplastids and plastids made their appearance as the large vacuole was replaced by more cytoplasm. To establish the identity of cytoplasmic inclusions received from the meiocyte and to elucidate the fine structure of the young spore, an electron micro- scope study was made of a sporangium shown by thick sections to contain young spores just separating from the tapetum (Fig. /, T).

Individual sporangia dissected from young fertile fronds were fixed in glutaral- dehyde followed by osmium tetroxide and embedded in Epon (Spurr, 1969). 0.5 mM sections were cut from individual sporangia and examined by phase contrast microscopy without staining. From a sporangium identified as having spores at the desired stage, thin sections were cut with a diamond knife, stained with uranyl acetate and lead citrate, and examined with an Hitachi HU-11A EM

Identifiable inclusions present in the young spore shown in Figs. / and 2 include lipid droplets (L), amyloplasts (A), and mitochrondria (M). Small vacuoles (V) are present, as well as a loosely organized endoplasmic reticulum.

At this stage, the spore nucleus appears to have not yet reached full interphase, since a nucleolus is not present and the nuclear membrane (Fig. 2, NM) is poorly defined. The mature spore of this species has a nucleolus occupying fully one- third of the nuclear cross-section (Marengo, 1956).

It is hoped that the optical disa

ppearance and re-appearance of inclusions in the enlarging

: spore can be followed with ultrastructural techniques. Properly buffered fixative may allow preservation of Stages plasmolyzed by the fixatives of light microscopy. Older sporangia are to be dealt with in the next phase of this study.

LITERATURE CITED

MARENGO, N. P. 1949. A study of the cytoplasmic inclusions during sporogenesis in Onoclea sensibilis. Amer. J. Bot. 36: 603-613. |

- 1956. The microscopic structure of the mature spores of the Sensitive Fern, the Ostrich Fern, and the Royal Fern. Amer. Fern J. 46: 97-104.

. 1973. The fine structu Bot. Club 100: 147-150. ETT,

re of the dormant spore of Matteuccia struthiopteris. Bull. Torrey

peta i oe features of the dividing meiocyte of Onoclea sensibilis. Amer. J. Bot. -601

64: 600 : SPURR, A. R. 1969. A low viscosity epoxy resin embedding medium for electron microscopy. J. Ultrastructural Res. 26: 31-43.

*C. W. Post College, Long Island University, Greenvale, NY 11548.

MARENGO & BADALAMENTE: FINE STRUCTURE OF ONOCLEA SPORE 53

FIG. 1. Longi siti section of a young Onoclea sisibile spore, newly separated from the tapetum (T). Identifiable inclusions are amyloplasts (A), lipid droplets (L), and mitochrondria (M). Vacuoles (V) are present, and a loosely organized endoplasmic reticulum is apparent. x 11,080.

*

AMERICAN FERN JOURNAL: VOLUME 68 (1978)

= OS ray ees = / eng # = we a get e: so

a

yo a Pe ie TNL.

IG. 2. Enlargement of the lower portion of the spore in Fig. 1. Organelles labeled as in Fig. /.

: : Nuclear membrane (NM) appears diffusely organized. Organelle adjacent to labeled amyloplast (A) 1s 2

probably the surface view of a mitoc

hrondrial crista. x 33,17

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 2 (1978) 55

The Anatomy of Equisetum diffusum Tubers 5.8 Bik"

Equisetum diffusum D. Don, which belongs to subg. Equisetum (Hauke, 1974), is characterized by similar fertile and sterile stems, and is distributed throughout the Himalayas from Kashmir in the west to Darjeeling-Sikkim and Khasya in the east, at elevations of 1,500-2,400 m. It inhabits moist, partially exposed, sandy- gravelly soil along roadsides or in ravines, and is fertile from July to September. Up to four underground tubers may be present per plant during March and April on collections made from the Dhobi ghat and Pari Tibba, at 1,500 m elevation near Mussoorie, a hill resort in the western Himalayas. Such plants were described briefly by Mehra and Bir (1959). Tubers also occur in E. arvense, E. palustre, E. sylvaticum, and E. telmateia (Campbell, 1918; Hauke, in litt.).

Although the anatomy of E. diffusum has been studied by Sen and Sen (1973), the tuber structure has remained undescribed until now. Material from Mussoorie was fixed in formalin-acetic acid-alcohol. Mostly free-hand transections were cut and stained with safranin and fast green.

Tubers on the rhizome appear to arise as a result of stunted growth of the lateral shoots arising from the nodes, and generally consist of one long, swollen inter- node. A median transection of the tuber is almost circular in outline and has a narrow zone of tracheary tissue composed of a ring of 10-12 small collateral vascular bundles embedded in a large matrix of ground tissue (Fig. /). The epidermis and 2 or 3 layers of outer cortex below it consist of somewhat thick- walled cells resembling cork. Usually these are devoid of any starch (Fig. 2). Often some long, papilliform epidermal hairs are present (Fig. 3). The inner cortex consists of thin-walled parenchymatous cells densely filled with simple, globular or oval starch grains having concentric striations and a well-marked, streak-like hilum. Each vascular bundle is demarcated by its own endodermal layer, the cells of which possess the usual casparian thickening on their radial walls. This is similar to the endodermis in E. arvense tubers (Barratt, 1920, fig. 9). A single-layered pericycle lies underneath the endodermis. The endodermis and pericycle have denser contents, compared with those of the adjacent tissue. The metaxylem elements are placed irregularly, with a few protoxylem elements somewhat mesarch in position. The carinal canal so characteristic of the rhizome and stem of this species is lacking in the tubers. Phloem has the usual structure (Fig. 4). Maceration of the xylem elements revealed only tracheids.

In contrast to the endodermis position in the tubers, that in the rhizome and stem of this species shows a common ring of endodermis surrounding all the vascular bundles. In my material, the root stele is di- or triarch, with two or three protoxylem groups surrounding a single axial metaxylem element, contrary to the earlier report of triarch roots by Sen and Sen (1973).

*Department of Botany, Punjabi University, Patiala 147002, India.

AMERICAN FERN JOURNAL: VOLUME 68 (1978)

FIGS. 1-4. Anatomical details of Equisetum diffusum. F\G. 1. Partial transection of tuber pres ie peripheral thick-walled cells and ring of vascular bundles. FIG. 2. Section of outer portion ° pees Showing epidermis and cortex. FIG. 3. Long, papilliform hairs. FIG. 4. Vascular bundle and s rounding tissue of tuber.

LITERATURE CITED nv BARRATT, KATE. 1920. A contribution to our knowledge of the vascular system of the genus Equisetum. Ann. Bot. 34: 201-235, t. VI-VII. sae CAMPBELL, D. H. 1918. The Structure and Development of Mosses and Ferns (Archegoniatae), 3rd ed. Macmillan, New York. HAUKE, R. L. 1974. The taxonomy of Equisetum: an overview. New Bot. 1: 89-95. ae MEHRA, P.N. and S. S. BIR. 1959. A note on chromosome numbers in some Indian species 0 Equisetum. Amer. Fern J. 49: 86-92. dk SEN, T. and U. SEN. 1973. Morphology and anatomy of Equisetum diffusum D, Don an ie ramoOsissimum Desf. subsp. debile (Roxb.) Hauke with a discussion on their taxonomy. Israel J. Bot. 22: 166-174,

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 2 (1978) 57

A New Species of Asplenium from Guatemala ROBERT G. STOLZE*

Asplenium is one of the largest and most interesting genera of ferns in the neotropics. Many of its species are distinctive and quite easy to identify, but a few others, notably A. auritum Swartz, A. cuspidatum Lam., and A. radicans L., are so highly variable that intensive monographic studies will be required to define their specific and infraspecific limits. During my studies of the genus for the ‘*Ferns and Fern Allies of Guatemala,’’ a total of 40 species (including a number of varieties) have been recognized in this small Central American country. One of these is new

Asplenium williemell Stolze, sp. Figs. 1- 4. Rhizoma erecta, paleacea; Tale lanceolatae vel lineares, fuscae, clathratae, 4-6 mm longae, 0. 5 0. 8 mm latae, plerumque attenuatae; folium pinnatum, 15-45

cm longum, 4- 6 cm m latum, 20 apicem pinnatifidum vel serratum gradatim decres- cens; petiolus 4-9 cm longus, plumbeus vel fuscus, anguste vel late alatus; pinnae 14- 20(22)-j -jugae, obtusae vel subacutae, serratae vel biserratae; venae acro- eee nei plerumque 1-furcatae, venae basiscopicae ie aa sori lineares, 3-8 mm longi, 0.5-0.8 mm lati, 1-2 sori proximales diplazioides

ers Terrestrial in cloud forest, Montana Canahui, eee, El Progreso, Guatemala, alt. 1,600-2,300 m, Steyermark 4379] (US; isotype F).

In wet forests, commonly on the forest floor, but rarely epiphytic, 1,250-2,300 m; Alta Verapaz; Baja Verapaz; El Progreso; El Quiché; San Marcos; Santa Rosa. Mexico (Chiapas).

Plants terrestrial, rarely epiphytic; rhizome stout, erect, amply provided with lanceolate or linear, lustrous, grey-brown, clathrate scales, these 4-6 mm long, 0.5-0.8 mm broad, mostly attenuate; leaves pinnate, subcaespitose, mature ones 15-45 cm long, 4-6 cm broad; petiole stout, 4-9 cm long, much shorter than the lamina, dull grey or grey-brown, glabrous, abaxially terete, adaxially flattened and narrowly to broadly green-alate (especially toward the lamina), each of the wings 0.3-0.8 mm broad; lamina linear to narrow-elliptic, glabrous, thin- to firm- membranaceous, slightly reduced at base, gradually reduced to a pinnatifid or serrate apex, not proliferous; rachis glabrous, dull grey or reddish brown, green- alate throughout; pinnae 14-20(22) pairs, the middle ones 2-3.5 cm long, 0.8-1.5 cm broad, sessile to short-stalked, spreading to slightly ascending, approximate to subdistant, oblong to lanceolate, obtuse to subacute, inequilateral at the base, basiscopically cuneate or excavate, acroscopically truncate and often auriculate or subauriculate, the margins obtusely or subacutely serrate to biserrate; veins on the acroscopic side commonly once-forked, the basal one twice-forked, the distal ones and those of the basiscopic side simple, distinct adaxially, indistinct abax- ially, the tips not or scarcely enlarged, ending well short of the margin; sori relatively long, often nearly reaching from midrib to margin (but tending to be more inframedial), linear, straight to slightly curved, 3-8 mm long, 0.5-0.8 mm broad, 1-2 proximal ones commonly double (diplazioid); indusium delicate, linear, pale yellowish to light brown, or hyaline, subentire.

*Department of Botany, Field Museum of Natural History, Chicago, IL 60605.

AMERICAN FERN JOURNAL: VOLUME 68 (1978)

R. G. STOLZE: NEW ASPLENIUM FROM GUATEMALA 59

SELECTED SPECIMENS EXAMINED:

MEXICO: Chiapas: Ridge along logging road from Las Margaritas to Campo Alegre, Municipio La Independencia, elev. 2,300 m, Breedlove 33685 (DS, F). GUATEMALA: Alta Verapaz: Epiphytisch, Coban, 1,350 m. Tuerckheim I1-1853 (US). Baja Verapaz: Pee Sie broadleaf montane cloud forest, Sierra de las Minas, 3 km SE of Purulha, alt. 1,800 m, L. O. Williams et al. 43279 ise El Quiche: San Miguel Uspantan, alt. 6,000 ft, Heyde & Lux 3235- B i San Da On forest floor; slopes of Tajumulco Volcano, 8-10 km west of San Marcos, alt. ca. 2,300 m, L. Wiliams et al. 26853 (F).

osa: Santa Rosa, alt. 4,000 ft, Heyde & Lux 3234 (US, in part; dB sheet of this at US is A. abscissum Willd.)

This is rather closely related to the neotropical species of A. harpeodes Kunze, A. miradorense Liebm., and A. pteropus Kaulf. All, in turn, form part of a larger complex of New and Old World species related to A. erectum Bory ex Willd., the latter reputed to be confined (at least in the strict sense) to the Old World. These taxa form a confusing tangle of species and/or varieties, which will be unraveled only when collections and types from around the world are brought together for comparison. So it is with some reluctance that I describe yet another species (albeit a distinct one), thus adding one more name to the complex.

Characters which are most useful in separating A. williamsii from its nearest relatives are the lustrous, grey-brown rhizome scales, which are 4-6 mm long, the conspicuously alate petiole and rachis, the relatively few (14-20) obtuse to sub- acute pinnae with mostly biserrate margins, veins which are commonly (acroscop- ically) once-forked, and the sori, most of which are very long and crowd the costa. An even more significant feature is the common occurrence of back-to-back (di- plazioid) sori, which are borne usually on the basal acroscopic vein.

Asplenium harpeodes has castaneous or reddish brown scales with usually filiform tips, nonalate petioles, and numerous attenuate pinnae with mostly simple veins and marginal serrations. Asplenium miradorense has dull, reddish brown scales only 2-3 mm long, 20-35 pairs of pinnae, and relatively short, medial sori. Asplenium pteropus has the conspicuously alate petiole and rachis of A. williamsii, but the rhizome scales are castaneous to dark brown, the 20-30 pairs of pinnae are simply serrate and simple-veined, and the sori are relatively short and medial.

The new species is named in honor of Dr. Louis O. Williams, former chairman of the Department of Botany at the Field Museum, whose field work and publica- tions form the backbone of the ‘‘Flora of Guatemala’”’ project.

FIGS. 1-4. Asplenium williamsii. ee 1. Habit, x 1/2. FIG. 2. Base of lamina showing reduced basal pinnae and alate petiole, x 3. FIG. 3. A central pinna with a double sorus, x 3. FIG. 4. Portion of rhizome and a chia of scales among sate bases, x 3

60 AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 2 (1978)

SHORTER NOTES

A NEW LOCATION FOR PELLAEA GLABELLA IN MINNESOTA.—During July 1977, while studying pteridology under Prof. W. H. Wagner, Jr., of the University of Michigan at the University of Minnesota Biological Station, I found a new locality in Minnesota for the Smooth Cliff Brake, Pellaea glabella Mett. ex Kuhn. The species was discovered on July 10 and a specimen collected on July 18. A colony of five small plants was found in a precarious location at the top of a rock outcropping about 30 m above Highway 61 and overlooking Lake Superior. The locality is in Lake County, approximately two miles north of Illgen City, and about 55 miles north of Duluth. A voucher, Weber ], has been deposited in the herbarium of the University of Minnesota (MIN). The northernmost station pre- viously known for P. glabella in Minnesota is about 130 miles to the southwest in Chisago County between Minneapolis-St. Paul and Duluth, according to Tryon’s ‘The Ferns and Fern Allies of Minnesota’ (1954, p. 52). According to Billington’s ‘Ferns of Michigan’’ (1952, p. 211), there is also a station in Ontonagon County in the upper peninsula of Michigan that is about 100 miles southeast across Lake Superior from the Lake County, Minnesota locality. —Larry A. Weber, 415 W. St. Louis St., Pacific, MO 63069.

SOME INSECT INTERACTIONS WITH AZOLLA MEXICANA.—It is com- mon knowledge that ferns are not very much affected by insect predation due to the high content of chemical repellents within the plants. Still, not enough is known about the relations of ferns and insects to make this statement an invari- ably valid generalization. In search of examples of insect-fern interactions, I came upon a mat of Azolla mexicana Schlecht. & Cham. and Lemna sp. in the Rio Potrero, Province of Guanacaste, Costa Rica, where a yellow solitary wasp, Polybia rejecta (F.) forma belizensis Cameron, was observed hovering and alight- ing on the Azolla plants, wandering about, and inserting its head among the com- pact leaves. When a suitable spot was found by the wasp, it went completely pail oi upturned the Azolla, and then searched among the roots of the fern. rage of the P olybia wasp immediately after this search-submerge-catch opera- rae rought to light an interesting case of interaction. Two genera of beetles of the atta breed their larvae among the Azolla roots, which afford shelter restricted = . eetles. The wasp preys on these larvae, although its diet is not Dryopidae fare em, as these wasps are rather opportunistic. Whether the asus ea ae are a natural control of Azolla populations in this habitat is not

wn. But if so, Polybia rejecta may be responsible for controlling the Dryopid

Seca and thus may affect the Azolla population dynamics. Or perhaps the

ne Sey p only an occasional factor in the biology of the Dryopidae

is known are Fas ‘a Apparently only one other record of submerging wasps

Washington igs rom eastern North America by Caudell (Proc. Entom. Soc.

Nacional d os oe 1922).—Luis D. Gomez P., Herbario Nacional, Museo onal de Costa Rica, Apartado 749, San José, Costa Rica.

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 2 (1978) 61

NOTES ON NORTH AMERICAN LOWER VASCULAR PLANTS.—Field work in Mexico and the examination of herbarium specimens at ARIZ, ASU, LL- TEX, and SRSC have revealed several new state records, a new record for the United States, and collections of apparently rare species.

Dryopteris cinnamomea C. Chr. has been found new to Texas and the United States. The collection data are: In a cave near Comstock, Val Verde Co., Texas, 980 ft elev., rare, 10 Sep 1965, C. Babcock 100 (SRSC, 2 sheets). These speci- mens were filed as Cystopteris fragilis (L.) Bernh. The species previously was known only from Mexico (Chihuahua, Durango, San Luis Potosi, Guanajuato, Hidalgo, Distrito Federal, Morelos, and Michoacan), according to Knobloch and Correll’s ‘‘Ferns and Fern Allies of Chihuahua’”’ (1962, p. 173). I have also seen material from Coahuila.

A second known collection of Notholaena jacalensis Pray and one new to San Luis Potosi has been made. The collection data are: Immediately N of Minas de San Rafael, San Luis Potosi, Mexico, ca. 22°13'N, 100°16’W, growing with Hech- tia, Agave lecheguilla, Helietta parvifolia, and Neospringlea integrifolia in highly mineralized soil, 1100 m elev., 30 Jun 1972, M. C. Johnston 8178C (LL-TEX). This species previously was known only from Jacala, Hidalgo, Mexico, according to Pray (Amer. Fern J. 57: 101. 1967).

Pellaea breweri D. C. Eaton has been found new to Colorado. The specimen data are: Fire Lookout, summit of Roundtop Mountain, Dinosaur National Monument, Moffat Co., Colorado, in rock crevices of N-facing cliff of saddle W of lookout, 2800 m elev., 27 Jun-1 Jul 1948, R. A. Wolf & K. S. Dever 5206 (LL-TEX). This species previously was known from Washington, Oregon, California, Idaho, Nevada, Utah, and Wyoming, according to A. F. Tryon (Ann. Missouri Bot. Gard. 44: 138. 1957). The NW Colorado locality represents only a short range extension from Wyoming stations.

Recently I made the second known collection of Selaginella macrathera Weath. in I. M. Johnst. The collection data are: Chihuahua Viejo, Sierra Mapula, Chihuahua, Mexico, ca. 28°33’N, 105°51'30’W, on N-facing, rocky slopes and summit, grassland with scattered oaks, and the ledges of outcrop cliffs, 5800-7300 ft elev., 20 Jul 1977, T. Reeves 5745B (ASU, GH). The species was known previously only from the type collection, according to R. M. Tryon, Jr. (Ann, Missouri Bot. Gard. 42: 42. 1955), which is about 90 mi ENE of the new station.

Selaginella leucobryoides Maxon has been found new to arizona and Nevada. The collection data are: Virgin Narrows, Mohave Co., Arizona, Sec. 32, T41N, R14W, N and E exposures on limestone cliffs and steep, rocky slopes, in desert shrub vegetation with Larrea, Ephedra, Thamnosma, Hilaria, Ferocactus, Echinocereus, Opuntia, Galium, and mixed with Cheilanthes parryi, Ca. 2000 ft elev., 10 Sep 1977, R. K. Gierisch 3983A (ASU); and Red Rock Canyon, Spring Mountains, Clark Co., Nevada, shaded and damp N-facing cliff, 4800 ft elev., 25 Nov 1967, V. Bostick s.n. (ARIZ). This species was previously known only from the Providence and Panamint Mountains of SE California, according to P. A. Munz (A Flora of Southern California, p. 14, 1974). The species is apparently endemic to the mountains of the Mohave Desert. This species should be added to

62 AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 2 (1978)

the list of lower vascular plants to be expected in the New York Mountains of SE California, according to C. D. MacNeill, W. Brophy, and A. R. Smith (Madrono 25: 54-57).

Travel was supported by NSF Dissertation Improvement Grant 77-00182 to Dr. D. J. Pinkava and the author. I thank Dr. A. F. Tryon for examining the Pellaea specimen and Dr. R. M. Tryon, Jr. for examining the material of Selaginella. 1 thank the curators of the cited herbaria for permission to examine their collections. —Timothy Reeves, Department of Botany and Microbiology, Arizona State University, Tempe, AZ 85281.

THE FERNS OF SAN SALVADOR ISLAND, II.—In the Shorter Notes of the American Fern Journal, vol. 65, p. 63, five species of ferns were reported from San Salvador Island, The Bahamas. These were Acrostichum danaeifolium Langsd. and Fisch., Asplenium dentatum L., Nephrolepis exaltata (L.) Schott, Pteridium caudatum (L.) Maxon, and Thelypteris kunthii (Desv.) Morton (as T. normalis),

Since that report, two arduous trips to the interior of the island have produced six more species not previously recorded for San Salvador Island. Five species were found in a coppice southeast of Guana Cay, the north central part of the island. The sixth species was found in the vicinity of Farquharson’s plantation ruins, the southeastern part of San Salvador Island.

Adiantum tenerum Swartz was infrequent on the wall of a limestone pit in the re east of the mangrove swamp and southeast of Guana Cay (R. R. Smith et al.

Campyloneurum phyllitidis (L.) Presl was found growing around the base of Bourreria ovata in the coppice southeast of Guana Cay (R. R. Smith et al. 56, 4071).

Phlebodium aureum (L.) J. Smith was locally frequent on the upper stems of Sabal palmetto located along the margins of sink holes in the coppice southeast of Guana Cay (R. R. Smith et al. 58).

_ Polypodium polypodioides (L.) Watt was occasional on the bases of tree trunks in the coppice southeast of Guana Cay (R. R. Smith et al. 76).

Tectaria lobata (Poir.) Morton was found only once in a limestone pit just off oe trail which connects Farquharson’s plantation ruins to the southern end of the eee The fern was collected about fifteen feet below the surface of a Smith et al. SER OO was approximately eight feet (R. R. Pikes gs Hee Meee he occasional on the upper stems of ‘i Guana Cay (R. R. Smith et a. 7). ee Oe ahi collections cited are deposited in the Hoysradt Herbarium of Hartwick

oNege (HHH), Oneonta, N.Y.—Robert R. Smith and Joyce E. Mauk, Depart- ment of Biology, Hartwick College, Oneonta, NY 13820.

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 2 (1978) 63

CHEILANTHES MICROPHYLLA, A GENUS AND SPECIES NEW TO THE BAHAMA ARCHIPELAGO.—The finding of C. microphylla (Swartz) Swartz on Grand (Middle) Caicos Island in the Bahama Archipelago raises to 44 the number of species of ferns and fern allies now known to occur in this region. This species also occurs in the southeastern United States, the Greater and Lesser Antilles, the Cayman Islands, and Mexico. The collection was made, in company with Ruben Sauleda and Patricia Adams, on rocks in partial shade about the mouth of caves on Village Hill, between the airstrip and Bambarra, Grand (Middle) Caicos Island, 12 Feb 1978, D. S. Correll 49461 (F, FTG, IJ, MBG, NY, US).— Donovan S. Correll, Fairchild Tropical Garden, Miami, FL 33156.

THE CHROMOSOME NUMBER OF NOTHOLAENA COCHISENSIS.—Love, Love, and Pichi Sermolli (Cytotaxonomical Atlas of the Pteridophyta, 1977) have recorded the chromosome numbers of several species of Notholaena. They synonymized N. pruinosa and N. integerrima under N. sinuata, both of which can be separated from N. sinuata. These species were dealt with by Knobloch, Tai, and Ninan (Amer. J. Bot. 60: 92-95. 1973), who stated that the chromosome number of N. cochisensis Goodding had not been ascertained.

oe

| he a

FIG. 1. Spore mother cell of Notholaena cochisensis (Knobloch 2523, MSC) at meiosis with about 87 bivalents, x 1666.

In 1977 we obtained a meiotic count shown in figure 1 of about 87 bivalents in a triploid plant of N. cochisensis collected in 1973 from McKelligan Canyon within the city limits of El Paso, Texas (Knobloch 2523, MSC). This plant had no more than 32 fertile spores in each sporangium and presumably is apogamous. The sporophytic number should be the same as the gametophytic number, as is the usual case in apogamous ferns. According to Foster and Gifford (Comparative Morphology of Vascular Plants, 1974), 32-spored sporangia may arise in at least three ways (two meiotic and one mitotic), but the type involved here is not known.

Traditionally, the nearest relatives of N. cochisensis are N. sinuata, N. integer- rima, and N. pruinosa. All of these are apogamous triploids and have n=2n ef 87 chromosomes. Whether these plants are hybrids between extant or extinct species cannot be stated with certainty until such hybrids have been synthesized. We thank Dr. Donald M. Britton for confirming our opinions.—I/rving W. Knobloch and William Tai, Department of Botany and Plant Pathology, Michigan State University, East Lansing, M1 48823.

64 AMERICAN FERN JOURNAL, VOLUME 68 NUMBER 2 (1978)

REVIEW

‘*THE PTERIDOPHYTE FLORA OF FIJI,’’ by G. Brownlie. J. Cramer, Post- fach 48, D-3301 Lehre, Federal Republic of Germany. 1977. 397 pp. DM 200.— The value of this new pteridophyte flora is threefold. It draws upon a considerable amount of recent field work in Fiji by the author himself, it updates the preceding account of Fijian pteridophytes by Copeland (1929) which appeared just before the explosion of taxonomic and nomenclatural modification in pteridophytes that continues today, and it complements the pteridophyte floras of Samoa (Christen- sen, 1943) and New Caledonia (Brownlie, 1969), which lie on either side of Fiji.

The work provides keys, descriptions, ecological information, line drawings, etc.; there are 32 new species or names and many range extensions and reinterpre- tations. Altogether it makes a very handsome volume and must be rated as a substantial contribution to pteridology.

Unfortunately it is not a polished work, although it would only have required a relatively small additional investment of effort to make it such. The introduction is inadequate, lacking any tabulation of Species or novelties, any phytogeographic information and analysis, and any general account of the vegetation. The area of the flora is not clearly delimited; Rotuma Island, at least politically part of Fiji, is ignored.

The treatment of Selaginella is admittedly tentative and incomplete. Marsilea is omitted, although it was collected in Fiji by Horne according to Baker. An addi- tional species is Pronephrium asperum (Presl) Holttum, represented by Gillespie 3880 (MICH).

A varietal name is elevated to replace the legitimate Dennstaedtia intermedia, despite the fact that names have no priority outside their rank. Dryopteris maxima Is transferred to Arachniodes while D. subarborea is maintained in Dryopteris, although the two are so closely related that Christensen opined they were only geographic variants. The separation of D. maxima is based solely on what in this Species complex is an unstable aspect of frond architecture and which in no way demonstrates a relationship with Arachniodes.

made as revisionary work proceeds on ‘‘Flora Malesiana.”’

Brownlie’s major error w as choice of publishers. The ti ; come for authors to abandon the profit- P ime has co

of printed matter,—. G. Price Ann Arbor, MI 48109.

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-_ ERICAN Volume 68 FERN ns J O U R N A L July-September, 1978

QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY

Spread of the Exotic Fern Lygodium microphyllum in Florida CLIFTON E. NAUMAN and DANIEL F. AUSTIN

Chlorophyll and Lipid Changes on Germination in the Non-green Spores of Thelypteris dentata ALLEN V. SEILHEIMER

Gametophytes of Botrychium multifidum as Grown in Axenic Culture ERNEST M. GIFFORD, JR. and DOROTHY D. BRANDON

Revision of the Genus Cochlidium (Grammitidaceae) L. EARL BISHOP

Shorter Notes: A Deletion from the Pteridophyte Flora of Nebraska; Cystopteris tennesseensis in Alabama; Equisetum x litorale Recorded for Minnesota; Lycopodium cernuum in Louisiana

MISSOURE: BOTARNGRL,

OCT 18 978

GARDEN LIBRARY

OS aA

nN ~

~J —

The American Fern Society Council for 1978

RICHARD L. HAUKE, Dept. of Botany, University of Rhode Island, Kingston, R.I. 02881. : President ROBERT M. LLOYD, Dept. of Botany, Ohio University, Athens, Ohio 45701. Vice-President TERRY R. WEBSTER, Dept. of Botany, University of Connecticut, Storrs, Conn. 06268. Secretary JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916. Treasurer JUDITH E. SKOG, Dept. of Biology, George Mason University, Fairfax, Va. 22030. ecords Treasurer DAVID B. LELLINGER, Smithsonian Institution, Washington, D.C. 20560. Editor-in-Chief

JOHN T. MICKEL, New York Botanical Garden, Bronx, N.Y. 10458. Newsletter Editor American Fern Journal EDITOR-IN-CHIEF

DAVID B. LELLINGER Smithsonian Institution, Washington, D. C. 20560 ASSOCIATE EDITORS

DAVID W. BIERHORST -.Dept. of Botany, University of Massachusetts, Amherst, Mass. 01002

GERALD J. GASTONY ............. Dept. of Biology, Indiana University, Bloomington, Ind. 47401

JOHN T. MICKEL .. New York Botanical Garden, Bronx, New York 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 Institu- tion, Washington, DC 20560. Second-class postage paid at Washington.

Matter for publication and claims for Missing issues (made within six months of the date of issue) should be addressed to the Editor-in-Chief.

Changes of address, dues, and applications for membership should be sent to Dr. J. E. Skog, Dept. of Biology, George Mason University, Fairfax, Va. 22030.

Orders for back issues should be addressed to the Treasurer.

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, $5.00; sustaining membership, $10.00; life membership, $100.00). Extracted offprints, if ordered in advance, will be furnished to authors at cost, plus postage

Back volumes $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. Ten percent discount on orders of six volumes or more; postage additional.

Library New York Botanical Garden, Bronx, New York 10458, is Librarian. Members any time, the borrower paying all shipping costs.

Newsletter Dr. John T. Mickel, New York Botanical Garden, Bronx, New York 10458, is editor of the newsletter “Fiddlehead Forum.”’ The editor welcomes contributions from members and non- mbers, including miscellaneous notes, offers to exchange or purchase materials, personalia, hor- tcultural notes, and reviews of non-technical books on ferns.

Spore Exchange

Mr. Neill D. Hall, 1230 Northeast 88th Street, Seattle, Washington 98115, is Director. Spores exchanged and collection lists sent on request.

Dr. John T. Mickel, may borrow books at

Gifts and Bequests

Gifts and bequests to the Society enable it t SETS siney ‘. 4 ta others interested

Pear ee books, back issues of the Journal, and cash or other gifts are always welcomed, and ax-deductible, Inquiries should be addressed to the Secretary

Se

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 3 (1978) 65

Spread of the Exotic Fern Lygodium microphyllum in Florida CLIFTON E. NAUMAN and DANIEL F. AUSTIN*

Lygodium microphyllum (Cav.) R. Br. (Schizaeaceae) was first reported as an adventive in Florida about a decade ago by Beckner (1968). The plants were known to Beckner from three collections in Martin County near Jonathan Dickin- son State Park and a single collection in Palm Beach County from Delray Beach. These climbing ferns should no longer be considered adventive, since they are now naturalized in both these counties. On the fringes of their range, the plants occur mostly in small clumps, while toward the center of the distribution near the Loxahatchee River and Loxahatchee Slough (Fig. /), Lygodium may cover acres. One colony in Palm Beach County (Sect. 12, T44S, R41W) was one-quarter of a mile long and about 200 yards wide in January 1978.

In Florida, the plants are confined to wet, disturbed sites. We have found them only near canals, rivers, ditches, in disturbed swamps, and other sites which have standing water for a large part of the year.

We have not determined when the plants were introduced, although the oldest collection we have seen was made in February 1958 (R. A. Long, FLAS). The apparently oldest center of dispersal for L. microphyllum is in the Loxahatchee River area. Large and seemingly old colonies are abundant, suggesting that the plants have been there longer than the past two decades. Another, apparently younger, focal point is in southeastern Palm Beach County. Perhaps the small colony (Fig. 1) was started by the plants being cultivated in nurseries in the 1950’s. If, as we have assumed, the ferns first became established in the Lower Loxahatchee River area, how they were dispersed upstream is unknown. Spores might have been spread accidently by birds, since young plants often appear first in small isolated patches on the margins of cypress heads.

Two other members of the genus have been reported in Florida, L. japonicum (Thunb.) Swartz and L. palmatum (Bernh.) Swartz. The former is an Asian fern naturalized from the Carolinas to Texas (Radford et al., 1968; Correll & Johnston, 1970). An old collection of L. japonicum in Dade County was considered to have escaped on vacant lots, but we suspect that it was only persistent from cultivation. We have not seen recent populations in southern Florida. Lygodium palmatum is considered a native of the eastern United States, ranging from Massachusetts to the Carolinas and Kentucky (Radford, et al., 1968). Although reported in Georgia and Florida (Small, 1938), we have seen no specimens.

*Department of Biological Sciences, Florida Atlantic University, Boca Raton, FL 33431.

Volume 68, number 2, of the JOURNAL was issued July 11, 1978.

66 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

At present the two Asian species L. japonicum and L. microphyllum are well established in Florida. Lygodium japonicum is naturalized only in northern Florida and L. microphyllum only in southern Florida. Both seem to be spreading and eventually may meet. In southern Florida, L. microphyllum has already af- fected the native vegetation by smothering shrubby and herbaceous plants.

ST. LUCIE CO.

LOCATION MAP

MARTIN CO.

PALM BEACH CO.

BROWARD CO. Set 1. Distribution of the Climbing Fern Lygodium microphyllum in pepe: Florida. Stars oe the points where the plants were known i in the late 1960's. Dots are sites for which voucher cena now exist. Open circles are where plants have been seen se no vouchers have been

ITERATURE CITED sai hres J. 1968 Lygodium nese, another fern escaped in Florida. Amer. Fern J. 58: CORRELL, D. S. and M. C. JOHNSTON. 1970. Manual of the Vascular Plants of Texas. Texas . Research Foundation, Renner, T sepiarn te . E., H. E. AHLES and C. R. BELL. 1968. Manual of the Vascular Plants of the arolinas. Univ. N. Carolina Press, Chapel Hill, NC.

SMALL, J. K. 1938. Ferns of the So . utheastern Stat Reprinted by Hitser, New Vork: inet; ates. Science Press, Lancaster, PA. (Rep

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 3 (1978) 67

Chlorophyll and Lipid Changes on Germination in the Non-green Spores of Thelypteris dentata’ ALLEN V. SEILHEIMER*

A number of morphological, biochemical, and cytochemical investigations comparing dormant fern spores with 2-celled germinated spores have been re- ported. Protein and lipid are the major energy reserves in dormant spores. Changes in storage protein (Towill & Ikuma, 1975), protein bodies (Gantt & Arnott, 1965), and lipid (Robinson et al., 1973; Towill & [kuma, 1975; Gemmrich, 1977) have been investigated during germination. However, most of these investi- gations have utilized a few species of taxonomically unrelated ferns which have chlorophyll-containing dormant spores (Lloyd & Klekowski, 1970). Proof of the presence of chlorophyll has been shown by the absorption spectra of intact Onoclea_ sensibilis spores (Towill & Ikuma, 1973) and by extraction of Polypodium vulgare spores (Robinson et al., 1973). Lloyd and Klekowski (1970) have shown that chlorophyll-containing fern spores are characterized by short viability and a relatively rapid germination rate.

A second—but much more widespread—type of fern spore, believed to be non-chlorophyllous, has remained largely unstudied. This type of spore is charac- terized by an absence of green pigmentation, long viability and a slow germination rate. The only evidence that non-green fern spores are devoid of chlorophyll is that they lack green pigmentation when observed in the light microscope (Lloyd & Klekowski, 1970)

This investigation was undertaken to determine if the non-green spores of Thelypteris dentata contain chlorophyll or lipids associated with the photosyn- thetic apparatus. Chlorophyll content, lipid classes, and fatty acid compositions of the dormant and germinating spores are compared.

MATERIALS AND METHODS

Fronds of Thelypteris dentata (Forssk.) E. St. John were collected from greenhouse-grown plants and placed abaxial surface down on paper. Spore release occurred within two hours. Spores were sifted through lens paper to remove sporangial debris. 200 mg of spores were surface sterilized with an 0.5-1% com- mercial bleach solution and sown on sterile, modified Knop’s medium (Gantt & Arnott, 1965). Germinated spores consisting of a prothallial and rhizoidal cell were obtained in 4 days under continuous fluorescent light (ca. 1500 lux) at about 2

Dormant spores weighing 200 mg, or the germinated spores derived from 200 mg of spores, were homogenized dry in a ground-glass tissue grinder for 5 min- utes, moistened with water for 5 minutes, and homogenized with chloroform-

oe ea gt of Botany, 220 Biological Sciences Center, University of Minnesota, St. Paul, MN

'This study i or : k erebape of the author’s Ph.D. thesis and was carried out at the Hormel

eS RS Austin, MN with the technical assistance xe L. Gellerman and W. H. Anderson. alg ony upport was Se ried te U. S. Public Health Service Grant AM 05165 from Ke N.I.H.

Schlenk. I thank my adviser, Dr. David 5 Mei nughtin for helpful discussions and soe Ae

68 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

methanol (2:1, v/v) for 5 minutes. After filtration, the residue was extracted again with 2:1 chloroform-methanol. Re-extraction of the residue with chloroform- methanol (1:1, v/v) was carried out two times to insure complete extraction of phospholipids. Filtrates were combined and purified according to the methods of Folch et al. (1957). The chloroform phase of the extract was evaporated to dry- ness under vacuum. Methanol was repeatedly added to the residue and re- evaporated to remove traces of water. The lipid yield of dormant spores was determined at dryness in tared glass tubes.

Thin-layer chromatography was used to identify and estimate the amounts of lipid classes present in spores. Reference compounds, solvents, spray reagents, and techniques were the same as those described by Gellerman et al. (1972). Known quantities of standards were compared with known quantities of samples (400 ug) to estimate the amounts of the lipid classes present.

Techniques used to determine the fatty acid composition of spores including saponification, esterification, preparative thin-layer chromatography of the lipids to remove pigments, and gas-liquid chromatographic analysis are described else- where (Gellerman et al., 1972). Results were checked against reference materials of known composition. Identifications and quantifications of methyl esters were made by measuring equivalent chain lengths and peak areas from gas-liquid chromatography.

Chlorophyll content relative to the amount of lipid present (weight per cent

lipid) was determined spectrophotometrically at 652 4m (Bruinsma, 1961). RESULTS AND DISCUSSION

_ The fresh dormant Thelypteris dentata spores contain approximately 50%

lipophilic material. Most of the lipid is located in large circular lipid droplets that

dominate the cytoplasm (Seilheimer, 1975). The large relative amount of tri-

TABLE 1. LIPID CLASSES AND CHLOROPHYLL CONTENT OF DORMANT AND GERMINATING SPORES OF Thelypteris dentata.

S % Lipid Lipid Classes , sitaie peek

inatin ce Lipids Dorman Germinating riglycerides Sterols and Diglycerides (free) ia apie Carotenes, qualene, Wax, and Esters a ea ws Pe onogalactosyl Diglyceride 5% Digalactosyl Diglyceri i 20% Siloings — ion Phospholipids a ; osphatidyl Glycerol Phosphatidy! Choline 0.2% ere Phosphatidyl Inositol ace 0.5% Chlorophyll (weight % lipid) ae 1.1%

glycerides (Table 1) present in dormant spores is presumably located in lipid droplets. Triglycerides have been reported as major energy reserves in other fern Spores (Robinson et al., 1973: Gemmrich, 1977), moss spores (Karunen, 1971; Gellerman et al., 1972), and certain seeds (Appelqvist, 1975).

A. V. SEILHEIMER: SPORE GERMINATION OF THELYPTERIS DENTATA 69

The fatty. acid of dormant spores consisted primarily of palmitic, oleic, and linoleic acids ( Table 2). The most abundant fatty acids of spores are believed to be components of the triglycerides similar to what was reported in spores of Polypodium vulgare (Robinson et al., 1973) and Anemia phyllitidis (Gemmrich, 1977).

Traces of the glycolipids, monogalactosyl diglyceride, and digalactosyl di- glyceride were detected in the dormant spores. The dormant spores also con- tained a trace of phosphatidyl! inositol and a low relative amount of phosphatidy! choline (Table 1). Phosphatidy! choline has been reported in envelope membranes of proplastids (Leese & Leech, 1976) and in mitochondria (Schwertner and Biale, 1973). This finding agrees with the ultrastructural observation of proplastids and mitochondria in the dormant spores (Seilheimer, 1975).

TABLE 2. FATTY ACID COMPOSITION OF TOTAL LIPID EXTRACTS

FROM DORMANT AND GERMINATING SPORES OF Thelypteris dentata. Spores (% of Total Fatty Acids)

Fatty Acids Dormant Germinating

Palmitic (16:0) 18.7 20.0 Palmitoleic (16:1) 0.3 0.6 Stearic :0) 3.8 4.6 Oleic (18:1) 46.9 45.0 Linoleic (18:2) 28.5 27.5

linolenic (18:3 6) 1.1 1.2 Linolenic (18:3 3) 0 1.8 Behenic (22:0) trace 0.1 Arachidonic (20:4 @ 6) h 0.6 Lignoceric (24:0) trace 0.2

No chlorophyll was detected in the dormant spores of T. dentata (Table 1), nor have chloroplasts been reported in the cytoplasm (Seilheimer, 1975). Further- more, no significant amounts of lipids associated with chloroplasts, such as monogalactosyl diglyceride, digalactosyl diglyceride, or phosphatidyl glycerol (Leese & Leech, 1976), were detected in the dormant spores ( Table 1). Analysis of non-green spore lipids from the fern Anemia phyllitidis also has shown a lack of diglycerides (Gemmrich, 1977). These results differ from those reported for the dormant spore of Polypodium vulgare, where chlorophyll, phospholipids, and glycolipids, which are normally associated with chloroplasts, are present (Robin- son et al., 1973)

Triglycerides (Table 1), presumably composed of palmitic, oleic, and linoleic fatty acids (Table 2), were also major components of the germinated spores. Similar results were reported in studies of lipid in other germinating fern spores (Robinson et al., 1973; Gemmrich, 1977). This finding suggests that large quan- tities of lipid reserves remain unutilized during the germination process. How- ever, a decline in triglyceride level was observed in 12 to 15 day old, multicellular gametophytes of Polypodium vulgare (Robinson et al., 1973) and Anemia phyl- litidis (Gemmrich, 1977).

Chlorophyll was present in the germinated spores ( Table 1), as were significant relative amounts of glycolipids and phospholipids (Table 1) that form structural components of chloroplasts and mitochondria. Monogalactosyl diglyceride and digalactosyl diglyceride are associated with grana formation in chloroplasts

70 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

(Leese & Leech, 1976) and chloroplast envelopes (Bahl et al., 1976). Their pres- ence is also reported in mitochondria (Schwertner & Biale, 1973). spe are found in chloroplast ehesieaes stroma, and grana (Bahl et al., 1976). phatidyl glycerol is a major component of chloroplast thylakoid cneiitbsilal (Leese & Leech, 1976). Bidepiatity) choline is reported in mitochondria (Schwertner & Biale, 1973) and chloroplasts (Leese & Leech, 1976). The only significant change in the fatty acid composition during germination of 7. dentata was an increase in linolenic acid (Table 2). Linolenic acid is a major fatty acid of photosynthesizing tissue and probably is a structural element of the chloroplast (Hitchcock & Nichols, 1971).

This investigation of T. dentata substantiates the light microscopic observa- tions and comments of Lloyd and Klekowski (1970) that the non-green, dormant spores of ferns lack both chlorophyll and lipid compositions associated with the photosynthetic apparatus. Chlorophyll and lipid composition indicative of chloro- plasts were observed after spore germination.

LITERATURE CITED APPELQVIST, L-A. 1975. Biochemical and structural aspects of storage and membrane lipids in 2a loping oil seeds. Jn T. Galliard and E. I. Mercer (eds.). Recent Advances in the Chemis- and Biochemistry of Plant Lipids. Academic Press, London. BAHL, “i "8. FRANCKE, and R. MONEGER. 1976. Lipid composition of envelopes, prolamellar bodies and other plastid membranes in etiolated, green and greening wheat leaves. Planta 129: 1

193-201.

BRUINSMA, J. 1961. A comment on the spectrophotometric determination of chlorophyll. Biochim. Biophys. Acta 52: 576-578.

FOLCH, J., M. LEES, and G. H. SLOANE-STANLEY. 1957. A simple method for the isolation and [pte of total lipids from animal tissues. J. Biol. Chem. 226 -509

GANTT, E., RNOTT. 1965. Spore germination and development of the young

patie. of the ostrich fern (Matteuccia struthiopteris). Amer. J. Bot. 52: 82-94.

GELLERMAN, J. L., W. H. ANDERSON, and H. SCHLENK. 1972. Highly unsaturated lipids of Mnium, Polytrichum, Marchantia, and Matteuccia. Bryologist 75: 550-557.

GEMMRICH, A. R. 1977. Mobilization of reserve lipids in germinating spores of the fern Anemia phyllitidis L. Pl. Sci. Letters. 9: 301-307.

Bip ee: C. and B. W. NICHOLS. 1971. Plant Lipid Biochemistry. Academic Press, New

or

KARUNEN, P. 1971. Lipid and pigment patterns in germinating Polytrichum commune spores. Phytochemistry 10: 2811-2812.

LEESE, B. M. and R. M. LEECH. 1976, Sequential gor in com lipids of developing proplastids isolated from green maize leaves. Pl. Physiol. 5 9-7

LLOYD, R.M., and E. J. KLEKOWSKI, JR. 1970. il wie and viability in Pteridophyta:

; Evolutionary signficance of chlorophyllous spores. Biotropica 2: 129-137.

eee a es L. SMITH, R. SAFFORD, and B. W. NICHOLS. 1973. Lipid metabolism in the fern Polypodium vulgare. Phytochemistry 12: 1377-1381.

SCHWERTNER, H. A. and J. B. BIALE. 1973. Lipid composition of plant mitochondria and of chloroplasts. J, Lipid Res. 14; 235-242,

Seon A. V. 1975. An ultrastructural study of dormant and germinating fern spores. Amer.

pA 62 Suppl.: 47.

TOWILL, * “ and H. IKUMA. 1973. Photocontrol of the germination of Onoclea spores. I. Actos spectrum, Pl. Physiol. 51: 973-978.

»and H. IKU 1975. Photocontrol of the germination of Onoclea spores. IV. Metabolic

changes during pirmiatiion, Pl. Physiol. 56: 468-473,

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 3 (1978) 71

Gametophytes of Botrychium multifidum as Grown in Axenic Culture ERNEST M. GIFFORD, JR. and DOROTHY D. BRANDON*

Mature gametophytes of Botrychium from nature have been described for sev- eral species, including B. dissectum (Bierhorst, 1958), B. japonicum (Nozu, 1954; Nishida, 1955), B. Junaria (Bruchmann, 1906), B. obliquum (Campbell, 1921), B. simplex (Campbell, 1922), and B. virginianum (Jeffrey, 1898; Bierhorst, 1958). The gametophytes are subterranean and are tuberous or somewhat elongate to button-shaped. They vary from 1-3 mm to 5-6 mm long, and even reach 1.5-2 cm in some species. The possession of a ‘‘dorsal ridge’? in which antheridia are embedded is a typical feature of all the known species. The gametophytes have an associated endophytic fungus, the presence of which presumably is essential for continued growth under natural conditions.

Botrychium spores appear to have a dormancy that is not easily overcome. To date, only the spores of B. virginianum (Campbell, 1895), B. ternatum (du Buys- son, 1889), and more recently of B. dissectum (Whittier, 1972, 1973a) have been germinated, the last species in axenic culture.

The morphology of the B. dissectum gametophytes cultured by Whittier (1972) fits the description for B. dissectum from nature, except for the presence of the endophytic fungus. Sucrose and other additives in the medium presumably re- placed the contribution of the fungus under natural conditions. The gametophytes, when sexually mature, were only a few millimeters long.

Whittier (1973a) has shown that light inhibits spore germination in B. dissectum. He found that a minimum of 3-4 weeks in darkness is necessary for germination to occur after eight weeks in culture. Increasing the length of the dark period in- creased the percentage of germination. It is interesting to note that Whittier (1973b) showed that six months of darkness were required for the germination of Psilotum spores.

In this paper we will describe the morphology of Botrychium multifidum (Gmel.) Rupr. gametophytes as grown on a defined medium in axenic culture. The gametophytes of B. multifidum have never been observed carefully in nature.

MATERIALS AND METHODS

Spores for the present study were obtained from a plant in a greenhouse at the Department of Botany, University of California, Davis. The specimen from which the spores were taken has been growing in the greenhouse for several years. It was identified originally as B. multifidum subsp. californicum. However, the results of a recent survey (Stevenson, 1975) have shown that leaves which fit the descriptions of two or more of the presently accepted subspecies of B. multifidum can occur in nature on one large, copiously branched plant (which may be more than 100 yr. old). Stevenson believes that the recognition of several subspecies of B. multifidum is unnecessary, and so we refer the material to B. multifidum.

*Department of Botany, University of California, Davis, CA 95616.

72 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

Spores were surface sterilized in 10% commercial sodium hypochlorite, washed in distilled water, and sown on a medium recommended by Whittier (1972), which is essentially that of Knudson, except that only 0.2% sucrose was added to the 0.6% agar. The pH was adjusted to 6.3. The culture tubes were plugged with

FIGS. 1-3. Various forms of Botrychi iff : 5 fi : : ychium multifid 7 tes arkness for nine months. FIGS. 1 and 3. x 15 F if gigs ala grown from spores in darkn

ePiG..2. x 20: . 4. Transection of antheridium from dark-grown

cmp of B. multifidum; the jacket is 2 or 3 cells thick, and arrows mark two visible opercular

cells, x 23 cotton and capped with polethylene. Knowin Ww . . . i ae Batata, te spore germination, the innoculated tubes were placed in a were ie er. ue to an extended leave of absence of the first author, the tubes removed from darkness for nine months. No light-grown controls were

attempted because it was kn : own from previous experi ; that the spores did not germinate in the light. 2 ee

g that a minimum period of darkness

GIFFORD & BRANDON: BOTRYCHIUM GAMETOPHYTES IN AXENIC CULTURE 73

RESULTS

Numerous spores germinated during the nine months in darkness. The resulting gametophytes ranged in size from 0.25 mm to 2-3 mm; in a few cases they were 0.5 cm long. The gametophytes were generally obovoid to club-shaped, but in some instances they were irregularly branched (Figs. 1-3). Rhizoids developed on the lower surface or, in some instances, only rhizoid primordia were formed. Except for the smallest gametophytes, a dorsal ridge was present upon which antheridia were embedded (Figs. ] and 2). The presence of a dorsal ridge is apparently one of the universal features of all Botrychium species thus far investigated. Antheridia were present on the ridge, often in groups (Fig. 1), which may be the result of periodic activity of the apical meristem located at the anterior end, toward the dorsal side (Bierhorst, 1958). No archegonia have been observed, either by in- spection of the surface of the gametophytes or in sectioned material. In two instances, we observed reduced, leaf-like structures (not illustrated) which may represent sporophytes, possibly of apogamous origin. In both instances, the leaves consisted of delicate petioles 1-2 cm long with reduced or abortive laminae. These were produced during the nine months that the cultures were in darkness. The leaves were dead when removed from the culture tubes. The gametophytes to which they were attached were friable, and very little information could be ob- tained as to the relationship or attachment of the structures. Comparable isolated, apogamously formed leaves have been found in B. dissectum (Whittier, 1976).

Some intact gametophytes that were transferred to fresh medium produced secondary outgrowths (Fig. 5). These newly formed branches developed chlorophyll and became yellowish-green. Branching was limited and the presence of chlorophyll was ephemeral, perhaps as a result of too high a light intensity.

Gametophytes transferred to fresh medium and placed in darkness underwent an extensive proliferation of new apices from the surface of the original gametophytes (Fig. 6). There was no definite pattern of branching, although in some instances the branching appeared to be dichotomous. The new outgrowths were somewhat friable and became dissociated rather easily. Antheridia de- veloped on the dorsal side of some of the new branches.

DISCUSSION

There are no descriptions of gametophytes of B. multifidum from nature. Milde (1858) depicited a young sporophyte with a bulbous base; the latter structure was interpreted as a gametophyte by Clausen (1938), although Milde did not describe it as such. Stevenson (1975) searched without success for gametophytes of B. mul- tifidum in several populations in the Sierra Nevada of California. In the absence of gametophytes from nature, it is difficult to know what morphological expres- sion is representative of the species. However, the gametophytes of species al- ready described to date share many similarities: tuberous to somewhat elongate shape, with a dorsal ridge, but varying from 1-3 mm to 0.5 cm or more long. For B. multifidum, perhaps the initial form of the gametophyte in culture (club-shaped with a dorsal ridge) is representative of the gametophyte in nature. Whether the

74 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

cultured gametophytes average larger or smaller than those growing under natural conditions is impossible to resolve at present.

FIG. 5: chi : eles :

sips eda multifidum gametophyte initially grown in the dark, then transferred to light;

PRE Ei tig Aer produced in light appears whitish, x 12. FIG. 6. Growth produced from - multifidum gametophytes transferred to fresh medium and maintained in the dark, x 15.

oe that, in the absence of fertilization and/or apogamous de- ee eeRea * site sas: the subterranean gametophytes proliferate profusely, ae east present study when they are grown in darkness. The clumps

roken up through action of soil insects, worms, or even larger animals,

thereby i asi . — increasing the dispersal of the gametophytes. Unfortunately, such obser- ations have not been made in nature.

GIFFORD & BRANDON: BOTRYCHIUM GAMETOPHYTES IN AXENIC CULTURE 75

In the absence of archegonia on cultured gametophytes of B. multifidum, one may assume that the two clusters of sporophyte-like structures were of apogam- ous origin.

LITERATURE CITED

BIERHORST, D. W. 1958. eas. on the gametophytes of Botrychium virginianum and B.

dissectum. Amer. J. 1-9.

SRUCHMANN, H. 1906. ‘ice ‘ini eid und die Sporenpflanze von Botrychium lunaria Sw.

Flora 96: 203-230. silat al R. du 1889. ll af cryptogames vasculaires d’Europe. II. Filicinées. Rev.

. Bourbonnais Cent. Fra 153-164. CAMPBELL, D: H, 1895. The Panty and Development of Mosses and Ferns, Ist ed. Macmillan, w York.

. 1921. The gametophyte and embryo of Botrychium obliquum Miuhl. Ann. Bot. 35: 141-158. ——_—_—. 1922. The gametophyte and embryo of Botrychium simplex Hitch. Ann. Bot. 36; 441-455. CLAUSEN, R. T. 1938. A monograph of the Ophioglossaceae. Mem. Torrey Bot. Club 19: 1-171. JEFFREY, E. C. 1898. The gametophyte of Botrychium virginianum. Trans. Roy. Canadian Inst. 5:

265-294.

MILDE, J. 1858. Die Gefasskryptogamen in Schlesien; und ueber Botrychium crassinervium Rupr.

und seine Verwandten. Nova Act. Acad. Caes. Leop.-Carol. Nat. Cur. 26: 371-753, 757-767.

NISHIDA, M. 1955. The morphology, gametophyte, young —— and systematic position of vail Acero eet ae sa Paytomerpaoigy 9: 6.

NOZU, Y. 1954. y young J §Soteiciien japonicum Und. Phytomor- phology ; vss 434.

STEVENSON, D. W. 1975. Taxonomic and morphological observations on Botrychium multifidum (Ophioglossaceae). Madrono 23: 198-204.

WHITTIER, D. P. 1972. Gametophytes of Botrychium dissectum as grown in sterile culture. Bot. Gaz. 133: 336-339 . 1973a. The effect of light and other factors on spore germination in Botrychium dissectum. Canadian J. Bot. 51: 1791-1794 . 1973b. Germination of Psilotum spores in axenic culture. Canadian J. Bot. 51: 2000-2001. . 1976. Tracheids, apogamous leaves, and sporophytes in gametophytes of Botrychium dis- sectum. Bot. Gaz. 137: 237-241

LUST NO MORE!!! after rare ferns, old-world epiphytes, all 18 sp. platyceriums & many cv’s, ferns of Malaysia, Thailand, Phillippines, W. Indies, China. Many unnamed sp. unknown to cultivation, and available here only at The ENDANGERED SPECIES, 842 Walnut Ave., Carpen- teria, CA 93013. $1.00/catalog.

eee

76 AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 3 (1978)

Revision of the Genus Cochlidium (Grammitidaceae) L. EARL BISHOP* # 00 2 .27/

The genus Cochlidium was first proposed by Kaulfuss in 1820. However, the name was ignored for more than a century after its publication. Most of the species now recognized as members of the genus were included in the later Pleurogramme (Blume) Pres! or, following Hooker (1864), were sunk into the broad concept of Monogramma. A detailed exposition of the generic concepts of the major nineteenth century workers would serve more to illustrate the confusion over the relationship of these ferns than to illuminate their taxonomic history. Some idea of this uncertainty can be gained by noting the number of genera to which certain of the species have been assigned.

Christensen (1929) revived Kaulfuss’ name, and with characteristic care and insight provided the basis for the modern concept of the genus. He included here all the New World grammitid ferns with coenosori and one species, C. furcatum, with discrete, polypodioid sori. A. C. Smith (1930) later transferred another such species, C. connellii, to the genus. Copeland (1947), however, removed such species to the genus Grammitis, basing his concept of Cochlidium strictly on the coenosoral character.

Morton (1967) rightly deplored the artifice of Copeland’s larger genera in the Grammitidaceae. His remedy was to dissolve his predecessor’s genera into a single, very large genus Grammitis, recognizing the former groupings as equally artificial sections. From my understanding of the family, I believe that consis- tency with such a practice would necessitate the inclusion of all species of the family into a single genus, for none of Copeland’s generic circumscriptions are without intimately related species that he included elsewhere. If a natural classifi- cation is the desired goal, the failure of another worker to define and differentiate his taxa properly is scarcely reason for their indiscriminate dissolution.

The genus Cochlidium, as here construed, consists of those neotropical grammitid ferns with simple and entire or at most sinuate fertile laminae, con- colorous scales, hydathodes, and 2-8-celled hairs which characteristically have prese intercellular walls and are frequently somewhat catenate. The presence ie peer o3i, will separate these species from nearly all those New World species f ammitidaceae with simple fronds, and concolorous scales will remove them rats pe ri Species formerly included in Xiphopteris. The typical Nasironus tk? © Re highly derived and completely glabrous species, o indication of the i a i a nese hans ee

Aldother uniPyiae cpap Ip of some superficially quite distinct plants. fon uf ie isu Ponape feature of the genus is the stelar organiZa- ule pend ae other Grammitidaceae exhibit a fundamentally soleno-

: s-section of the Cochlidium rhizome shows 1-3 vascular

*1543 F Street, Anchorage, AK 99501.

L. E. BISHOP: REVISION OF COCHLIDIUM 77

bundles, each with a straight or lightly arced row of tracheids. This contrasts sharply with the basically circular arrangement seen in most other grammitid ferns.

The presence of a coenosorus is a conspicuous feature in the genus. In most species in which it occurs, however, the coenosorus is occasionally or even fre- quently interrupted. On the other hand, in C. jungens the sori are in some plants entirely separate, although normally there is much soral fusion in most popula- tions, occasionally to the point of forming a complete coenosorus. This shows that the coenosorus, considered an all-important generic character by Copeland and earlier authors, is not necessarily stable on even the species level. All stages leading to the compound sorus from free veins and separate sori are indicated by these variations. Also, the free receptacles of C. serrulatum are basal on the veins, parallel to and frequently almost fused with the costa, and are terminated by the abruptly geniculate sterile portion of the vein. This arrangement strongly suggests this species’ derivation from a coenosoral ancestor.

Another character notable in the group with compound sori is the tendency of the fertile lamina to fold conduplicately over the sporangia. This feature is charac- teristic of certain species (C. pumilum and, interestingly, C. serrulatum), but occurs in at least some individuals of most species. My own observations in Costa Rica suggest that in C. rostratum folded fronds occur in plants inhabiting more stressful environments. The fronds of those species with polypodioid sori borne distally on the fertile veins seem never to exhibit conduplicate folding.

I wish to thank the staff of the U.S. National Herbarium for making this facility fully available to me, and to express my appreciation to John T. Mickel and Bruce McAlpin for their aid in my work at New York Botanical Garden. All specimens not otherwise indicated are at US.

Cochlidium Kaulf. Berlin. Jahrb. Pharm. Verbunden. Wiss. 1820: 36. 1820.

Xiphopteris Kaulf. Berlin. Jahrb. Pharm. Verbunden Wiss. 1820: 35. 1820. LECTOTYPE: Acro- stichum serrulatum Swartz [= Xiphopteris serrulata (Swartz) Kaulf.], chosen by J. Smith, Hist. Fil. 179. 1875.

Micropteris Desv. Mém. Soc. Linn. Paris 6: 217. 1827. LECTOTYPE: Acrostichum serrulatum Swartz [= Micropteris serrulata (Swartz) Desv.], chosen by Copeland, Gen. Fil. 215. 1947.

Antrophyum sect. Pleurogramme Blume, FI. Jav. Fil. 69. 1829. LECTOTYPE: Taenitis linearis Kaulf. [=Pleurogramme linearis (Kaulf.) Presl], chosen by J. Smith, Hist. Fil. 177. 1875.

Pleurogramme (Blume) Presl., Tent. Pterid. 223. 1836.

(and sole original species): Acrostichum graminoides Swartz

[=Cochlidium graminoides (Swartz) Kaulf.].

The date of the lectotypification of Xiphopteris is somewhat open to question. Kaulfuss erected the genus on the basis of two species, both of which had been described by Swartz in the same publication. Fée (Gen. Fil. 100. 1852) accepted Kaulfuss’ genus but reduced X. myosuroides to synonomy with X. serrulata. It might be argued that since either name was available for application to his broader concept, Fée’s selection of X. serrulata constituted implicit lectotypification. I maintain this not to be the case, as the type of neither species was excluded from

78 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

the genus. Had one name had priority, none would consider that the synonymiz- ing of a later name constitutes typification. Alternately, if Kaulfuss had designated a type species, Fée would not have been obliged to select that name for his

broader species concept. Therefore, conversely, his selecting a name should not imply the selection of a lectotype.

KEY TO THE SPECIES OF COCHLIDIUM 1. Sori compound, the receptacles linearly confluent and closely parallel to the costa 2. Sori superficial, although at times protected by the conduplicately folded fertile lamina, 3. Fertile portion of the frond abruptly contracted in fwie and long-acuminate; "<a lamina usually 3 mm or more wide (West Indies, South America) minudum 3. Lae a of the frond neither contracted nor acuminate; sterile lamina mostly ste than 3

4. diate 4 veins forked; midrib not evident dorsally; fronds usually 2-3 mm wide ype.

. punctatum 4, — veins simple; midrib at least somewhat evident dorsally; fronds seri less than 2 mm

e

5. Sterile veins short, reaching less than halfway to the margin; fronds 1 mm or less wide, more than 2 cm lon 6. Fronds dichotiiens (Jamaica) . 1. C. graminoides 6. Fronds simple (South America) . 2. C. pumilum 5. Sterile veins reaching more than aa to the margin; fronds 1-3 mm wide, or if narrower than 1 mm, then less than 2 cm lon 7. Laminar margins sinuate; sterile yes s

omewhat evident ventrally, cee: in conspicu- ous hydathodes tanitg Pain arer a: repr aminar Pe ds hudatt t but jae conspicuous. 8. Midribs distinctly ninaiieaioas sb ae

laminar trichomes 2~ asco not clavate; fronds than 2 cm long (Puerto Rico, Domi . C. jungens 8. Midribs not prominulous ventrally; laminar trichomes 2- 3-celled, the Sail cells enlarged; fronds mostly 1-1.5 mm wide and less than 2 cm long (Jamaica, Cuba)

2. Sori deeply immersed in a central groove 9. Hydathodes conspicuous; s

mostly 1.5-3 mm wide and more

Renee ewee

S ney Mere Re ee Sercuen Ve OR ee et eee 1 ee - Sori separate, round or hy often confluent at maturity but with discrete receptacles easily —_

with the sporangia rem

10. Sterile fronds and ae ft

portions of fertile fronds distinctly serrate or acutely lobed (wide- sprea OG) iia ae a. ©

serrulatum

(emeine: stone atone distant on slender, creepin seein) stomata “most less than 57 x 45 um. (Venezuela ee 16.

we

C. furcata donsaiy with midrib h bs inar tissue, fi Prominulous but covered with lam ronds mostly unforked; Sporangial capsules usually more than 140 xm long.

L. E. BISHOP: REVISION OF COCHLIDIUM 79

14. Fronds 2.5-4 mm wide, entire, mostly i é regular rows of sori which at maturity are not

confluent across the midrib (South Amer 13. C. tepuiensis

14. Fronds 1-3 mm wide, entire or sinuate, ae with sori irregularly disposed or all com-

pletely confluent at maturi

15. Margins of the sterile emingé sinuate; fronds 1-2 mm wide (Venezuela) . 14. C. acuminata 15. Margins of the sterile laminae entire; fronds mostly 1.5-3 mm wide.

16. Mature sporangial capsules obpyriform, mostly 180-200 um long; fertile laminae flat

2 America) 15. C. connellii

16. Mat sporangial capsules globose, mostly 150-180 4m long; fertile laminae at times

spaginiicialy folded over the sori (Puerto Rico, Dominica) ................. 6. C. jungens

- 1. Cochlidium ee enerets | (Swartz) Kaulf. Enum. Fil. 86. 1824. Acrostichum graminoides Swartz, Prodr. Fl. Ind. Occ. 128. 1788. "TYPE: Jamaica, Swartz s.n. (S not seen; isotypes C, L-these yeh de Cc Chr. 1929, BM not seen, photo and fragment at US!). G : :

Asplenium graminoides (Swartz) Swartz, Fl. Ind. Occ. 3: 1608. 1806.

Monogramma ? furcata Desv. Ges. Naturf. Freunde Berlin Mag. 5: 303. 1811 /nom. nov, et superfl. TYPE: Based on Acrostichum graminoides Swartz.

Pleurogramme graminoides (Swartz) Fee, Mem. Foug. 3: 37. 1852.

56. Monogramma graminoides (Swartz) Baker in Hook. & Bak. Syn. Fil. 375. 1868, as Monogramme. Pleurogramme furcata (Desv.) J. Smith, Hist. Fil. 178. 1875. RANGE AND HABITAT: Jamaica. According to Proctor (1953), this species occurs locally on the eastern slope of the John Crow Mountains, at elevations of 2000-3000 ft

Proctor’s collection represents the only reported gathering of this species in this century. It is otherwise known only from Swartz’s original collection and from a few plants collected by James Wiles around the turn of the nineteenth century. As I have not seen the Proctor plants and since the few specimens available to me are fragmentary, I have little to add to Christensen’s (1929) account of this species. However, it should be noted that despite the implication to the contrary in Christ- ensen’s key, the fertile lamina is at times conduplicately folded over the sori.

SPECIMEN EXAMINED:

JAMAICA: ‘‘Near the top of the blue-mountain,’’ Wiles (BM not seen, K not seen, photo US, US!).

— 2. Cochlidium pumilum C. Chr. Dansk Bot. Ark. 6(3): 19. 1929.

AY PE: Chuquiribamba, Ecuador, 9 Sept 1868, Massee s.n. (K not seen).

RANGE: Colombia, Ecuador, Bolivia, Brazil.

Although I have not seen the type, the collections cited closely match Christen- sen’s description. The species seems to be rare, but is clearly wide-ranging.

I agree with Christensen that this species must be very closely related to C. graminoides, but the discovery of additional material reinforces the assumption that the two are distinct. Cochlidium pumilum generally exhibits unforked fronds and strongly conduplicate fertile laminae. Also, judging from the material seen, the coenosorus is often interrupted in this species, but is rarely so in C. graminoides.

SPECIMENS EXAMINED:

COLOMBIA: Acosta-Arteaga 689 (NY). BOLIVIA: New Brazil, 5500 ft, R. S. Williams 1171. Songo, Bang 907B (NY, US). BRAZIL: A. Richard in 1834 (NY).

80 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

A. Cochlidium serrulatum (Swartz) L. E. Bishop, comb. nov.

Acrostichum serrulatum Swartz, Prodr. Fl. Ind. Occ. 128. 1788»TY PE: Jamaica, Swartz s.n. (S not seen; isotype US!).

Grammitis serrulata (Swartz) Swartz, J. Bot. (Schrader) 1800(2): 18. 1802.

Gymnopteris serrulata (Swartz) Bernh. Neues J. Bot. 1(2): 48. 1805.

Asplenium serrulatum (Swartz) Swartz, Fl. Ind. Occ. 3: 1607. 1806.

Xiphopteris serrulata (Swartz) Kaulf. Farrnkr. 87. 1827.

Micropteris serrulata (Swartz) Desv. Mém. Soc. Linn. Paris. 6: 217. 1827.

Micropteris orientalis Desv. Mém. Soc. Linn. Paris 6: 217. 1827. TYPE: Mascarene Islands, without collector (presumably P not seen).

Polypodium serrulatum (Swartz) Mett. Fil. Lips. 30. 1856, non Swartz, 1802, nom. illeg.

Xiphopteris extensa Fée, Mém. Foug. 11: 14, t. 19, f. 3. 1866. TYPE: Guadeloupe, L’ Herminier in 1864 (presumably RB or P not seen).

Xiphopteris orientalis (Desv.) Fourn. Compt. Rend. Hebd. Séances Acad. Sci. 81: 1140. 1875.

Polypodium duale Maxon, Contr. U.S. Natl. Herb. 16: 61. 1912, nom. nov. TYPE: Based on Acrostichum serrulatum Swartz.

Xiphopteris auyantepuiensis Vareschi, Fl. Venez. 1(2): 879. 1969. TYPE: Auyan-tepui, Edo. Bolivar, Venezuela, 1800 m, Vareschi & Foldats 4806 (VEN not seen). ISOPARATYPE: Auyan- tepui, Edo. Bolivar, Venezuela, 1800 m, Vareschi & Foldats 4813 (US!).

RANGE AND HABITAT: Ranges throughout the West Indies and from the Mexican state of Puebla south to Bolivia. In the Old World it is known from the wetter parts of tropical Africa, Madagascar, the Mascarene Islands, and Amster- dam Island. Epiphytic and epipetric on mossy rocks. It is most frequently encoun- tered at altitudes of 500-1500 m.

Cochlidium serrulatum is probably the most common species of the family in the New World. In spite of its small size, it is one of the most abundant in collections, and no species has a wider range. As is often the case with wide- spread species, considerable variability can be detected within the taxon as it is generally circumscribed. However, to date no one has successfully demonstrated the distinctiveness of any proposed segregates.

The type of Micropteris orientalis is uncertain. However, a collection in the de Jussieu herbarium at Paris (Ile de F rance et Bourbon, de Petit Thouars s.n. in 1808 (photo US) May represent an isotype at least. But in any case, no Indian Ocean specimens I have seen warrant taxonomic distinction on the basis of the characters that have been studied,

I suspect that Fée described X. extensa because he had previously confused Swartz s Acrostichum serrulatum with the same author’s Polypodium myosuroides and had combined the two names under X. serrulata (Swartz) Kaulf.

The original descripti collections. No type w publication cited above.

on of X. auyantepuiensis Vareschi was based on two as selected, and so the name remained invalid until the The duplicate of the paratype at US seems well within the € species as a whole. Vareschi emphasized the dimorphic X. auyantepuiensis. But in most specimens of C. se”

rulatum, the sterile fronds are shorter and more deeply lobed than the fertile ones.

L. E. BISHOP: REVISION OF COCHLIDIUM 81

In considering the morphological plasticity of this species, it should be pointed out that Walker (1966) indicated his Jamaican sample of the species to be an apogamous tetraploid. This account alone is sufficient to suggest that such a wide-ranging and variable taxon might be biosystematically complex. It would not be surprising if a carefully detailed study demonstrated that our present “‘species”’ consisted of two or more cryptospecies and hybrids between them.

To transfer the type species of a genus so well-known as X iphopteris demands more than ordinary justification. Copeland (1947, 1952) strongly emphasized the restriction of the sori to a distal, nearly entire portion of the lamina in proposing an intimate relationship of X. serrulata with X. myosuroides and related species. But as I have discussed elsewhere (Bishop, 1974), laminar dissection is a weak charac- ter for the determination of species relationships in the Grammitidaceae. The X. myosuroides group shows clathrate scales, a solenostelic rhizome and a peculiar type of foliar trichome very different from that of C. serrulatum or other species of Cochlidium. In addition, the sori are merely impressed into the fertile lamina; the laminar margins are never conduplicately folded around the soral region. The necessary nomenclatural adjustments for the species normally included in Xiphopteris are being made elsewhere.

Aside from the conspicuously serrate or serrate-lobate fronds, C. serrulatum differs further from the other species of Cochlidium. The rhizome is relatively long-creeping with somewhat distant fronds, a character found otherwise only in C. wurdackii. The 2-4-celled laminar hairs are unusual in the genus in that the terminal cell is considerably elongated and the intercellular walls are somewhat less thickened. These trichomes seem to be nearly or quite lacking in many speci- mens. The hydathodes are much less conspicuous than in other species. In fact, they have been visible to me with certainty only in microscopic preparations. As they frequently are not situated over the terminus of the veins, the hydathodes may be vestigial in this species.

In view of the distinctiveness of this species, it might reasonably be retained in its own monotypic genus. However, since it is clearly related to the other species of Cochlidium, 1 prefer to include it here. But in any case, I feel sure that C. serrulatum cannot be retained justifiably in any generic concept which includes other species usually referred to Xiphopteris unless the entire family be construed to constitute a single genus.

One specimen from Venezuela merits special mention (Steyermark & Nilsson 264-A). This plant reveals its affinity to C. serrulatum through its elongate rhizome and the low serrations on some of the leaves. Other fronds, however, are quite entire. More significantly, the fertile lamina is scarcely expanded or condup- licate and is conspicuously narrower than the mature sori. These characters re- move the plant from the usual range of variation of the species and suggest possi-

ble hybridization with one of the Venezuelan species with polypodioid sori.

REPRESENTATIVE SPECIMENS EXAMINED: ; CUBA: Oriente: Sierra Nipe, near Woodfred, 450-550 m, Shafer 3432. HAITI: Massif du Nord, Anse-a-Foleur, top of Morne Colombeau, 900 m, Ekman H4368. DOMINICAN REPUBLIC: Loma de la

82 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

Sal, Jarabacoa, 1100 m, Liogier 11356, 1100-1400 m, Liogier 11691. JAMAICA: Portland: E slope of John Crow Mountains ca. 5 mi SW of Priestman’s River, Proctor 4294. PUERTO RICO: Serra de Naguabo, Monte El Duque, Shafer 2246. ST. KITTS: Slopes of Mt. Misery, Britton & Cowell 500. NEVIS: Nevis Peak, 900-1095 m, A. C. Smith 10552. MONTSERRAT: Top of Chaners Mountain, 3000 ft, Shafer 292. GUADELOUPE: Bains Jaunes, 700 m, Questel 1797. DOMINICA: Lower slope of Morne Micotrin, Laudat, 700 m, Hodge & Hodge 2095. MARTINIQUE: Calebasse, Deux Choux, Duss 1609. ST. LUCIA: Vicinity of Mt. Gimie, ca. 1500 ft, Webster, Ellis & Miller 9416. ST. VIN- CENT: Soufriére, 2400 ft, Eggers 6709. GRENADA: Vicinity of Grand Etang, 1600-1800 ft, Proctor 17049. TRINIDAD: Fendler 8] in 1877-78.

MEXICO: Veracruz: Zacuapan, Purpus 302]. Puebla: Barranca below Honey Station, Pringle 13853. Oaxaca: Dto. Teotitlan, Mun. de Tenango, First half of trail from S. Martin Caballero to Tenango, 4300-5000 ft, Hallberg 1437. Chiapas: Mun. de Ocosingo, near Laguna Ocotal Grande, ca. 25-30 km SE of Monte Libano, ca. 950 m, Dressler 1641. GUATEMALA: Huehuetenango: Sierra de los Cuchumatanes, vicinity of Maxbal, 1500 m, Steyermark 48782, 48886. Alta Verapaz: Cerro Sillab, Senahu, Hatch & Wilson 169. Izabal: Montafia del Mico, between Milla 49.5 and ridge 6 mi from Izabal, 65-300 m, Steyermark 38627. San Marcos: S-facing slopes of Volcan Tajumulco, along the Rio Chopal, Finca El Porvenir, 1300-1500 m, Steyermark 37465. Chiquimula: Volcan Ipala, near Amatillo, 900-1510 m, Steyermark 30511. BELIZE: Mountain Pine Ridge, Bartlett 11642. HONDURAS: Cortes Lake Yojoa, near Agua Azul, 650 m, Williams & Molina 17877. Atlantida: Lancetilla Valley near Tela, Standley 56353. COSTA RICA: Guanacaste: Vicinity of Tilaran, 500-650 m, Standley & Valerio 44565.

tarenas: Biological Field Station at Finca Wilson, 5 km S of S. Vito de Java, 1300-1400 m, Lellinger 807. Alajuela: Near Artezalea and Methodist Rural Center, ca. 8 km NE of Villa Quesada, 550 m, Molina et al. 17748. Limon: Alto Lari, 600 m, Jiménez M. 1888. Heredia: Rio Sarapiqui near Cariblanco, N of Volcan Pods, 850 m, L. O. Williams 20246. San José: Near Tarbaca, ca. 15 km S of S. iene 1900 m, L. 0. Williams 19441. Cartago: Estrella, 20 mi S of Cartago, 5200 ft, Stork 3297.

ANAMA: Chiriqui: Vicinity of El Boquete, 1000-1500 m, Cornman 910, 952, 1028, 1109, 1238, 1239. a: La Eneida, region of Cerro Jefe, Dressler 3315. Darien: Cerro Mali base camp in the Serrania

del Darién, 1400 m, Gentry & Mori 13776. Powe gominie Nueva Esparta: Island of Margarita, Juan Griego trail, 450 m, Johnston 144. Aragua:

ween El Portachuela and Ocumare, Pittier 11384. Faleon: Near Santa Ana, Cerro Santa Ana, 750

oo and Raudal Yayacopi), ca. 700 ft, Schultes & Cabrera 16182. Huila: , t, Little 7715. Chocé: Between Palestina and Agua Negra, Rio S. Jua , Cua

m trecasas 21555. El Valle: La Cumbre, 1700- lli if na Island, Om, Killip & Garci En re, bai 2200 m, Killip 1 1318. Narino: S end of Gorgona Islan¢, ¢

Martin: Top of ridge E of Tingo Maria, 1020 m, Allar » 800 m, Mexia 6287. Huanuco: Tingo Maria, 700 m, Tryon & Ine

_/

—

L. E. BISHOP: REVISION OF COCHLIDIUM 83

Paulo: Morro Jaragua, 1000 m, A. C. Brade 6590. — Serro do Embeque, 1200 m, Hatschbach 24951. Sta. Catarina: Morro do Bau, I[tajai, 700 m, z 3017. FRENCH GUIANA: May 1831, Le- prieur. SURINAM; Brownsberg, 400 m, Gonggerijp ‘ Sickel ong GUYANA: Trail from Kaieteur Falls to Tukeit, Potaro River Gorge, Maguire & Fanshawe 2348

LIBERIA: Vicinity of Mt. Coffee, ca. 40 mi inland from ene 50-150 m, Cook 122. MAURITIUS: Pike in 1867. MADAGASCAR: Mont Papanga, 1600-1700 m, Humbert 6944.

4. Cochlidium minus (Jenm.) Maxon, Sci. Surv. Porto Rico and Virgin Is. 6: 407.

1926. Monogramma minor Jenm. Bull. Bot. Dept., n. ser. ae rigs 1897. TY PE: Murray’s Flat, above Mt. Moses, St. Andrew Parish, Jamaica, Jenman s.n. in 6. (NY!; isotype US!).

RANGE AND HABITAT: Jamaica aa eet Proctor (1953) describes this species as epipetric. One Cuban specimen is cited as ‘‘Epiphytic or terrestrial in dense clumps.’’ Elevations on two sheets from Cuba are 1350 and 1400 m.

Of C. minus from Jamaica, I have seen only the type material. Evidently rare and local, this is also the smallest known species of the genus. Both Maxon (1926) and Christensen (1929) included here a number of specimens from Cuba and Puerto Rico that to me clearly belong to two other species, both undescribed.

SPECIMENS EXAMINED:

CUBA: Oriente: Palma Mocha, 1400 m, Ekman 521]. Crest of Sierra Maestra between Pico Tur- quino and La Bayamesa, 1350 m, Morton & Acufia 3563. § side of the crest of the Sierra Maestra W of Aserradero S. Antonio de las Cumbres, region of La Bayamesa, 1400-1500 m, Morton 9591.

5. Cochlidium repandum L. E. Bishop, sp.

Filix parvula quae in saxibus arDorbuagne colonialiter crescens invenitur. Rhizoma ascendens, frondes approximatas et paleas 2-4 mm longas apicibus acuminatis ad basin cordatam per 8-12 cellulas lids fone wid Supmes tenues atrobrunnei teretes 1-15 mm longi atque diametro 0.2-0.3 mm. Lamina

-0-2.5 mm, chartacea vel subcoriacea, patula vel suberecta, ad apicem rotun- data, basin versus acuminata, costa dorsaliter parum prominula vagina sclerotica eius ‘plus minusve hic evidenti at ventraliter costa praecipue per longitudinem fertilem prominenti edenti venulas visibiles quae in hydathodos claras terminant, margine repando aut interdum subin pre pilis 2-4 cellularibus in margine cos- taeque faciebus ambabus instructa, stomatibus abaxialibus 55-67 x 47-53 um, sorifera per partem usque tertiam asia: hic limbo laminali plano aut circum soros conduplicato. Venae steriles simplices, illae ee seca receptaculares eee furcam itidem fertilem ath venam proximam distalem jungit, hoc sorum efficientem. Coe rus in lam isienin ag Demian aliquando nliquantin interruptus. S okangicwtih eapttias subglobosae 100-120 x 95-110 um, cellulae annuli 10-12, sporae 22-25 um diametro. “HOLOTYPE: On tree fern bases, Arroyo del Medio, above the falls, Sierra de Nipe, Oriente, Cuba, 450-550 m, Shafer 3473 (US)

PARATYPES: The following collections from Cuba at US are referable here: Clément 4631, 4978, 6724, Acuna 12348, Pollard et al. 240, Shafer, 3475, 4110, 8072, 8159.

RANGE AND HABITAT: Apparently confined to the eastern end of Cuba, Pcia. Oriente, from Sierra de Nipe to Baracoa. Judging from the number of collections, it must be at least locally common at middle altitudes. It is recorded growing on both trees and rocks.

84 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

Confounded by Christensen and Maxon with C. minus and C. jungens, this species is easily separated by the repand margin and other characters mentioned in the key. Cochlidium minus occurs in Cuba only in the Sierra Maestra, where C. repandum is absent. Apparently the two species grow at different elevations as well.

‘ 6. Cochlidium jungens L. E. Bishop, sp. nov.

Epiphytum parvum in sylvis muscosis vigens. Rhizoma ascendens vel erectum nonnunquam ramosum ac caespitosum, frondibus stipitibusque confertis onus- tum, paleis 1.5-3 mm longis e basi truncata aut subcordata 8-12 cellulas lata usque ad apicem filiformem acuminate angustatis praeterea instructum. Stipites obsoleti vel usque 10 mm longi alati atrobrunnei 0.2-0.4 mm diametro. Lamina 2-8 cm x 1.5-3 mm, chartacea vel subcoriacea, vulgo erecta, ad apicem rotundata basin versus acuminata, costa utrinque prominula sed vagina atrofusca rarenter evidenti, in margine integro costaque utrinsecus pilis ex 2-8 cellulis constantibus illis interdum raris vel sub aetate provecta carentibus, stomatibus dorsalibus 65-82 x 52-62 wm, sorifera ad apicem vel frondis medio tenus, hac lamina fertili expansa aut super sporangias conduplicata. Venae steriles plerumque simplices, illae fer- tiles saepe furcatae, receptaculares ad basem venae furcataeque acroscopicae, increbre furca receptaculari adjecta ad venam proximam jungenti et coenosorum facienti. Sporangia capsulis 150-180 x 150-170 wm, cellulis annuli 8-11, sporisque papillosis 26-31 wm diametro

“HOLOTYPE: Epiphytic in mossy forest, top of Morne Micotrin, St. George Parish, Dominica, 3800 ft, Nicolson 1975 (US).

PARATYPES: The following are also at US: Dominica: Webster 13376; Puerto Rico: Britton 7917, Hess 313, Hioram 335, Shafer 2245, 3647.

RANGE AND HABITAT: Dominica and Puerto Rico. Epiphyte in cloud

forest, 800-1500 m elevation. This is yet another species formerly confounded with C. minus. Actually, with its larger, erect and often crowded fronds, it is hardly even superficially like that species, and finer characters clearly separate the two. Nor does C. jungens show any more than a casual resemblnce to C. repandum of Cuba.

Of interest in this species is the variation with respect to fusion of the sori. In some plants coenosoral development is complete. But in most the sori are sepa- hag or else show various degrees of soral coalescence. Since some workers have ‘ t the coenosorus to be critical in the generic delimitation of Cochlidium, 1 have c — ts, epithet to emphasize that with respect to this character, the species with polypodioid sori are joined to those with coenosori by this species. However,

this should not be taken to indicate that C. jungens is phylogenetically inter- mediate between the groups.

=, 7. Cochlidium seminudum (Willd.) M . : ie 6: 407. 1926. -) Maxon, Sci. Surv. Porto Rico and Virgi

Blechnum seminudum Willd. Phytogr. 13, 1. 8, f. 2. 1794. TYPE: Martinique, /sert s.n. in 1787 (B

L. E. BISHOP: REVISION OF COCHLIDIUM 85

Taenitis pumila Kaulf. Enum. Fil. 132. 1824, nom. nov. et superfl. TYPE: Based on Blechnum

seminudum Willd. aenitis linearis Kaulf. Enum. Fil. 131. 1824. TYPE: Guadeloupe, without collector (Herb.

Sprengel, presumably destroyed).

Micropteris blechnoides Desv. Mém. Soc. Linn. Paris 6: 217. 1827, nom. nov. et superfl. TYPE: Based on Blechnum seminudum Willd.

Grammitis isertii Swartz, Adnotat. Bot. 65. 1829, nom. nov. et superfl., as “‘iserti.’’ TYPE: Based on Blechnum seminudum Willd

Pleurogramme graminifolia (Hooker) Presl, Tent. Pterid. 223. 1836.

Pleurogramme linearis (Kaulf.) Presl, Tent. Pterid. 223, 1. 10, f. 2. 1836.

Pleurogramme pumila (Kaulf.) Presl. Tent. Pterid. 223. 1836.

Monogramma graminifolia (Hooker) Hooker, Sp. Fil. 5: 124. 1864, as ‘‘Monogramme.”

Taenitis seminuda (Willd.) Kuhn, Fil. Afr. 59. 1868.

Pleurogramme seminuda (Willd.) J. Smith, Hist. Fil. 178. 1875.

Pleurogramme nuda Goebel, Flora 117: 119. 1924.

RANGE AND HABITAT: Jamaica, Hispaniola, Puerto Rico, Lesser Antil- les, Trinidad, and Venezuela. Christensen (1929) also cites specimens from French Guiana. Due to the usual confusion about Isert’s specimens, nineteenth century authors also assigned the species to Africa, where it does not exist.

Cochlidium seminudum is a fairly common, widespread, and larger member of the genus. As as result, it has had an appropriately checkered taxonomic history.

Hooker described Taenitis graminifolia without reference to Willdenow’s species, suggesting only that it might be identical with Grammitis graminoides (=Cochlidium) of Jamaica. Although I have not seen type material, the illustra- tion is good and no species is known from St. Vincent superficially similar to C. seminudum.

Willdenow’s type is a small plant of this species. Kaulfuss assigned the name Taenitis pumila to this specimen and differentiated his Taenitis linearis as being twice as large and stipitate. The type was undoubtedly destroyed along with the rest of the ferns of Sprengel’s herbarium, so that the application of the name can never be certain. However, assuming that it represents an otherwise known tax- on, one part of Kaulfuss’ description, ‘‘Frons.. . hinc [e medio] angustata in caudam elongatam producta,’’ could only apply to this species.

Presl erected his genus Pleurogramme to include both of Kaulfuss’ species as well as Hooker’s T. graminifolia. Therefore, all three names in Presl’s genus are reducible to a single species concept.

As pointed out by Christensen, Goebel (1924) sought to clear the confusion of this species with C. punctatum. The Brazilian plants frequently had been included in the same species concept as the Caribbean C. seminudum by nineteenth cen- tury workers, but Goebel clearly established the two populations as distinct. Unfortunately, he applied the name Pleurogramme seminuda to the Brazilian species and gave the new name Pleurogramme nuda to the Caribbean population. No type was cited, but the intended application of the name is obvious.

The most distinctive characteristic of this species is the constriction of the lamina through the fertile portion. No other species exhibits this feature. The sori are impressed into the dorsal surface, but the limb of the fertile lamina 1s usually

/

AMERICAN FERN JOURNAL: VOLUME 68 (1978) 86

flat. Only rarely is it pepe folded ie the sporangia. In such cases, fertile portion is generally falcate as well. OLEPRESUNTATIVE SPECIMENS EXAMINED: : ‘sede JAMAICA: Portland: E slope of the John Crow Mountains 1.5-2.5 mi SE of rig i Hie ft, Proctor 10491. HISPANIOLA: Samana: Top of Pan de Azucar, ca. 510 m, Ekman Pek: PUERTO RICO: El Yunque, Serra de Luquillo, 2500-3000 ft, Chase 6730c. rare pis pase Naguabo, 210-675 m, Shafer ea Re cot ea Raia pe = i i jacent ridge, 900- ,A.C. . ‘ ed ee Rei 300) ft, Proctor 19102. GUADELOUPE: Bains ae by Questel 2003. DOMINICA: Vicinity of Fresh Water Lake near Laudat, 450-600 a : : . oc Se MARTINIQUE: Deux Choux, 460 m, Stehlée ean oe Soufriere, » £aZ2 : i : Bot. Gard. Herb. 5 Setar. Be ee cas a Mountain, Island of Margarita, 600 m, Johnston 186. — NE of Giliria, N of Puerto de Hierro, Cerro Patao, Peninsula de Paria, 800-825 m, Steyermar: Agostini 91125. : idi m (Raddi) L. E. Bishop, comb. nov. ee ee bens : Ht 225, 1. Set yrr: Brazil, Raddi s.n. (presumably nea re i) Hooker, Sp. Fil. 4: 172. 1862.

sic ica teconan ne Vise. Brés. 2: 51, 1. 96, f. 1. 1873. TY PE: Pico da Tijuca, ~ Rio de Janeiro, Brazil, Glaziou 5384 (presumably RB not seen; isotypes B, C-fide C. Chr., NY!).

Monogramma rudolfii Rosenst. Festschr. Alb. v. Bamberg 63. 1905. SY NTYPES: Tresbarras- serra, Est. Sta. Catharina, Brazil, 900 m, Schmalz 162 (S; isosyntype NY!); Rio Grande, Est. Sao Paulo, Brazil, Wacket 45 (S; isosyntype NY!).

Polypodium paucinervatum (Fée) C. Chr. Ind. Fil. 332. 1905.

Cochlidium paucinervatum (Fée) C. Chr. Dansk Bot. Ark. 6(3): 22. 1929. f

RANGE AND HABITAT: Southeastern Brazil, known from the states 0 Minas Gerais, Rio de Janeiro, Sao Paulo, Parana, and Sta. Catarina. Epiphytic or occasionally epipetric, at 400-1300 m elevation. ; :

It is unfortunate that Christensen (1929) adopted Fée’s epithet for this species, even though admitting the probable identity of Raddi’s earlier species. He states, “As to some points Raddi’s description and figure do not agree perfectly with C. paucinervatum, and I therefore prefer not to use Raddi’s specific name although it is fully available and although I believe that his species is identical with Gr. paucinervata.”’ Although I sympathize with Christensen’s reluctance to adopt species whose type he had not seen, in this case I believe it to be justified an necessary.

Raddi’s plate is quite good. It shows well the general habit of this species. The only abnormality exhibited, and the one which no doubt disturbed Christensen, 1S that one of the three fertile fronds is shown with a discontinuous coenosorus. Bu this is not uncommon in species of the genus with coenosori, and the other i: fronds show continuous soriation. Another point concerning the drawing 1S - the fronds in ventral view show conspicuous hydathodes; indeed, these are Cele brated in Raddi’s name. These dark, very evident hydathodes constitute a con- Spicuous feature of many, but not all, specimens of this Brazilian species. Geographical evidence for the identity of Raddi’s species may also be cited. No other remotely similar species is known to occur in southeastern Brazil, but ms Present one must be fairly common. Raddi did much of his collecting in the area 0

L. E. BISHOP: REVISION OF COCHLIDIUM 87

Rio de Janeiro, well within the species’ range. It seems most unlikely that he gathered a very similar species collected by no subsequent worker but failed to notice the one in question.

In contrast to Raddi’s drawing, that of Fée is almost unidentifiable. Even the Glaziou collection on which it is based is so depauperate that attention to mi- crocharacters and a consideration of range are necessary to make a reasonable assignment to species. Interestingly enough in the light of the above discussion, the NY plant has round sori that appear superficially polypodioid, and Fée’s species was so described. This condition, though, seems best explained by con- sidering the material subfertile.

Monogramme rudolfii Rosenst. was based on two syntypes. Duplicates of each at NY belong clearly to the present species concept. But since I have not seen the original material and since the selection seems not immediately necessary for nomenclatural purposes, I decline to designate a lectotype.

Goebel (1924) presented a detailed comparison of this species with the Carib-

bean C. seminudum. To his distinctions I would add that the rhizome scales of C. punctatum are generally 20-30 cells wide at the base, whereas those of C. seminudum are half as many cells wide. The lamina of the latter species is abruptly contracted through its fertile length, whereas that of the former is of uniform width. The sporangial capsules of the Brazilian plant are somewhat ob- pyriform and 200-250 um long; those of C. seminudum globose and 100-150 7m long. As Goebel pointed out, the fertile veins generally do not quite fuse in this species. This provides yet another example of imperfect coenosoral development. However, this discontinuity is usually discernible only in cleared specimens and is scarcely detectable externally.

REPRESENTATIVE SPECIMENS EXAMINED:

BRAZIL: Minas Gerais: Bello Horizonte, Serra do Piedade, 1300 m, Foster & Foster 556. Rio de Janeiro: Itatidia National Park, S face of Mt. Itatiaia, Eiten & Eiten 7524-C. Sao Paulo: Rio Grande, Aug 1913, A. C. Brade. Parana: Serra do Capivary Grande, Hatschbach 8235. Sta. Catarina: Tresbar- raserra, 1000 m, Schmalz (Ros. Fil. Austrobras. Exs. 73).

9. Cochlidium linearifolium (Desv.) Maxon ex C. Chr. Dansk Bot. Ark. 6(3): 23. 1929

Monogramma linearifolia Desv. Ges. Naturf. Freunde Berlin Mag. 5: 302. 1811.°TY PE: French Guiana, without collector (P not seen).

Grammitis linearifolia (Desv.) Steudel, Nomencl. Bot. 2: 187. 1824.

Pleurogramme immersa Fée, Mém. Foug. 3: 37. 1852, nom. nov. et superfl. TYPE: Based on Monogramma linearifolia Desv.

Pleurogramme linearifolia (Desv.) Moore, Ind. Fil. xxvii. 1857.

Monogramma immersa (Fée) Hooker, Sp. Fil. 5: 125. 1864, as ‘‘Monogramme.” :

RANGE AND HABITAT: I have seen specimens from Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Guyana, and Brazil. Christensen (1929) reports it from Tovar, Venezuela. Although this species 1s to be expected in Venezuela, I suspect any specimen from the elevation of Tovar would likely be C. rostratum. I have seen no material from the Caribbean, where Christensen reports specimens from Cuba, Haiti, and Guadeloupe. The Ekman

88 AMERICAN FERN JOURNAL. VOLUME 68 (1978)

collection from Haiti is definitely C. rostratum, and the others need re- examination. Cochlidium linearifolium is an epiphytic species of the lowlands, attaining a maximum elevation of 800 m.

Neither Christensen nor I have seen Desvaux’s type. However, because of the preference of C. rostratum for high elevations, I think it unlikely that the speci- men represents that species. Another possibility is that of confusion with C. seminudum, which is reported from French Guiana. There is nothing in Des- vaux’s description to indicate difference from this species, except the note that the fronds are subfalcate. This is very rarely the case in C. seminudum, whereas falcate fronds are common in C. linearifolium.

This species has been difficult to distinguish from C. rostratum. By recognizing the altitude difference, it has been possible to correlate the prominent hydathodes with smaller sporangial measurements, and therefore to give a better circumscrip- tion of the concept than did Christensen, whose venation characters do not hold in my preparations. But the situation is not so clear-cut as would be liked. For instance, the two Costa Rican specimens I have assigned here are really inter- mediate, both in characters and elevation.

The capsules of specimens included here are often little more than half the size of those of C. rostratum, and the stomata generally are smaller as well. This Suggests a difference in chromosome number between the two species. I suspect that cytological investigation might show more biosystematic complexity in this area of the genus than can be detailed from herbarium material alone.

REPRESENTATIVE SPECIMENS EXAMINED:

(GUATEMALA; Alta Verapaz: Cubilquitz, 350 m, von Tuerckheim 8372. BELIZE: Mountain Pine Ridge, Bartlett 11762. HONDURAS: Cortés: E. shore of Lake Yojoa, 360 m, Williams & Molina 1141. NICARAGUA: SE of La Tronquera, 50 m, Molina 15003. COSTA RICA: Puntarenas: Osa Peninsula pray sity = sah de Osa, 600 m, Lellinger 676. San José: S. Isidro del General, 2500 ft, be say fave Z anama: Cerro Jefe, 10 mi S from Goofy Lake in Cerro Azul, Correa &

arien: Summit of Cana, 2500 ft, Stern et al. 533. ore tim aca ar carga oo : , - m, Killip 35341. Valle: Buenaventura, 0-10 m, Killip

17, 23 - BRAZIL: Amapa: Rio laue 0.5 km E of confluence with the Rio Oiapoque, Irwin et al. 47926.

10. — rostratum (Hooker) Maxon ex C. Chr. Dansk Bot. Ark. 6(3): 23:

oc niles sitesi Hooker, Sp. Fil. 5: 122. 1864, as ‘‘Monogramme.’’“ TY PE: Omotepee,

Omotepec. ve aati (K not seen; isotype US!). Hooker cites the location as ‘Island on the Lake PPleurcsramine P Ne Petit tog spelled Ometepe) is an island in Lago de Nicaragua.

his! Genend: Pax 3 : exa Christ in Pittier, Prim. Fl. Costar. 3(1): 10. 1901. TYPE: Valley of the eho tatat is . ie Costa Rica, 700 m, Pittier 12061 (BR or P not seen). Cochlidiu 8yroflexa (Christ) C. Chr. Ind. Fil. 430. 1906.

Ona "e m rostratum var. areolatum C. Chr. Dansk Bot. Ark. 6(3): 25. 1929. TYPE, Morne RA ie pee ee Haiti, Ekman H-4376 (BM not seen; isotype US!).

(Oaxaca) ae HABITAT: Cuba, Jamaica, Hispaniola, Guadeloupe, Mexico

Venez , prepoinien Honduras, EI Salvador, Nicaragua, Costa Rica, Panama,

uela, and Colombia. Epiphytic, at 700-2600 m elevation.

L. E. BISHOP: REVISION OF COCHLIDIUM 89

The type.represents an extreme form. The sterile lamina is narrow and erect, the fertile strongly conduplicate, falcate, and with a long, sterile apex or “‘ros- trum.’’ I agree’ with Christensen (1929) that there seem to be all intermediates between this and the forms with wider, laxer, and flat leaves. He took plants of the latter sort to be representative of Pleurogramme gyroflexa Christ, although he did not indicate that he had seen the type. This type needs checking; the elevation given in the original description is low for C. rostratum, and most workers have not distinguished this species well from C. linearifolium.

hristensen removed all Caribbean material to his variety areolatum, which was characterized solely by the usually anastomosing veins. The venation is visi- ble only in cleared specimens, and I have found such areolation to be rather casual in my preparations. Since these plants are not otherwise different from those of continental populations, I believe a variety based on such a weak, occult charac- ter to be trivial.

The single Mexican specimen that I have seen (Mickel 914) needs special men- tion. The hydathodes are quite conspicuous, as in C. linearifolium. In addition the sporangia are a bit small for C. rostratum (240-270 um). However, I retain it here on the basis of the large spores (50-55 ym) and the elevation (ca. 7000 ft.). Certainly the population merits closer study.

REPRESENTATIVE SPECIMENS EXAMINED:

BA: Oriente: Crest of the Sierra Maestra between Pico Turquino and La Bayamesa, 1350 m, Morton & Acufia 3549. JAMAICA: Portland: John Crow Peak, 6000 ft, Fawcett. HAITI: Marmelade, Morne Belle-Terre, Massif du Nord, ca. 1100 m, Ekman H8211. GUADELOUPE: Bains Jaunes, 950 m, Stehlé 341, 800 m, Stehlé 1096.

CO: Oaxaca: Valley of the Yelagago River ca. 20 mi NE of Villa Alta, 6800-7200 ft, Mickel 914. GUATEMALA: Alta Verapaz: Senahu, Cerro Sillab, Hatch & Wilson 164. HONDURAS: Francisco Morazan: Rancho Quemado, above S. Juancito, 2100 m, L. O. Williams 18518. EL SALVADOR: Santa Ana: El Trifinio, Cordillera Miramundo, 2000-2200 m, Molina, Burger & Wallenta 16796. COSTA RICA: Alajuela: Vicinity of Fraijanes, 1500-1700 m, Standley & Torres 47486. Heredia: Vara Blanca de Sarapiqui, 1750 m, Skutch 3322. San José: Las Nubes, 1500-1900 m, Scamman 7260. Cartage: Near La Sierra, ca. 25 km S of Cartago, 2000 m, L. O. Williams et al. 28060. PANAMA: Chiriqui: Between Alto de las Palmas and the top of Cerro de la Horqueta, 2100-2268 m, Maxon 5514.

VENEZUELA: Lara: Fila de Terepaima between Alto de Chuparral and la Loma Redonda, 21-23 km S of Cabudare, 1300-1460 m, Steyermark et al. 103382. COLOMBIA: Meta: Renjifo Massif, Cordillera La Macarena, 1300-1900 m, Idrobo .& Schultes 974.

Q c

-A1. Cochlidium proctorii (Copel.) L. E. Bishop, comb. nov.

Grammitis proctorii Copel. Philipp. J. Sci. 80: 121. 1952, as “‘proctori.’’ TYPE: East slope of John Crow Mountains, ca. 2 mi SW of Ecclesdown, Jamaica, 2000 ft, Proctor 5642 (MICH not seen; isotypes LJ-fide Proctor 1953. US}). aoe

RANGE AND HABITAT: Known only from the type locality. Epiphytic.

This is the only species of the genus outside South America with sori borne distally on the veins. Among this group of primarily Venezuelan species, it shares with C. tepuiense two regular rows of sori which are more or less separate at maturity. Like C. attenuatum it has a lightly repand margin, and the thinner texture and dorsally black midrib are similar to C. furcatum. Cochlidium proctor has more abundant marginal hairs than other species of the genus. In C. jungens there are occasionally fairly numerous hairs, but this species Is easily separable

90 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

from C. proctorii by its erect habit, thicker texture, green midrib, entire margin, and larger sporangial capsules (length 150-180 vs. 100-140 4m).

12. Cochlidium furcatum (Hook & Grev.) C. Chr. Dansk Bot. Ark. 6(3): 20. 1929. Grammitis furcata Hook. & Grev. Ic. Fil. t. 62. 1828’TYPE: Guyana, Parker s.n. (KK not seen). ?Mecosorus nudus Klotzsch, Linnaea 20: 405. 1847. SYNTYPES: Guyana, Rich. Schomburgk

1187, 1851 (B neither seen).

Polypodium furcatum (Hook. & Grev.) Mett. Abh. Senckenberg Naturf. Gesell. 2: 34. 1856, non Swartz, 1801 nec Desv., 1827 nec Roxb., 1844.

Polypodium dicranophyllum C. Chr. Ind. Fil. 331. 1905, nom. nov. TYPE: Based on Grammitis furcata Hook. & Grev.

Grammitis dicranophylla (C. Chr.) Vareschi, Fl. Venez. 1: 874. 1969, nom. illeg.

RANGE AND HABITAT: Guyana, Trinidad, Venezuela, Colombia, Brazil. Epiphyte at 600-2000 m elevation. Reports from Jamaica exist, but I have seen no specimens, Proctor (1953) does not mention it, and I doubt that it occurs there.

This and the following four species constitute a group whose taxonomic disposi- tion is still unsatisfactory. The taxa remain refractory in spite of considerable effort to delimit the species on the basis of microscopic characters. I believe that hybridization is occuring here. This, coupled with the reduction of already small plants in exposed habitats, has resulted in a number of collections which I am unable to place with assurance. My treatment here is tentative and one which does least violence to existing species concepts and nomenclature.

Cochlidium furcatum and C. tepuiense are the two commonest, most distinc- tive, and most stable entities, although even they seem to intergrade with one another and each with other species. Cochlidium attenuatum and C. connellii exist as identifiable concepts, but are less discrete and more variable. I accept them as species partially because both names are available in the genus. Cochlidium wurdackii is a rare but distinct entity worthy of specific recognition. I feel strongly that only careful field observation, perhaps in conjunction with cytological data, will resolve the biosystematic problems with these species.

Although I have not seen the type of the present species, the specimen is fairly well illustrated and is from Guyana, where the species is widespread. In addition, this is the species most regularly having dichotomous fronds. Therefore, I believe the name to be properly applied.

Mecosorus nudus Klotzsch is more problematic. Copeland (1952) with good reason referred the name to C. furcatum, but neither of us have seen the type an the original description is hardly definitive. Schomburgk did collect from Mt. Roraima where C. attenuatum and C. connellii occur. Since the fronds are de- ae as very rarely bifid, it is possible that the type material represents one of

pecies. It should be noted that Copeland (1947) suggested this species for the type of Me

Schrad., as

which Klot

Cochlidi free, dista

this was the type and sole species of the earlier Microgramma Presl, zsch illegitimately adopted as a section of his new genus.

um furcatum is the only South American member of the genus with lly borne sori which has been found outside the Guyana Highlands. ther workers have emphasized the simple fertile veins and forked fronds to

cosorus. The type, of course, must be Polypodium persicariifolium |

L. E. BISHOP: REVISION OF COCHLIDIUM 9]

define this species. Unfortunately, both simple fronds and forked fertile veins occur here, and forked fronds are found occasionally in the other species. Several other features help characterize this concept. The small sporangial capsules, mostly 110-140 um long, well separate C. furcatum from all but C. tepuiense, which has capsules mostly 140-170 wm. The present species typically has the sori irregularly disposed and the lamina expanded around the receptacle, resulting in the fertile portion of the frond being irregularly repand. Cochlidium tepuiense has entire fronds bearing two regular rows of sori. The lamina of C. furcatum is much thinner than that of the following four species, and the dark perivascular scleren- chyma of the midrib is typically visible on the back. This character especially removes the species from C. connellii, which has thickly coriaceous fronds. The repand sterile lamina and very narrow fronds distinguish C. attenuatum and C. wurdackii respectively from C. furcatum. I have seen plants apparently inter- mediate between C. furcatum and each of the next three species. REPRESENTATIVE SPECIMENS EXAMINED:

VENEZUELA: Bolivar: Salto de Iwaracart-mert, at the W end of Sororopan-tepui, 1615 m, Steyermark 60227. Amazonas: Vicinity of Base Camp, Cerro Sipapo (Paraque), Maguire & Politi 27943. COLOMBIA: Cundinamarca: Las Cascades, S side of Guavio River 18 km NE of Gachala, 2070 m, Grant 10517. Vaupés: Rio Piraparana, Schultes & Cabrera 17069. TRINIDAD: Las Lapas Road, 2000 ft, Hombersley 277. SURINAM: Hendriktop, 1080 m, B. W. 5709a. GUYANA: Below — Tukeit, Potaro River Gorge, Maguire & Fanshawe 23484. BRAZIL: Glaziou 12368.

-13. Cochlidium tepuiense (A. C. Smith) L. E. Bishop, comb. nov.

Polypodium tepuiense A. C. Smith in Gleason & Killip, Brittonia 3: 148. 1939.°TY PE: Mt. Auyan- tepui, Edo. Bolivar, Venezuela, 1850 m, Tate 1248 (NY!; isotype US!).

Grammitis tepuiensis (A. C. Smith) Vareschi, Fl. Venez. 1: 871. 1969.

RANGE AND HABITAT: Widespread in the highlands of SE Venezuela. Epiphytic, at 1000-2000 m elevation.

This is the largest of the Guyana Highlands group of species. It is best charac- terized by the broad fronds up to 3 mm wide bearing two regular rows of sori which at maturity may be confluent lineally but are separate across the midnb. The entire margin separates it from C. furcatum and C. acuminatum. The larger

ae and somewhat separate sori remove it from the present concept of C. connel- li.

SPECIMENS EXAMINED:

VENEZUELA: Bolivar: Headwaters of the Rio Venamo, northern slopes of Cerro La Danta, NW of Cerro Venamo, 1040 m, Steyermark & Nilsson 53. Headwaters of the Rio Chicanan 80 km SE of El Dorado, Sierra de Lema, 700 m, Steyermark 89412. NE portion of Cerro Sarisarifama, Meseta del one 1400 m, Steyermark et al. 109184. Amazonas: Cerro Sipapo, Maguire & Politi 27615-B, 27708,

14. Cochlidium attenuatum A. C. Smith in Gleason, Bull. Torrey Bot. Club 58: 308. 1

Grammitis tatei Copel. Philipp. J. Sci. 80: 120. 1952, nom. nov. TYPE: Based on Cochlidium attenuatum A. C. Smith in Gleason, non G. attenuata Kunze.

TYPE: Mt. Duida, Venezuela, 5500-6000 ft, Tate 428 (NY! isotype US!)

RANGE AND HABITAT: Fairly widespread through the Guyana Highlands of Venezuela and Guyana. Epiphytic or epipetric at 1500-2000 m elevation.

92 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

This species is somewhat marginally represented by the type of C. attenuatum. The smaller plants of the type collection show very narrow fronds with essentially entire sterile laminae, superficially resembling C. wurdackii. However, the specimens have large sporangia, long scales, large stomata, and the larger plants do have repand margins. So I feel that this type is for now sufficiently well placed with, if not especially typical of, this species concept.

I have applied this name to plants whose most conspicuous and distinctive feature is the lightly repand sterile portion of the frond. Such plants agree in other characters as well. The fronds are narrow, often 1 mm or less in width; in large plants up to 15 cm they may reach 2 mm across. The sori are usually confined to the distal quarter of the frond, which is here a bit expanded, and are some what irregularly disposed, but more or less confluent at maturity. The sporangial cap- sules are larger than in the last two species, 160-220 »m long, and the stomata and scales are larger than those of C. wurdackii (65-90 x 55-65 zm and 2-3 mm long, respectively, in C. attenuatum).

REPRESENTATIVE SPECIMENS EXAMINED:

VENEZUELA: Bolivar: Cerro Guaiquinima, Rio Paragua, open savanna 1 km S of Cumbre Camp, 1800 m, Maguire 32796. SW portion of Cerro Jaua, 2228-2250 m, Steyermark et al. 109604. Mt. Roraima, im Thurn 166. Upper cumbre of the NW cumbres of Churi-tepui (Muru-tepui), 2250-2300 m, Wurdack 34242. Headwaters of the Rio Chicanan SW of El Dorado, Serra de Lema, 500 m, Steyer- mark 89598. Amazonas: Rio Cunucunuma, Cerro Huachamacari, 1800 m, Maguire et al. 30305.

15. Cochlidium connellii (C. H. Wright) A. C. Smith, Bull. Torrey Bot. Club 57:

Polypodium connellii Baker ex C. H. Wright in N.E. Brown, Trans. Linn. Soc. London, Bot. 6: 82. 1901. SYNTYPES: Summit, Mt. Roraima, British Guiana, McConnel & Quelch 570 (K not seen, isosyntypes NY! US!), /1/ (K not seen), 1/8 (K not seen).

RANGE AND HABITAT: Guyana Highlands of Venezuela and Guyana. Epipetric, terrestrial, or occasionally epiphytic; at 1600-2300 m elevation.

This is manifestly and admittedly the least satisfactory species concept in this treatment of the genus. I have followed general usage and, I hope, the original description by applying the name to those plants with erect, coriaceous, congested fronds growing in tufts at higher elevations. These typically have fronds 3-6 cm long and 1.5-2 mm wide (Steyermark 59673, Vareschi & Foldats 4777, 4866, 4887). Except for the size, shape, and habit of the fronds, these are otherwise similar to C. attenuatum in their larger sporangial capsules 170-220 wm long, stomata 70-95 um long, and distal, somewhat irregular sori. Also, the sterile laminae sometimes are very slightly repand. Contracted plants of C. attenuatum, such as Steyermark 109604, differ only in their narrower, more repand fronds. On the other hand, occasional plants with large sporangia seem very close to reduced examples of C. furcatum or C. tepuiense. As mentioned earlier, I believe that es field knowledge of these plants can clarify the problems here. so < seme of the name C. connellii and the present species concept must aie a snare japon: The protologue included three different numbers. I have wale ae es of only one of these, and even the plants included there are not

orm. For instance, the sheet at US has three mounted plants. One has lightly

L. E. BISHOP: REVISION OF COCHLIDIUM 93

repand fronds ca. 1 mm wide and does not differ from the concept of C. at- tenuatum as accepted here. The other two have entire, coriaceous fronds ca. 1.5 mm wide and up to 7 cm long with large sporangia. The sori are irregularly distant with the lamina expanded around them, much as in C. furcatum. These plants do not fit well into any of the species here recognized. However, until all of the original material can be examined and until a greater understanding of the Guyana Highlands Cochlidium species is attained, I think it unwise to disrupt current usage of the C. connellii.

REPRESENTATIVE SPECIMENS EXAMINED:

VENEZUELA: Bolivar: Upper part of Auyan-tepui, 2300 m, Vareschi & Foldats 4887. Plateau portion of SE-facing slopes of Ptari-tepui, 1600 m, Steyermark 59673. Cumbre of the east-central portion of the Meseta de Jaua, ca. 60 Km. NW of the mision de Campamento Sanidad on the Rio Kanarakuni, 1922-2100 m, Steyermark 97988. Amazonas: Cerro Sipapo (Paraque), 5500 ft, Maguire & Politi 27615-A. SE escarpment of Cerro Huachamacari, Rio Cunucunuma, 1900 m, Maguire et al.

h nov. Filicula gracillima tegetiformis quae ripas alpinas incolit. Rhizoma tenue repens frondes paulo dissitas gerens paleisque minutis triangularibus 0.4-0.7 mm longis

acroscopico distaliter recto neque non ramo breviore basiscopico patenti. Sori distales, pauci 1-5 in quaque fronde fertili, inordinate dispositi, lamina in area sor latitudine expansa. Sporangiorum capsulae 160-190 x 130- 170 wm, annuli cellulae 9-11, sporae 28-36 wm diametro.

HOLOTYPE: Abundant in mats on rocks along Rio Sarvén, 1 km downstream from Camp 9, Sarvén-tepui, Edo. Bolivar, Venezuela, 1200 m, Wurdack 34349 (US).

PARATYPE: Terrestrial, frequent on river banks, along Rio Tirica below Upper Falls above summit camp, central section of Chimanta Massif, Edo. Bolivar, Venezuela, 1940 m, Steyermark & Wurdack 577 (US).

This apparently localized species differs from all others of the genus in its very small rhizome scales. Other characters which distinguish it from species with discrete sori include the slender, creeping rhizome, the very narrow fronds, and the smaller stomata. The habitat is also unusual in the genus. Both collections show the plants forming mats in sandy soil. All other species are epiphytic or grow among epipetric bryophytes. Ba

The species is named for John J. Wurdack, not only for his responsibility for both gatherings, but also in recognition of the effect his considerable field effort has had in the advance of our knowledge of neotropical floras.

94 | AMERICAN FERN JOURNAL: VOLUME 68 (1978)

SPECIES IGNOTA NECNON DUBIA Pleurogramme myrtillifolia Fee, Mem. Foug. 3: 38. 1852; 5: t. 10c. 1852.

Monogramma myrtillifolia (Fée) Hooker, Sp. Fil. 5: 125. 1864, as ““Monogramme.”’

TYPE: Habitat in monte Tisis [Mexico?], collector unknown (possibly RB not seen).

This species is described as having small, spathulate, very thick leaves scat- tered on a long-repent rhizome. The locality is quite unknown to me, and it may not be American. Fée’s illustrations of the whole plant very much suggest one of the smaller species of Microgramma or Lemmaphyllum, and the details could conceivably be juvenile or misinterpreted. Despite its inclusion in Pleurogramme, I very much doubt that this plant is a Cochlidium, but only by the rediscovery of Fée’s type can the matter be settled.

LITERATURE CITED BISHOP, L. E. 1974. Revision of the genus Adenophorus (Grammitidaceae). Britonnia 26: 217-240. CHRISTENSEN, C. 1929. Taxonomic fern studies I. Dansk Bot. Ark. 6(3): 1-93. COPELAND, E. B. 1947. Genera Filicum. Ronald Press, New York. . 1952. Grammitis. Philipp. J. Sci. 80; 93-276. GOEBEL, K. 1924. Archegoniatenstudien. Flora 117: 91-132. HOOKER, W. J. 1864. Species Filicum, vol. 5. Dulau & Co., London. MAXON, W. R. 1926. Pteridophyta. Sci. Surv. Porto Rico Virg. Isls. 6(3): 373-521. MORTON, C. V. 1967. The genus Grammitis in Ecuador. Contr. U.S. Natl. Herb. 38; 85-123. PROCTOR, G. R. 1953. A Preliminary Checklist of Jamaican Pteridophytes. Institute of Jamaica,

>

Kingston. SMITH, c C. 1930. Notes on Pteridophyta from Mount Roraima. Bull. Torrey Bot. Club. 57:

WALKER, T. G. 1966. A cytotaxonomic survey of the pteridophytes of Jamaica. Trans. Roy. Soc. Edinb. 66: 169-237,

SHORTER NOTES

A DELETION FROM THE PTERIDOPHYTE FLORA OF NEBRASKA.—In the ‘*Atlas of the Flora of the Great Plains” (Great Plains Flora Association, lowa State University Press, Ames, 1977), Lycopodium annotinum was listed for the first time for Nebraska. The report was based on the following specimen: ppcnosiag Cherry County, Nebraska, Rev. John M. Bates s.n. in 1892 (NEB : 907). Recently I had the opportunity to examine this specimen and found it to

€ correctly determined. However, debris at the base of the plant indicated that this specimen could not have been collected in Nebraska. Par earies = debris were leaves and cone scales, both staminate and pistil- Le . ze auca, which does not occur in Nebraska. In addition, three MOS- Thuidi scovered: Polytrichum commune, Rhytidiadelphus triquetrus, and idium minutulum. The latter two species are not known in Nebraska. Consid-

stern United States, where Bates is known to have

coll ; “i imen’ lected on several occasions. In addition, the handwriting on the specimen $

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 3 (1978) 95

label is not that of Bates. For these reasons, it is assumed that there was a mixup of one sort or another which caused this error, and, therefore, Lycopodium an- notinum must be eliminated from the flora of Nebraska and the Great Plains. For making the necessary moss determinations I thank Steve Churchill, Botany De- partment, University of Kansas.—Ralph E. Brooks, University of Kansas Her- barium, 2045 Avenue A, Campus West, Lawrence, KS 66044.

CYSTOPTERIS TENNESSEENSIS IN ALABAMA.—The Tennessee Bladder Fern, C. tennesseensis Shaver, has been listed as occurring in northeastern Alabama by Dean (Ferns of Alabama, 1969). It was found at the mouth of Nick- a-Jack Cave in Jackson County, just within the state line. Now two additional localities can be reported from the north-central part of the state.

In August 1975, C. tennesseensis was found growing around a sinkhole in Morgan County. This locality is in Newsome Sinks, a large lime-sink valley about 20 miles south of Huntsville. The specimens lack the foliar bulblets usually pres- ent on plants of C. tennesseensis and were identified initially as C. fragilis var. mackayi Lawson. However, samples were identified recently as C. tennesseensis by Dr. W. H. Wagner, Jr. Vouchers (Short 386) are deposited at AUA and MICH, and later collections (Short 887) from the same locality will be distributed by AUA.

In the summer of 1976, a fern gametophyte with young sporophyte was found growing in damp soil under shrubs at the residence of Dr. William A. Short of Athens, in Limestone County. The development of this plant was observed, and by June 1977 the sporophyte had produced sori characteristic of Cystopteris and bulblets characteristic of C. tennesseensis. The habitat was unusual for this species; the fern grew in deep soil containing only a few pieces of limestone gravel from a nearby driveway. A voucher (Short 905) is deposited at AUA.

Shaver (Ferns of Tennessee, 1954) suggested that C. tennesseensis may have originated from hybridization between C. bulbifera (L.) Bernh. and C. protrusa (Weath.) Blasdell, since many of its characteristics are intermediate between these species. It reproduces by spores as well as bulblets, and probably is an allopolyploid, according to Blasdell (Mem. Torrey Bot. Club 21(4): 51. 1963).

Cystopteris tennesseensis generally grows on damp, shady, calcareous-rock outcrops. It ranges from northern Alabama to Michigan westward to Kansas, mainly in upland physiographic provinces, according to Wherry (The Fern Guide, 1961). It also has been reported on marl outcrops in the coastal plain of North Carolina, according to Radford, Ahles, and Bell (Manual of the Vascular Flora of the Carolinas, 1968), where we observed it in October 1977. The knowledge of this species’ distribution in Alabama is still incomplete, but suitable habitats are frequent in the northern counties and occasionally occur in the coastal plain. It can be inferred that C. tennesseensis probably is more widely distributed in Alabama than previously has been believed.—John W. Short and John D. Freeman, Department of Botany and Microbiology, Auburn University Agricul- tural Experiment Station, Auburn, AL 36830.

96 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

EQUISETUM x LITORALE RECORDED FOR MINNESOTA.—On 8 August 1977, while preparing an inventory of the floodplain flora of parts of the upper Mississippi River and its tributaries, one of us (SDS) collected a specimen of Equisetum from a large colony that was growing under a stand of Salix interior on dredge spoils and disturbed floodplain at the confluence of Valley Branch Creek and the St. Croix River (Mile 11.75) in Washington County, MN. The specimen was identified by one of us (JHP) as Equisetum x litorale Kuhl., which is the hybrid between E. arvense and E. fluviatile. Dr. Richard L. Hauke, of the Uni- versity of Rhode Island, verified the determination. A voucher (Swanson 2878) is deposited at University of Wisconsin-La Crosse (UWL). According to Dr. Gerald B. Ownbey, of the University of Minnesota, this is the first record for Minnesota.—James H. Peck and Steven D. Swanson, Dept. of Biology, Univer- sity of Wisconsin-La Crosse, La Crosse, WI 54601.

LYCOPODIUM CERNUUM IN LOUISIANA.—The Nodding Club-moss, re- cently reported by Eleuterius (Castanea 41: 180-181. 1976) from Jackson County, Mississippi, has recently been collected in central and southeastern Louisiana.

On 13 Sept 1975, Thomas and an aquatic plants class found several plants along a west-facing bank of the Pearl River Canal east of Louisiana route 41 and Evans Creek in St. Tammany Parish, Sec. 41, T6S, RI4E (Thomas 46276, NLU). The area was revisited by Thomas and Allen later in the same month, and three populations of L. cernuum were observed (Thomas & Allen 47241, NLU). No other ferns or fern allies were found. In the spring of 1977, another specimen of L. cernuum from St. Tammany Parish was collected by a student at Louisiana State University. Thomas, Landry, and others visited this site in the fall of 1977. The plants were in the bottom of an abandoned gravel pit near Hickory, La. Lycopodium cernuum, L. appressum, and L. carolinianum all were abundant, along with Burmannia capitata.

On 15 Oct 1976, Holmes co

det 1 llected two specimens of L. cernuum in a small roadside ditch in a seepage are

; : a in a longleaf pine woods along Middle Branch ——? - Red Dirt Game Management Area of Kisatchie Seunvine Forest in pee gay arish (Holmes 2902, NATC). This area is the largest hillside bog in L li ee abundance of bog plants, including Lycopodium appressum, .

- Cdrolinianum, Burmannia capitata, Pinguicula pumila, and Sarracenia alata. sively by Holmes and by Thomas on separate occa- 0 plants of L. cernuum could be located. The winter lled these plants, which are at the north- range. If this location represents a perma- » then L. cernuum should also be found in Vernon Parish and e . le Thomas, Dept. of Biology, Nort’ : onroe, LA 71209; W. C. Holmes, Dept. of Biologica St hertwvenens State University of Louisiana, a etiackek LA 71457; pie ae 5 Allen, 1] u Lucas Circle, Lafayette, LA 70508; and Garrie Landry,

P!. of Botany, Louisiana State University, Baton Rouge, LA 70803.

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Trichomanes Gametophytes in Massachusetts

BRUCE MCcALPIN and DONALD R. FARRAR 97

A New and Unique, Mat-forming Merlin’s-grass (Isoetes) from Georgia

State and Local Fern Floras of the United States, Supplement II

Shorter Notes: Athyrium filix-femina New to skatchewan; New Combinations in the Fern Flora of Venezuela; Trismeria. . .trifoliata?

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AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 4 (1978) 97

Trichomanes Gametophytes in Massachusetts BRUCE McALPIN* and DONALD R. FARRAR**

The final event in the 1976 New England Fern Conference convened by area pteridologists was a morning field trip to Mt. Toby, in Franklin County, Mas- sachusetts. An unexpected find on this foray was the filamentous gametophytes of an unknown species of Trichomanes. These were growing far back in crevices in non-calcareous, conglomerate rocks (pH 7) exposed along a small creek at the base of Mt. Toby. This is the habitat typical of the Trichomanes and Vittaria gametophytes found commonly in the southern Appalachian Mountains (Farrar, 1967, 1971). By shining a flashlight into the dark recesses, plants were seen to be relatively abundant, although individual clones were not extensive. The largest clone occupied an area approximately 50 cm?, whereas most were less than two cm in diameter. A subsequent foray provided additional specimens from this site.

The discovery of Trichomanes gametophytes in Massachusetts is of consider- able significance. This site is over 400 miles northeast of the most northerly stations previously known in Hocking County, Ohio and Pendleton County, West Virginia. It is also the first collection of Trichomanes from a site well within the area glaciated during the Pleistocene. All previous collections, which extend from West Virginia and Georgia westward to northwest Arkansas, are from unglaciated areas which have existed as dissected uplands at least since the late Tertiary.

Throughout their range in the temperate United States, the gametophytes are sterile. They often lack sex organs and apparently never produce sporophytes. Their reproduction is by vegetative buds, called gemmae (Fig. 4). Sporophytes of two species of Trichomanes, T. boschianum Sturm ex v. d. Bosch and T. petersii

Gray, occur in the same non-glaciated range as the Trichomanes gametophytes. A few diploid populations of 7. boschianum produce viable spores, but most populations of T. boschianum and possibly all populations of T. petersii in this area are polyploid and produce aborted spores. Sporophytes of both species produce dispersable vegetative buds.

Although the Trichomanes gametophytes in the temperate United States have generally been assumed to be of the same species as these sporophytes, this has not been proven. The occurrence of gametophytes in Massachusetts, over 400 miles from either sporophyte species, decidedly increases the probability that species other than T. boschianum and T. petersii may be represented. This collec- tion also points out the need for further search in the northeastern United States. It is likely that discovery of Trichomanes gametophytes will enlarge the pteridophyte flora of most New England states. We thus encourage field workers in this area to search for these small, alga-like plants (Figs. 1-4) and to report to us any additional discoveries. Specimens from Mt. Toby have been sent to the Gray Herbarium, the U. S. National Herbarium, and the Herbarium of the New York Botanical Garden.

*The New York Botanical Garden, Bronx, NY 10458.

’

*Department of Botany and Plant Pathology, lowa State University, Ames, IA 50011. Volume 68, number 3, of the JOURNAL was issued October 2, 1978.

AMERICAN FERN JOURNAL: VOLUME 68 (1978)

i 4

er ~

FIGS. 1-4. Trichomanes gametophytes. FIG. 1. Dark, overhanging rock outcrop nearly covered 2

clones . gametophytes, x 1/5. FIG. 2. Clone of gametophytes (arrow) surrounded by Mire 2. FIG. 3. Clone of gametophytes demonstrating growth habit, 10. poe

Gasciohen filaments with characteristic gemma (double arrow) and gemmifers (arrow), X |

LITERATURE CITED heir FARRAR, D. R. 1967. Gametophytes of four tropical fern genera reproducing independently oft Be oa in the southern Appalachians. Science 155: 1266-1267. United 971. The biology of ferns with asexually reproducing gametophytes in the eastern Mth Ph.D. Thesis, University of Michigan, Ann Arbor, MI.

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 4 (1978) 99

A New and Unique, Mat-forming Merlin’s-grass (Isoétes) from Georgia PHILLIP M. RURY*

The quillworts, Isoétes (Lycopsida, Isoétales), constitute a heterosporous fam- ily of about 60 species recognized by botanists since the time of Linnaeus as having fleshy, two-, three- or four-lobed corms with a crown of spirally arranged, ligulate microphylls and numerous dichotomizing roots that emerge in series along the median furrow(s) of the corm.

Vertical stem growth in Isoétes results from a shoot apical meristem that pro- duces sterile and fertile leaves and a central core of primary vascular tissue. Expansion growth is effected by the activity of a lateral meristem or ‘‘cambium”’ that encloses the primary vascular cylinder. During growth in girth, the original corm structure is maintained by radial displacement and seasonal loss of the outer, older leaves, roots, and corm tissue.

Sexual reproduction involves the formation of micro- and megasporangia, the spores of which develop endosporically into male and female gametophytes. Water-mediated cross fertilization produces daughter sporophytes.

Developmentally, young sporophytes exhibit a distinct change in their overall morphology (Paolillo, 1963). During the first 10-15 plastochrons, leaves are pro- duced alternately on either side of the embryonic shoot apex. During this stage, the apical meristem produces neither cauline vascular tissue nor primary stem growth; thus the vasculature of these distichous juveniles consists of a sympodium of traces to leaves and roots on either side of the shoot apex and median furrow. The resultant two-ranked leaf arrangement is lost, however, as soon as the apical meristem becomes large enough to initiate leaf primordia in a spiral sequence. Apparently, this apical enlargement also is responsible for the differentiation of the vascular core and the concomitant inception of vertical stem growth.

Vegetative propagation is rare in Isoétes. Goebel (1879) reported it in J. echinospora Dur. Goebel (1905, p. 431) also reported adventitious plantlets on /. lacustris leaves in a sporangial or sub-sporangial position in sterile specimens from the Vosges Mountains of France. Similar sterile but viviparous plants of /. lacustris were reported from Windmere, England by Manton (1950, p. 255). The developmental origin of such epiphyllous plantlets is not understood and is in need of investigation.

Branching in the upper portions of Jsoétes axes, another mechanism of vegeta- tive reproduction, also has been reported (Motelay & Vendryes, 1882; Solms- Laubach, 1902; Eames, 1936, p. 51), and a recent developmental study has re- vealed basal branching in two species of Isoétes (Karfalt & Eggert, 1977).

With the exception of branching and bud-formation, the quillworts traditionally have been considered as a morphologically uniform family of pteridophytes. The discovery of the Peruvian genus Stylites (Amstutz, 1957), however, has expanded

LL “Department of Botany, University of North Carolina, Chapel Hill, NC 27514.

100 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

our biological and morphological concept of the Isoétaceae. Detailed develop- mental and morphological studies of Isoétes (e.g. Paolillo, 1963), Stylites (Rauh & Falk, 1959 a, b), and the structural intermediate of these two genera, I. triquetra A. Br. (Kubitzki & Borchert, 1964), have indicated that the family Isoétaceae actually represents a polymorphic, clinal complex probably best treated as a single genus.

The profound influence of microhabitat upon Jsoétes morphology (Kubitzki & Borchert, 1964; Matthews & Murdy, 1969) must be considered in order to under- stand the complex polymorphism encountered within this family. The recent rec- ognition of the following new and unique species of Isoétes that is a narrow endemic in the Piedmont of Georgia further emphasizes the influence of habitat upon /soétes morphology and supports the concept of the extant Isoétaceae as a polymorphic, monogeneric family of pteridophytes.

a He

/ : : ce J

(OD “Be ~

1 Saf SY ' 2 GO on

FIG. 1. Habit of Isoétes fegetiformans. Triple plant with roots trimmed exhibiting seven micro- *porangia and distal, adventitious plantlets, x 3. [Drawing courtesy of Ms. Betsy Birkner, Staff Artist, Dept. of Botany, U.N.C., Chapel Hill]

Isoetes tegetiformans Rury, sp. nov. Figs. 1-8. _ Plantae amphibiae, sine os pape gerentes. Caulis prostratus, centraliter sulcatus, 3-35 mm longus,

ict rig folio Consociatae; folia pauca (4-8), disticha, subulata, 2-4 cm longa,

cin a et equitantia, ae peripheralibus absentibus, tomate ae

supremam dispositis, ligula minus quam 1 mm longa, :

a beth aire megasporangia elliptico-reniformia, ca. 1 mm longa, pete

microsporangis cil tice om diam., triletis, demisse tuberculatis, brunneis: oa giae Iptico-reniformia, ad 1 mm longa, microsporis multis, —_s

gis, monoletis, Spinulosis, brunneis.

P. M. RURY: A NEW MERLIN’S-GRASS FROM GEORGIA 101

TYPE: Heggies Rock, 3.8 mi E from Columbia Junior High School along Georgia Rte. 232 and County Road 2122, Columbia County, Georgia, 110 m elev., 33°32'30"N Lat., 82°15'05"W Long., 21 Jan 1978; solitary, dense, mat- forming population growing in a 2-4 cm thick layer of fine, siliceous soil in a S-facing pool ca. 20 m? and 30 cm deep containing water with a circumneutral pH, P. M. Rury & M. Treiber 259 (NCU; isotypes B, BH, DUKE, F, GA, GH, K, KYO, MASS, MICH, NY, UNCC, US).

This unique, mat-forming population of Jsoétes occurs in the largest of many pools on a 90-acre granitic flatrock. This pool is unique among the pools at Heg- gies Rock with respect to its location, aspect, size, structure, and resultant soil and water depths: The Jsoétes population dominates this large pool, nearly to the exclusion of other vascular plants, by virtue of its vegetatively reproductive, mat-forming (‘‘tegetiformans’’) growth habit.

The individual plants possess a slender, centrally furrowed, prostrate axis ca. | mm thick which exhibits bidirectional, intercalary-extension growth from the me- dian furrow. The resultant axes are 3-35 mm long and possess both a linear phyllo- and rhizotaxis. The shoot apical meristem overlies the median furrow and de- velops typical Isoétes leaves and sporophylls that are distichously disposed. However, unlike all other members of the Jsoétaceae and vascular plants in general, this shoot apex produces no cauline vascular tissue or vertical (primary) growth of the axis, and apparently has been reduced to a mere phyllogenic role. This type of distichous phyllotaxis and concomitant anatomy have been described by Paolillo (1963, p. 17) as a juvenile stage in Isoétes corm development which typically persists only for the first 10-15 plastochrons of the young sporophyte.

The rhizotaxis of these plants is distinctly tristichous, a feature previously reported only for the genus Stylites, with two lateral rows of slender roots emerg- ing proximal to the insertion of the leaf bases and one basal row of stout, often coiled roots which emerge proximal to the median furrow along the bottom of the axis. The roots of all specimens examined exhibit typical Jsoétes root anatomy but, as in J. triquetra and Stylites, are devoid of dichotomous branching. Usually three roots are produced per leaf, one from each row, and the resultant leaf-root complex becomes displaced along the axis and apparently becomes senescent as a unit. Developmentally, these three roots arise endogenously from a tracheary plexus in the vascular continuum, from which the single leaf trace departs to the associated, superadjacent leaf. This structural and developmental relationship between the roots and leaves of I. tegetiformans supports the contention of Liebig (1931) that the roots of Isoétes are actually adventitious, as in other vascular cryptogams where roots arise in association with leaves.

The vasculature of the axes consists entirely of a sympodium of leaf and root traces which is continuous across the median furrow. The vasculature of these Plants, therefore, supports the argument of both Farmer (1890) and Stokey (1909) that the vascular tissues of the Isoétes stem represent a sympodium of leaf traces and that there is no true cauline portion to the stele. This sympodium, its ensheathing stem tissue, and the associated leaf-root complexes are displaced bidirectionally as a result of the sub-apical, intercalary-extension growth from the

102 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

median furrow. The occurrence of a ‘“‘typical’’ isoétaceous cambium in the region of the median furrow suggests that the intercalary growth results from cell divi- sions of this cambium that are restricted to the plane of the furrow. The often contorted morphology of the prostrate axes may be the result of differential cam- bial activity and/or the crowded conditions of their microhabitat. Additional study of this phenomenon is clearly necessary.

recs ee eee a ' et = © © © F eerererePFYPTPRET : ; ' J i 2 Fe ££ S FEE

. IG. 2. Habit of Isoétes tegetiformans. Double and triple plants with roots trimmed. Note the pair of Stal, adventitious plantlets in the plant on the right. Scale is in centimeters.

Adventitious buds which develop into daughter plantlets are found distally on the axes at an observed minimum of 1-2 mm from the parental ‘‘leaf-producing apex.’’ These buds arise de novo from the cauline storage parenchyma and form a vascular continuum that is identical to, but non-contiguous with, that of the paren: tal axis. This bud formation potential is exemplified by the regeneration of seemingly dead stem fragments and by the remarkable ability of the entire popula- tion to survive repeated summer droughts and to revegetate after the water supply is replenished.

Sexual reproduction in this new species is typically isoétaceous, and the adaxial sporangia, which do not exceed 1 mm in breadth, are completely covered a subligular velum. The phenology of these plants apparently is seasonal, Sse PUCEOEPORES being produced in the winter-spring months and megaspores beans = in the summer-fall months. Although microsporogenesis was gee micro » mature microsporangia were seen to contain numerous, brown, spinu ee

Spores 25-31 wm long. Megasporogenesis is typically isoétaceous and begin

P. M. RURY: A NEW MERLIN’S-GRASS FROM GEORGIA 103

with the formation of four separate megasporocytes. Four to twenty megaspores were observed per sporangium; these exhibit a low-tuberculate sculpture, are dark brown, and are 275-370 um in diameter. The occurrence of numerous young sporelings of typical Isoétes-Stylites morphology indicates that sexual reproduc- tion is successful within the population.

Isoétes tegetiformans exhibits a combination of characteristics that is unique among the known species of Isoétaceae, namely: sexual maturity in permanently distichous plants; a shoot apex which produces leaves and sporophylls only; a tristichous rhizotaxis of demonstrably adventitious roots; unbranched, frequently dimorphic roots; sympodial stem vasculature; a prostrate, intercalary growth habit; and the formation of cauline, adventitious buds.

TABLE 1. COMPARATIVE FEATURES OF I. tegetiformans AND I. melanospora.

I. tegetiformans I. melanospora Population structure Dense, mat-forming Solitary, non-gregarious Juvenile morphology Fixed ‘‘neoteny”’ Environmentaly induced

and anatom Adventitious buds Cauline Absent Root morphology Non-dichotomizing, Profusely dichotomizing, frequently dimorphic consistently isomorphic Sporangia width Up to 1 mm 1-2 mm Velum coverage Consistently complete Complete to slight Megaspores per sporangium 4-20 8-32 (greater for plants ignable to J. piedmontana)

Megaspore size 275-370 um 350-480 um Megaspore sculpturing Low tuberculate Tuberculate to ridges Microspore length 26-31 zm 25-30 wm Microspore sculpturing Spinulose Smooth to papillate

Although unique with respect to this combination of features, this new quillwort can readily be recognized as a member of sect. Tuberculatae (sensu Pfeiffer, 1922) on the basis of its megaspore sculpture. It is interesting to note that the South American Stylites (2 spp.) and I. triquetra (the Stylites-Isoétes intermediate) also are assignable to this section and are the only members of the Isoétaceae which resemble J. tegetiformans in possessing non-dichotomizing roots. Within sect. Tuberculatae, I. tegetiformans is most similar to the members of the J. melano- spora Engelm. complex of the southeastern United States. Unbranched speci- mens of I. tegetiformans are superficially similar to those of the incompletely described J. melanospora on the basis of plant size, distichous leaf arrangement, degree of velum coverage, and megaspore features. However, upon closer com- parative study, a clear distinction between these two granitic endemics becomes apparent (see Table 1).

Isoétes melanospora was originally described by Engelmann (1877, p. 395) from Stone Mountain, a large granitic dome in DeKalb County, Georgia. Engelmann described this Quillwort as a distichous plant with a compressed, bilobed corm inhabiting muddy pools at the summit of the granitic dome. The major reproduc-

VOLUME 68 (1978)

AMERICAN FERN JOURNAL

P. M. RURY: A NEW MERLIN’S-GRASS FROM GEORGIA 105

tive features considered diagnostic for this species are sporangia completely cov- ered by the subligular velum and black, tuberculate megaspores. Subsequent to its discovery, J. melanospora has been collected from several additional sites on the granitic flatrocks of the southeastern Piedmont.

In his revision of the Isoétaceae of the southeastern United States, Reed (1965) included a species, J. piedmontana (Pfeiffer) Reed, which is closely related to and sympatric with J. melanospora. It is interesting to note in this regard that the narrow range of the granitic J. melanospora is included within the much broader range of J. piedmontana, a ubiquitous southeastern species which frequently in- habits the peripheral seepage areas and deeper soils (in pools) of these granitic outcrops. Subsequent comparative ecological and morphological study of /. melanospora and I. piedmontana has revealed the complete morphological inter- gradation of these two species (Matthews & Murdy, 1969). Furthermore, these authors uncovered a direct correlation between microhabitat features and the observed morphological intergradation of the populations investigated. Within a single population, they discovered that the smaller, distichous J. melanospora individuals are restricted to the shallower soils, whereas the larger, spiral-leaved (‘‘typical’’) specimens of J. piedmontana consistently inhabit the areas with greater soil depths. These authors suggested that introgression of these two sym- patric species might be responsible for their morphological intergradation. They apparently did not consider the possibility that these two ‘‘species’’ might actually represent different ontogenetic phases of the same species.

The neglect of developmental evidence in isoétaceous taxonomy has been acknowledged by Reed (1965), who stated that ‘‘no attempt has been made to collect any one species in various phases of its growth from the same colony.” I have observed both spiral and distichous individuals that are clearly assignable to I. melanospora (sensu Engelmann, 1877) within single populations at Stone Mountain and at Mt. Arabia, both in DeKalb County, Georgia. Distichous specimens were collected from both localities for comparative, uniform- environment growth studies in the laboratory. In 4-6 weeks, all of the distichous specimens from Stone Mountain and a majority of those from Mt. Arabia in- creased in stature and attained a spiral phyllotaxis characteristic of J. piedmon- tana and all other ‘‘typical,’’ sexually mature Isoétaceae. These observations, along with those of Matthews & Murdy (1969), strongly suggest that the distich- ous, juvenile form (sensu Paolillo, 1963) of J. melanospora merely represents an

FIGS. 3-8. Morphology and anatomy of Isoétes tegetiformans. FIG. 3. Triple, root-trimmed plant, following one year of ‘‘optimal” indoor cultivation, with two distal, adventitious plantlets, 1.4. FIG. 4. Same as Fig. 3, with arrows indicating the daughter plantlets, x 3. FIG. 5. Median longitudinal ‘ities SE'uis calize Clune: at vigke anaios tad 13 £ (mf). Note the distichous phyllotaxis and the vascular continuum (vc), x 6. FIG. 6. Detail from Fig. 5. Note the leaf producing apex (1pa), leaf Primordium (1p), ligule (1g), and the primordia of basal roots (brp). The leaf traces (It) and root traces (1g) are components of the vascular continuum (vc), x 15. FIG. 7. Scanning electron micrograph of several monolete, spinulose microspores, x 700. FIG. 8. Scanning electron micrograph of a trilete, low-tuberculate megaspore, x 87

106 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

arrested (neotenic) stage in the development of plants assignable to [. piedmon- tana. Although the present evidence seems convincing, further comparative biological studies of additional populations of these two ‘‘species’’ are desirable in order to confirm their synonymization. Nevertheless, it can be concluded that I. melanospora is not a strictly distichous species, as originally described by Engel- mann (1877), and thus is in need of redefinition.

Having clarified reasonably well the morphological nature of J. melanospora and J. piedmontana, a clear distinction between I. tegetiformans and I. melano- spora sensu lato can be made. Isoétes tegetiformans resembles the distichous phase of J. melanospora with respect to plant size, leaf arrangement, and repro- ductive features. However, I. tegetiformans can be distinguished easily on the basis of its dense population structure; growth habit, stem morphology and anatomy, permanent distichy, unbranched, often dimorphic roots, and cauline adventitious buds. A comparison of the characteristics of J. tegetiformans and I. melanospora is presented in Table 1.

It is evident that the distichous, prostrate habit shared by these two granitic quillworts is merely the result of a neotenic convergence in form that is temporary in J. melanospora but permanent in I. tegetiformans. Indoor cultivation of the latter species for more than 18 months under the same growing conditions experi- enced by specimens of J. melanospora resulted in no significant morphological changes. This suggests that the neotenic form of I. tegetiformans is an immutable, species-specific characteristic, whereas a similar neotenic phenomenon in I. melanospora merely results from microhabitat influence upon the stature and correlative degree of pre-sexual morphological development of the individuals. In addition to being a fixed neotenic form, I. tegetiformans is also very distinct from I . melanospora (and all other North American species of /soétes) in having non- dichotomizing, frequently dimorphic roots and in forming numerous cauline, ad- ventitious buds (plantlets). Beyond such general considerations, however, the precise nature of the relationship between I. tegetiformans and I. melanospora is enigmatic, and probably shall remain so until a thorough systematic study and taxonomic revision of the entire J. melanospora complex is completed.

a epee causing the unique morphology of this plant are presently conjec- rural and require further investigation. However, Lammers (1950) revealed that Bee Se hi I. melanospora subjected to full sunlight at ca. 10,000 ft-c exhibit a lc rate nearly twice that of their dark respiration rate. If the photo- eee response curve and relative rate of dark respiration are comparable iF is Pied on the prolific growth habit of this species is easily understood inhi ca a sae = and microhabitat. The pool it inhabits is sou nth fe dese cee ank, and has a maximum soil and water depth that ene ihe Syélntinsias ate : natural, south-facing spillway from the pool. Throug ee the accuraiiaten of * Sitka Fe ores Ue Spa ay — webs er the potential stature of - ie, ee eel . me lant states ale ai | e ae The long-term effect of such restrictions on P ps sible for ns potential optimal photosynthetic efficiency, may be resp € unique morphology and prolific growth habit of this Merlin’s-grass-

P. M. RURY: A NEW MERLIN’S-GRASS FROM GEORGIA 107

Therefore, this spreading, carpet-forming species might best be regarded, evolutionarily, as an Jsoétes which ‘‘grew-out”’ like a grass instead of ‘‘growing- up”’ like a Quillwort.

ACKNOWLEDGMENTS

I would like to thank Dr. Albert E. Radford (Univ. of North Carolina, Chapel Hill) for taking me and others to Heggies Rock, the site of the present discovery, and for his diligent personal efforts to establish this outstanding granitic flatrock as a “national natural landmark area.”’ I also wish to express my deep appreciation and gratitude to Dr. David W. Bierhorst (Univ. of Massachusetts), Amherst) for his encouragement, technical assistance, and consultation during the course of this study, and to Dr. Warren H. Wagner (Univ. of Michigan, Ann Arbor) for his encouragement and advice. Many thanks are also extended to Dr. William C. Dickison (Univ. of North Carolina, Chapel Hill) for his technical assistance and for reviewing the manuscript and to Dr. Patricia G. Gensel (Univ. of North Carolina, Chapel Hill) for her assistance in scanning electron microscopy and for her critical evaluation of the manuscript. I would also like to express my gratitude to Dr. William J. Dress (Bailey Hortorium, Comell University) for his assistance in preparing the Latin species description and to Dr. Timothy C. Plowman (Bo- tanical Museum, Harvard University) for his helpful comments.

LITERATURE CITED

AMSTUTZ, ERICA. 1957. Stylites, A new genus of Isoetaceae. Ann. Mo. Bot. Gard. 44: 121-123.

EAMES, A. H. 1936. Morphology of Vascular Plants, ed. 1. McGraw-Hill, New York, NY. ;

ENGELMANN, G. 1877. About the oaks of the United States. Trans. St. Louis Acad. Sols: 3: 2-400.

FARMER, J. B. 1890. On Isoetes lacustris L. Ann. Bot. 5: 37-62, t. V-VI.

GOEBEL, K. 1879. Ueber Sprossbildung auf Isoétesblattern. Bot. Zeit. 37: 1-6.

———. 1905. Organography of Plants, part II. Clarendon Press, Oxford.

KARRFALLT, E. E. and D. A. EGGERT. 1977. The comparative morphology and development of Isoetes L. II. Branching of the base of the corm in I. tuckermannii A. Br. and I. nuttallii A. Br. Bot. Gaz. 138: 357-368.

KUBITZKI, K. and R. BORCHERT. 1964. Morphologische Studien an Isoetes triquetra A. Br. und Bemerkungen uber das Verhaltnis der Gattung Stylites E. Amstutz zur Gattung Isoetes L. Ber. Deutsch. Bot. Ges. 77: 227-234. :

LAMMERS, W. T. 1950. A Study of Certain Environmental and Physiological Factors Influencing the Adaptation of Three Granite Outcrop Endemics: Amphianthus pusillus Torr., Isoetes melanospora Engelm., and Diamorpha cymosa (Nutt.) Britt. Unpublished Ph.D. Thesis, Emory University, Atlanta, GA. 85 pp.

LIEBIG, JOHANNA. 1931. Erganzungen zur Entwicklungsgeschicte von Isoétes lacustre L. Flora 125; 321-358. :

MANTON, IRENE. 1950. Problems of Cytology and Evolution in the Pteridophyta. University Press, Cambridge. :

MATTHEWS, J. F. and W. H. MURDY. 1969. A study of Isoetes common to the granite outcrops of the southeastern piedmont, United States. Bot. Gaz. 130: 53-61. .

MOTELAY, L. and A. VENDRYES. 1882. Monographie des Isoétaceae. Actes Soc. Linn. Bor- deaux 6: 309-409.

PAOLILLO, D. J. 1963. The Developmental Anatomy of Isoetes. Illinois Biol. Monogr. 31: 1-130.

PFEIFFER, NORMA E. 1922. Monograph of the Isoetaceae. Ann. Mo. Bot. Gard. 9: 79-232.

108 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

RAUH, W. and H. FALK. 1959a. Stylites E. Amstutz, eine neue Isoetaceae aus den Hochanden Perus. I. Teil: Morphologie, Anatomie, und Entwicklungsgeschichte der Vegetationsorgane. Sitzungsber. Heidelberger Akad. Wiss. 1959: 1-83.

——.. 1959b. Ibid. II. Teil: Zur Anatomie des Stammes mit besonderer Berucksichtigung der Ver- dickungsprozesse. Ibid. 1959: 87-160.

REED, . 1965. Isoetes in southeastern United States. Phytologia 12: 369-400.

SOLMS-LAUBACH, H. GRAF zu 1902. Isoétes lacustris, seine Verzweigung und sein Vorkommen in den Seen des Schwarzwaldes und der Vogesen. Bot. Zeit. 60: 179-206, t. VII.

STOKEY, ALMA G. 1909. The anatomy of Isoetes. Bot. Gaz. 47: 311-335, t. XIX-XXI.

|

REVIEW

coe PTERIDOPHYTORUM GENERA IN TAXONOMICUM OR- DINEM REDIGENDI,” by R. E. G. Pichi Sermolli, Webbia 31: 313-512. 1977.— Professor Pichi Sermolli has written a most detailed and exhaustive treatment of fern classification down to the level of genus. His treatment is illustrated with a table of families and tree-like diagrams of the genera or groups of genera within each family, and includes a conspectus of the families and genera with generic numbers allowing for intercalation of additional genera, an index of the genera and their synonyms, and discussions of the genera and their characteristics.

In contrast to most linear arrangements of genera proposed in the past, Pichi Sermolli’s treats all the genera of a single line of evolution from primitive to advanced before passing on to another line of evolution. In my opinion, this is a more natural and satisfactory method than is treating first all primitive genera and = all advanced ones, regardless of which of several phyletic lines they belong

His classification groups 443 genera of living pteridophyta into 64 families— both record high numbers. Many of the genera I consider subgenera at best. Judging by the lengthy discussion of the genera, Pichi Sermolli gives more weight re differences in morphology than do most authors; he is a self-admitted

er.

There is one good reason for such splitting in proposing a classification: it forces users to consider the differences between finely split genera which otherwise might be ignored. The disadvantage is that in using one level of classification sortie than two (genus and subgenus), Pichi Sermolli’s classification contains less peor about relationships of genera within his families. Fortunately, his sie Ike lagrams convey this information, although his conspectus and any her-

arlum arrangement based on it would not. Whether one adopts finely split gener@ or —o comprehensive ones is to some extent a matter of personal preference OF pesca way Pichi Sermolli’s work will be very useful for a long time to

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 4 (1978) 109

State and Local Fern Floras of the United States, Supplement II MERYL A. MIASEK*

This list continues the two publications of S. F. Blake (Amer. Fern J. 31: 81- 90, 131-143. 1941; 40: 148-165. 1950) to bring to the attention of JOURNAL readers recent Flofas, manuals, and checklists dealing with ferns and fern-allies of the United States” Supplement II covers the years 1950-1977. The list is or- ganized alphabetically by state and author, with regional works at the end. Mimeographed lists and other ephemera are excluded. Works dealing with Cana- dian ferns are represented only when pteridophytes of the United States also are treated. Many short papers dealing with range extensions and locality records also are excluded; some of these are given in a list titled ‘“‘Regional, State and Local Fern Floras, Manuals, Checklists and New Fern and Fern-ally Records for the U.S. and Adjacent Canada, Since 1950,’ which I compiled. This list is available from the Library of the New York Botanical Garden, Bronx, NY 10458.

ALABAMA DEAN, BLANCHE E. 1964. Ferns of Alabama and Fem Allies. American Southern, Northport, AL. 232 p

Pp. . 1969. Ferns of Alabama, rev. ed. Southern Univ. Press, Birmingham, AL. 214 pp. ALASKA

ANDERSON, J. P. me Flora of Alaska and Adjacent Parts of Canada. lowa State Univ. Press,

Ames, IA. 543 p GIAEREVOLL, O. “1958. Botanical investigations in central Alaska, especially in the White Moun-

tains. Part I. Pteridophytes and Monocotyledons. K. Norske Vedensk. k. Selsk. Skr. 1958(5): 1-74. uaa E. 1968. Flora of Alaska and Neighboring Territories. Stanford Univ. Press, Stanford,

008 p WELSH, s | ibe cen Anderson’s ee of Alaska and Adjacent Parts of Canada. Brigham Young . Press, Provo, UT. 724 WIGGINS, I. L. and J. H. rows S. 1962. A Flora of the Alaskan Arctic Slope. Division I. Pteridophytes, pp. 33-44. eg Institute of North America Spec. Publ. No. 4.) Univ. of Toronto Press, Toronto. 425 p ARIZONA BOHRER, VORSILA L. and MARGARET BERGSENG. 1963. An Annotated Catalogue of Plants from Window Rock, Arizona. Navajo Tribal Museum, Window Rock, AZ. 29 pp. MORTON, C. V. 1964. Pteridophyta, pp. 27-49. In : H. Kearney and R. H. Peebles. Arizona ora. Univ. of California Press, Berkeley. 1085 p PASE, C. P. and R. R. JOHNSON. 1968. Flora and 5 fee oa of the Sierra Ancha Experimental Forest, Arizona. (U.S.D.A. For. Serv. Res. Paper RM-41.) Rocky Mountain Forest and Range Experiment Station, Ft. Collins, CO. 20 pp. ARKANSAS THOMPSON, R. L. 1977. The vascular flora of Lost Valley, Newton County, Arkansas. Castanea 42: 61-94.

*Library, New York Botanical Garden, Bronx, NY 10458.

110 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

CALIFORNIA FERLATITE, W. J. 1974. A Flora of the Trinity Alps of Northern California. Univ. of California Press. Berkeley. 206 pp. GRILLOS, S. J. 1966. Ferns and Fern Allies of California. Univ. of California Press, Berkeley. 104

Pp. HOOVER, R. F. 1966. An annotated list of the pteridophytes of San Luis Obispo County, California. AFJ 56: 17-26. —. 1970. The Vascular Plants of San Luis Obispo County, California. Univ. of California Press, Berkeley. 350 HOWELL, J. T. ad Mars Flora, 2nd ed. Univ. of California Press, Berkeley.

——, P. H. RAVEN, and P. RUBTZOFF. 1958. A Flora of San lone ome oe printed Mane Wasmann J. Biol. 16.) Univ. of San Francisco Press, San Francisco, CA. 157 p KIEFER, L. L. and BARBARA JOE. 1967. Check list of California pteridophytes. Madrofio 19:

73.

65- MUNZ, A “e we A Flora of Southern California, Univ. of California Press, Berkeley. 1086 pp. meonaane es D. D. KECK. 1959, A California Flora. Univ. of California Press, Berkeley. 1681 pp. bei Ps 1952. A Flora of Santa Barbara. Santa Barbara Botanic Garden, Santa Barbara, CA. 100

THOMAS. J. H. 1961. Flora of the Santa Cruz Mountains of California. Stanford Univ. Press, Stanford, CA. 434

TRUE, G. H. 1973. The Fens and Seed Plants of Nevada County, California. California Academy of Sciences, San Francisco, CA, 62

TWISSELMAN, E. C. 1967. A Flora of cae County, California. oe 2d from Wassmann J. Biol. 25.) Univ. of San Francisco Press, San Francisco, CA. 395

WITHAM, HELEN. ae Ferns of San Diego County. San — Natural History Museum, San Diego, CA. 72 p

COLORADO

BARRELL, J. 1969. Flora of the Gunnison Basin aye Saguache, and Hinsdale Counties,

Colorado). Natural Land Institute, Rockford, IL. 4 p.

car aca H. D. 1964. Manual of the Plants of eee. 2nd ed. Sage Books, Denver, CO, Pp

d L. W. DURRELL. 1950. Colorado Ferns and Fern Allies-Pteridophyta. Colorado Agricultural Research Foundation, - Collins, CO. 96 WELSH, S. L. and J. A. ERDMAN. 1964. Annotated checklist . ms plants of Mesa Verde, Colorado. Brigham Young aa Sci. Bull., Biol. Ser. 4(2): CONNECTICUT

BRADLEY, L. J. 1955. The Ferns and Flowering Plants of the Audubon Center, Greenwich, Con- necticut. Audubon Center, Greenwich, CT. 100 pp.

DELAWARE REED, C. F. 1953 (see MARYLAND)

DISTRICT OF COLUMBIA BLAKE, S. F. 1957. Ferns and fern-allies of the District of Columbia. AFJ 47: 149-155. REED, C. F. 1953 (see MARYLAND) FLORIDA LAKELA, OLGA and F. C. CRAIGHEAD. 1965. Annotated Checklist of the Vascular Plants of

Collier, Dade and Monroe Counties, Florida. Fairchild Tropical Garden and the UR % Miami Press, Coral Gables, FL. 95

» and R. W. LONG. 1976. Fems of Florida. Banyan Books, Miami, FL. 178 pp-

M. A. MIASEK: STATE AND LOCAL FERN FLORAS 111

LONG, R. W. and OLGA LAKELA. 1971. A Flora of Tropical Florida; a Manual of the Seed Plants and Ferns of Southern Peninsular Florida. Univ. of Miami Press, Coral Gables, FL. 962 pp

GEORGIA McVAUGH, R. and J. H. PYRON. 1951. Ferns of Georgia. Univ. of Georgia Press, Athens, GA. 195 pp HAWAII

HUBBARD, D. H. 1952. Ferns of Hawaii National pee peor Nature Notes 5(1).) Hawaii Natural History Assn., Hawaii National Park, HI.

IDAHO

DAVIS, R. J. 1952. Flora of Idaho. Wm. C. Brown, serie oi IA. 828 pp. FLOWERS, S. 1950. A list of the ferns of Idaho. AFJ 40: -131 ST. JOHN, H. 1963. (see WASHINGTON)

ILLINOIS DOBBS, R. J. 1963. Flora of Henry County, Illinois. Natural Land Institute, Rockford, IL. 350 pp. +

ap. FELL, E. W. 1955. Flora of Winnebago County, Illinois. Nature Conservancy, Washington, DC. 207

Pp.

JONES, G. N. 1963. Flora of Capps ed. (Amer. Midl. Nat. Monogr. 7.) Univ. of Notre Dame Press, Notre Dame, IN. 401 p ile D. FULLER. 1955. (arene Plants of Illinois. Univ. of Illinois Press, Urbana, IL.

593 pp. MOHLENBROCK, R. H. 1954. Flowering Plants and Ferns of Giant City State Park. Division of Parks and Memorials and Illinois State Museum, Springfield, IL. 24 p ————. 1955-56. The pteridophytes of Jackson County, Illinois. AFJ 45: 143-150; 46: 15-22. . 1967. es Illustrated Flora of Illinois-Ferns. Southern Illinois Univ. Press, Carbondale, IL. 191 p e ADT. ok to the Vascular Flora of Illinois. Southern Illinois Univ. Press, Carbondale, IL. 494 pp. , and J. W. VOIGT. 1974. A Flora of Southern Illinois. Southern Illinois Univ. Press, Carbon dale, IL. 390 pp. SWINK, F. 1974. Plants of the Chicago Region, 2nd ed. Morton Arboretum, Lisle, IL. 474 pp.

INDIANA

CLEVENGER, SARAH. Hh bg distribution of the ferns and fern allies found in Indiana. Butler Univ. Bot. Stud. 10:

DEAM, C. C. 1970. shed a sot (Reprinted from the 1940 edition.) S-H Service Agency, New York, NY. 1236

MOHLENBROCK, RK. H. ne JANE HINNERS ENGH. 1964. Fems and fern allies of Pine Hills Field Station and environs (Illinois). AFJ 54: 25-38.

IOWA COOPERRIDER, T. S. 1959. The ferns and other pteridophytes of Iowa. State Univ. of lowa Stud, Nat. Hist. 20(1): 1-66. . 1960. The Lycopodiaceae and Selaginellaceae of lowa. AFJ 50: 267-268. - 1962. The vascular aes of Clinton, Jackson and Jones Counties, lowa. State Univ. lowa St ud, Nat. Hist. 20(5): EILERS, L. : 1971. The ae ts of the Iowan area. State Univ. Iowa Stud. Nat. Hist. 21(5): 11-1

112 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

GULDNER, L. F. 1960. The Vascular Plants of Scott and Muscatine Counties. (Davenport Public Mus. Publ. Bot. 1.) M. L. D. Putnam Memorial Fund, Davenport, IA. 228 pp.

PECK, J. H. 1976. The pteridophyte flora of lowa. Proc. lowa Acad. Sci. 83: 143-160.

THORNE, R. F. 1964. Relict nature of the flora of White Pine arenas Forest Reserve, Dubuque County, Iowa. State Univ. lowa Stud. Nat. Hist. 20(6): 1-33.

KANSAS BARKER, W. T. 1969. The flora of the Kansas Flint aoe ean Kansas Sci. Bull. 48: 525-584. BROOKS, R. 1967. Ferns in Kansas. Kansas School Nat. ~15: HUMFELD, P. H. 1951. A checklist of Kansas sihaeneal any 41: 53-60, 79-85. McGREGOR, R. L. 1960. Ferns and allies in Kansas. AFJ 50: PETRIK-OTT, ALETA JO. 1975. A county checklist of the Ans and fern allies of Kansas, Ne- braska, South Dakota, and North Dakota. Rhodora 77: 478-511.

KENTUCKY

WHARTON, MARY E. and R. W. BARBOUR. 1971. A ni to the Wildflowersand Ferns of Kentucky. Univ. of Kentucky Press, Lexington, KY.

LOUISIANA MAPLES, R. S., JR. and D. D. LUTES. 1966. A checklist of ferns in Lincoln Parish, Louisiana, AFJ 56: 33-36.

MAINE WISE, D. A. 1970. The flora of Isle Au Haut, Maine. Rhodora 72: 505-532.

MARYLAND REED, C. F. 1953. The Ferns and Fern-allies of en and Delaware, Including District of Columbia. Reed Herbarium, Baltimore, MD. 286 p . 1960. (see VIRGINIA)

MASSACHUSETTS ARNOLD, H. J. and S. W. BAILEY. 1957. Bartholomew’ s Cobble, Sheffield, Massachusetts. Eagle Print and Bindery, Pittsfield, MA. Unpaged. EATON, R. J. 1974. A Flora of Concord. (Mus. Comp. Zool. Spec. Publ. 4.) President and Fellows of Harvard College, Cambridge, MA. 236 wowse* S. . 1975. The Flora of Essex County, Massachusetts. Peabody Museum, Salem, MA.

269 p KNOWLTON, C. H. 1950. Flora of Rocky feces Reservation, Medfield, Massachusetts. Trustees of Public Reservations, Boston, MA. MacKEEVER, F. 1968. Native and Notice ours of Nantucket. Univ. of Massachusetts Press, Amherst, MA. 132 pp.

MICHIGAN

BILLINGTON, C. 1952. Fems of Michigan. (Cranbrook Inst. Sci. Bull. 32.) Cranbrook Institute of Science, Bloomfield Hills, MI. 240

HAGENAH, 1963. Pteridophytes of the Huron Mountains, Marquette County, Michigan.

Michigan Bot. 2: 78-92.

HALL, M. T. and P. W. THOMPSON. 1959. ~ annotated list of the plants of Oakland County Michigan. Cranbrook Inst. Sci. Bull. 3

VELDMAN, LORRAINE C. and D. E. WUIEK. 1971. Pteridophytes of Beaver Island, Charlevoix County, Michigan. Michigan Bot. 10: 194-196.

M. A. MIASEK: STATE AND LOCAL FERN FLORAS 113

MINNESOTA BUTTERS, F. K. and E. C. ABBE. 1953. A floristic study of Cook County, northeastern Minnesota. odora 55: 21-55, 63-101, 116-154, 161-201. LAKELA, OLGA. 1952. A list of ferns from the north shore of Lake Superior, Minnesota. AFJ 42: 16-18. . 1953. A checklist of ferns and flowering plants of the Quetico-Superior Wilderness Re- search Center. Amer. Midl. Nat. 50: 488-500, . 1965. A Flora of Northeastern Minnesota. Univ. of Minnesota Press, Minneapolis, MN.

54 1 pp.

MOORE, J. W. 1973. A Catalog of the ait of Cedar Creek Natural History Area, Anoka and Isanti Counties, Minnesota. (Occ. Pap. Bell Mus. Nat. Hist. 12.) Bell Museum of Natural History, University of Minnesota, Palit MN. 28

MOYLE, J. B. 1964. Northern Non-woody Plants; a Field Key to the More Common Ferns and aig Plants of Minnesota and Adjacent Regions, rev. ed. Burgess, Minneapolis, MN.

TRYON, R. iL 1954. The Ferns and Fern Allies of Minnesota. Univ. of Minnesota Press, Min- neapolis, MN. 166 pp. MISSISSIPPI JONES, S. B., Jr. 1969. The pteridophytes of Mississippi. Sida 3: 359-364. PULLEN, T. M. 1966. Additions to the fern flora of Mississippi. AFJ 56: 37. MISSOURI

HANEBRINK, E. L. 1958. The Flora of Southeast Missouri. Throw Print, Kennett, MO. 78 pp. STEYERMARK, J.A . 1963. Flora of Missouri. lowa State Univ. Press, Ames, IA. 1725 pp.

MONTANA

DORN, R. D. and JANE L. DORN. 1972. The Ferns and Other Pteridophytes of Oa Wyo- ming, and the Black Hills of South Dakota. The Authors, Laramie, WY. 94

NEBRASKA PETRIK-OTT, ALETA JO. 1975. (see KANSAS)

NEVADA BEATLEY, JANICE C. 1969. Vascular Plants of the Nevada Test Site, Nellis Air Force Range, and Ash Meadows. Univ. of California Laboratory of Nuclear Medicine and Radiation Biology, Los Aides, CA. 122 p

County, and Adjacent Parts of Clark, Lincoln, and Esmeralda Counties, Nevada. Univ. of California Laboratory of Nuclear Medicine and Radiation Biology, Los Angeles, CA. 49 at

CLOKEY, I. - 1951. Flora of the Charleston Mountains, Clark County, Nevada. Univ. Calif. Publ. Bot. 24: 22-27

NEW HAMPSHIRE

PEASE, A. S. 1964. A Flora of Northern New Hampshire. New England Botanical Club, Cambridge, MA. 278 pp.

NEW JERSEY MONTGOMERY, J. D. 1972. Pteridophytes of the Jockey Hollow section of Morristown National Historical Park, New Jersey. Bull. Torrey Bot. Club 99: 139-142

114 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

NEW MEXICO

DITTMER, H. J., E. F. CASTETTER, and O. M. CLARK. 1954. The Ferns and Fern Allies of New Mexico. (Univ. New Mexico Publ. Biol. 6.) Univ. New Mexico Press, Albuquerque, NM. 139 pp

NEW YORK

BROOKS, K. L. 1956. Notes on the pteridophytes of Delaware County, New York. AFJ 46: 109-121.

BYE, R. A., Jr. and F. W. OETTINGER. 1969. Vascular Flora of Onondaga County, New York. State Univ. College of Forestry, Syracuse Univ., Syracuse, NY. 248

CROCKETT, L. J., ed. 1967. Some contributions to the floristics, plant ecology, and geology of the Hudson Highlands section of the lower Hudson Valley. Sarracenia 11: 1-89.

DOMVILLE, MARY and H. F. DUNBAR. 1970. The Flora of Ulster County, New York; an ee List of Vascular Plants. John Burroughs Natural History Society, New Paltz,

NY. 136 DUNBAR, H. F., a. 1961. Ferns of Ulster County, New York. John Burroughs Natural History Society, New Paltz, NY. 8 pp. —— “§ Sei M. WEINGARTNER. 1962. Ferns of Staten Island. Bull. Torrey Bot. Club 89: 335.

ie E. 1975. Ferns along the Bronx River Parkway. Fiddlehead Forum 2(1): 2. sacha H. A. 1962. Plants of the Vicinity of New York, rev. ed. Hafner, New York, NY. 307

G0LbEN. D. P. and E. B. EHRLE. 1971. Studies on the plants of the Genesee Country, 7. The ferns of Livingston County, New York. Proc. Rochester Acad. Sci. 12: 139-145. GRELLER, A. M. 1977. A vascular flora of the forested portion of Gapping Park, Queens

County, New York, with notes on the vegetation. Torreya 104: 6. HEHRE, E. J., Jr. 1977. The flora of Gardiners Island. Rhodora 79: HL pee N MIANUS RIVER GORGE CONSERVATION COMMITTEE OF THE NATURE CONSER Manet 1961. Flora and Fauna of the Mianus River Gorge Wildlife Refuge and Botanical reserve. Bedford Village, NY. 35 pp. SMALL, 3 K. 1975. Ferns ng see of New York. (Reprinted from the 1935 edition.) Dover, WweYor NY. w4.2

NORTH CAROLINA BLOMQUIST, H. L. and H. J. OOSTING. 1953. A Guide to the Spring and igs Summer Flora of the Piedmont, North Carolina, Sth ed. The Authors, Durham, NC. pp. BROWN, C. A. 1957. Botanical Reconnaissance of the Outer Banks of hie? Carolina. (Coastal Stud, “pe Tech. Rep. 8(C).) Louisiana State Univ. Coastal Studies Institute, Baton Rouge,

LA. 17

EVANS, A. M. 1964. Pteridophytes, pp. 3-34. In A. RADFORD et al. Manual of the esos Flora of the Carolinas. Univ. of North Carolina Press, Chapel Hill, NC. Ixi + 1183 p

FREEMAN, O. M. 1953. Preliminary report on the ferns and their allies in Polk cone North Carolina. seers 18: 60-63.

earessncts - J., D. L. NICKRENT, and G. F. LEVY. 1977. A contribution towards a

ascular Ne of the Great Dismal Swamp. Rhodora 79: 240-268.

PALMER. PATRICIA G. 1970. The sii He Overton Rock Outcrop, Franklin County, ane Carolina. J. Elisha Mitchell Soc. 86: 8

NORTH DAKOTA PETRIK-OTT, ALETA JO. 1975. (see KANSAS) OHIO

ANLIOT, S. F. 1973. The Vascular Flora of Glen Helen, Clifton Gorge, and John es State Park. (Ohio Biol. Surv. Biol. Notes 5. ) Ohio State Univ., Columbus, OH. 162

M. A. MIASEK: STATE AND LOCAL FERN FLORAS 115

BRAUN, E. LUCY. 1969. An ecological survey of the vegetation of Fort Hill State Memorial, Highland County, Ohio and annotated list of vascular plants. Bull. Ohio Biol. Surv. 3(3): -134. CHUEY, a F.and N. STURM. hee iby oe checklist of Ashtabula, Trumbull and Mahoning nties, Ohio. AFJ 59: 4 VANNORSDALL, H. H. 1956. oe a Ohio. The Author, Wilmington, OH. 298 pp.

OKLAHOMA WATERFALL, U. T. 1952. A Catalogue of the Flora of Oklahoma: a List of the Ferns and Flowering Plants Growing Native or Naturalized in the State, with Reference to Pertinent Literature. Research Foundation, Stillwater, OK. 91 pp

OREGON

BAKER, W. H. 1951. Ferns of Iron Mountain, Rogue River Range, Oregon. AFJ 41: 20-23. and G. = RUHLE. 1957. Ferns of Oregon Caves National Monument and hare AFJ

7: 98-1

caorearin EXTENSION SERVICE, OREGON STATE agp 1971. Ferns to Know in as di (Reprinted from the 1914 edition.) Corvallis, OR. 16

FRANKLIN, J. and C. T. DYRNESS. 1971. A Checklist of Vascular Pauts on the H. J. Andrews Eaten Forest, Western eae a S.D.A. For. Serv. Res. Note PNW- 138.) Forest Service, U.S.D.A., Portland, OR. 3

IRELAND, O. L. 1968. Plants of the Three Sisters nan Greis Cascade Range. (Mus. Nat. Hist. Bull. 12.) Univ. of Oregon, Eugene, OR. 130 p

PENNSYLVANIA

HENRY, L. K. 1971. An ee list of the vascular flora of Butler County, Pennsylvania. Ann. arnegie Mus. 43: 178. , and W. E. BU nt ion Check list of the vascular flora of Allegheny County, Pennsyl- vania. Trillia 11: 3-128. LEWIS, J. F. 1968. Guide to Plants—Bear Run Nature Reserve, the Kaufman Conservation on Bear Run: Fayette County. Western Pennsylvania Nature Conservancy, Pittsburgh, PA. 55 pp. LOVELL, J. F. 1965. Guide to Plants—Jennings Blazing Star Nature Reserve, Butler County. Western Pennsylvania Nature Conservancy, Pittsburgh, PA. 47 pp. REED, C. F. 1951. (see MARYLAND)

RHODE ISLAND

CRANDALL, DOROTHY L. 1965. County distribution of ferns and fern allies in Rhode Island. AFJ 55: 97-112

SOUTH CAROLINA

FITZPATRICK, J. W., J. R. CLOUTS, and W. T. BATSON. 1977. The vascular flora of two north-facing bluffs in Calhoun County, South Carolina. Castanea 42: 50-5

SOUTH DAKOTA

BROOKS, R. 1969. The ferns of the Black Hills. Trans. Kansas Acad. Sci. 72: 109-136.

DORN, R. D. and JANE L. DORN. 1972. (see MONTANA)

PETRIK- OTT, ALETA JO. 1975. (see KANSAS)

THILENIUS, J. F. 1971. Vascular Plants of the Black Hills of South Dakota and Adjacent Wyoming. (U.S.D.A. For. Serv. Res. Pap. RM-71.) Rocky Mt. Forest and Range Experiment Station, Ft. Collins, CO. 43 pp.

VAN BRUGGEN, T. 1967. The pteridophytes of South Dakota. Proc. South Dakota Acad. Sci. 46

44.

116 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

WINTER, J. M., C. WINTER, and T. VAN BRUGGEN. 1959. A Checklist of the Vascular Plants of South Dakota. Dept. of Botany, State University of South Dakota, Vermillion, SD. 176 pp-

TENNESSEE

BAILEY, J. K. 1969. A seared study of the ferns on the limestone bluffs on Norris Lake. J. Tennessee Acad. Sci. 44: 92-95.

prices te D. 1955. An annotated list of pu vascular plants of the gorges of the Fall Creek Falls State Park. J. Tennessee Acad. Sci. 108.

SHAVER, Fr M. 1954. Ferns of Tennessee, wn Fern Allies Excluded. Bureau of Publications, George Peabody College for Teachers, Nashville, TN

SMITH, C. R. and R. W. PEARMAN. 1971. A survey of the mioridaphytes of northeastern Tennes- see. Castanea 36: 181-191.

TEXAS ee D. S. 1956. Ferns and Fern ee : Texas. (Contr. Texas Res. Found. 2.) Texas earch Foundation, Renner, TX. 188 "1955. Pteridophyta, pp: 3-121, t. 1-39. tn C. L. LUNDELL et al. Flora of Texas, vol. 1. Univ. of Texas Press, Dallas, TX. , and M. C. JOHNSTON. i ae of the Vascular Plants of Texas. Texas Research Penindation. Renner, TX. 1881 GOULD, F. W. 1969. Texas a A Checks and Ecological Summary. Texas A&M University, College erg EX. 121 JONES, F. B. et al. 1961. Poweniis Plants and Ferns of co oe Coastal Bend Counties. Rob and Bessie rhs Wildlife Foundation, Sinton, TX 6 pp. McDOUGALL, W. B. and O. E. SPERRY. 1951. a of yogh Bend Viggen: Park. (Reprinted 1957.) Government Printing Office, Washington, DC. x MAHLER, W. F. 1973. Botanical Survey of the Lake Monti. Area. PiSMU Contr. Anthro. 9.) Southern Methodist University, Dallas, TX. 25 ROACH, fs W. and B. B. HARRIS. 1952. Sand Hill Hise of Henderson County, Texas. AFJ 42: 5.

UTAH LINDSAY, D. W. 1959. Vascular Plants Collected in Glen Canyon, 1958, pp. 63-72. In A. M. WOODBURY, ed. Ecological Studies of Flora and Fauna in Glen Canyon, Appendix A.

(Univ. Utah Dept. Anthro. Anthro. Pap. 40.) University of Utah, Salt Lake City, UT. 226 pp.

VIRGINIA FREER, R. - 1950. A preliminary checklist of plants of the central Virginia Blue Ridge. Castanea *

MASSEY. ‘ B. , comp. 1961. Virginia Flora. (V.P.I. Techn. Bull, 155.) V.P.1. Agricultural Exten- sion Service, Blacksburg, VA. 258 p. = The Ferns and Fern Allies of Virginia, 4th ed. (V.P.I. Agr. Ext. Serv. Bull. — -I. Agricultural Extension Service, Blacksburg, VA. 63 pp. ica P. M. 1972. An Illustrated Guide to the Ferns and Fern Allies of Shenandoah National

ark, Virginia. (Shenandoah Nat, Hist. Assoc. Bull. 6.) Shenandoah Natural History AS sociation, Luray, VA. 5 52 p

REED, C. F. 1959. Ferns sad vhs sities - 1960. The ferns and ‘‘fern-allies”’

-36, of — and Northampton Counties, Virginia and

ue 1963. Pteridophytes a ta Mountain Lake area, Giles County, Virginia, including notes from Whitetop Mountain. Castanea 28: 113-150.

M. A. MIASEK: STATE AND LOCAL FERN FLORAS 117

WAGNER, W. H., JR. and FLORENCE S. WAGNER. 1966. Pteridophytes ern Mountain Lake area, Giles Co., sigelasne Biosystematic studies, 1964-65. Castanea 31: 121-140.

MUSSELMAN, L. J., D. L. NICKRENT, and G. F. LEVY. 1977. (see nore CAROLINA)

TRUESDELL, H. (ser Feris in Clark and Loudoun Counties. Castanea 15: 51-55

WASHINGTON

JONES, G. N. 1960. The Flowering Plants and Ferns of Mount Rainier. (Reprinted from the 1938 edition.) (Univ. Wash. Publ. Biol. 7.) University of Washington, Seattle, WA. 192 pp.

og i a tot H. 1963. Flora of Southeastern Washington and of Adjacent Idaho, 3rd ed. Outdoor

ures, Escondido, CA. 583 pp.

eee: 7. R. 1967. A key to the ferns and allies of the state of Washington. Occ. Pap. Univ.

Puget Sound Mus. Nat. Hist. 31/32: 287-292. 1967. More on fern distribution in amas State. Occ. Pap. Univ.Puget Sound Mus.

Nat. Hist. 31/32: 293-310.

———., H. M. JENSEN, and W. E. MOON. 1965. Distribution of ferns on vor islands of Washington State. Occ. Pap. Univ. Puget Sound Mus. Nat. Hist. 28: 258-275.

, and ae M. JENSEN. 1966. Distribution of ferns on enn more islands of Washington State.

Occ . Univ. Puget Sound Mus. Nat. Hist. 29/30: 27

pbemeereT. OF BOTANY, UNIVERSITY OF wAsHiNGTOr 1969. A Checklist of Vas- cular Plants of Pines Washington, Pacific Coast to Columbia River. Univ. of Wash- ington, Seattle, WA. 33

WEST VIRGINIA CLARKSON, R. B. 1966. The vascular flora of the Monongahela National Forest, West Virginia. Castanea 31: 1-120 CORE, E. L. 1960. Plant Life of West Virginia. Scholar’s Library, New York, NY. 224 pp. ———.. 1966. Vegetation of West Virginia.McClain Printing, Parsons, WV. 215 pp. STRASBAUGH, P. D. and E. L. CORE. 1952. Flora of West Virginia (Part 1). West Virginia Univ. Bull., Ser. 52(12-2): 2-43.

WISCONSIN HARTLEY, T. G. 1966. The flora of the ‘‘Driftless Area.” Univ. lowa Stud. Nat. Hist. 21: 21-27. SEYMOUR, F. C. 1960. Flora of Lincoln County, Wisconsin. The Author, Taunton, MA. 363 pp. TRYON, R. M. 1953. The Ferns and Fern Allies of Wisconsin, 2nd ed. Univ. of Wisconsin Press, Madison, WI. 158 pp.

WYOMING DORN, R. D. and JANE L. DORN. 1972. (see MONTANA) PORTER, C. L. 1962. A Flora of Wyoming, part 1. (Univ. Wyoming Agr. Exp. er Bull. 402.) Agricultural Experiment Station, University of Wyoming, Laramie, WY. THILENIUS, J. F. 1971. (see SOUTH DAKOTA)

REGIONAL WORKS BAILEY, VIRGINIA L. and H. EB. BAILEY. 1955. A guide to the flowering plants and ferns of the western national parks, part 1. Amer. Midl. Nat. 54: 1-32. COBB, B. 1956. A Field Guide to the Ferns...of Northeastern and Central North America. Houghton ifflin, Boston, MA. 281 pp. (Reprinted 1963.) 270 CRONQUIST. A. et al. 1972. Intermountain Flora, vol 1. Hafner, New York, NY. iii + pp. EIFERT, VIRGINIA L. 1963. Native Ferns of Eastern North America, 3rd ed. Canadian Audubon Society, Toronto. 64 pp. FERNALD, M. L. 1950. Gray’s Manual of Botany, 8th ed. American Book, New York, NY. 1632 Pp.

118 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

FOSBERG, F. R. 1961. Southern distribution of Botrychium oneidense and B. multifidum, AFJ 51: 175-180.

Shee H. A. 1952. The New Britton and Brown Illustrated Flora of the Northeastern United States and Adjacent Canada, vol. 1. New York Botanical Garden, New York, NY. Ixxvi + 482 pp. (Reprinted 1963.)

,and A. CRONQUIST. 1963. Manual of ——, ne of Northeastern United States and

Adiacent Canada. Van Nostrand, Princeton, NJ.

HITCHCOCK, C. L. and A. CRONQUIST. hi Flora i i Pacific Northwest. Univ. of Wash- ington Press, Seattle, WA. xix + 230 p

HOLMGREN, A. H. and J. L. REVEAL. 1966. ‘Checklist of the Vascular Plants of the Intermoun- tain Region. (U.S. For. Serv. oy Pap. INT-32.) Intermountain Forest and Range Experi- ment Station, Ogden, UT. 160

MICKEL, J. T. 1974. Checklist of Ce cuits of North America north of Mexico. Fiddlehead Forum 1(3): 1-4.

RODIN, R. J. 1960. Ferns of the Sierra. Yosemite Nature Notes 394): 45-124

SHAVER, J. M. 1954. Ferns of the Eastern Central States, with Special Reference to Tennessee. Dover, New York, NY. 502 pp. (Reprinted 1970.)

SHREVE, F. and I. L. WIGGINS. 1964. Vegetation and Flora of the Sonoran Desert, vol. 1. Stanford Univ. Press, Stanford, CA

SMALL, J. K. 1964. Ferns of the Southeastern States. Hiatoer. New York, NY. 517 pp. (Reprint of the 1938 edition.)

TAYLOR, T. M. 1970. Pacific Northwest Ferns and Their Allies. Univ. of Toronto Press, Toronto.

pp. WHERRY, E. T. 1961. The Fern Guide: Northeastern and Midland United States Bi Adjacent Canada. (Doubleday Nature Guides 9.) Doubleday, Garden City, NY. 318 oe The Southern Fern Guide: Southeastern and South- a United seek (Double- y Nature Guides 10.) Doubleday, Garden City, NY. 349 p WILEY, pepe A. 1973. Ferns of Northeastern United States. eke New York. NY. 108 pp. (Reprint of the 1964 edition.)

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 4 (1978) 119

SHORTER NOTES

ATHYRIUM FILIX-FEMINA NEW TO SASKATCHEWAN.—In eastern North America, the Northern Lady Fern, Athyrium filix-femina var. michauxii (Spreng.) Farw., reportedly ranges from central Labrador, Newfoundland, Nova Scotia, and Maryland westward to central and southeastern Manitoba, South Dakota, and Missouri. This variety overlaps and apparently intergrades with the Southern Lady Fern, A. filix-femina var. asplenioides (Michx.) Farw., which extends from Florida and western Texas northward to New England, New York, Pennsyl- vania, West Virginia, Kentucky, Indiana, Missouri, and Oklahoma. Variety michauxii has been separated by a gap of 700 miles or more from the Western Lady Fern, A. filix-femina var. sitchense Rupr. (syn. subsp. cyclosorum (Rupr). C. Chr.), which ranges from southem Alaska to northern Mexico and eastward to western Alberta, Montana, the Dakotas, Nebraska, Colorado, and New Mexico. When considered as subsp. cyclosorum, a southern ecogeographical var. califor- nicum Butters has been distinguished by some authors. In Canada, the midconti- nental gap which has been apparent between the Northern and the Western Lady Fern has been thought to extend from central Manitoba across Saskatchewan to westernmost Alberta.

Recently we discovered var. michauxii at four separate localities in northeast- ern Saskatchewan. The collection data are as follows: 14 mi. W of Numabin Bay of Reindeer Lake, mile 1.25 of Highway 105 (Wollaston Lake Road), 56°16'N Lat., 103°36’W Long., lush white birch/river alder/willow woods along stream, Ternier & Jasieniuk 2113 (SASK); 3.5 mi. N of Courtenay Lake, mile 98 of Highway 105, 57°30’N Lat., 103°58’W Long., moist white birch woods at creek mouth, Ternier & Jasieniuk 1420 (SASK); 5 mi. S of Geikie River bridge, mile 110 of Highway 105, 57°37’N Lat., 103°54’W Long., moist white birch woods along small creek, Ternier & Jasieniuk 2352 (SASK); and Parks Lake inlet to Hidden Bay of Wollaston Lake, 58°08’N Lat., 103°41’W Long., lush, moist gallery white birch/black spruce/river alder mixed woods, Harms 21605, 21606 (SASK), Harms & Wright 23757 (SASK). The plants were common to moderately abundant at each of these sites.

These collections represent the first records of A. filix-femina in Saskatchewan and constitute a northwestward range extension of approximately 250 miles for var. michauxii from its known occurrence at Swampy Lake on the Hayes River in central Manitoba. Since the present range extension is northwestward, rather than due west, it is largely parallel to the eastern limit of var. sitchense in the Rocky Mountains, and so the apparent gap between the Northern and Westem Lady Ferns in Canada remains at over 600 miles. ree

The sterile fronds of the Saskatchewan specimens seem remarkably similar to those of the Spinulose Shield Fern, Dryopteris austriaca (Jacq.) Woynar var. spinulosa (Muell.) Fiori, one of our more common boreal ferns, and could easily be confused with it. In my opinion, the likeness is more apparent in the Sas-

120 AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 4 (1978)

katchewan specimens than it is in specimens from eastern Canada and the United States. I suspect that the Lady Fern may have been overlooked previously in Saskatchewan and western Manitoba by collectors who mistook it for D. au- striaca. However, an apparently consistent vegetative character to distinguish the two is the venation of the pinnules. In Athyrium, the veins reach the pinnule margin, whereas in Dryopteris they stop short and end in elongate hydathodes that are best seen on the adaxial surface of the frond.—Vernon L. Harms, Fraser Herbarium, Department of Plant Ecology, University of Saskatchewan, Saska- toon, Sask. S7N 0WO0, Canada.

NEW COMBINATIONS IN THE FERN FLORA OF VENEZUELA.—While we were revising a checklist of the ferns of the state of Mérida, Venezuela, we found some entities that needed a new combination or a new name. The nomenclatural changes for these are as follows:

Lycopodium nesselianum Duek & Lellinger, nom. nov.

Urostachys moritzii Nessel, Rev. Sudam. Bot. 6: 160, t. 9, f. 29. 1940, non Lycopodium moritzii

Keng Bot. Zeit. 19: 65. 1861. HOLOTYPE: Sierra Nevada, Venezuela, Greven 61 in 1898 (not

Trichomanes radicans var. kunzeanum (Hooker) Duek & Lellinger, comb. nov. Trichomanes kunzeanum Hooker, Sp. Fil. 1: 127, t. 39D. 1844. SYNTYPES: Pangoa, Depto. Junin, Peru, Mathews 1088 (K not seen Morton photo 19051); Pampayacu, Depto. Huanuco, Peru, Poeppig (K not seen); and Caracas, Distr. Fed., Venezuela, Linden 176 (K not seen; isosyntype FI not seen Morton photo 16586). The isosyntype is labelled ‘‘Hautes Andes de Truxillo et de Mérida.” Grammitis amphidasyon (Kunze ex Mett.) Duek & Lellinger, comb. nov. seeds amphidasyon Kunze ex Mett. Abhandl. Senckenb. Naturforsch. Gesell. 2: 49. 1856. SYNTY PES: Mérida, Edo. Mérida, Venezuela, Funck & Schlim 959 and 1100 (both B neither seen). — gameriana (Vareschi) Duek & Lellinger, comb. nov. : olypodium gamerianum Vareschi, Acta Bot. Venez. 1: 117, f. 16. 1966. Cerca de Laguna de los nteojos, Edo. Mérida, Venezuela, 4100 m elev., Vareschi & Maegdefrau 6839 (VEN). ee xanthotricha (Klotzsch) Duek & Lellinger, comb. nov. g plipedie xanthotrichum Klotzsch, Linnaea 20: 376. 1847. SYNTYPES: Mérida, Edo. Mérida, a a, Moritz 250? (B not seen); and Guyana, Rich. Schomburgk 1172 (B not seen). lie rams mortonianum Duek & Lellinger, nom. nov. ; ait ilo attenuatum Kunze, Linnaea 36: 56. 1869, non Fée, Icon. Esp. Nouv. [Mém. 6] 1,1. 1, peas SYNTYPES: Mérida, Edo. Mérida, Venezuela, Funck & Schlim 970 (LZ destroyed); se on sais Edo. Aragua, Venezuela, Moritz 125 (LZ destroyed), 441 (LZ destroyed; isosyntyPé er orton photo 7129), and Fendler 290 (LZ destroyed; isosyntypes US, Y not seen photo : a CIDIA T, Universidad de Los Andes, Apartado Postal 219, Mer ida, enezuela, and David B. Lellinger, U.S. National Herbarium, Smithsonian In- Stitution, Washington, DC 20560.

AMERICAN FERN JOURNAL: VOLUME 68 NUMBER 4 (1978) 121

TRISMERIA. . .TRIFOLIATA?—Since having been described in 1753 by Lin- naeus as Acrostichum trifoliatum, the Split-pinna Fern, as Wherry called it in his ‘*Southern Fern Guide,’’ was segregated into its own genus Trismeria by Fée in 1852, and more recently was transferred to Pityrogramma by Tryon (Contr. Gray Herb. 189: 68. 1962), who also indicated that the pinnae may be simple or 2- to 7-foliolate.

During a population sampling, mass field collections were made for flavonoid analysis and it was observed that the typical form, with 3-foliolate pinnae, was scarcely represented in the collections compared to other types of pinnae. A total of 4716 fronds from twelve populations made from sea level to 1,200 m altitude along the Caribbean-facing side of Costa Rica were examined. Percentages of the pinna types were as follows: 25.1696% simple; 55.1738% 2-foliolate; 18.7234% 3-foliolate; 0.9329% 4-foliolate; and none 5- to 7-foliolate. Trend analysis to estab- lish a possible correlation between frequency of x-foliolate pinnae along an altitud- inal gradient proved negative.

TABLE 1. DISTRIBUTION OF PINNA TYPES IN FIVE Trismeria trifoliata POPULATIONS.

I- 2- 3- 4- 5- 6- 7- Population foliolate foliolate _foliolate foliolate foliolate _foliolate _foliolate A 25.9825 : : 0. 0. 0. B 17.1521 35.2750 33.6569 13.5922 0.3236 0. 0. C 28.1833 50.0848 19.6943 1.5280 0.1697 0.1697 9.1697 D 17.0774 75.0000 6.8661 1.0563 0. 0. 0. E 20.0000 45.3333 29.8666 4.8000 0.2666 0. 0.

Another set of five populations growing at 800-1,200 m elevation in the Meseta Central of Costa Rica was sampled. The results from 2608 fronds are shown in Table 1.

Linnaeus apparently chose the epithet trifoliatum because of the generally trifoliolate condition of the frond illustrated in plate 45, figure 2 of Sloane's Ney age.’’ However, this generalized condition certainly is not the case in Costa Rican material.

The composition of the farina deposited on the undersurface of fertile T. trifoliata pinnae is very similar to that of Pityrogramma calomelanos and some other species of this genus (Wollenweber, E. 1976. Zeitschr. Pflanzenphysiol. 78: 344-349). The main component is a dihydrochalcone, which is accompanied by varying amounts of minor flavonoids, some of which are new products (V. Dietz, unpubl.). A detailed analysis of T. trifoliata population samples has not yet been accomplished, but our impression is that only rather small variations of a mostly quantitative nature occur. Such variations can be observed in P. calomelanos even on different fronds of one plant Hence, it obviously would be idle to search for correlations between farina composition and pinnule number in T. trifoliata.—Luis Diego Gomez P., Herbario Nacional, Museo Nacional de Costa Rice, Apartado 749, San José, Costa Rica and Eckhard Wollenweber, Technische Hochschule Darmstadt, D-6100 Darmstadt, Federal Republic of Germany.

122 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

/ REVIEW

‘‘FERNS OF HONG KONG,” by Harry H. Edie. xvii + 285 pp. Hong Kong University Press, 1978. ISBN 962-209-002-8. HK$40.00 (Ca. US$8.00).—This paperback book is a comprehensive introduction to the ferns (but not fern allies) of Hong Kong, Kowloon, and the New Territories. It is the only account of the ferns published for the region in over 50 years, and so is indispensible, especially because it incorporates the most recent generic concepts and alignments. The introduction is useful for beginning students of ferns; the part on fern ecology is especially interesting. The taxonomic part of the book includes keys, descrip- tions, distributions, and notes on the species. Synonyms are listed in a separate checklist that provides a useful overview of the colony’s ferns. Many of the species are illustrated with the author’s own line drawings. A glossary, a list of Chinese names for the ferns, and an index conclude the volume. This work is highly recommended for amateur and professional students of Asiatic ferns.—

AMERICAN FERN JOURNAL

Manuscripts submitted to the JOURNAL are reviewed for scientific content by one or more of the editors and, often, by one or more outside reviewers as well. During the past year we have received the kind assistance of J. G. Bruce, C. Haufler, R. L. Hauke, R. M. Lloyd, J. Montgomery, R. Oliver, G. R. Proctor, J. E. Skog, A. R. Smith, R. G. Stolze, W. C. Taylor, J. Utley, and R. A. White, to whom we are deeply indebted. We welcome suggestions of other reviewers and offers of assistance. —D. B. L.

INDEX TO VOLUME 68

— attenuatum, 120; danaeifolium, 62; graminoides, 77, _ Baskin, Carol C. (see J. M. Baskin) 9; serrulatum 77, 80; trifoliatum, 121 Baskin, J Carol C. Baskin. Geographical Distribution of = var. sulphureum, 13, 19; sul- Isoétes butleri in the Southeastern United States, 7

5 ecu, Bir, S. S. The Anatomy of Equisetum diffusum tubers, 55 Additions and Corrections to the Pteridophyte Flora of Chihuahua, Bishop, L. E. Revision of the Genus Cochlidium (Grammitidaceae)

6 Blechnum seminudum, 84, 85 : ewe Bolbitis, 31; bernoullii, 31; ser. Egenolfianae, 31; lancea, oe? ' 32; x prolifera, 32; quoyana, ; The Anatomy of Equisetum diffusum Tubers, 55 i ‘ . pees, 11; phyllitidis, 69; tomentosa var. mexicana, Bommeria knoblochii, 11; subpaleacea, 11 ne Botrychium, 71, 73; dissectum, 71, 73; japonicum, 71; seep 11:

Anogram ee

hieualine a SR RE menor; 3,5 multifidum, 71-75, subsp. californicum, 71; obliquum hniodes asia le simplex, 71; termatum, 71; virginianum,

Aspidotis meifolia, 12 Brandon, Dorothy D. (see E. M. Giff

ord, Jr.) Asplenium, 57; aucitam, 5: cusridatam Brooks, R.E. A Deletion from the Pteridophyte Flora of Nebrask

59; graminoides, 79; harpeodes, 59;

57: dentatum. 6?: erectum

miradorense, 59; pteropus, | Brownlie, G. The Pteridophyte Flora of Fiji (rev-), 64

59; radican : : . Athyrium me ek ae ppraee um, 11; williamsii, 57-59 Buck, W. R. The Taxonomic Status of Selaginella eatoni, 33 asplenioides, 119 ce Bry 2, 4%, 51; filix-femina, 119, var. Buck, W. R. (see J. H. Peck) wick iki ect ars . fornicum, 119, subsp. cyclosorum, 119, | Campyloneurum phyllitidis, 62 3 oe ——— 119; thelypterioides, 2 Cheilanthes, 13, 15, 18, 19, 21-24; albomarginata, 18, 215 6 Austin, D. F. are. E ica atchewen, 119 Aleuritopteris, 23; aurea, 23; aurantiaca, 23; argentea, 20; bul is. Azolla, @; mexicana, : 18; chrysophylla, 23; grisea, 18, 21; kuhnii var. pret . i issi ifolia, 12; micro) ‘ alamente (see N. P. Marengo) farinosa, 15, 18, 20, 21; longissima, 18; me a

63; mossambicensis, 21, 23; parryi, 61; rufa, 1 23; viscida, 18

nente, Marie A. Baroutsis, Judith G. & G. J. Gastony. Chromosome Numbers in ; Cheilanthes microphylla, a Genus and Species

the Fem Genus Anogramma, Il, 3 New to the Bahama

ANNUAL INDEX

rchipelago, 63 chon and Lipid Changes on Germination in the Non-green of Th — wey 67 some of Notholaena cochisensis, 63

— By

um, 77-79; 8 .

7 Poste ee Euge ; Daichsians (1925-1977), 6 Cystopteris, 95; bulbifera, 95; — 30, 61, var. mackayi, 95; protrusa, 95; tennesseensis

Cystopiecis tennesseens agnicaey 95 No

Death Notice: amps, Delchamps, C. E oo. notice), 6 A Deletion from the Pteridophyte Flora of Nebraska, 94 Dennstaedtia intermedi

The Distributi parent Constituents of the Farinose Exu- dates in G aad Ferns, The Distribution and Ecology of Thigh as in Southeastern Vir- and North Carolina, 4 , R. D. Polystichum auegie Found in don “— Hills, 29 aC 45, 46, 49, 50, 120; 46, 49; austriaca, 120, var. spinulosa, 119; x ce “4 ces pepe 9-51; celsa

cristata, ‘6. 49, 50; celsa x einen, 46; Siege: x tudovican, 49 , 50;

ristat) sa, 50; poaeeacs 46, 49, 50; intermedia x apa 46; ludoviciana, 49; maxima 64; x separabilis, 46, 47, ulosa, 46, 47, = 50; pc 64; x triploidea, 46, 49; x ia 50 Duek, J.J. & D. B. Lellinger. New Combinations in the Fern Flora of Venezuela, 120 Edie, H. H. Ferns of Hong Kong (rev.), 122 31

Egenolfia,

Elaphoglossum, mort :

Equisetum, 37, 96; arv 30, 5, 96; diffusum, 55, 56; subg. Equisetum, 37, 55; x ferrisii, 12; fluviatile, 96; giganteum, 37; subg. Hippochaete, 37; | ae rale, 96; myriochaetum, 37; “aye 38, 39, 55; pratense, 38, 39; ramosis- simum subsp. debile, 37; pre it oh sylvaticum, 55; tel-

mateia, 5. Equisetum litorale Recorded for Minnesota, 96 Rerahiich rp 1 Call . 2g Ri in T I Habitats, » D. R. (see B. McAl in) ca of Hong Kong (rev.), 122

The Ferns of San Salvador Island, II, 62

= Structure of the Newly Formed Spore of Onoclea sen- sibilis

Freeman com 2, D. (See J. W. Shor)

Game

Grown in Axenic Cul- ture, 71

Gastony, G. J. (see J. G. Baro Geographi

Sis) Phical Distribution of ele butleri in the Southeastern United angel

Gifford, E. M., Jr. & Dorothy D. Brand hytes of B Sa multifidum as Grown in Axenic ‘ete Iture, nm uses ssman, S. R. The Establishment of Bracken Following Fire in

Topical Habitats, 4]

123

Gomez ro we D. Some —e avernctions even psa oa 60; T

Gomez P., ‘: D. & E. Woltearetea: Trismeria.. Arifoliata?, si Grammitidaceae, 76 Grammitis, 76; amphidasyon, 120; attenuata, 91; pre anes furcata, 90; gameriana, 120; inoides rtii linearifolia, 87; paucinervata, spate minuda, 84; serrulata, 80; stipitata, 64; tate, 91: tepulensis, 91: hotricha, 120

ymnogramma, 13, 16, 22; sect. Ceropteris, 23; sulphurea, 14 Gymnopteris sierdlate Harms, V. L. Athyrium filix-femina New to Saskatchewan, 119 Hauke, R. L. Microreplicas as a Technique Cie sa Surface Silica Micon ology in Equise' Hennipman, E. grains re of the (Lomariopsidaceae) (rev.), 3 &A

acs rn Genus Bolbitis

Henry, R. D. . Scott. “ the Distribution of Lycopodium flabelliforme in Illinois, 30

Holmes, W. C. (s D. Thomas)

meta 100, 101

Isoat

s, 99 ey "pl, 103, 106, 107; butleri, 7, 8; echinospora, 99; lacus tris, 99; melanospora, 103, 105, 106; piedmontana, 103, 105, 106; egetiformans 100-103, 105, 106; triquetra, 100, 101, 103; sect. eauaee , 103 Knobloch, I. W. & D. S. C ci idaringe ae ve of paae ain Mexico. Knobloch, I W. Tai. The Fonte oe of Notho-

papenipits to the

as appressum, 96; carolinianum, itzii, 120; nesselianum, 120

Lycopodium cernuum in Lou —_ ida, 99 — 65; japonicum, 65, 66; microphyllum, 65, 66; palmatum,

areng . P. & Marie A. Badalame: he Fine Structure of the siti ‘toc Spore of Onoclea aro 52

Marsilea, 64 Matteuccia struthiopteris, 52 (

Mauk, Joyce E. (see R. R. Smith cAlpin, B. & D. R. Farrar. Trichomanes Gametophytes in Mas- sachusetts, 97

Mecosorus nudus, 90 ee ie ia , Me yl A.S dl F FA th it tates Supplement II, 109

Mic , 94

accion as a Technique for Rapid Evaluation of Surface Silica

Micromorphology in Scan ae

Micropteris, 77; blechnoides, 85; cttiintele. 80; paar TI, 3

Monogramma. , 76; furcata, 79; graminifolia, ides, 79;

gyroflexa, 88; immersa, 87; linearifolia, 87; minor, 83; aby lifolia, 94; rostrata, 88; rudolfii, 86, 87

A Monograph of the Fem Genus Bolbitis (Lomariopsidaceae)

(rev.), 31 —— i. Me gone L. Nickrent) “sah & D. F. Austin. Spread of the Exotic Fern Beetle in Florida, 65 ae scioana, 19 is exaltata, 62 and Unique, Mat-forming Merlin’s-grass (Isoétes) from

124

New Combinations in the Fern Flora of Venezuela, 120

A New tion for Pellaea glabella, A New Species of Asplenium from Guatemala, 57 Ni L., L. J. Musselman, Laura A. Pitchford n. The Distribution and Ecology of Dryopteris in se tern Virginia and Adjacen' ak Carl , 45 Notes on North American Lower cul s, 61 Notholaena, 13, 15, 19, 21-24, 63; aliena, 14; sltiis, 24; Poa iana, 15; aurantiaca, ryopoda, 24; candida var. us: i 21, 24, var. copelandii, 20, 21, 24; ee 24 - 25; sect Cincinalis, 23; cochisensis, 63; galeottii, 15; greyii, 14, 21, 24 greggii, 21, 24; incana, 24; integerrima, a: jacalensis, 61; leonina, 14; nivea var. flava, 23; pruinosa, 63; rosei, 14; schaffneri, 14, 25, var. neallii, 19, 25, var. schaffneri, 25; as 63; standleyi, 19, 21,2 2; tichomano ides,

4; sulphure Onoclea sensibilis, s wee On the War ibcalen of hs soe peat in linois, 30 Onychi eb auratum, 19; siliculosum, 15, 19-21 Osmunda cinnamomea, 45; re Peck, J. H. 4W. R. Buck. the Silngiosiin apoda Complex in Iowa,

Peck, J. H. & S. D. Swanson. Equisetum litorale Recorded for Minnesota, 96 Pellaea, 62; breweri, 61; glabella, 60 i re

ium au 2 Pichi Sermolli, R. E. G. Tentamen eid ies Genera in onomicum Ordinem Redigendi (rev.), 108

Pitchford, ae D. L. Nickrent)

Pityro; , 13, 16, 21-24, 121; calomelanos, 121; chrysophylla, 13; cer 20, 22, 23, 25, var. heyderi, 17, var. marginata, 18; austroame' tae 18, 20; thesia: jo a Ag calomelanos, 15, 17, 18, bts ar. ava, 25 v. niger vi ea, 25; dealbata, 24; lehmannii, 18, 20; s 13, 17; t Remmi 13, 16, 18, 22, 23, var. tria’

sulphure neti slits 15; tartarea, 14, 18, 25, var. aurata, 25, var.

Megat — 11; semicordata, 11

Pleur: Th 8S: en 79; graminifolia, 85; parse les, '- eaiens: 88, 89; i rsa, 87; linearifolia, 87; linearis, 77, 85; ria 94; ge 85; pumila, 85; seminuda,

Polybotrya, 31

“encoehigdels ie subfam. Cryptogrammeae, 22, sect. Adianteae,

AMERICAN FERN JOURNAL: VOLUME 68 (1978)

togramma pilosa in Alabam Reeves, T. Notes on North red erican Lower Vascular Plants, 61 Reviews: Ferns be Hong ern 122; A M raph of the Fem

Genus Bolbit eae e Pteridophyte Flora of oh 64; Tentamen a rake ‘Genet in Taxonomicum Ordin bydooneemnet

pera of the Genus Cecio vesiomiennss Rury, P. M. A New and Unique, Mat-forming ane S-grass fisciees) from soma D. W. (see D. L. Nickrent)

z

in the Ne -green Spores of Thelypteris dentata, 67 Selaginella, 33, 34, 36, 62, 64; armata 2. 2s 33, 34; eatonii, 33-36; eclipes leucobryoides, 61; ludoviciana, 33, 36; macrathera, 61; mutica var. mutica, 11; pallescens, 11; pincola, 11; subg. Stachygynand m, 3 The Selaginell, a Complex in si Short, J. W. & J. D. Freeman. Cystopteris tennesseensis Alabama

topter: in 95; Rediscov nega Distribution and Phytogeographic Af- finities of Lept ma pilosa in Alabama, 1

inopteris, ped eateasa, 19

Smith, R. R. & Joyce E. Mauk. The Ferns of San Salvador Island,

II, 62 Some Insect Interactions sh Azolla mexicana, 60 Spend of the Exotic Fern Lygodium microphyllum in Flo rida, 65 Ss and Local Fem cic i the United States, conlaile II,

9 Stolze, R. G. A New Species of Asplenium from Guatemala, 57 ses, 99-101, 10 Sw n, S. D. (see J. H. Peck) Teenitis| graminifolia, 84, 85; graminoides, 79; linearis, pumila, 85; seminuda, 85

TI, 85;

Tai, W. (see I. W. Knobloch)

The Taxonomic Status of Selaginella eatonii, 33

Tectaria lobata, 62 ; Tentamen Pteridophytorum Genera in Taxonomicum Ordinem Re-

mauropelta, 9; a se 70; subg. Gonio- pteris, 9; uri 62 peepee 9: oroniensis, 9, 10; ovata var. lindheimeri 11; pilosa var. alabamensis, 1; veer’ var. sonoren-

Be th resniera age S sect. Chei — 22, subfam. Gymnogrammoideae, 22, New sine from rs Rica, 9 eS nog Thomas, R. D., W. C. Holmes, C. M. Allen & G. Landry Pavan ‘anphidasyon, 120; connellii, 92; dicranophyllum, 90; Lycopodium cemuum in siiaalnal ,% ‘ e, 80; furcal W; ni ; myosuroides, 80; Trichomanes, 97, 98; boschianum, 2, 97; kunzeanum, 120; petersil Paucinervatum, 86; persicariifolium, 90; polypodioides, 62; 2, 97; radicans var. kunzeanum thatch 86; fn , 80; tepuiense, 91; vulgare, 67, 69; Trichomanes Gametophytes in Massachusetts, 97 eeseap Trismeria, 121; trifoliata, 121 stichoides, 45; lonchitis, 29 Urostachys moritzii, 1 stemerah — Arad in the Black Hills, hones ia, 97; li i ie E, ‘ tee t jaea glabella in Minnesota, Puritan 7; mad i Wallner, E. The Distribution a ‘chee cal Constituents © ch aah aquilinum, 41; mrinose Exudates in nogrammoid Ferns, 13 pide “stg Flora of Fiji Ga. ‘3 eae E. (see L. D. Gomez =ioshonasn Dy, 22; vdistone 19 Xipho; ak 6, om a om yantepuiensis, 80; extensa, 80; scovery, Distributio ytogeographic Affinities of Lep- myosuroides, 77, 81; aicuaate: 80; serrulata, 77, 80, 81 ERRATA P. 23, fio aoe e 12, For ' “chryosophylla”’ read ‘‘chrysophylla.”’ 4 - title. For ‘‘Selanginella’’ read ‘*Selaginella.”’ i ‘6 ”” <é P. 79, - haf furcata’”’ read ‘‘furcatum.”’ « a” e , ‘ or ‘tepuiensis read “‘tepuiense.’

. 79, line 5. For ‘‘acuminata’’

read ‘‘acuminatum.’’

TRIARCH Over 5© Wears

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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:

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AMERICAN FERN JOURNAL

Volume 69

1979

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 69, Number 1, Pages 1-32, Issued March 29, 1979

Equisetum ramosissimum in North America RICHARD L. HAUKE

Notes on the Dispersion of Dryopteris Spores in the Great Dismal Swamp THOMAS W. MAGRAW and LYTTON J. MUSSELMAN

An Artificial Crossing Technique for Selaginella TERRY R. WEBSTER

A Tropical Fern Grotto in Broward County, F DANIEL F. AUSTIN, GRACE SE CARD IVERSON, and CLIFTON E. NAUMAN

Studies of the Azolla-Anabeana Symbiosis Using Azolla mexicana, I. Growth in Nature and Laboratory ROBERT W. HOLST and JOHN H. YOPP

Thelypteris in Arkansas W. CARL TAYLOR and DAVID M. JOHNSON Robert J. Rodin (1922-1978)

Shorter Notes: Vittaria lineata Rediscovered in Georgia; The Distribution of Dryopteris goldiana and D. — in Missouri; Gymnogramma ys. Gymnogramm

Reviews 5, 16,

Volume 69, Number 2, Pages 33-64, Issued June 26, 1979

A Community of Lycopodium Gametophytes in Michigan JAMES G. BRUCE and JOSEPH M. BEITEL Acrostichum in Florida DAVID C. ADAMS and P. B. TOMLINSON

Phyllitis scolopendrium Newly Discovered in Alabama JOHN W. SHORT

Studies in Lycopodiaceae, II. The Branching simian and Infrageneric Groups of Lycopodium B. @LLGAARD

Shorter Notes: Isoétes butleri in Georgia Juvenile Leaves of the A pogamous mous Fern Notholaena cochisensis; A Thelypteris New to Florida

Review

1

6

9

14

17

26

31

33

42

47

49

Volume 69, Number 3, Pages 65-96, Issued October 5, 1979

A New Nephrolepis Hybrid From Florida CLIFTON E. NAUMAN 65 Spore Morphology of Anemia subgenus Anemia STEVEN R. HILL 71

The Development of Plantlets from Strobilus Branches in Lycopodium phlegmaria ¥-C WEE &

Incidence of Epiphytism in the Lycopsids JOSEPH M. BEITEL 83

Cyrtomium fortunei in Lousiana and Mississippi GARRIE P. LANDRY, MICHAEL ISRAEL, ROBERT SCHWARZWALDER, JR, and R. DALE THOMAS 85

The Fine Structure of the Pre-meiotic Stages of Sporogenesis in Onoclea sensibilis NORMAN P. MARENGO 87

Apical Dominance in Anemia phyllitidis Gametophytes THOMAS L. REYNOLDS 92

Shorter Notes: A New Combination in eee Cheilanthes alabamensis New to Sexuality in Asplenium resliens 94 Reviews 70, 84, 91 Volume 69, Number 4, Pages 97-124, Issued December 31, 1979

Disjunct Populations of Isoétes macrospora in Southern Tennessee W. MICHAEL DENNIS, A. MURRAY EVANS, and B. EUGENE WOFFORD 97

Three New Elaphoglossums from Guatemala JOHN T. MICKEL 100

The Role of Temperature in the Vegetative Life Cycle of Isoétes butleri JERRY M. BASKIN and CAROL C. BASKIN 103

Gametophyte Morphology of the Fern Genus Drynariopsis (Polypodiaceae) SUBHASH CHANDRA 111

A New Speci , : C. B. GENA, T..N. BHARDWAJA, oe and A. K. YADAV 119

Shorter Note: A Simplified Nutrient Medium

for Growing Fern Prothallia 122

Review 102 American Fern Journal

Index to Volume 68

AMERICAN Ea F BF R N ee JOURNAL =

QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY

Equisetum ramosissimum in North America RICHARD L. HAUKE

Notes on the Dispersion of Dryopteris Spores in the Great Dismal Swamp THOMAS W.MAGRAW and LYTTON J. MUSSELMAN

An Artificial Crossing Technique for Selaginella TERRY R. WEBSTER

A Tropical Fern Grotto in Broward County, Florida DANIEL F. AUSTIN, GRACE BLANCHARD IVERSON, and CLIFTON E. NAUMAN

Studies of the Azolla-~Anabaena Symbiosis Using Azolla mexicana, I. Growth in Nature and Laborato ROBERT W. HOLST and JOHN H. YOPP Thelypteris in Arkansas W. CARL TAYLOR and DAVID M. JOHNSON Robert J. Rodin (1922-1978) Shorter Notes: Vittaria lineata Rediscovered in Geo

Distribution of Dryopteris goldiana and D. ahaa n Missouri; Gymnogramma vs. Gymnogramme

Reviews 5, 16,

WISSOURT BOTANICAT

APR : 23 1979

GARDEN LIBRARY

=]

te

31

The American Fern Society Council for 1979

RICHARD L. HAUKE, Dept. of Botany, University of Rhode Island, Kingston, R.I. 02881.

President ROBERT M. LLOYD, Dept. of Botany, Ohio University, Athens, Ohio 45701.

Vice-President

LESLIE G. HICKOK, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916

Secretary JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916.

Treasurer JUDITH E. SKOG, Dept. of Biology, George Mason University, Fairfax, Va. 22030.

Records Treasurer

DAVID B. LELLINGER, Smithsonian Institution, Washington, D.C. 20560. Editor-in-Chief

JOHN T. MICKEL, New York Botanical Garden, Bronx, N.Y. 10458. Newsletter Editor American Fern Journal ED

OR DAVID B. LELLINGER Smithsonian Institution, Washington, D. C. 20560

ASSOCIATE EDITORS DAVID W. BIERHORST ..Dept. of Botany, University of Massachusetts, Amherst, Mass. 01002 GERALD J. GASTONY ............. Dept. of Biology, Indiana University, Bloomington, Ind. 47401 JOHN T. MICKEL New York Botanical Garden, Bronx, New York 10458

The ‘‘American Fern Journal’ (ISSN 0002-8444) is an illustrated quarterly devoted to the general study of fems. It is owned by the American Fern Society, and published at the Smithsonian Institu- tion, si niphinaens DC 20560. Second-class postage paid at Washington.

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General inquiries concerning ferns should be addressed to the Secreta

Subscriptions $9.00 gross, $8.50 net if paid through an agency jeceney fee $0.50); sent free to members of the American Fem Society (annual dues, $8.00: life membership, $160.00). Extracted offprints, if ordered in advance, will be furnished to authors at cost, plus postage.

Back volumes $5.00 to $6.25 each; single back numbers of 64 pages or ae. $1.25; 65-80 pages, $2.00 sae hae 80 aot $2.50 each, plus shipping. Ten percent discount on orders of six volumes or more;

tiona

Library Dr. — Mickel, New York Botanical Garden, Bronx, New York 10458, is Librarian. Members may borrow books at any time, the borrower paying all shipping costs. Newsletter : Dr — met Mickel, New York Botanical Garden, Bronx, New York 10458, is editor of the sie ter a Forum.”’ The editor welcomes contributions from members and non- ncluding miscellaneous notes, offers to exchange or purchase materials, personalia, hor- ear notes, and reviews of non-technical books on fems Spore Exchange

Mr. Neill D. Hall, 1230 Northeast 88 ~ ae Seattle, tor. Spores exchanged and collection lists sent on re ee :

Cite Py ot arena Git and Bequests to members and to others interested

in ferns. Botanical books, back issues of the Joumal, and cash - = gifts are always we Icomed, and are tax-deductible. Inquiries should be addressed to the Secre

AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 1 (1979) l

Equisetum ramosissimum in North America RICHARD L. HAUKE*

Equisetum ramosissimum Desf. is widely distributed in the Eastern Hemi- sphere, ranging through southern, eastern, and mediterranean Africa, the Canary and Azores islands, Europe north to southern Germany, and Lithuania and east through the Balkans and Asia Minor to Mongolia, Korea, and Japan. A sub- species, E. ramosissimum subsp. debile (Roxb.) Hauke, is found in India, southeastern Asia, and the south Pacific Islands to Fiji.

This species is a member of subg. Hippochaete and, as the specific epithet indicates, is much branched, unlike most of the other members of the subgenus. It is also unlike them in often having double or triple lines of stomata between the ridges along the stems. In these characters it can readily be mistaken for EF. giganteum, E. x schaffneri, or, when with only single lines of stomata, E. myriochaetum

As with other species of Equisetum, E. ramosissimum subsp. ramosissimum is variable (Milde recognized 34 varieties). In addition to the species mentioned above, it also can be mistaken for E. hyemale, E. laevigatum, E. x ferrissii, or E. x moorei (in Europe). But the combination of regular branching, long internodes, long sheaths, brownish sheath coloration with a black girdle, flattened sheath segments, persistent but shriveling sheath teeth, often multiple lines of stomata, prominent vallecular and carinal collenchyma, silica surface with numerous pilules, and green spores tends to distinguish this species from all the others, which may show one or more of the above characters, but not most or all of them.

Under the distribution of E. ramosissimum, Milde (1867, p. 461) listed British Columbia 49° n Br. (Dr. Lyall).’’ Since there has been no further record of this species in North America, it has not been included in the flora of North America. In the last few years, however, a few collections from North America have come to my attention:

NORTH CAROLINA: New Hanover Co.: Dense population of branching plants on sandy ditchbank through ballast dump on east side of Northeast River, south of the old US-17 bridge, Wilmington, 22 Aug 1970, S. W. Leonard 3557 (KIRI). Ditch and sand flats on ballast area, east side of Northeast River, Wilmington, 3 Sep 1970, 8. W. Leonard & A. E. Radford 3716 (TENN, KANU).

FLORIDA: Escambia Co.: Plants to 2.5 m tall, supported by thicket of wax myrtle and willow (much lower and unbranched in adjacent mowed field), continuous over broad field, moist to wet clayey soil, NW corner of Main and Donelson, S side of Pensacola, T2S, R30W, 24 Jun 1975, D. B. Ward 8968 (FLAS, KIRI). Vigorous colony, on rich, black soil, with Eupatorium, pose Fumaria, Plantago, Rumex, etc., at NW corner of Main St. and Donelson St., Pensacola, 2 May 1975, J. R. Burkhalter 2254 (FLAS). : :

When Leonard sent me a specimen in August, 1970, he described it as ever- green, mostly branched, and at its reproductive peak, and he noted that the area was rich in European records. I wrote to him that it looked to me like E. x ferrissii (known from North Carolina), but because of the time of coning and presence of green spores, it must be an odd form of E. hyemale var. affine.

* Department of Botany, University of Rhode Island, Kingston, RI 02881. Volume 68, number 4, of the JOURNAL was issued December 29, 1978.

AMERICAN FERN JOURNAL: VOLUME 69 (1979)

Q0Giewwm a

. Herbarium specimen of Equisetum ings Desf. from Florida, Ward 8968 (KIRID).

FIG.

PGs. 2-6. Nodal sheaths (mm scale included). 2 UE. ‘sae atic peng Ward 8968. FIG.

ramosissimum, Jordan, Hauke s.n. (KIRI). FIG. 4 4. E. x ferri . Michigan, Hauke 56 RI). FIG, § ramosissimum, North Carolina. Le eae & Radiwe oe (TENN).

ramosissimum, Jorden: Hauke

R. L. HAUKE: EQUISETUM IN NORTH AMERICA 3

In June, 1975, Daniel Ward sent me some specimens of Equisetum collected in Florida. Most were obviously E. hyemale var. affine, but Burkhalter 2554 ap- peared different. Ward wrote, ‘‘The area is fill, on the site of the ballast dumps of the early seaport, Pensacola having been during the heyday of the yellow pine days, the second most active port on the Gulf. The Equisetum, although seemingly reproducing only by vegetative means (no outlying stations are known), occurs on both sides of a 30 ft. paved road, suggesting a very early station since bisected.’’ I identified the odd specimen as E. x ferrissii and commented on the excessive length of the internodes and sheathes. A later collection by Ward at the same station is illustrated (Fig. /).

A. Murray Evans sent me a specimen in January, 1978 of what he thought was E. laevigatum collected at Wilmington, NC by Leonard and Radford. At first it appeared to me to be E. x ferrissii, but the stomatal lines were frequently dou- bled, and this together with the green spores and regularity of branching made me realize that it represented E. ramosissimum subsp. ramosissimum.

Since E. ramosissimum and E. x ferrissii (Fig. 4) have similar internal anatomy and sheath appearance, I obtained scanning electron micrographs of the surface silica micromorphology to confirm this identification. Equisetum ramosissimum (Fig. 9) is characterized by having numerous pilules of silica over the surface, including the silica overarching the stomates. Equisetum X ferrissii (Fig. 7) has the surface pitted between transverse plates of silica, and completely lacks the pilules so characteristic of E. ramosissimum. The Wilmington (Fig. 10) and Pen- sacola (Fig. 8) collections display numerous pilules, thus confirming their identity as E. ramosissimum.

How can we account for the sudden appearance of E. ramosissimum subsp. ramosissimum in southeastern North America? The North Carolina locality sug- gests that it might have been brought in with the ballast many years ago and Overlooked until recently. Vegetative transport of Equisetum with fill has been noted previously (Hauke, 1963, pp. 28-29). The presence of E. ramosissimum in Pensacola, also a seaport, could be similarly explained. However, for a number of years Equisetum was known from only one locality in Florida (E. hyemale, along the Appalachicola River), and it seemed unlikely that in a metropolitan area like Pensacola a population of Equisetum would have escaped detection for long. Ward and Burkhalter (1977) discussed another plant from the ballast area of Pen- sacola, Acacia smallii, a small tree collected in 1901 and 1903, then not again until they collected it in 1975. Of the area, they say, ‘“The quantities of ballast have been leveled and distributed to fill much of the old harbor area, and warehouses, parks, and roadways are now scattered where once sailing ships were at anchor. But waste areas have persisted, and with them many of the early introductions. If a tree could escape detection in Pensacola for nearly three quarters of a cen- tury, it is not surprising that Equisetum ramosissimum also escaped detection until recently ,

The possibility that E. ramosissimum is more widespread in North spagiaiel ‘ than the two known localities cannot be denied. The occurrence of E. ramosis-

AMERICAN FERN JOURNAL: VOLUME 69 (1979)

FIGS. 7-10. Scanning electron micrographs of Equisetum stem surfaces. FIG. 7. E. x ferrissil,

Kansas, Hartman 1090 (KIRI). F simum, Jordan

line on FIG. 10 is

FIG. 8. E. ramosissimum, Florida, Ward 8968. FIG. 9. E. ramosts- eed sn. FIG. 10. E. ramosissimum, North Carolina, Leonard 3557 (KIRI). Scale

R. L. HAUKE: EQUISETUM IN NORTH AMERICA 5

simum in the Pacific Northwest remains a distinct possibility. In an extensive survey of herbarium specimens from throughout North America, I never detected any E. ramosissimum. But since | did not at first recognize the two presently known populations as FE. ramosissimum, that is not proof of its absence. How- ever, | am confident that under conditions of systematic comparison of many specimens, I would have detected it if present, even though individual specimens sent at widely separate times were not immediately recognized. LITERATURE CITED HAUKE, R. L. 1963. A taxonomic monograph of the genus Equisetum subgenus Hippochaete. Nova Hedwigia Beih. 8:1-123, r. /-22. MILDE, J. 1867. Monographia Equisetorum. Nova Acta Acad. Caes. Leop. 32(2): 29-607, 1. 1-35. WARD, D. B. and J. R. BURKHALTER. 1977. Rediscovery of Small’s Acacia in Florida. Florida Sci. 40:267~-270.

REVIEW

*‘A TAXONOMIC REVISION OF THE NEW ZEALAND SPECIES OF ASPLENIUM”’ and ‘‘ASPLENIUM HYBRIDS IN THE NEW ZEALAND FLORA,” by P. J. Brownsey, New Zealand J. Bot. 15: 39-86, 601-637. 1977.— Because of its propensity to form interspecific hybrid complexes, Asplenium is a taxonomically troublesome genus everywhere it occurs. The species in New Zea- land are no exception. Brownsey’s revision is an excellent integration of classical taxonomy and cytotaxonomy. Besides a key to the species, synonymies, descrip- tions, statements of habitat and distribution, and comments, silhouettes and draw- ings of rhizome scales are given for each of the species, and all are mapped in detail. The photographs of spores taken through the light microscope are not so revealing as scanning electron microscope pictures would have been, but even so, obvious differences between some of the species can be seen. The revision treats 16 basic species and subspecies, and a crossing polygon shows hybridization between them. Especially A. bulbiferum, A. flaccidum, and A. obtusatum each form several different hybrids. Nineteen hybrids are described and illustrated. At. least as many more other combinations also occur, but their characterization and analysis awaits further cytotaxonomic study. The species complexes, so far as currently known, are analyzed with genome diagrams. Brownsey sensibly has chosen to refer to the hybrids with formula names, rather than to coin new epithets. Since the number of possible hybrids is nearly infinite, it is best to use formula names for all hybrids except those.that are unusually important because of their wide range, abundance, horticultural value, or role as parents of other hybrids. —D. B.L

5 AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 1 (1979)

Notes on the Dispersion of Dryopteris Spores in the Great Dismal Swamp THOMAS W. MAGRAW* and LYTTON J. MUSSELMAN**

During the past few years, considerable attention has been paid to Dryopteris in the Dismal Swamp. However, no studies on spore dispersion have been con- ducted in the swamp, although such studies are important in understanding the life histories of these ferns.

The Dismal Swamp contains a large number of plants of sexual Dryopteris species plus some hybrid plants (Nickrent et. al., 1978). The hybrids are of par- ticular interest because some of them, like D. celsa X cristata are able to repro- duce and so form large stands. In the hybrids, the production of giant spores, considered to be a form of apomixis, is well known (De Benedictis, 1969). In addition to giant spores, other abnormal spore types (diads, triads, and fused tetrads) are produced in a manner apparently similar to that described by Hickok and Klekowski (1973).

The objectives of this study were to examine the dispersion of spores within the fern’s immediate habitat, to assess spore drift above the canopy, and to determine the dispersability of hybrid spores.

MATERIALS AND METHODS

The spore traps used in this study consisted of a 4.0 x 1.3 cm strip of double- Stick transparent tape attached to one side of a microscope slide. The slides of ground-level traps were held with their long axis vertical just above the soil by the handle of bacteriological slide holders. Slides above ground level were positioned similarly by suspending the slide holders from a monofilament line placed 1.5 m above ground level so that the slides faced south. Pairs of traps were placed vertically and facing west on a fire tower at 5.33 m intervals starting just below the top of the tree canopy at 21.33 m above the ground. All of the spore traps actually may have under-collected fern spores, compared with the true number in the air. Other sampling machinery, e. g. “‘rotoslide samplers’’ (Raynor et al., 1976), may be more effective in trapping spores.

Spores were counted at two contrasting sites about 25 miles SSW of Norfolk, VA. A fire tower located in the Dismal Swamp National Wildlife Refuge was used to collect data from above and just below the tree canopy. No large Dryopteris populations occur within a mile of the tower. A ground-level station just south of U.S. highway 158 in Gates County, NC served to make ground-level observa- tions. Dryopteris is the most abundant herbaceous plant at this site. Both D. celsa and its hybrid with D. cristata are present at this site.

MAGRAW & MUSSELMAN: DISPERSION OF DRYOPTERIS SPORES 7

Dryopteris plants in the Dismal Swamp release their spores at slightly different times. Dryopteris intermedia is first, followed by D. spinulosa, D. cristata, and D. celsa. The peak of spore release by the latter species for the 1978 season was in the last week of June. Because of time limitations on our research, spore trapping began at that time and continued into August.

TABLE 1. AVERAGE DAILY SPORE COUNT PER CM? BETWEEN JUNE 27 AND JULY 1, 1978 FOR FIVE LOCATIONS 1.5 M DISTANT IN A STRAIGHT

E.

Number of fronds/m? Ground level 1.5 m above ground level Total 15 0.14 34.14

12 4 0.34 4.34

l ee 0.24 1.44

0 9 0.53 9.53

0 4 0.53 4.53

Total Dave 1.78

Athyrium asplenioides, Asplenium platyneuron, Osmunda cinnamomea, and Woodwardia areolata all are very common throughout the swamp. Voucher slides of spores of each were made in order to identify spores on the traps accurately. Dryopteris spores were distinguished easily from other fern spores by their reniform shape and distinctive perispore.

TABLE 2. AVERAGE DAILY SPORE COUNT PER TRAP AT 1.5 M ABOVE GROUND AND AT GROUND LEVEL. Spore type June 27-July 1 July 1-3 July 21-27

1.5 m above ground level 10 8

Normal 6 Hybrid 0 0 0 Total 10 6

0 m above ground level Normal 71 11.33 Hybrid 0 0 0.67 Total 271 60 12

Judging by the spore counts on traps set at the ground-level (Table 1), there Is little vertical or lateral movement of fern spores from their point of origin. Most Spores fall to the ground close to the fronds which produced them, presumably because of the relatively mild air currents within the dense forest. There Is a rough correlation between frond density and the number of spores trapped. Relatively few spores are wafted into the air and caught on traps 1.5 m above ground level. It is possible that the results in Table / are biased because the data were collected at Stations spaced 1.5 m distant along a straight line, five at ground level and five 1.5 m above the ground. A grid pattern containing more stations might have been more revealing.

8 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

Hybrid spore dispersion is of particular interest since it has been suggested that Dryopteris hybrids like D. x separabilis may have invaded at least one of the study sites in the form of an unreduced giant spore (Nickrent et al., 1978). The present study indicates that hybrid spores are only infrequently collected (Table 2). This may be an indication of the low number of spores dispersed from hybrid plants, or perhaps giant spores are even more prone to fall directly beneath the plants that produced them than are normal spores.

TABLE 3. TOTAL COUNTS OF SPORES TRAPPED BELOW AND ABOVE THE

TREE CANOPY IN THE DISMAL SWAMP.

Height above ground (m) July 7-14 July 14-21 July 21-27 July 29-Aug 5 Total 42.67 1

0 1 on 0 l 37.33 02 5 2 0 7 32.00 0 12 2 0 3 26.67 12 02 3 0 4 21.33 02 Sie 32 0 3 Total 1 7 10 0

Both traps lost. ?One trap lost.

A small number of spores do manage to drift on air currents above the tree canopy, clearing it by even as much as 18.29 m (Table 3). The top of the canopy is about 24.38 m above the ground. There appears to be no significant difference in the number of spores trapped at any level between 21.33 and 37.33 m. About 60% of the fern spores recovered from the fire tower traps were not Dryopteris, which ls reasonable because large populations of Athyrium asplenioides, Osmunda cin- namomea, O. regalis, Woodwardia areolata, and W. virginica are nearby.

Future studies should be conducted over the entire sporulating season and beyond to examine more carefully the presence of spores carried high in the air. Likewise, traps should be set at various distances from the main Dryopteris cen- ters within the swamp. Additional work also is needed on the number and types of viable spores produced by hybrids, as well as their dispersion in the air.

This work was supported by NSF grant SMI-76-0123. Preliminary collections were made by Ms. L. A. Pitchford and were supported by a grant from the National Geographic Society.

LITERATURE CITED

u ferns. Amer. J. Bot. 60:1010-1022. eg D. L., L. J. MUSSELMAN, LAURA A. PITCHFORD, and D. W. SAMPSON. - The distribution and ecology of Dryopteris in southeastern Virginia and adjacent North

Carolina. Amer. Fern. J. 68:45-51.

RAYNOR, G. S.,E.C. OGDEN, and JANET V. HAYES. 1976. Dispersion of fern spores into and within a forest. Rhodora 78:473-487,

AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 1 (1979) 9

An Artificial Crossing Technique for Selaginella TERRY R. WEBSTER*

Various methods for obtaining microgametophytes, megagametophytes, and sporelings of Selaginella have been described. Slagg (1932), in a study of micro- gametophyte development in S. kraussiana (Kunze) A. Br., germinated micro- spores on plaster of Paris blocks. A similar method for germinating both micro- spores and megaspores was described by Bold (1967). Bierhorst (1964) described methods for obtaining reproductive stages of Selaginella for classroom use. Web- ster (1967) described the induction of sporelings under greenhouse and field condi- tions. Wetmore and Morel (1951) germinated megaspores of S. pallescens (Presl) Spring in Mart. and S. flabellata (L.) Spring on nutrient agar under sterile condi- tions. Through the use of a medium supplemented with various growth factors, they were able to grow gametophytic tissue for several months.

Despite the above procedures for germinating spores and obtaining sexual ma- terial of Selaginella, there is but one report of artificial crosses in the genus. Burgeff and Filippi (1957) made crosses between S. martensii Spring var. marten- sui and S. martensii var. variegata Hort. They sowed surface-sterilized mega- Spores and microspores together on nutrient agar in culture tubes and, after gametophytes had formed, flooded the cultures with water to achieve fertilization. After 30-40 days, sporelings appeared. Using this method, they studied the inheri- tance of variegation in S. martensii var. variegata.

In this paper, a crossing technique differing in several respects from the one used by Burgeff and Filippi (1957) is described. Although it has been used suc- cessfully to repeat the crosses by Burgeff and Filippi, the technique was de- veloped for making crosses with S. kraussiana and its varieties, and the following description is based on work with these taxa. Application of this technique toa study of inheritance of pigmentation in S. kraussiana var. aurea W. Bull will be the subject of a later report.

DESCRIPTION OF THE TECHNIQUE

Plants were grown in flats or pots in the greenhouse. Under natural daylengths, abundant spores were produced from spring through fall, but spore production was much diminished during the winter. Supplemental light from 6PM to 6AM was provided by three 110 w, incandescent bulbs suspended 50 cm above the plants; this was found to increase the period of spore production.

Strobili of S$. kraussiana and its varieties typically bear a single basal mega- Sporangium; the remainder of the strobilus is microsporangiate. The ripe mega- sporangia protrude noticeably from the strobilus, and can be removed with a pair of fine forceps. The excised megasporangia are placed in a petri dish where, upon drying, they dehisce, shedding their spores. The megaspores are collected and Stored in vials. At room temperature, stored megaspores remain viable for several months. According to Robert (1971), megaspores of S. kraussiana exhibit 90%

Leese ; : ; : 268. “Botany Section, Biological Sciences Group, University of Connecticut, Storrs, CT 06268

10 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

germination if cultured immediately following harvest and less than 50% if culture is delayed for three months. A detailed study of viability of stored megaspores of Selaginella is needed.

Megaspores of S. kraussiana and its varieties exhibit a prominent reticulate sculpturing. Microspores possess long spines and tend to adhere to the sculptured surface of megaspores. Any adhering microspores can be removed by washing the megaspores thoroughly prior to incubation. The washing apparatus developed for this purpose consists of a short length of plastic tubing fitted at each end with a polyethylene stopper modified to hold a fine nylon screen cloth (Fig. /). Tygon

FIG. I. Apparatus (side and end views) for washing megaspores and microsporangia. FIG. 2. Self- watering incubation device.

tubing 1.5 cm in diameter and 2.0-2.5 cm long was found convenient. The stop- pers used were those supplied with Kimble Opticlear vials, size 19 x 48 mm. The bottom of each stopper was cut off, and a rod was inserted into the hollow interior to push out the top insert. The center of the insert was cut out, leaving a thin ring. Nylon screen cloth with a mesh opening of 240 zm was laid over the top of the stopper and secured by snapping the ring back into place. The washing apparatus containing megaspores is immersed in a container through which there is a stream of running water. A 30-40 minute rinse is sufficient to remove any adhering microspores and to pass them through the mesh, leaving only clean megaspores inside. Megaspores of S. kraussiana measure 800-1500 zm in diameter (Robert, 1971), and microspores 50-70 xm (Robert, 1973), thus the reason for using a 240 #m mesh opening in the washing apparatus.

After washing, megaspores are removed and incubated on layers of filter paper in a Sself-watering device (Fig. 2). The device consists of a 100 ml polypropylene beaker, on top of which rests a plastic petri dish with a hole drilled in the bottom. One end of a cotton wick is inserted into the hole with the opposite end in contact with distilled water in the beaker. The layers of filter paper in contact with the wick remain constantly moist. A similar device was used by Robert (1971) to germinate megaspores of S. kraussiana.

T. R. WEBSTER: CROSSING TECHNIQUE FOR SELAGINELLA 11

Unlike megasporangia, microsporangia do not protrude from the strobilus at maturity and must be selected with the aid of a dissecting microscope. In dissec- tions of strobili, mature microsporangia can be detected by their amber color and granular texture. Typically a mature strobilus bears one or two ripe microsporan- gia. Since intact microsporangia containing microspores are incubated, dehis- cence of microsporangia must be prevented. Therefore, the dissection is carried out on wet filter paper. The microsporangia are then washed using the same procedure as described for the megaspores, and are then sown on filter paper kept moist by the self-watering device described above. Separate cultures are main- tained for microgametophytes and megagametophytes.

The author maintains cultures in a growth chamber at a temperature of 21+ 2°C and a light intensity of 125 ft-c obtained with fluorescent illumination. A 12-hr light/12 hr dark cycle is used. Under these conditions, megaspores germinate in 8-14 days, and the resulting megagametophytes can be maintained for several months with no exogenous nutrients. The endogenous food reserve, composed of lipid and protein (Robert, 1971), is sufficient to support the slow-growing megagametophytes. Although some gametophytes may become contaminated, infection by microorganisms is usually minimal, so that an occasional change of filter paper and water is sufficient for maintaining the gametophytes.

For fertilization experiments, the author uses megagametophytes which are several weeks old. Such gametophytes possess numerous archegonia and tufts of thizoids. Microsporangia incubated for 4-12 weeks contain mature microgameto- phytes. Microsporangia cultured for longer than 12 weeks often contain micro- Spores which have already shed their contents. To obtain swimming sper- matozoids, 5-10 microsporangia are placed in the cavity of a depression slide containing a drop of distilled water. The sporangia are teased apart with fine dissecting tools, thus dispersing their endosporic microgametophytes. Within a few minutes, spermatocytes are released. A short time later, each spermatocyte releases a single biflagellate spermatozoid which swims away rapidly. Slagg (1932) and Robert (1973, 1974) have described the details of microgametophyte and spermatozoid development in §. kraussiana. Since each microspore contains ap- proximately 250 spermatocytes and each sporangium contains approximately 600 microspores (Slagg, 1932), large numbers of sperm are contained in a single water droplet. The shedding of sperm can be hastened by chilling the microsporangia Prior to teasing them open. The chilling can be accomplished by placing the depression slide containing the sporangia in a refrigerator (7°C) for 4-6 minutes.

Two methods have been used to achieve fertilization. The megagametophytes may be placed directly in the drop of water containing sperm. Upon microscopic €xamination, a dramatic chemotactic response can be seen as the male pegs are attracted to the archegonial pad. After several minutes, a dense a Spermatozoids is visible in the vicinity of the archegonia. As previously noted by Slagg (1932), the spermatozoids are active for approximately 30 minutes.

Another method for achieving fertilization is to transfer the layers te containing megagametophytes from the self-watering device to a petri dish.

12 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

sperm-containing droplet is then added to the megagametophytes. To ensure adequate moisture for the swimming spermatozoids, additional water is added. Since some microgametophytes may not release their gametes immediately, the megagametophytes are left in the sperm bath overnight. Fertilization treatments may be repeated at weekly intervals. In experiments with S. Araussiana and its varieties, two or three treatments resulted in 60-90% successful fertilizations. After the fertilization procedure, the megagametophytes are again incubated on filter paper in the self-watering device. To check for the possible occurrence of apogamy, some unfertilized megagametophytes are incubated separately as con- trols. Repeated tests have shown no evidence for apogamy in S. kraussiana and its varieties.

Sporelings appear in fertilized cultures 4-6 weeks after fertilization. Sporelings will grow for several weeks on filter paper moistened with distilled water, the endogenous food reserve of the megagametophyte serving to nourish the young sporophyte.

Eventually the sporelings can be transferred from filter paper to soil in a pot. Fine forceps should be used to avoid damaging the delicate roots. The soil mixture used by the author consists of a finely sifted sandy loam. To prevent dislodging the delicate sporelings, the soil is watered by setting the pot in a tray of water. To maintain the sporelings in a moist atmosphere, the surface of the pot is covered with plastic or glass. The pots are maintained in the growth chamber until the sporelings are firmly established, after which the pots are transferred to the greenhouse. Each sporeling eventually develops into a mat of intertwined stems which form at their tips spore-bearing strobili. For §. kraussiana and its varieties, the time required to complete the life cycle, from spore to spore, is 9-11 months.

Crossing techniques have been widely used for homosporous pteridophytes (Lovis, 1968). Through their use, much has been learned about the genetics and breeding systems of ferns (Klekowski, 1971). However, with the exception of the study by Burgeff and Filippi (1957), the heterosporous genus Selaginella has been neglected in this regard. It is hoped that the method described here will encourage further work into the genetics and breeding systems of Selaginella.

The author wishes to thank Mr. James A. Tanno for his helpful suggestions and Ms. Virge Kask for preparing the illustrations.

LITERATURE CITED

BIERHORST, D. 1964. Suggestions and comments on teaching materials of the non-seed-bearing vascular plants. Amer. Biol. Teacher 26: 105-107. ninco H. C. 1967. A Laboratory Manual for Plant Morphology. Harper and Row, New York. es ito H. FILIPPI. 1957. Analyse und Erblichkeit der Panaschierung bei Selaginella ensil Spring var. variegata Hort. Biol. Zentbl. 76:637-680. oe ee E. J., Jr. 1971. Ferns and genetics. BioScience 21:317-322. an eh 8 1968. Fern hybridists and fern hybridising. Il. Fem hybridising at the University of Leeds. Brit. Fern Gaz. 10:13-20. ROBERT, D. 1971. Le gametophyte femelle de Selaginella kraussiana (Kunze) A. Br. I. Organisation

générale de la mégaspore. Le diaphr: : ; Cytol. Biol. Vég. 34:96-— 164. phragme et l’endospore. Les réserves. Rev. Cy

T. R. WEBSTER: CROSSING TECHNIQUE FOR SELAGINELLA 13

ROBERT, D. 1973. Le gamétophyte male de Selaginella kraussiana (Kunze) A. Br. Organisation et développement. Etude en microscopie électronique. Ann. Sci. Nat. Bot. (Paris) XII, 14:465-5S04.

. 1974. Etude ultrastructurale de la spermiogenése, notamment de la différenciation de l’ap- pareil nucléaire, chez le Selaginella kraussiana (Kunze) A. Br. Ann. Sci. Nat. Bot. (Paris) XII, 15:65-118.

SLAGG, R. A. 1932. The gametophytes of Selaginella kraussiana. |. The microgametophyte. Amer. J. Bot. 19:106-127.

WEBSTER, T. R. 1967. Induction of Selaginella sporelings under greenhouse and field conditions. Amer. Fern. J. 57:161-166.

WETMORE, R. and G. MOREL. 1951. Sur la culture du gametophyte de Séelaginelle. Compt. Rend. Acad. Sci. 233:430-431.

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14 AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 1 (1979)

A Tropical Fern Grotto in Broward County, Florida

DANIEL F. AUSTIN,* GRACE BLANCHARD IVERSON,** and CLIFTON E. NAUMAN*

John K. Small (1931, p. 1) once wrote that Florida should be known as the *“‘land of ferns.” Several areas throughout the state have become famous for their abun- dance of these plants. Most notable are the rocklands of Dade County (Small, 1920a, b), the Pineola Grotto of Citrus County (Harper, 1916; Lakela, 1964), and the lakes region of Central Florida (Small, 1932, p. 15; Wherry, 1964, p. 28; Lakela & Long, 1976, e. g., p. 123). No one has even suspected until recently (Nauman, 1978) that Broward County boasted ferneries. Few even believe that a site remains unreported that contains the most impressive display of ferns in the southern end of the state.

The site we hereby record is located in Sects. 5,6, T49S, R42E (Fig. /) and contains almost half of the pteridophyte species known in tropical Florida. Fur- thermore, it appears to have the largest and healthiest populations in the state of some of the rarer species. We call the site the ‘‘Cypress Creek Hammock” because it represents a remnant of the Cypress Creek drainage system (Steinberg, 1976). Vegetation in the area is complex, with swamp forest, hammocks, pine- lands, and fallow fields.

A limestone outcrop of undetermined affinity (J. E. Hoffmeister, 1978, pers. comm.) forms a ridge on the western limit and islands near the eastern side. This broken ridge system was dissected by Cypress Creek before the area was drained by canals, and the central swamp is overlain with various depths of peat and sand.

We have been studying the site since September 1976. During that time, we

have discovered that it contains over 200 species of plants, and the list grows with each Visit. Not only is the site the last remaining stronghold of ferns in southeast- ern Florida, but it contains range extensions for several plants (e.g. Asplenium trichomanes-dentatum, Catopsis floribunda, Polypodium ptilodon, Tectaria heracleifolia, Tectaria incisa, Tillandsia valenzuelana), and several species on the Smithsonian and Florida Committee Rare and Endangered Plant lists (€.8- Asplenium trichomanes-dentatum, Asplenium serratum, Epidendrum nocturnum, Ophioglossum palmatum, Tillandisa flexuosa). _ Cypress Creek Hammock is unique in several ways: in its number of ferns, for its range extensions, and for existing as an “‘island”’ of native vegetation in met- ropolitan Broward County. We believe that any one of these factors makes the site worthy of preservation, and that their combination makes it imperative. Some our colleagues have been helpful toward this end by writing letters. We solicit €tters or support in other ways from all those interested in ferns, plants, and native Florida habitats.

* : : ; : oe of Biological Sciences, Florida Atlantic University, Boca Raton, FL 33431. ence Department, Broward Community College, North Campus, Coconut Creek, FL 33066.

D. F. AUSTIN ET AL.: A TROPICAL FERN GROTTO 15

We have received help and encouragement from many individuals in our study. In particular, we wish to thank the following: G. N. Avery (Everglades National Park), E. S. Ayensu (Smithsonian Institution), D. S. Correll and Helen Correll (Fairchild Tropical Garden), R. A. De Fillips (Smithsonian Institution), A. M Evans (University of Tennessee), J. Greis (formerly Broward Co. forester), D. Lellinger (Smithsonian Institution), J. Popenoe (Fairchild Tropical Garden), A. R. Smith (University of California, Berkeley), and R. G. Stolze (Field Museum, Chicago).

FERNS OF CYPRESS CREEK HAMMOCK

Acrostichum danaeifolium Langsd. & Fisch. (Pteridaceae). Leather Fern, Frequent in swampy

Anemia adiantifolia (L. ‘ Pods (Schizaeaceae). Pine Fern. Occasional colonies in pineland areas on the fringe of the hamm

Asplenium trichomanes- dotenin bs (Aspleniaceae). pag neti Several small colonies following a limestone outcrop along the swamp/hammock interface; rare.

Asplenium serratum L. (Aspleniaceae). Bird’s-nest Fern. seve plants in leaf litter and at bases of hammock trees; rare

Blechnum sevealats mL. C. Rich. (Blechnaceae). Swamp Fern. Common throughout the swamp.

Si bea oa phyllitidis (L.) Presl (Polypodiaceae). Strap Fern. Abundant as an epiphyte on fallen lo

Sinha souiniei (Poepp.) Morton (Aspidiaceae). Florida Tree Fern. At least six colonies occur; one contains 20 adults with fronds exceeding | m and more than 30 sporelings. This is probably the largest population of this species in the United State

Ctenitis submarginalis (Langsd. & Fisch.) Tun (Aspidiaceae). Brown-hair Comb Fern. This species is Nia as new for Broward County, having been found there since Nauman (1978) was in press. Several rare

Nephrolepis bserat (Swrata) Schott (Davalliaceae). Giant Sword Fern. Abundant in various Parts of the ham ie

Nephrolepis erat (L.) Schott (Davalliaceae). Sword Fern. Common througho

Ophioglossum palmatum L. (Ophioglossaceae). Hand Fern. Rare in Florida, yet i site contains at least 30 individuals.

Osmunda regalis var. spectabilis (Willd.) A. Gray (Osmundaceae). Royal Fern. Occasional in the

swamp.

Phlebodium aureum (L.) J. Smith (Polypodiaceae). Cabbage Palm Fern. Common wherever cab- bage palms grow.

Polypodium Sshednal var. michauxianum Weath. (Polypodiaceae). Resurrection Fern. Com- mon, especially o

Polypodium inion var. caespitosum (Jenm.) Evans (Polypodiaceae). Plume Polypody. Ra epiphytes; the plants are smaller than the more typical size in other parts of their range. : ;

Psilotum = (L.) Pal. Beauv. (Psilotaceae). Whisk-fern. Common; mostly on the bases 0 cabbage palm

iat “aulinum var. latiusculum (Desv.) Underw. (Pteridaceae). Bracken. Shaded ar

Preris Satis Swartz (Pteridaceae). Giant Brake. Occasional in distu

Pteris vittata L. (Pteridaceae). Chinese Brake. Occasional along paths an the hammock: usually on rocks.

Salvinia i sank S. rotundifolia auctt. non Willd.) (Salviniaceae apa al but |

Tectaria hrc (Willd.) Underw. (Aspidiaceae). Halberd Fern. Occasion limestone outer

Infrequent in

rbed parts of the swamp. d other disturbed parts of

). Water Fern. Occasional

ocal on

16 AMERICAN FERN JOURNAL: VOLUME 69 (1979) Tectaria incisa Cav. f. incisa Seog At least two agen are known from this location.

Known elsewhere from a few sites in Dade County (Naum Thelypteris dentata (Forsk.) E. St. John Caiadiaceae). me coe in disturbed parts of the ham-

mock. Thelypteris interrupta (Willd.) lwatsuki (= T. totta (Thunb.) Schlepe) (Aspidiaceae). Common in the swamp Theluieris kunthii (Desv.) Morton (= T. normalis (C. Chr.) Moxley) (Aspidiaceae). Marsh Fern. Frequent in the hammock.

Thelypteris ovata R. St. John var. ovata (Aspidiaceae). Occasional on limestone outcrops. Thelypteris reptans (Gmel.) Morton (Aspidiaceae). Creeping Fern. One small colony known. Thelypteris torresiana (Gaud.) Alston (= Macr other torresiana (Gaud.) Proctor) (As-

pidiaceae). Occasional in disturbed parts of the ham

Vittaria lineata (L.) J. E. Smith ( Vittariaceae). Sicbanine Fern. Frequent on cabbage palms. LITERATURE CITED HARPER, R. M. 1916. The fern grottoes of Citrus County, Florida. Amer. Fern J. 6:68-81 LAKELA, OLGA 1964. Fewer Florida rarities: changing flora of Pineola Grotto, Citrus County. Sida 299-305.

, an LONG. Ferns of Florida. Banyan Books, Miami, FL.

NAUMAN, C. i 1978. A check-list of the ferns of Se a. hens Castanea 43:155-162.

SMALL, J. K. 1920a. A journey to the fern grottoes. J. N.Y. . Gard. 21:25-39. or Of grottoes and ancient dunes. J. N.Y. Bot. Ga. 21:45-S2.

————. 1931. Ferns of Florida. Science Press, New Yor

STEINBERG, B. 1976. Vegetational analysis of the Ailenitic — ed of Broward County, Florida. Master's Thesis. Florida Atlantic University, Boca ;

WHERRY, E. T. 1964. The Southern Fern Guide. Doubleday, reel a NJ.

REVIEW

“EVOLUTIONARY PATTERNS AND PROCESSES IN FERNS,” by J. D. Lovis in R. D. Preston and H. W. Woolhouse (eds.). Advances in Botanical Research, vol. 4, pp. 229-415. 1977. Academic Press, London and New York. ISSN 0-12-005904-5. $30.00.—Professor Lovis has written a tour de force that is the most important summary of evolutionary mechanisms in ferns since Professor Manton’s seminal book on fern cytology published over 25 years ago. Lovis’ paper begins with a detailed consideration of the fossil record and how it relates to the classification and phylogeny of recent ferns. A careful list of chromosome numbers in fern genera is given. A classification of living ferns by family and subfamily is proposed that takes both fossil and chromosome data into full ac- count. Polyploidy is discussed at length and is analyzed by its systematic and geographic distribution. Present knowledge of genome analysis, cytogenetics, in- trogression, breeding systems, and apomixis in ferns is all summarized. An exten- sive list of references concludes the work. Unfortunately, they are cited without nig titles, and so are of limited use apart from their mention in the text.—

AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 1 (1979) 17

Studies of the Azolla—Anabaena Symbiosis Using Azolla mexicana, I. Growth in Nature and Laboratory ROBERT W. HOLST! and JOHN H. YOPP*

The symbiosis of Azolla and Anabaena is the only known association of a pterophyte with a cyanophyte. Anabaena azollae is found in all species of the aquatic heterosporous fern Azolla. It is enclosed within a small, basal cavity of the upper green lobe of the bilobed, alternate leaves. Very rapid propagation of the intact Azolla is generally achieved by fragmentation, but may occur by germina- tion of algal-infected sporocarps (Rao, 1935; Smith, 1955, pp. 371-383). In many temperate and tropical regions, Azolla is considered a pest because it can cover a farm pond or rice paddy rapidly (Sculthorpe, 1967). However, it has long been cultivated as a green manure for rice crops in southeast Asia (Moore, 1969). This latter use provides the basis for the recent intense interest in this unique sym- biosis. The algal symbiont is a nitrogen-fixing organism that behaves much like Rhizobium species do in their symbioses with legumes.

Studies on the factors affecting the growth of the Azolla-Anabaena symbiosis are relevant to research on nitrogen-fixation and also weed control. Previous laboratory studies on nitrogen-fixation have employed A. filiculoides Lam. or A. caroliniana Willd. (Ahmad, 1941; Peters & Mayne, 1974). However, of the six New World species, A. mexicana Presl is the major representative of the central and western United States (Svenson, 1944). The present study was conducted to characterize the optimal environment for growth of A. mexicana. By virtue of environmental preferences or a more controllable or rapid growth rate, this species might provide a better and more convenient model than larger land plants do for studies of symbiotic nitrogen fixation.

MATERIALS AND METHODS

Studies in the natural environment.— Growth of Azolla mexicana was studied . a small, 2 ha pond in southwestern Jackson County, Illinois. This pond, in the Mississippi River flood plain, is fed by runoff from low alluvial plains and swamps delimited by levees.

The environmental factors recorded at each observation were light intensity (Weston photometer), light quality (IL 150 Photometer, International Light, Newburyport, MA.), photoperiod, air temperature in the shade and in the sun, and water pH and surface and bottom temperature. Total plant cover of the pond Surface and the percentage contributed by A. mexicana were also recorded. Plant cover was determined by the line intercept method from photographs of the pond surface (Canfield, 1941).

Laboratory studies. — Plant material was collected from the pond on 18 June 1975. It was separated and washed clean of other aquatic plants. All subsequent

2901.

i Ee ae ; ale, IL 6 Department of Botany, Southern Illinois University at Carbondale, Carbonda Ce iuieion,

De nt address: Office of Pesticide Programs, U.S. Environmental Protection Agency,

18 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

experiments were performed with material vegetatively reproduced from this col- lection.

Several media successfully employed for culture of aquatic plants (including Azolla spp.) were tested singly and in various combinations for their capability to support the growth of A. mexicana (Johnson et al., 1966; Olsen, 1970; Lahdes- maki, 1968; Ahmad, 1941; Nickell, 1958; Peters & Mayne, 1974). Buffers were not used unless specifically noted in the tables or figures. The pH was adjusted with either N KOH or HC1.

Plants were maintained in deep, rectangular, polyethylene containers partially covered with plate glass to obtain a humidity higher than ambient. The number of initial plants per container was approximately one per 10 cm”. Culture density was selected from preliminary experiments, which indicated that crowding is an im- portant variable.

The containers were placed in walk-in growth chambers (Sherer- Gilette Model CEL 511-38) that were set for different light intensities, photoperiods, and day/ night temperatures.

All growth studies were conducted using 5 plants per replicate. Each set of five plants was grown in a 125 ml Erlenmeyer flask containing 50 ml of culture medium.

Growth is expressed as a fold change which was determined from the formula: fold change = Wt/Wo, where Wt is fresh weight at time of sampling and Wo is initial fresh weight of the culture. A fold change of 2 indicates doubling in fresh wre The number of days or hours to attain this value is designated the doubling

ime.

Chlorophyll content was determined by the measurement (Beckman DBGT spectrophotometer) of the optical density (O.D.) at 652 nm of an 80% acetone extract of a plant homogenate. The method of Arnon (1949) was used in the following form:

O.D.652nm x ml of liquid = mg chlorophyll/g FW 3.45

All experiments were replicated three times. Means, standard variations, and F test probability were determined using multiple regression analysis (Kelley et al., 1969). Student’s t-test and correlations were obtained using the IBM 5100 Basic Statistic Program.

RESULTS AND DISCUSSION

Natural environment study.—The appearance of A. mexicana in its natural habitat was strongly correlated with surface and bottom temperatures of the pond ( Fig. 1). The critical temperature (approximately 15° C) was attained early in April of 1975, but not until after 1 May 1976 due to cooler and fluctuating tempera- tures, which delayed the appearance of the fern. Light intensity and, of course, photoperiod were not greatly different for the two spring seasons. Numerous observations indicated that floating sporophytes resulted from the temperature- dependent germination of zygotes in the bottom mud.

HOLST & YOPP: AZOLLA- ANABAENA SYMBIOSIS,

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20 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

The earlier and more extensive vegetative growth of the fern during the spring of 1975, as compared to that of the following spring, also paralleled the much more rapid rise and higher water temperatures attained in 1975. After the water temper- atures rose to 20° C, A. mexicana became the dominant, floating, emergent plant in the study location. The total plant cover was almost 60%, with the fern account- ing for 80-90% of that total.

The fern showed signs of increasing red pigmentation and senescence in the open areas of the pond. Light intensity there was 95 to 120 klux ( Table 1), and temperatures remained fairly constant from May through August of 1975. Photo- period was apparently not a factor because A. mexicana was seen to increase in May and June of 1976, a period of decline in 1975. The green plants along the shaded shoreline accounted for the fern’s presence in the summer months, even when water temperatures exceeded 25°C. The light intensity never exceeded 3 klux in these shaded areas. High light intensity, therefore, appears to be the factor most responsible for senescence of this fern. Ahmad (1941) gave a similar explana- tion for senescence of A. pinnata R. Br. Light quality did not significantly change throughout the study period (Table /).

TABLE 1. TOTAL ILLUMINATION (klux) AND SPECTRAL INTENSITIES (uw/cm? - nm) OF LIGHT ON CLOUDLESS DAYS AT 10AM LOCAL TIME, SUMMER 1975, AT BRUSHY BAYOU POND.

Area Total light Red light Far red light Blue light Direct sun 95-120 110-125 90-100 15-115 Shade 3-0 57 6-9 5-12

Sexual reproduction, as shown by mega- and microsporocarp production, oc- curred in both years of the study in June (Fig. 1, arrows). Since the populations were decreasing in June 1975 and increasing in June 1976, photoperiod may be the principal trigger of sporulation.

A second growth of A. mexicana was noted in fall of 1974, when water tempera- tures remained between 18 and 25°C. However, September 1975 was very cool, water temperatures fell below 15°C, and A. mexicana did not reappear.

During its season, the fern grew well in the shaded areas of the pond and even on the moist bank. There was, however, an apparent correlation between soil moisture and the survival of the A. mexicana, even on obviously moist soil. There — . relatively distinct level in the moist soil bank above which the fern was not ound.

The pH of the pond ranged from 6.3 through 6.8 throughout the study.

Previous studies on other species of Azolla indicate that the intact fern grows well under reduced light conditions, at a PH of 5 to 7, and at temperatures not exceeding 23 to 25°C (Ahmad, 1941; Moore, 1969; Olsen, 1970). Azolla mexicana appears to have similar environmental requirements.

Laboratory studies.— Preliminary experiments based on the natural environ- mental study Showed that a photoperiod of 14 hours light/10 hours dark, a 12 klux light intensity (cool white fluorescent), and a 23°C day/18°C night thermoperiod were optimal for growth. These values were employed in laboratory experiments

HOLST & YOPP: AZOLLA-ANABAENA SYMBIOSIS, |. 271

unless otherwise noted. Temperature, pH, light intensity, osmotic stress, nutrient element concentration and salinity were the experimental variables in the labora- tory studies.

Growth medium.—Although A. mexicana grew on all the media tested. a medium was devised by extensive experimentation that gave superior results (Table 2). The doubling rate in this medium was 68 to 75 hours if other conditions were optimal. This is about 48 hours less than that reported by Nickell (1958) for A. mexicana and for A. caroliniana (Dr. Gerald Peters, pers. comm.). It is similar to that reported for A. pinnata (Brotonegoro & Abulkadir, 1976).

PO,4 + TRIS

CITRATE

TRIS

GROWTH (FOLD CHANGE IN FW)

ou + oO o+ “I @ o

FIG. 3. Effect of pH on A. mexicana growth under environmental conditions as in Fig. 2. Buffers (10 mM) were POs (potassium phosphate), citrate (citric acid-sodium citrate), TRIS (Tris [Hydroxy methyl]-amino methane), and MES (2-[ N-Morpholina] ethane sulphonic acid).

Crowding effect.—A typical growth curve for A. mexicana obtained under op- timal conditions is given in Fig. 2. The culture generally reached its most rapid growth rate after five days. The rapid rate continued for up to 12 days and if crowding was noted, decreased rapidly thereafter. Crowding was scored when Overlapping of fronds between plants occurred. Mats of ferns up to 2 cm thick frequently were observed. Cultures could be thinned and the remaining plants fragmented by hand without experiencing a reduction in growth. Similar mat ormation and results from fragmentation have been reported for A. filiculoides in African waters (Ashton & Walmsley, 1976).

PH of medium.—The pH of the unbuffered medium was set at 6.5. . after two to three days the pH dropped to 4.2 and remained constant thereafter. I the medium was not changed biweekly, A. mexicana began to produce yellow- edged fronds. A variety of buffers (phosphate, Tris, MES, citrate) were used at 0.5 mM to determine accurately the pH range of A. mexicana. It grew well fas the pH range of 4.2-8.0 (Fig. 3). Below pH 4.2, growth decreased dramatically.

AMERICAN FERN JOURNAL: VOLUME 69 (1979)

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re) (M4 NI 3ONWHDO GQ104) HLMOND

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(M4 NI SONVHS a0 4) HLMOUD

HOLST & YOPP: AZOLLA-ANABAENA SYMBIOSIS, |. 23

The peak noted at 6.5 was not significant (P=0.01). The decline in growth above 5.1 with citrate and Tris buffers may be due to the buffers per se. Other buffers used were not toxic above 5.2. This optimal pH range agrees with that reported by Nickell (1958) for A. mexicana and for other species of the fern (Moore, 1969: Olsen, 1970). Ashton and Walmsley (1976), however, reported dual, light intensity-dependent pH optima. This was not observed in the present study.

Temperature effects.—When A. mexicana was cultured under low light inten- sities (less than 13 klux), it grew well at temperatures up to 30°C (Fig. 4). Growth slowed at 35°C and the fern died within seven days at 40°C. Growth ceased but the plants remained green and viable at 5°C.

A similar temperature range and optimum were reported for A. pinnata in Asia by Moore (1969). Azolla filiculoides in Africa has a higher temperature optimum (27.5°C) and a broader range (5 to 45°C), according to Ashton and Walmsley (1976). The same species in India possesses a temperature range and optimum similar to that of A. mexicana (Ahmad, 1941).

TABLE 2. BASIC MEDIUM FOR MAXIMUM GROWTH OF INTACT Azolla mexicana IN VITRO.

Macronutrients mg/l Micronutrient stock’ CaCh - H20 176 H3 BOs 286 mg K H2POs 53 CoChk - 6H20 5m l 78 MnCh - 4H20 181 mg MgSOs - 7H20 240 ZnSOs4 - 7H20 22 mg Fe stock! CuSO. - 5H20 8 mg FeChel? 250 mg NaMoOs - 2H20 13 mg FeSO. - 7H2O 250 mg NaCl 100 mg 50 m1 H2O 100 ml

‘Use 1 ml/1 of medium. late (10% *FeChel is sodium ferric diethylenetriamine penta-acetate-Sequestrene 330 Fe Iron — : metallic Fe) sold by Ciba-Geigy.

Solute tolerance.—Addition of NaCI to produce a salt concentration of 2000 ppm in the medium had no significant effect on growth (Fig. 5). At 4000 ppm solutes, growth was slightly reduced. Over the range of 556 (basal medium) to 4000 ppm solutes, only a slight decrease in chlorophyll was observed (700 to 500 Hg chlorophyll/g FW). ;

The review by Moore (1969) indicated that Azolla species were killed after three weeks in Knop’s solution (1500 ppm salts). Haller et al. (1974) found that 4. Caroliniana grew well in a sea and pond water mixture of 3000 ppm solute and was not killed by a salt content of 16,000 ppm. :

Osmotic effects.—In order to separate the effect of the salts themselves from the Osmotic effect they exert, A. mexicana was grown in media of sion polyethylene glycol (PEG) content. The failure of the fern to grow on a . in the natural environment suggested inhibition due to low water eae oe srowth occurred in 4000 ppm NaCl, which is equivalent to -1.69 bars ( pate : 1974), a range of 0 to -10 bars was selected. As seen in Fig. 6, growth ag uy decreased down to -5 bars. There was no growth below -5 bars, but the fern

24 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

remained green and viable. The effect of lowered water potential due to PEG parallels that due to salt, indicating more of an osmotic than an ionic effect pro- duced by salts.

Light intensity.—Light intensity was a major growth regulator. At intensities above 20 klux, orange-red pigmentation developed rapidly. Within seven days, plants ceased growth and entered into a senescent phase. Growth did not occur below 0.75 kluy

GROWTH (FOLD CHANGE IN FW) AFTER 7 DAYS

0 -2 -4 - 8 -10 5 (BARS)

FIG. 6. Effect of osmotic stress on A. mexicana growth. The values are for seven days’ growth in

media containing appropriate amounts of polyethylene glycol (PEG 6000). All other environmental conditions as in Fig. 2.

The present study has revealed the major environmental variables affecting the growth of A. mexicana. These data and the formulation of a growth medium that Supports one of the most rapid growth rates reported for a species of Azolla will enable A. mexicana to be used as an experimental organism.

LITERATURE CITED AHMAD, G. 1941. Effect of light intensity and temperature on the growth of Azolla filiculoides. J. Indian Bot. Soc. 20:313-326. ARNON, D. I. 1949. Copper enz Plant Physiol. 24:1-15. ASHTON, P.J., and R. D. WAL Endeavour 35:39_43. BRONTONEGORO, S., and S. ABDULKADIR. 1976. Growth and nitrogen-fixing activity of Azolla pinnata. Ann. Bogor. 6:69-77. CANFIELD, R. 1941. Appl Forestry 39:388-394

ymes in isolated chloroplasts. Polyphenol oxidase in Beta vulgaris.

MSLEY. 1976. The aquatic fern Azolla and its Anabaena symbiont.

ication of the line interception method in sampling range vegetation. J.

HOLST & YOPP: AZOLLA-ANABAENA SYMBIOSIS, |. 25

HALLER, W. T., D. L. SUTTON and W. C. BARLOWE. 1974. Effects of salinity on growth of several aquatic macrophytes. Ecology 55:891-894.

jigea — V., P. A. MA EU X, bigs = J ai ddaclte Aesbit A cobalt requirement for symbiotic

of Azolla filicul Plant Physiol. 41:852-855.

RELLY. F. 5 D. L. BEGGS, K. A. McNEIL, J. EICHELBERGER, and J. LYON. 1969. Multi- ple es hoard Approach. Southern Illinois University Press, Carbondale, IL.

LAHDESMAK . 1968. Free amino acids in the leaves of Salvinia natans and Azolla filiculoides grown in eats and — — Plant. 21:1097-1103.

MOORE, A. W. 1969. Azolla: Biology and agronomic significance. Bot. Rev. 35:17-34.

NICKELL. L. G. 1958. Physiological studies with sof: Ps aseptic conditions. I. Isolations and preliminary growth studies. Amer. Fern J. 3

NOBEL, P. S. 1974. Introduction to Biophysical eet : Beker: Freeman, San Francisco.

OLSEN, Ae 1970. On biological nitrogen fixation in nature, particularly in blue-green algae. C.

av. Lab. Carlsberg 37:269-283.

aa 4 A., and B. C. MAYNE. 1974. The Azolla, carr Azollae relationship. |. Initial characterization of the association. Plant Physiol. 53:813-81

RAO, H. A. 1935. The structure and life history of Azolla pinnata R. sere with remarks on the fossil history of the & fon ecto Proc. Indian Acad. Sci., sect. B, 2(2):175-200.

SCULTHORPE, C. D. 1967. The Biology of Aquatic Vascular Plants. Edward Armold, London.

SMITH, G. M. 1955. tiie Botany, vol. II], 2nd edition. McGraw-Hill, New York.

SVENSON, H. K. 1944. The new world species of Azolla. Amer. Fern J. 34:69-84.

a

pH ANALYZER pH controls: | e microorganisms in sol ENDS SOIL e soil fertility GUESSWORK e plant's ability to use fertilizer e plant's resistance to disease e mature development, growth, and yield

pH ANALYZER $19.95 prepaid ows PH status instantly.

recision instrument sh No Lon boonbood Simply insert probes into

No soil samples, chemicals, or dyes. il eee tool for growing all plants. Free e guidebook. Send check, money order, Mastercharge, or Environmental Concepts, Dept. FJ shay N.W. 57th auderdale, FL 33309 nies ~ FREE catalogue.

26 AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 1 (1979)

Thelypteris in Arkansas W. CARL TAYLOR* and DAVID M. JOHNSON**

Recent herbarium studies and field work indicate that the genus Thelypteris in Arkansas is represented by four species. Two of these, 7. kunthii (Desv.) Morton and T. torresiana (Gaud.) Alston, are reported here for the first time in the state.

In the United States, T. kunthii, a neotropical species, is found in open or partially shaded, moist depressions on the coastal plain from South Carolina to Texas (Smith, 1971). Specimens of 7. kunthii have been collected from Lee and Pulaski Counties (central and east central Arkansas), but both collections were made from plants persisting in garden plots. These plants do not appear to have escaped cultivation sufficiently to be considered naturalized. However, numerous plants recently have been found in Ashley County (southeast Arkansas) which are scattered over several square miles of cut-over pine-hardwood forest northwest of the town of Crossett (Fig. ]). These plants undoubtedly are naturalized and prob- ably became established as a result of new habitats created by lumbering opera- tions in the area. Common associated species in these open, disturbed habitats include Rhus copallina, Liquidambar styraciflua, Callicarpa americana, Quercus stellata, Acer rubrum, Diospyros virginiana, Berchemia scandens, Pteridium aquilinum var. pseudocaudatum, Onoclea sensibilis, Polystichum acrostichoides, and Asplenium platyneuron var. incisum.

Thelypteris torresiana, a species native to Asia, was first discovered in the United States in 1904. An interesting historical account of its collections is given by Leonard (1972), who also postulates the occurrence of this species in south- eastern Arkansas. The discovery of T. torresiana in virtually the same area of Ashley County and in habitats like those of T. kunthii suggests that disturbance from lumbering is also responsible for the establishment of T. torresiana (Fig. 2). The discovery of naturalized plants of T. kunthii and T. torresiana extends the known range of both these species north into the coastal plain of southeastern Arkansas (Fig. 6).

Thelypteris noveboracensis (L.) Nieuwl., which in Arkansas inhabits moist, rocky soils of woods and thickets along streams, reaches the southwestern extent of its range in the Ouachita Mountains (Fig. 3). The known Arkansas populations of T. noveboracensis are over 200 miles from the nearest reported stations in the Ozark Hills of southern Illinois and the Tennessee River Hills of northeastern Mississippi.

— Var. haleana Fern. has been reported in Drew and Bradley Counties. €se two populations also contain var. pubescens and forms that appear to be

« \ A agape Department, Milwaukee Public Museum, Milwaukee, WI 53233. Botany Department, University of Michigan, Ann Arbor, MI 48109.

TAYLOR & JOHNSON: THELYPTERIS IN ARKANSAS

ee eiciral

= a

.)

ae Panes °

Lee

°

nt ak on

ee

2 = a >

— @ 2s 8

is. FIG. 6. Physiogra

FIGS. 1-5. Distribution maps of Arkansas Thelypter

28 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

We wish to thank Dr. Alan R. Smith for checking the identifications of T. kunthii and T. torresiana.

REPRESENTATIVE COLLECTIONS:

Thelypteris kunthii: Ashley Co.: Along timber access road SE of Lake Georgia-Pacific; T17S, ROW, sect. 35, 25 Oct 1975, Johnson 281 (Hendrix College).

Thelypteris torresiana: Ashley Co.: Along timber across road W of Crossett; T18S, ROW, sect. 9, 25 Oct 1975, Johnson 283 (Hendrix College).

Thelypteris noveboracensis: Cleburne Co.: Johnson 393 (Hendrix College). Garland Co.: Moore 6278 (APCR). Montgomery Co.: Taylor 1084 (SIU). Pike Co.: Taylor 2903 (SIU). Polk Co.: Moore 410261. (UARK). Pulaski Co.: Johnson 112 (Hendrix College). Saline Co: Moore 480/54 (UARK).

Thelypteris palustris var. pubescens: Ashley Co.: Johnson 276 (Hendrix College). Bradley Co.: Demaree 19445 (MO). Drew Co.: Moore 420048 (UARK). Greene Co.: Moore 480685 (UARK). Hempstead Co.: Moore 480353 (UARK). Izard Co.: Johnson 422 (Hendrix College). Lawrence Co.: Taylor 1794 (S1U). Little River Co.: Palmer 8359 (MO). Polk Co.: Moore & McWillliam s.n. (U ARK). Sharp Co.: Wade 167 (UARK). Washington Co.: Henbest 16 (UARK).

Thelypteris palustris var. haleana: Bradley Co.: Cypress swamp near Warren, Demaree 19445 (SMU). Drew Co.: Swamp near Wilmar, Demaree 24624 (SMU).

LITERATURE CITED

DEMAREE, D. 1943. A catalogue of the vascular plants of Arkansas. Taxodium 1:1-88.

LEONARD, S. W. 1972. The distribution of Thelypteris siana in the southeastern United States. Amer. Fern J. 62:97-99.

SMITH, A. R. 1971. Systematics of the neotropical species of Thelypteris section Cyclosorus. Univ. Calif. Publ. Bot. 59:1- 143.

ROBERT J. RODIN (1922-1978)

Robert Joseph Rodin was born in Sacramento, California, on 15 July 1922. He lived in Eureka and in Turlock in his youth. During World War II service in the lS. Marine Corps, he was stationed in Guam and China. In 1948 he married Elva Bain, who, with three daughters, survives him. He received a Ph.D. degree in botany from the University of California at Berkeley in 1951. After graduation, he accepted a position as a professor of biology at Forman Christian College, Lahore, West Pakistan. He collected ferns in the Himalayas extensively during this time. From 1953 until his retirement in 1976, Rodin was a professor of biology at California Polytechnic University in San Luis Obispo. His research specialties included the anatomy of the Gnetales, ethnobotany of Ovamboland, and the taxonomy of the pteridophytes of California, an interest which led to his publish- ing the *‘Fems of the Sierra’ for the Yosemite Natural History Association. In recent years he was active in the conservation movement in California. In 1966— 67 he was a Fulbright Professor at the University of Delhi. He also took part in botanical expeditions to South Africa in 1947 and to southwestern Africa in 1973. He died in San Luis Obispo on 27 June 1978.—D.B.L.

AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 1 (1979) 29

SHORTER NOTES

VITTARIA LINEATA REDISCOVERED IN GEORGIA. — Vittaria lineata (L.) J. E. Smith, the Shoestring Fern, is primarily restricted to peninsular Florida reaching as far north as Citrus Co. on the Gulf coast and Duval Co. on the Atlantic coast (E. T. Wherry, The Southem Fem Guide, Doubleday, 1964). However, a disjunct population had been known since 1938 in Lincoln Co., Geor- gia. This piedmont population grew in pits in south-facing sandstone cliffs. Unfor- tunately, this area was quarried and the original locality destroyed. Thus, as far as known, Vittaria has been extinct in Georgia for some years.

This note reports the rediscovery of V. lineata in Georgia on Cumberland Island, Camden Co. The plants were observed at three localities on the island some distance apart. Two sites were bark fissures of Live Oak trees (Quercus virginiana Miller); the third was in a bark fissure of Sweet Bay (Magnolia vir- giniana L.) All sites were approximately one meter above the ground on the east and southeast exposures of the tree boles. One specimen (Worthington 336) is on deposit at the University of Georgia herbarium, and another (Stoneburner s.n., 19 Sept 1978) is in cultivation at the University of Georgia Plant Growth Facilities.—James G. Bruce, Department of Botany, and D. L. Stoneburner, James I. Richardson, and Joanne Worthington, Institute of Ecology, University of Georgia, Athens, GA 30602.

THE DISTRIBUTION OF DRYOPTERIS GOLDIANA AND D. MARGINALIS IN MISSOURI. — Dryopteris goldiana (Hooker) Gray is one of the rarest ferns in Missouri. The only known occurrences have been early collections made in two isolated locations along the eastern border of the state. These were noted by Steyermark in ‘* Flora of Missouri” (1963, p. 40). In July, 1976, I discovered the first locality for this species on the western side of the state. Plants were found growing in rich loess deposits underlain by local limestone formations In a ravine tributary to an unnamed intermittent drainage north of the Missouri River in south-central Clay County, in a city park within the city limits of Kansas City. This particular site is 200 miles from either of the two previously recorded Mis- souri locations, and suggests that perhaps D. goldiana also occurs in the interior sections of Missouri, although it has not been recorded from there. Habitat infor- mation listed in Gleason and Cronquist’s ‘* Manual of Vascular Plants of North- eastern United States and Adjacent Canada’”’ (1963, p. 24) noted a preference for moist woods in circumneutral soil. The Kansas City site is Knox Silt Loam oe PH 5.6-7.3. Other fern species found in association with D. goldiana show the richness of the site: Adiantum pedatum var. pedatum, Athyr um PyYEOCATP a Botrychium virginianum, Cystopteris fragilis, Dryopteris mar ginalis, Onoclea Sensibilis, and Woodsia obtusa, as well as liverworts. The richness of the fern flora at this site is unique in northwestern Missouri. bois + tox thie

The presence of Dryopteris marginalis (L.) Gray at this site 1s unusual, see Species is commonly found in shaded crevices of rock ledges and bluff sites

30 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

Missouri where soils are generally derived from sandstone, chert, or granite. While in the same vicinity of D. goldiana, the plants of D. marginalis were found at a slightly higher elevation on a north-facing slope. Dryopteris marginalis also was located on the thick loess deposit, an unusual substrate for this species in Missouri. This particular location is the northwesternmost occurrence of D. mar- ginalis in Missouri; Steyermark’s ** Flora of Missouri’ (1963, p. 40) recorded it from Saline County, 75 miles east of this latest discovery. The species is fairly common south of the Missouri River in most counties of the Ozarks, but has not been found in the northwestern part of the state.

The site was nominated as one of Missouri’s State Natural Areas, representing the rich river hills community type as well as preserving the habitat for a rare Missouri species, D. goldiana. Through a cooperative agreement between the Kansas City Parks Department and the Missouri Department of Conservation, State Natural Area status was approved in February 1978.

Documentary specimens have been filed in the herbaria of the University of Missouri at Columbia (UMO) and at Kansas City. — Greg F. Iffrig, Missouri Cooperative Wildlife Research Unit, Stephens Hall, University of Missouri, Co- lumbia, MO 65211.

GYMNOGRAMMA VS. GYMNOGRAMME. — Confusion has existed over the proper spelling and gender of the generic name Gymnogramma. Although this name is illegitimate (see Underwood, Bull. Torrey Bot. Club 29:617-634. 1902), its wide use in the past has made it often quoted in current literature. The name has been considered to be either feminine or neuter and has been alternatively spelled Gymnogramme.

In originally naming the genus Gymnogramma, Desvaux (Berl. Mag. 5:304- 1811) cited the derivation of gramma as from the Greek meaning ‘“‘line.’’ In this sense, the word gramma (ypauud) is the Doric form of the more usual Attic Greek gramme (ypaupr) (Pichi-Sermolli, Webbia 21:487-505. 1966). Both words have the Same meaning and both are feminine in gender. The problem arises from the similar-looking Attic Greek word, gramma (ypappa), which is neuter and means “that which is drawn’’ or ‘‘letter.’’ Some authors, unaware of the feminine Doric form gramma and/or confused by the identical spelling of the different neuter Attic word gramma, have decided that Desvaux’s use of gramma as feminine was improper and that the word should either be considered as neuter oF should be changed to gramme. However, since Desvaux specified that gramma means *‘line,”’ and treated it as feminine, he clearly was intentionally and cor rectly employing the Doric form of gramme. Modifications of the generic name to Gymnogramme or specific epithets to a neuter ending are not correct.

This clarification has a rather wide application since there are names of many fern genera based on gramma or gramme (e.g. Anogramma, Coniogramme, etc.) In all cases when these are derived from the Greek word meaning ‘‘a line,”’ the original spelling, whether as gramme or gramma, should be maintained and epithets should have a feminine termination. If there are any derived from the

AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 1 (1979) 31

Greek word meaning “that which is drawn”’ or ‘‘a letter,”’ these also should retain their original spelling but with epithets of a neuter termination. Where a derivation is not included, gender of the name should follow that published with the name.—Christopher H. Haufler and Rolla M. Tryon, Gray Herbarium, Harvard University, 22 Divinity Avenue, Cambridge, MA 02138.

REVIEW

“FERNS,” by Philip Perl and the Editors of TIME-LIFE Books, Alexandria, Va. 1977. 159 pp. + many photographs; numerous watercolors by Richard Crist. $8.95.—To say that this volume is beautifully illustrated and contains a wealth of information for the amateur fern grower is true, but it also is an oversimplification that does not do justice to the term ‘‘review.”’ In fact, this book is a good example of a modern, mass-produced encyclopedia volume. It is an attractively packaged hodge-podge of enticing tidbits of information which are priceless for beginning fern growers. Unfortunately, the editors were too intent on interjecting poetic and artistic touches, rather than on fulfilling their implied purpose of giving com- prehensive information on fern growing, including recent and advanced knowl- edge of the subject.

The introductory chapter is a delightfully written historical sketch, but is bro- ken up by the inappropriate interposition of photographic essays having little orno relationship to the text. The remaining chapters fare a bit better in this regard. There is much to be gleaned from the horticultural information in chapter four, although I doubt that many readers will find a ready supply of builder S sand. Perlite, which is available nearly everywhere and which is included in most soil Preparations based on the Cornell mix, is omitted here, although many of the cultural suggestions in chapter five include it.

The photographic illustrations are superb, especially those in the essay on identifying ferns by family. On the other hand, I found the delicately attractive watercolors in chapter five to be little more than decorative. They have ye ethereal quality not always representative of the stark greens and bold forms - the species they pretend to illustrate. For example, the excellent photographs 0 the Hacksaw fern on pages 70 and 72 bear little resemblance to the illustration on Page 110. A number of the watercolors appear so similar as to be of little value in distinguishing between superficially similar but distinct genera.

32 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

The fern encyclopedia comprising chapter five is the meat of the volume, and occupies the second half of the book. It contains some identifying characteristics of a variety of ferns, mostly cultivated, their provenance, and hints on their culture, with recommended temperature, foot-candles of light, and pH readings ad infinitum, as if the latter two were all that important. I am not convinced that the average fern grower will ever determine his foot candles or pH, or even care. Elsewhere in this work the ubiquitous Asparagus-fern is defrocked, but in chapter five, Selaginella and Equisetum are treated as if they were legitimate ferns. An extensive and useful bibliography is given on pages 152-153.

Considered as a whole, even with its idiosyncracies, this is a handy and attrac- tively priced volume for fern growers.—Robert Read, Department of Botany, Smithsonian Institution, Washington, DC 20560.

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Plant Classification Second Edition

Lyman Benson, Pomona College, Emeritus 1979 Casebound 736 pages

This classic text has been revised to include recent research: new examples for major manuals on floras in the Pacific Northwest, California, Northeast, and South; reassignment of the members of the Amentiferae to new positions in the Thalamiflorae and Calyciflorae; revisions in the tribes of the grasses; recent findings in evolution; and classification systems of flowering plants. Plant Classification, Second Edition is a superbly illustrated text/reference book written to help readers: acquire essential vocabulary for describing characteristics of plant groups; identify plants by applying the use of keys and descriptions; gain knowledge of plant.taxa through preparation and preservation of specimens to form an ordered collection; develop an under- standing of the basis for classification of plant groups; and gain an appreciation of the association of species in natural vegetation.

Section Heads:

Flowering Plants Gymnosperms

erns

Psilophytes and Horsetails Club Mosses

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Natural Floras HEATH

Appendix Glossary

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AMERICAN ee c E ee N ee 2 JOURNAL ope

QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY

A Community of Lycopodium Gametophytes in Michigan JAMES G. BRUCE and JOSEPH M. BEITEL 33

Acrostichum in Florida DAVID C. ADAMS and P. B. TOMLINSON 42

Phyllitis scolopendrium Newly Discovered in Alabama JOHN W. SHORT 47

Studies in Lycopodiaceae, II. The Bran

ranching Patt erns and Infrageneric Groups of Lycopodium B.@LLGAARD 49

Shorter Notes: Isoétes butleri in Georg Juvenile Leaves of the ipuaa? = Notholaena 62 cochisensis; A Thelypteris New to Florida

MISSOURI BOTANICAL

JUL 10 BP

GARDEN LABRARY

The American Fern Society Council for 1979

RICHARD L. HAUKE, Dept. of Botany, University of Rhode Island, Kingston, R.I. 02881. President ROBERT M. LLOYD, Dept. of Botany, Ohio University, Athens, Ohio 45701. Vice-President LESLIE G. HICKOK, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916. Secretary JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916. Treasurer JUDITH E. SKOG, Dept. of Biology, George Mason University, Fairfax, Va. 22030. Records Treasurer DAVID B. LELLINGER, Smithsonian Institution, Washington, D.C. 20560. Journal Editor ALAN R. SMITH, Dept. of Botany, University of California, Berkeley, Calif. 94720

Memoir Editor JOHN T. MICKEL, New York Botanical Garden, Bronx, N.Y. 10458. Newsletter Editor American Fern Journal EDITOR DAVID B. LELLINGER Smithsonian Institution, Washington, D. C. 20560

ASSOCIATE EDITORS DAVID W. BIERHORST ..Dept. of Botany, University of Massachusetts, Amherst, Mass. 01002 GERALD J. GASTONY ...........,, Dept. of Biology, Indiana University, Bloomington, Ind. 47401 JOHN T. MICKEL New York Botanical Garden, Bronx, New York 10458

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Memoir Dr. Alan R. Smith, Dept. of Botany, Univ. of California, Berkeley, California 94720, is Editor of the memoir series “ Pteridologia.”’ Newsletter = eateries Mickel, New York Botanical Garden, Bronx, New York 10458, is editor of the sien omega Forum.”” The editor welcomes contributions from members and oe hembers, including miscellaneous notes, offers to exchange or purchase materials, personalia, hor- ticultural notes, and reviews of non-technical books on ee. : Library New York Botanical Garden, Bronx, New York 10458, is Librarian. Meare any time, the borrower paying all shipping costs. Spore Exchange

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AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 2 (1979) 33

A Community of Lycopodium Gametophytes in Michigan JAMES G. BRUCE* and JOSEPH M. BEITEL**

Since Fankhauser’s discovery of Lycopodium annotinum gametophytes in 1873, the sexual plants of Lycopodium have been sought repeatedly, and yet they remained elusive. There are now only 27 species for which gametophytes (19 subterranean, 8 surficial) have been found, representing fewer than 7% of the species.

The continuing rarity of gametophytes cannot be attributed to a lack of search- ers in temperate areas, although since most club-mosses are tropical it does point in part to the geographical separation between most botanists and most species of Lycopodium. But the gametophytes of even such common temperate species as L. tristachyum, L. digitatum, L. appressum, as well as those of the less common North American plants such as L. porophilum an L. prostratum, either never have been described or have been described so poorly as to render discussion almost meaningless. This lack of knowledge is due partly to the infrequent occurrence of the gametophytes and further to the fact that the more frequently encountered temperate species have subterranean gametophytes. Detection of the latter is dependent upon finding attached emergent sporophytes (Fig. 2). That they can be found and sometimes even in great numbers is amply illustrated by Spessard (1922) and Eames (1942).

In this paper we report finding a community of nearly 500 gametophytes in a Jack Pine plantation in Michigan representing five (or possibly six) species of Lycopodium: L. annotinum, L. clavatum, L. digitatum, L. lucidulum, and L. obscurum. Lycopodium dendroideum also may be present but is apparently indistinguishable gametophytically from L. obscurum. A key to these species is given below. Lycopodium digitatum has not been reported from nature previously (Wilce, 1965; Bruce, 1976) and its naturally occurring gametophytes are described here for the first time. Whittier (1977) has reported gametophytes of this species from culture.

KEY TO THE en ottec AND/OR YOUNG SPOROPHYTES F LYCOPODIUM SPECIES

—_

: Paraphyses (multicellular uniseriate sterile hairs around the sex organs) present, thallus generally Oerer ot branihed axigl ee aac ennoniassees L. lucidulum Paraphyses absent; thallus usually not axial, but if axial, then tapering and carrot- senghon Ss + Gametophytes carrot-shaped Pree oes Sas ictal eC 4 2. Gametophytes disc- ehiiake sometimes with convoluted edges, occasionally nearly Se com

3. Leaves of young sporophytes with long, bristle tipS ....-----++:eeessrrrrrrreert 3. Leaves of young sporophytes merely acuminate. : hel: of ak aie zis than 7 mm long; gametophytes nade co

nvoluted nnotinum

BS

and frag ile Pei cially ater ne ST Ee Ea LY ha a duty artnet me ametophytes coarsely convolu : Leaves a Se es — obsc ais Se ee um i Perret eS gh a ale

eens “Department of Botany, University of Georgia, Athens, GA 3

4giqyePartment of Botany, Division of Biological Sciences, watt a re)

Volume 69, number 1, of the JOURNAL was issued 29 Mar 1979.

f Michigan, Ann Arbor, MI

AMERICAN FERN JOURNAL: VOLUME 69 (1979)

FIGS. 1-3. Habitat and habit of gametophytes and young sporophytes of Lycopodium. FIG. 1. Jack Pine plantation with opening occupied by several Black Cherry trees. FIG. 2. Young ¢ ae sporophyte of L. obscurum. ca. XI FIG 3 Young sporophyte and gametophyte ( (white tuberous Structure embedded in sandy soil at end of sporophytic stem) of L. lucidulum, X1.5.

BRUCE & BEITEL: LYCOPODIUM GAMETOPHYTES 35

The subterranean gametophytes of other species that have been reported for North America are as follows: L. annotinum (Spessard, 1917), L. clavatum (De- gener, 1924; Gauthier & Dumais, 1938; Spessard, 1917; Stokey & Starr, 1924), L. complanatum (Degener, 1924; Eames, 1942; Spessard, 1917: Stokey & Starr, 1924), L. dendroideum (Ames, 1926; Degener, 1924; Spessard, 1917, 1922), L. lucidulum (Eames, 1942; Spessard, 1917, 1922), and L. obscurum (Eames, 1942; Dore, 1945; Gauthier & Dumais, 1938: Stokey & Starr, 1924).

TABLE 1. NUMBER OF LYCOPODIUM GAMETOPHYTES AND ATTACHED SPOROPHYTES AT THE JACK PINE PLANTATION SITE.

Total Number No. of Attached Sporophytes Species of Gametophytes 0 l >I

L. annotinum l ai | ma L. clavatum 70 19 26 25

L. digitatum 136 18 117 I L. obscurum/dendroideum 144 13 82 49 L. lucidulum 125 39 80 6 Totals: 476 89 306 81

The habitat of the gametophyte we studied is a shrubby JackPine (Pinus banksiana) plantation in Mecosta County, Michigan. The plantation was approx- imately 30 years old in 1977 as determined by tree corings. The trees are spaced 3—4 m apart in rows 4-5 m apart. Occasional irregular spacing apparently is due to the early death of some trees. Black Cherry (Prunus serotina) trees, the only other tree associate in the study area (Fig. 7), occur in some of the larger spaces. These trees may have persisted since before the pines were planted. The flora 1s depau- perate in understory species. The shrubby species include scattered Rubus idaeus and Comptonia peregrina. The herbaceous understory consists of Preridium aquilinum, Carex pensylvanica, Rumex acetosella, Hieracium sp., Monarda fis- fulosa, and Verbascum thapsus. Associated bryophytes are Polytrichum Juniperinum, Brachythecium curtum, Pleurozium — schreberi, Dicranum polysetum, D. scoparium, and Rhynchostegium serrulatum. The —. Were associated mainly with the more open areas, and particularly with the Blac Cherry trees.

The uneven forest floor consists of small knolls and depressions aie co €nce in elevation of approximately 15-20 cm. The gametophytes ees ie associated with the small depressions and the slopes leading into them. In e an the substrate is needle and leaf duff covering a humic layer 1-2 —— sot ‘urn overlies the sandy soil with some humic materials mixed in (Fig. 3). . . pa in this habitat have been classified as Coloma sands, a transitional soil . the true podsol soils in the northern part of the state and the gray-brown ape Soils farther south (Wildermuth & Fonder, 1931). The pH of this soil 1s 1.1.

36 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

BRUCE & BEITEL: LYCOPODIUM GAMETOPHYTES 37

The number of gametophytes found in each species is listed in Table 1. All of the gametophytes appeared to be quite old. Frequently they showed evidence of earlier sporophyte production. In addition, all that we found were relatively large (Figs. 4-6, 8, 10, 12, 14; Table 2); and, as shown by Bruchmann (1910), Lycopodium gametophytes are believed to grow slowly. Gametophytes and juvenile sporophytes of each species are briefly described below.

Lycopodium annotinum.—Only one gametophyte of L. annotinum was found. It was large (Fig. 6), highly convoluted, and relatively fragile. Gametophytes of what Bruchmann (1898) called Type I, including in this paper L. annotinum, L. clavatum, L. dendroideum, and L. obscurum, all assume a disc shape relatively soon after germination. Then, due to a poorly understood growth mechanism, the edge of the disc convolutes and the gametophyte finally resembles a flat walnut meat. Continuation of this convoluted growth yields the essentially spherical or ovoid masses seen in Figs. 4-6, 8, and 10. The juvenile sporophytes of L. an- notinum are characterized by fairly elongate leaves (Fig. 7) and, in those older than one year, the annular constrictions for which the species is named become readily apparent.

TABLE 2. MAXIMUM DIAMETERS (AND LENGTHS IN L. FLABELLIFORME) OF LYCOPODIUM GAMETOPHYTES IN mm.

Species N! x Range S.D. L. annotinum 1 1 19 ibe L. clavatu 48 11.2 6-22 3.6 L. dendroideum/obscurum 90 14.2 7-26 ag L. digitatum 117 i 2.5-13 1.8

"Apparent discrepancies between the number of gametophytes analyzed in this Lawley — reported in Table 1 result from omissions in the analysis of injured or broken gametophytes. *Values for gametophyte length.

Lycopodium clavatum.—Numerous gametophytes and alonihi snonieiely 4 clavatum were found (Table 1). The gametophytes were similar . ane annotinum in being highly convoluted and relatively fragile (Fig. 8). za gametophytes had multiple sporophytes; indeed, one had is evo sporophytes! One albino sporophyte ca. 4. cm long, emergent, and tota é is of chlorophyll was found (Fig. 4). The juvenile sporophytes of L. clava i . easily recognized by the characteristic long hair-tip on their leaves, a aes found in the adult sporophytes (Fig. 9).

oe btivte of L. annotinum, x3. FIG. 7. Young — cei ametophyte of L. clavatum, x3. FIG. 9. Young sporophyte of L. clavarum, hytes, x 2. leaves. FIG. 10. Gametophyte of L. obscurum/dendroideum with four attached sporophytes, FIG. 11, Young sporophyte of L. obscurum/dendroideum, x4.5.

AMERICAN FERN JOURNAL: VOLUME 69 (1979)

BRUCE & BEITEL: LYCOPODIUM GAMETOPHYTES 39

Lycopodium obscurum and L. dendroideum.—The gametophytes of L. den- droideum and L. obscurum are here treated together. Although the adult sporophytes of these species are readily distinguishable (Hickey, 1977), we could not distinguish their gametophytes or juvenile sporophytes. Adult sporophytes of both species occurred within the locality, but L. obscurum was by far the more common. The gametophytes were similar to those of the two preceding species in that they were convoluted (Fig. 10), but the convolutions were very coarse and the gametophytes, in contrast to those of L. annotinum and L. clavatum, were firm and sturdy.

The young sporophytes’ leaves lacked the bristle tip of those of L. clavatum and generally were shorter than those of L. annotinum. However, the young sporo- phytes were surprisingly similar to those of L. digitatum (cf. Figs. 11 and 13).

Lycopodium obscurum/dendroideum yielded the most abundant gametophytes and young sporophytes (Table 1). Multiple sporophytes were common, as in L. clavatum, and in one case ten sporophytes were attached to a single gametophyte. Also as in L. clavatum, one albino sporophyte was found (Fig. 5). It was emer- gent, ca. 6.5 cm long, and totally devoid of chlorophyll.

Lycopodium digitatum.—Numerous specimens of L. digitatum' gametophytes and young sporophytes were found (Table 1). The distinctive carrot shape of the gametophytes (Fig. /2) makes them readily identifiable. All intermediate stages in the development of sporophytes from very young with little or no chlorophyllous tissue to those exhibiting the characters of the adult flat-branched sporophytes were present. Although these plants could have been the similar L. complanatum, no sporophytes of that species were seen at the locality, and the nearest known locality is well over 100 miles north of the site. We cannot rule out the possibility that some of the carrot-shaped gametophytes without attached sporophytes might have been L. tristachyum, but we consider this to be unlikely. Only one unat- tached, possibly three- or four-year old sporophyte plant of the closely related L. fristachyum was seen. Its distinctive, bluish-green color was in conspicuous con- trast to the darker green of L. digitatum. :

Some L. digitatum gametophytes appeared to have been partly eaten (Fig. 16). Most gametophytes had only a solitary attached sporophyte, although one sp ge men had five. Orientation of the gametophyte was apparently random an seemingly did not affect sporophyte production.

ae. ‘Hickey and Beitel (1979 i ‘ igi A. Braun is the correct name for

: ) recently determined that L. digitatum A. ; *pecies formerly called L. fabelitotme (Fern.) Blanch. or L. complanatum vat. flabelliforme Fern

FIGS. 12-17. Gametophytes and young sporophytes of Lycopodium. FIG. 12. —— .

digitatum, x4. FIG. 13. Young sporophyte of L. digitatum, XS. FIG. AM pigeeh a He 16.

lucidulum, dorsal surface, x5. FIG. 15. Young sporophyte oF i Gann x ane pe ametophyte of L. digitatum damaged by predator. Axis of gametophyte oriented mune aun

a8€ on flank below meristem which is at left, x 16.5. FIG. 17. Gametophyte of L. ag

Circular scar Showing old connection of foot to main axis of sporophyte, x25.

40 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

Young sporophytes were essentially indistinguishable from those of L. obscurum/dendroideum (cf. Figs. 11 and 13). However, as soon as the young sporophytes began to branch, phyllotactic changes took place that culminated in plants with essentially adult foliage, going from spiral to falsely opposite.

Lycopodium lucidulum.—The gametophytes of L. Jucidulum, although generally axially elongated, had definite upper and lower surfaces (Fig. 14). A peculiar feature of these gametophytes was that all 125 of them came from a small square of ground roughly 45 cm ona side. Furthermore, these gametophytes occurred in an upland, well drained site, whereas the nearest adult sporophytes were in a white cedar bog approximately one half mile away. The gametophytes were particularly distinctive at the microscopic level because of the presence of multicellular, uni- seriate paraphyses associated with the gametangia. The young sporophytes had rather boat shaped leaves with minutely toothed margins (Fig. 15).

GAMETOPHYTE SIZE AND LONGEVITY

One striking characteristic of the gametophyte community in the JackPine woods was the total absence of reproductively mature sporophytes. The habitat is several acres in size and is bordered by areas which do possess mature sporo- phytes.

Coincident with the lack of mature sporophytes was the apparent old age of the gametophytes. This observation is inductive and is based on the large size of the gametophytes, their presumed slow rate of growth, and the evidence of previous sporophyte production (Fig. 17). The assessment of old age is bolstered by Bruchmann’s observations (1910) on European species with subterranean gametophytes in which he was able to ascertain ages of 3-8 years preceding reproductive maturity. Our observations over a four year period show that the gametophytes produce numerous sporophytes. These results suggest premature death of the young sporophytes, more of which are produced each year. Based on long observations, Eames (1942) allowed ten years for gametophyte production in the habitats he studied. Yet, the presumed old age of the gametophytes suggests

One further observation of relevance here is the fact that no small gametophytes were found (Table 2). In other sites that we have investigated, numerous gametophytes have been recovered that were not only producing young sporophytes, but were all smaller than any of the gametophytes at the present site.

These observations suggest that current conditions are not favorable for spore arrival and/or germinatio

theless, the present conditions are ideal for continued gametophyte growth and

The habitat is succe

Spessard (1917) and Eames (1942) noted that successional habitats are the most frequent kind of Site

BRUCE & BEITEL: LYCOPODIUM GAMETOPHYTES 41

Gametophyte clustering was seen for all species studied here, with the excep- tion of L. annotinum, which was found in numbers too small to be relevant. The most spectacular case was L. lucidulum, with all 125 gametophytes from a square 45 cm on a side. Spessard (1922) noted similar occurrences of L. lucidulum, as well as less spectacular finds of other species. The clustering of gametophytes has also been noted in Botrychium (W. H. Wagner, U. of Michigan, pers. comm.). While this is regarded as a feature of special note, it should be remembered that pteridophytes with surficial gametophytes also illustrate striking clustering. How- ever, in the case of the surficial gametophytes, the microenvironmental features controlling population size and distribution, such as moisture, protection, and shading, frequently are more obvious. It seems reasonable that microenvironmen- tal features might also control population distribution in subterranean gameto- phytes as well. However, in the latter case, the environmental features are much less obvious, such as distribution of mycorrhizal fungi.

We wish to express our appreciation to W. H. Wagner, Jr., for help and support during portions of this study, to William R. Buck for verifying the identifications of the Bryophytes, to Randy O. Wayne for numerical analysis of the gametophytes of L. digitatum, and to Ellen and Lee Weatherbee without whom we would never have made this discovery.

LITERATURE CITED

AMES, R. S. 1926. Another station for Lycopodium prothallia. Emer. Fern J. 16: 26. BRUCE, J. G. 1976. Gametophytes and subgeneric concepts in Lycopodium. pees J. Bot. 63:919-

924. BRUCHMANN, H. 1898. Uber die Prothallien und die Keimpflanzen mehrerer europaischen Lycopodien, und zwar iiber die von Lycopodium clavatum, L. annotinum und L. selago. F. A. Perthes, Gotha. ‘ ie Die Keimung der Sporen und die Entwicklung der Prothallien von Lycopodium clavatum L., L. annotinum L. und L. selago L. Flora 101:220-267. DEGENER, 0. 1924. Four new stations of Lycopodium prothallia. Bot. Gaz. 77: 89-95. DORE, W. G. 1945. Site for club-moss prothallia in Nova Scotia. Canadian Field-Nat. 59:172-173. EAMES, A. J. 1942. Illustrations of some Lycopodium gametophytes. Amer. ap ae 32:1-12. FANKHAUSER, J. 1873. Ueber den Vorkeim von Lycopodium. Bot. Zeit. 31: GAUTHIER, R., and R. DUMAIS. 1938. Les prothalles de Lycopodes . = Québec. Nat. Canadien 65:280-284. ; HICKEY, R. J. 1977. The Lycopodium obscurum complex in North America. Amer. Fern J. 67:45- 48.

Lycopodium flabelliforme. Rhodora

ae J. M. BEITEL. 1979. A name change for —140

SPESSAR p, ESA: 1917. Prothallia of Lycopodium in America. Bot. Gaz. 63:66-76. :

1922. Prothallia of Lycopodium in America. II. L. lucidulum and L. obscurum var. den

droideum, Bot. Gaz. 74:392-413.

STOKEY, A. G. and A. M. STARR. 1924. Lycopodium prothallia in western Massachusetts. Bot.

Gaz. 77:80-88. She WH La D. P. 1977. Gametophytes of Lycopodium complanatum in axenic culture. Bot.

Amer. Misc. Ser. Publ. 154:56 (abstract). WILCE, J. ae Section complanata of the genus WILDERMUTH. R., and J. F. FONDER. 1931. U.S.D.A. Soil tes. Rep. 1927(18):1-3.

Lycopodium. Nova Hedwigia 19:1-233, ¢ I-XL. Soil survey of Mescosta County, ! eau

42 AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 2 (1979)

Acrostichum in Florida DAVID C. ADAMS® and P. B. TOMLINSON**

This article is the result of field study of several Acrostichum aureum L. and A. danaeifolium Langsd. & Fisch. populations in the swamps of South Florida. It was made with the object of establishing consistently applicable diagnostic differ- ences between these oftimes confused species. Our observations will facilitate field identification, and we also have found some pertinent ecological differences between the two species.

No attempt has been made to resolve the possibly more complex problem of Acrostichum in the Asian tropics, where in addition to two distinct and widely distributed species (Troll, 1933), there may be locally differentiated forms which have remained insufficiently examined, as in New Caledonia.

The differential characteristics which our study has elucidated are summarized in the following key:

Axis usually creeping, relatively frequently branched. Fertile fronds with only the upper pinnae (up to 5 pairs and the terminal pinna) fertile (Fig. /A). Pinnae few, usually not more than 30, rather distant and often irregularly distributed, usually not overlapping, the lowest pinnae always distant, long-stalked (up to 3 cm) (Fig. /C). Areoles next to the midrib narrow, always 3 times longer than wide. Rachis rounded below, decidedly grooved above with the margin of the groove acute, and with short spines (the midribs of aborted pinnae) frequent on the lower part of the rachis (Figs. 1B, D). Basal scales not leaving prominent scars (Fig. /E). Paraphyses with a slender stalk, ending ina single, + isodiametric, irregularly much-lobed cell (Fig. / F).

: A. aureum

Axis usually erect, relatively infrequently branched. Fertile fronds with all or most of the pinnae ertile, rarely only a few upper ones fertile (Figs. 2G, H). Pinnae many, usually 40-60, closely set, overlapping and regularly arranged (Fig. 2A), often subopposite, the lowest pinnae relatively short-stalked (less than 2 cm) (Fig. 2E). Areoles next to the midrib broad, never more than 3 times longer than wide (Fig. 2D). Rachis with several shallow grooves below, flat or scarcely grooved above, with the margins of the groove blunt (Fig. 2E), and with basal spines or aborted pinnae absent (Figs. 2B, H). Basal scales leaving prominent scars (Fig. 2F). Paraphyses with the stalk ending in a horizontally extended, smooth or little-lobed cell (Fig. 2/) . A. danaeifolium

The best characters’ for distinguishing the species are found in features of the whole frond and in the difference in distribution of fertile pinnae. These differ- ences and general features of ecology were well appreciated by Small (1931). Jenman (1909) also gave a good account of the two species, and many contrasting features are clearly recognized by Garcia de Lopez (1978).

The marginal spines of A. aureum are very distinctive when present. Paraphy- ses provide an excellent diagnostic feature for fragmentary fertile herbarium specimens. Neither areole size and overall orientation nor leaflet shape seem consistently useful, although these characters were emphasized both by Small (1931, p. 237) and by Wherry (1964), except for the difference in length of areoles

*P. O. Box 155, Teton Village, WY 83025. Harvard University, Harvard Forest, Petersham, MA 01366.

ADAMS & TOMLINSON: ACROSTICHUM IN FLORIDA 43

joy Fae

at

d. FIG. 1C. FIG. 1. Acrostichum aureum. FIG. 1A. Upper part of frond. FIG. a gion SS aan FIG. Middle of frond showing pinnule attachment. FIG. 1D. Part of leaf axis with a marginal : IE. Leaf base with adventitious roots and scales. FIG. IF. Paraphyses.

AMERICAN FERN JOURNAL: VOLUME 69 (1979)

Ss

ES

Ee Be

FIG. 2. Acrostichum dan sterile frond. FIG. 2

aeifolium. FIG. 2A. Upper part of sterile frond. FIG. 2B. Unarmed pps se - Single pinna. FIG. 2D. Areoles in midrib region. FIG. 2E. Middle o Fea Showing pinnule attachment. FIG. 2F. Base of leaf axis with persistent scale scars. FIG. 2G. UP part of fertile frond. FIG. 2H. Lower part of fertile frond. FIG. 21. Paraphyses.

ADAMS & TOMLINSON: ACROSTICHUM IN FLORIDA 45

next to the midrib. The leaflets in A. aureum seem somewhat more thickly coriaceous than in A. danaeifolium, but textural differences are difficult to assess in herbarium specimens. There are no immediately obvious differences in sporan- gium size or spore diameter. The SEM photographs of Garcia de Lopez (1978) indicate that the wall sculpturing of the spores also is identical.

Leaf dimensions are variable, but A. aureum seems to be the smaller species. In the populations examined, the more erect axis of A. danaeifolium leads to a closer aggregation of leaves, which seems consistent but requires verification in other parts of the species’ range. In this respect, A. aureum seems better adapted to vegetative persistence because of its more frequent branching.

° ie ie \ —~

FIG. 3. Distribution of Acrostichum in South Florida based on specimens in the Harvard University Herbaria and field observations.

In South Florida, A. aureum is the less common species and is seemingly restricted to the saline muck of back-mangrove communities, as suggested by its coastal distribution (Fig. 3). Acrostichum danaeifolium is the more common species and occurs both in saline substrates of back-mangroves and also extends inland to fresh-water swamps, commonly in sink-holes in hammocks. It also is a plant of disturbed marl sites and has weedy tendencies. The greater range (Fig. 3) and wider edaphic tolerance of this species may be due in part to - greater resistance, which in turn may be related to its more condensed crown 0 een

In our experience, both species, but especially A. danaeifolium, have a sb tolerance for light conditions but seem most vigorous in full sun. They ak oc : together at the same site, but in our experience show no evidence of shot Gi However, Garcia de Lopez (1978) mentions intermediate forms which she sae €rs to be hybrids in the Dominican Republic. Sown spores of both gua wel nate readily and produce sporelings without manipulation. Since the se cepha 2 Parts are large and easily studied microscopically, Acrostichum wou €xcellent subject for studies of the fern life cycle.

46 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

This study was initiated as a class project in the *‘ Plants of the Tropics’’ biology class taught at Fairchild Tropical Garden in 1976, with support from the Atkins Garden Fund of Harvard University. We are indebted to Don Evans, George Avery, and Rolla Tryon for comments and field assistance and to the Director, Gray Herbarium of Harvard University, for access to the collections.

LITERATURE CITED

GARCIA de LOPEZ, IVONNE. 1978. Revision del género Acrostichum en la Reptblica eS as Moscosoa 1:64—70.

JENMAN, G. S. The Ferns and Ferm-allies of the British West Indies and Guiana. Gov't. Printing Sia Port-of-Spain, Trinidad.

SMALL, J. K. 1931. Ferns of Florida. Science Press, New York.

TROLL, Pie ee Botanische Mitteilungen aus den Tropen VIII. Uber Se aureum L.,

ichum speciosum Willd. und neotone Formen des letzteren. Flora, n.f. 28:301—328. Winey, £ en 1964. The Southern Fern Guide. Doubleday, Garden City, NY

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AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 2 (1979) 47

Phyllitis scolopendrium Newly Discovered in Alabama JOHN W. SHORT*

Long known as one of the rarest ferns in North America, the American Hart’s- tongue, Phyllitis scolopendrium var. americanum Fern., has been reported from New Brunswick, Ontario, New York, Michigan, and Tennessee (Wherry, 1961). It has been reported from two localities in Tennessee (Shaver, 1954). The first plants were found at a cave mouth in Roan County in 1849, but they had disap- peared by 1900. The second site, discovered in 1878 a few miles north of Bridgeport, Alabama, in a lime sink in Marion County, Tennessee, contained plants which are surviving but are underdeveloped. The Hart’s-tongue has not been found elsewhere in the southeastern United States until now.

In early October 1978, E. Batchelder, a co-worker of the author and an avid spelunker, told of a caving expedition to Jackson County, Alabama, the previous weekend. While entering a cave through a deep sink, he saw an unusual-looking fern and photographed it. Two weeks later the photograph had been developed, and the fern proved to be P. scolopendrium.

The site was visited on October 21 by a group consisting of the author, Mr. Batchelder, C. Batchelder, J. Shearon, M. Shearon, and D. Gazaway. The sink is about a mile from the village of Paint Rock on the western flank of Nat Mountain at about 1200 feet elevation. The sink is about 40 feet deep and its walls are vertical or overhanging, except for a narrow cleft through which passes a path so steep that a rope is necessary to enter the sink. There are also two cave mouths at the bottom.

A small but thriving colony of P. scolopendrium was found at the bottom of this sink. Twenty plants were observed on this and subsequent visits. During a second visit on November 24, it was found that one of the plants had been destroyed by some person who apparently had slid off the main path to one of the caves and slid down the steep slope where the ferns were growing. Athyrium pycnocarpon, Asplenium rhizophyllum, and Cystopteris tennesseensis were also found in this sink.

Eight of the Hart’s-tongue plants were mature adults, including a few which were quite robust. The rest were juveniles of various apparent ages. The ferns Were growing on rocks or in soft, black soil. One adult was on a ledge a few feet above the sink bottom. The Phyllitis plants were photographed and a number of leaves were collected as vouchers (Short 1187). These specimens are being tributed to various herbaria, including AUA, MICH, TENN, UNA, US, an VDB.

On the slopes of the mountain and in a nearby sink were found a ngs “4 other fern species, including Adiantum pedatum, Asplenium platyneuron, ae liens, Pellaea atropurpurea, Polypodium polypodioides, Polystichum acros ichoides, and Woodsia obtusa.

*4309-B Boxwood Ct., Huntsville, AL 35805.

48 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

Nat Mountain is about 40 miles southwest of the Marion County, Tennessee, locality in a large area of rugged terrain formed by stream dissection of the Cum- berland Plateau. The mountains, actually mesas, rise to elevations of 1600-1700 feet, about 1000 feet above the surrounding flat-bottomed valleys. The slopes are limestone with thin sandstone caps. This heavily forested, largely inaccessible area comprising the section of Alabama east of Huntsville and north of the Ten- nessee River has been little explored by botanists, even though some nearby areas have received considerable attention. It would seem that P. scolopendrium may be present at other suitable locations in this area. Indeed, E. T. Varnedoe, another experienced spelunker, saw Mr. Batchelder’s photograph and was of the opinion that she had seen the fern in several such places in Jackson and Madison Counties.

LITERATURE CITED SHAVER, J. M. 1954. Ferns of Tennessee. Bureau of Publications, Geo. Peabody College for

Teachers, Nashville, TN. WHERRY, E. T. 1961. The Fern Guide. Doubleday, Garden City, NY.

REVIEW

*“SYNAPTOSPORY: A HYPOTHESIS,” by K. U. Kramer, Gard. Bull. 30:79- 83. 1977.—It has been known for a long time that certain pteridophytes shed their spores in small groups or in tetrads and that others disperse whole sporangia, some with the spores already germinating inside. Not much thought has been given to the biological or evolutionary importance of this phenomenon, which Kramer has termed synaptospory.

Kramer suggests that the function of sculptured perispores is to increase the chance of synaptospory. This makes it likely that gametophytes will grow close together and that inter-gametophytic selfing will occur. Such cross-fertilization tends to maintain genetic diversity and to reduce the expression of recessive deleterious genes.

Synaptospory tends to be found in terrestrial ferns, which are long-lived and grow mostly in extensive, stable habitats. Dispersal of such species at a distance is of relatively little consequence to their evolutionary success because of their stable habitats.

Epiphytic ferns, on the other hand, commonly lack a visible perispore and do not exhibit synaptospory. (The perispore of these ferns is not really lacking, but rather is tightly adherent to the exine, as demonstrated by electron microscopy-) The habitat of epiphytic ferns is minute, short-lived, and unstable. Epiphytic ferns tend to occupy their immediate habitat by means of creeping rhizomes. But their need over the long term is to spread to similar but non-adjacent habitats, and single-spore dispersal accomplishes this better than synaptospory.—D. B. L.

AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 2 (1979) 49

Studies in Lycopodiaceae, II. The Branching Patterns and Infrageneric Groups of Lycopodium sensu lato B. OLLGAARD*

Several contributions of Lycopodium taxonomy on the infrageneric level have been made recently (Wilce, 1965, 1972; Chu, 1974; Ollgaard, 1975: Bruce, 1976a, b, c). Earlier literature on the classification of this group was summarized by Wilce (1965) and Bruce (1976b) and need not be repeated here. The present study includes new observations on branching patterns and attempts to correlate these with infrageneric classifications of Lycopodium, about which there has been some disagreement. Only Troll (1937, pp. 465-482) has dealt with the subject at some length considering the genus as a whole; other students (Holloway, 1916, 1919; Wilce, 1965; Primack, 1973) have treated smaller groups in greater detail.

MATERIALS

The present study is based on field observations made in various parts of Europe and during two expeditions to Ecuador in 1973 and 1976 (Holm- Nielsen et al., 1975; Ollgaard & Balslev, 1979). In addition, herbarium specimens represent- ing a large number of species, mainly from Ecuador, have been studied from several herbaria (A, B, BM, CAS, DS, E, F, G, GB, GH, K, MSC, NY, OXF, P, S, UC, and US) in preparing a treatment of the Lycopodiaceae for the ** Flora of Ecuador.’’ Material from other parts of the world has mainly been studied in the Botanical Museum, Copenhagen (C), the Herbarium Jutlandicum (AA U), and the Museum of Natural History, Stockholm (S). I am indebted to the curators of the institutions mentioned above for permission to study the specimens in their care. Special thanks are due to Dr. D. R. Given, Christchurch, New Zealand, for material of Lycopodium laterale R. Br., and to Mr. John Woodhams, Royal Botanic Gardens, Kew, for a specimen of L. nummularifolium Blume. I wish to express my gratitude to the Danish Natural Science Research Council for support for my field work in Ecuador in 1976.

In the following text, the infrageneric groups correspond to those of Wilce (1972), unless otherwise stated. For nomenclatural reasons, some of the names used by Wilce have been changed: Subg. Urostachya Pritzel is the original spell- ing, and should be retained, not Urostachys.' Subgenus Lycopodiella (Holub) Ollgaard? comprises the species included in subg. Lepidotis (Palisot) Baker by Wilce. Since the name Lepidotis is a nomen confusum, as stated by Holub (1964), the generic name Lycopodiella is here made available at the subgeneric level.

“Botanical Institute, University of Aarhus, 68 Nordlandsvej, DK-8240 Lema ‘aaa subg : pet eR alt ycope ‘ ‘Herter (Bot. Jahrb. Engler 43, Beibl. 98:5, 29, 30. 190%) gry eltbctivaly made a new genus s.—Ed. 21 ycopodium subg. Lycopodiella (Holub) B. Oligaard, comb. nov. Basionym: Lycopodiella Holu Preslia 36:20, 22. 1964.

50 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

BRANCHING PATTERNS IN LYCOPODIUM

The genus Lycopodium commonly is used as an illustration of dichotomous branching in vascular plants. Indeed, dichotomy is the basic mode of branching in this genus, although lateral branching is a regular event in at least one species and occurs less regularly in others. Trichotomy, which is similar to dichotomy but results in three branches, occurs more randomly.

Usually the descriptions of the vegetative body of Lycopodium apply a number of terms which were originally coined for plants with lateral branching, such as sympodial and monopodial branching, main and lateral branches, or principal and secondary branches. Such terms are in principle inaccurate or incorrect for plants in which minor and major branches must be considered ontogenetically equiva- lent. However, the establishment of a terminology which is both accurate, cor- rect, and practical is hardly possible in a genus with such diverse and plastic patterns. In this paper, I use the term main to indicate a function of an axis, but avoid the terms principal, secondary, and lateral, except where they are properly applied in cases of true lateral branching.

The variation in Lycopodiaceae branching patterns is mainly due to three vari- able characters, as enumerated below.

Relative size of branches of dichotomies.—The branches of a dichotomy may have equal or unequal diameters. The terms isotomy and anisotomy for these conditions were introduced by Troll (1937, pp. 468-469). Since equal branch diameter is the sole criterion for isotomy, it follows that isotomous branches may differ in length, function, or both. Correspondingly, anisotomy was defined by Troll as dichotomy leading to unequally thick branches resulting from unequal branch primordia. A precise distinction between anisotomy and isotomy some- times is difficult, although the smaller branch of an anisotomy usually is recog- nized by its reduction in both diameter and length.

A general feature of anisotomous species is the formation of one or more main axes functioning as rhizomes or aerial main stems. This is achieved through the regular alternation of major and minor branch primordium positions in successive dichotomies. This is the dichopodial branching of Bock (1962); more commonly it is termed pseudomonopodial branching. Overtopping in the major branch and development of the minor branches into shorter and more or less determinate branches or branchlet systems complete the development.

Orientation of successive branching planes.—Branchings may occur in planes successively perpendicular to one another, in a single plane, or may form other angles in relation to each other. Following Troll (1937, p. 474), the first two conditions are termed cruciate and flabellate, and the third is here termed incli- nate. Many species display combinations of these conditions, often in fixed se- quences In connection with certain events in their development.

Observations of these characters are made with some difficulty. Herbarium material is often inadequate because of distortion as a result of pressing. Living Soe also may be distorted. The so-called erect species usually are erect only

en quite young. Later they overturn under the increasing weight of the plant,

B. OLLGAARD: STUDIES IN LYCOPODIACEAE, II. 5)

the stem base becomes decumbent, and the branch apices turn upwards. Ih specimens with long, lax, pendant stems, the orientation is affected by contact with adjacent stems. In both cases, precise observation of branch orientation is complicated or impossible, but often it is possible to find short sequences of branches which are characteristic.

Functional differentiation.—The branches of a dichotomy may be completely similar in function, or a branch-pair may differentiate into one vegetative and one fertile branch, into one rhizomatous and one aerial branch, and so on.

SUBGENUS UROSTACHYA

This group contains probably more than 400 mainly tropical, entirely isotomous species. About a third are terrestrial; the others are mostly erect or pendant epiphytes.

It has been argued that the bulbils formed in L. selago L., L. lucidulum Michx., etc. are examples of extreme anisotomous branching (Bierhorst, 1971, p. 9). But according to other views (reviewed in Cutter, 1966), they are foliar in origin. The capacity to form bulbils may be a character of taxonomic significance in subg. Urostachya, but it is not further considered in this study.

In the majority of species, branch length and development is almost identical. This is especially true for a great number of terrestrial species. Many pendant epiphytes vary with respect to branch length and corresponding branching inter- vals. This probably is an adaptation of individual branches to different light inten- Sities in large plants, which may develop hundreds of branches from a single basal stem. However, the branch diameters are the same in corresponding long and short branch intervals of such individuals.

The orientation of successive branchings in a small whole plant of the terrestrial L. selago is illustrated diagrammatically in Fig. / . The diagram indicates that most branchings occur in planes inclinate to the preceding branching plane. Cruciate sequences are frequent, mostly in the younger parts, but they hardly ever occur In both branches of a pair; one usually is inclinate and the other perpendicular to the preceding branching plane. In the living plants, the branches usually are more or less ascending, and the inclination of branching planes tends to result in a concen- tric and concave arrangement of branches, like portions of funnels inserted se each other. Such an arrangement probably catches light more ehechively co regular cruciate branch systems would. The orientation of successive branc | ing planes is variable in detail, but in all Ecuadorean species of subg. Urostachya tt generally corresponds to the pattern found in L. selago. :

elena ofl sanoutinien te , a species which was described 7 Lim (1937, p. 476, fig. 36611) as flabellate, was found not to be so. It has both fla ° et and cruciate sequences (Fig. 2). The flattening plane of the whole shoot is ae leaf orientation and does not coincide with the branching planes. In the diagram, the longest diameter of the ellipses indicates the leaf plane

In most species of subg. Urostachya, the branches devel with minor variations of length. In the L. phlegmaria group, the Contraction (change from expanded leaves to reduced, claspin

op identically, usually the position of branch g, and appressed

52 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

leaves) is variable in individual branches. The contracted branches in this group are very frequently termed strobili, and the leaves sporophylls. This usage is largely inadequate, as the constricted branches usually are only partially sporan- giferous. Furthermore, sporangia often are found in the axils of expanded leaves.

The L. saururus group, most of which are high- Andean species, has differentia- tion of the branches resulting in the development of horizontal rhizomatous and erect aerial branches (Fig. 5). The rhizomatous branches produce roots along

FIGS. 1-3. Branching patterns in Lycopodium. F1G. 1. Orientation of successive branchings of a whole plant of L. selago. Godhavn. Greenland, Laegaard s.n. (AAU). FIG. 2. Orientation of succes- sive branchings of L. nummularifolium. The longest diamater of the ellipses indicates the shoot flattening plane. New Hebrides, Braithwaite s.n. cult. Kew. FIG. 3. Generalized diagram of the rhizome branching in subg. Lycopodium.

their underside and may be either superficial or subterranean. In a common yet undescribed species from Ecuador, the production of elongate, subterranean rhizomatous branches enables it to form relatively large clones and to be rather successful in the lower paramo bogs (Fig. 4). Apparently this type of branch differentiation does not represent a stable character in all the species in which it occurs; In some of them it is occasional or even rare.

Adventitious shoots may be formed by lateral branching at the base of senile or ee plants in some terrestrial or epiphytic species, and so may rejuvenate

B. OLLGAARD: STUDIES IN LYCOPODIACEAE, II.

s subterranean,

- /

Sl »y 8378

& Balslev 8

Ecuador, Qllgaard & Balsle

FIG. 4. Lycopodium sp. with it

Pcie

es in the L. crassum group.

FIGS. 4-5. rhizomatous

Qligaard

Ecuador

ered, Paramo de Tufino, Pcia. Carchi, ia. Carchi,

crassum 5.1. Paramo de Tufino

Habit of speci s branches uncov

(AAU). FIG. 5. L.

(AAU).

,

54 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

SUBGENUS LYCOPODIELLA

This subgenus includes about 40 species, all terrestrial. They can be divided into three groups discussed below on the basis of habit, including branching pat- tern. In contrast to subg. Urostachya, the species are all basically anisotomous.

Group of L. inundatum and L. carolinianum.—In fertile plants, the first branch- ing from previously dormant buds of L. inundatum L. takes place in the vertical plane and produces an erect, strobilus-bearing branch and a creeping and rooting vegetative branch (Fig. 8). Sterile individuals branch only in the horizontal plane. The erect branch usually remains simple, but approximately isotomous branch- ings may occur as abnormalities. In the large Brasilian plant here tentatively named L. alopecuroides var. furcatum Fée, the erect branch normally is up to three times branched (Fig. 7). The orientation of these branchings is unknown. In several species, the creeping branch may produce up to several erect, strobilus- bearing branches by branching in the vertical plane, whereas all branchings in the horizontal plane result in the formation of creeping, vegetative branches. No inclinate branchings have been found in this group.

Group of L. cernuum.—This group consists of profusely branched plants, some of which are reminiscent of small Christmas trees. Although they have a common basic pattern, their branching patterns are exceedingly variable, even within one species. The widespread L. cernuum L. can serve as a model for the group (Fig. /1). Young plants of this species produce an arching or looping main stem that roots upon soil contact and forms a new loop. This may be repeated until the plant is sufficiently strong to branch in the vertical plane, producing an erect branch on the upper side of the loop and a continuing looping branch. In theory, this may continue indefinitely.

The branching pattern of the erect branch of L. cernuum was excellently de- scribed by Troll (1937, pp. 477-478). The erect branch has a distinct pseudomonopodial main axis which may bear a few arrested buds below the subopposite pairs of horizontal flabellate branchlet systems. The pairs of branch- let systems alternate regularly by ca. 90°, creating an illusion of decussate ar- rangement. The branch pairs arise by two anisotomies in rapid succession in approximately the same plane. This characteristic difference of branching interval length in the main axis is gradually lost towards the apex of the erect branch. The flabellate branchlet Systems ultimately may form small, drooping strobili.

In addition to the erect branches just described, the continuous looping stems produce flabellate anisotomous branchlet systems in the horizontal plane, and sometimes divide nearly isotomously, thus forming a new continuing looping stem. Sometimes the branchlet systems borne on the looping stem may forma few strobili. Sometimes one or two of the lowest branchlets on the erect branch, which mostly remain as arrested buds. develop into new looping stems.

Other taxa of the group differ from L. cernuum in branching pattern, mainly with respect to the length of the looping branches and the compoundness and branching sequence of the erect branch.

B. OLLGAARD: STUDIES IN LYCOPODIACEAE, II. 55

FIGS. 6-11. Branching in Lycopodium. Rhizome branching in L. complanatum. The xylem (a ay lateral marks indicate the horizontal plane. Pcia. Loja, Ecuador, Qllgaard s. n. (AAL ). FIG. ‘ dlopecuroides var. furcatum. Sao Paulo, Brazil, Brade 5131 (S). FIG. 8. L. inundatum. ee Nordstrand, Denmark, Qllgaard s. n. (AAU). FIG. 9. Subterranean lateral branching in t. ei fe New Zealand, Given 8837 (AAU). FIG. 10. Aerial branching in L. laterale.. Charleston, Ww. te a New Zealand, Wardle s. n. (AAU). FIG. 11. L. cernuum. Above Balzapamba, Pcia. bollvar, Ecuador. Ollgaard photo.

56 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

Group of L. laterale.—This small group is found mainly in Australia and New Zealand. I have had access only to material of L. ramulosum Kirk and L. laterale. The branching pattern of L. ramulosum strongly resembles that of L. inundatum, one difference being that the erect, strobiliferous branch is not produced at the first branching, but follows after a sequence of nearly isotomous, horizontal, and flabellate branchings of vegetative stems. I have seen no subterranean parts of this species.

Holloway (1916, p. 255) described the subterranean branching of L. laterale (Fig. 10) by saying that ‘‘The adult plant consists of an irregular and much branched colourless rhizome, which ramifies through the soil in all directions. The shorter branches emerge at the surface to form the erect aerial shoots.’’ The fresh sterile material I have studied of this species has subterranean, ascending rhizomatous stems which branch a few times nearly isotomously in the horizontal plane when they emerge on the surface, almost like in L. ramulosum. No further dichotomous development was found in the sterile material. However, the subter- ranean branching indicated by Holloway is due to true lateral branching. Lateral branches and dichotomous ones as well are formed from superficial and subterra- nean stems. In the subterranean stems, they usually arise from the undersides of stems in a way and position comparable to roots, breaking through the stem cortex (Fig. 9). The anatomy of these branchings remains to be studied more closely. In this connection, it is interesting to note Holloway’s (1916, p. 299) remark that “The more deeply growing almost naked rhizome of L. ramulosum is interesting as Suggesting an intermediate form between the typical scaly rhizome and the leafless adventitious root.’’ The secondary branches may behave like the primary branch, giving rise to tertiary branches, etc.

Erect, strobilus-bearing shoots arise similarly, but ascend more steeply to the surface. Their initial aerial development resembles that of sterile shoots, being flabellate and nearly isotomous. However, after a few dichotomies, orientation of the branching planes becomes irregular. Only two herbarium specimens have been boiled for examination of this character. They gave no idea of a fixed se- quence (Fig. /0). Strobilar branches apparently originate in almost any position, and thus no definite branching plane was found in which strobili arise, as seems tO be the case for the L. inundatum and L. carolinianum groups and for L. ramulosum.

The strobili of L. laterale are not lateral in origin as commonly described and indicated by the specific epithet. They are completely differentiated dichotomous branches, or the basal part of the branch may bear normal leaves.

SUBGENUS LYCOPODIUM

This subgenus includes perhaps 60 species which are all terrestrial with subter- ranean or superficial, creeping, scandent, or climbing pseudomonopodial main stems here termed rhizomes. Wilce (1972) arranged these species in seven groups, some of which are treated as genera by Holub (1975), Love, Love and Picht Sermolli (1977), and Pichi Sermolli (1977). ,

B. OLLGAARD: STUDIES IN LYCOPODIACEAE, II. 57

The basic branching pattern of the groups is very uniform. All species are anisotomous or occasionally isotomous in their rhizome branching. In L. spurium Willd., isotomy occurs with fairly regular intervals in the rhizomes, although anisotomy is the prevalent mode of branching. In other species, isotomy seems to occur more randomly. The result of anisotomy is the production of a pseudomonopodial rhizome that gives off minor branches alternately to the left and right.

The orientation of the branching planes in the rhizome is inclinate throughout the subgenus, although my material of L. casuarinoides Spring did not show this character with certainty; it had very shallow inclination angles or possibly flabel- late branching. The minor branches of the rhizome develop into variously differ- entiated shoot systems. A main axis may become more or less distinct and bear branchlet systems which in turn may produce strobili. The orientation of branch- ing planes in these shoot systems is invariably inclinate, although the inclination angle is variable, even within the same plant. Frequently it is impossible to deter- mine the inclination angle exactly, particularly in the species with radial branch symmetry. In major branches, the plectostele provides help in discerning the inclination angle of successive branchings, as the orientation of the stelar plates 1s fairly constant throughout a branching interval, and the stelar plates in derived branch steles are tilted towards each other (Fig. 6). In minor branches, the stelar patterns are not helpful for orientation because the steles are radially symmetrical. The effect of inclinate branching of branchlet systems is particularly well dis- played in some species of the L. complanatum group in which the upper sides of the flattened or quadrangular branchlets are tilted towards each other, although sometimes only slightly so. In some species (e.g., L. tristachyum Pursh), the regular inclination of all branchings of the branchlet systems creates a more OF less complete funnel-shaped arrangement of branchlets, whereas in other species (¢.8.. L. complanatum L.) the branchlet systems are often flat because the branches bend backwards after their inclinate origin.

DISCUSSION branching

Tie wa: ; sent study is that three types of branchi ain conclusion of the present 7 deviating

Patterns can be distinguished in Lycopodium sensu lato, —

pattern of L. /aterale, which is incompletely understood. —— gee Species, there seems to be perfect correlation of branching pattern types - oe : subgenera proposed by Wilce (1972), and thus the branching patterns give Turthe

ter have not been recognized previously. Also, the branching in L. /aterale and in some species of subg.

Troll (1937, p. 474ff.) supported a theory of correla the orientation of successive branching planes. He ¢

Urostachya is new. tion of shoot symmetry and laimed that branching in

58 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

lycopsids is of two basic kinds, cruciate and flabellate. According to this theory, cruciate branching is correlated with radial symmetry and flabellate branching is correlated with bilateral or dorsiventral symmetry. As examples of the former, he mentioned L. selago and other species of subg. Urostachya. Lycopodium num- mularifolium was mentioned as the striking exception in this subgenus; it exhibits correlation of flattened shoots (due to leaf orientation) and flabellate branching pattern. The shoot systems of L. clavatum L. and L. annotinum L. were claimed to be dorsiventral, even if superficially they appeared to have radial symmetry, and the shoot systems were said to be flabellate. The apparent radial symmetry of the rhizome of L. complanatum L. (in correlation with Goebel’s observation that in this species branchings do not take place in one plane) was used to support the theory, considering that the aerial branchlet systems of the same species are flabellate and dorsiventral.

The results of the present study contradict Troll’s theory of correlations. The arguments against it are as follows. (1) Branching in L. selago and its allies was not in any case found to be entirely cruciate. The majority of branchings are inclinate, while some apparently are cruciate, and a few are flabellate. (2) In L. nummularifolium, branchings are partly flabellate, but also can be cruciate. When they are flabellate, the branching plane does not coincide with the flattening plane of the leafy shoot (Fig. 2). (3) Whether or not the shoots of L. clavatum and L. annotinum are “‘truly’’ radial or dorsiventral, the branching (Fig. 3) is inclinate to sometimes indistinctly cruciate. This applies also to all portions of L. com- planatum plants. While a correlation of shoot symmetry with type of branching orientation seems absent, the flattening of dorsiventral branchlets in the

Subgenus Lycopodium seems to be uniform in its inclinate-cruciate branchings. The inclination angle is variable, even within a single specimen. Occasionally it can be 90°, and in this case should be termed cruciate. In subg. Lycopodiella, at least some cruciate branchings initiate the production of branches with different functions, but a similar consequence is not found in subg. Lycopodium. The change from inclinate to cruciate in the latter subgenus is part of a continuously variable character, and only the descriptive terms used have rigid definitions. The branching type of L. casuarinoides has not yet been determined because of in- adequate material.

Subgenus Lycopodiella has anisotomy in common with subg. Lycopodium, but in contrast is flabellate in the horizontal plane in looping branches, with a perpen- dicular branching plane in which originate the fertile branches of L. inundatum, L. carolinianum L., and their relatives, and in the L. cernuum group the erect and ultimately fertile branches. The rather fixed pattern of purely flabellate and cru- ciate branching sequences is here interpreted as a distinct subgeneric character.

B. OLLGAARD: STUDIES IN LYCOPODIACEAE, II. 59

Lycopodium laterale and the closely related L. ramulosum pose a puzzling problem. Whereas L. ramulosum seems to have a definite, predictable pattern, L. laterale is highly irregular. The growth habit of L. ramulosum and sterile creeping shoots of L. /aterale are rather similar, and so are the branching patterns. This points to the assumption that the irregular branching of the fertile shoot system in L. laterale is due to its erect position. In contrast, the flabellate pattern of L. ramulosum and sterile prostrate branches of L. laterale might be due to their prostrate habit. Experiments and further field studies are needed to examine this possible correlation.

In spite of the difference in habit between L. inundatum (Fig. 8) and L. cernuum (Fig. 11), there also is a striking similarity. Usually the first unbranched stem formed by the hibernating bud in L. inundatum is more or less erect until the first dichotomy, by which the strobilar branch and the creeping branch are formed. This is rather like the first stage of looping stem development in L. cernuum. The situation in L. alopecuroides var. furcatum (Fig. 7), in which the ‘*creeping’’ stem forms a high loop, is even more similar to L. cernuum. If the erect branch of L. cernuum is reduced to a simple branch, the similarity in habit is striking indeed.

Lycopodium ramulosum could easily be fitted into this common pattern, assum- ing an essential vegetative development before the formation of strobili. How- ever, its subterranean stems are insufficiently known, and its close alliance with the irregular L. laterale makes the homology of its branching pattern with the other groups questionable.

Descriptions of subg. Urostachya usually indicate that the roots form a basal tuft. This is the most common situation. However, in the L. crassum group, additional roots emerge from the underside of prostrate or subterranean shoots. As indicated above, the distinction between these rooting shoots and erect ones Is weak and variable. It is believed that perhaps all species of the subgenus possess the capacity to root as a consequence of soil contact. Internal or cortical roots are Produced acropetally at regular intervals in L. pithyoides Schlecht. & Cham., according to Stokey (1907). Under certain circumstances, the roots may emerge and penetrate the substrate, instead of penetrating the cortex to the stem base. This capacity is used for vegetative propagation of pendant epiphytic lycopodiums in greenhouse cultivation. Unfortunately, Pritzel’s figure 372 (1900, p. 593) a Saururus Lam., which belongs to the same group as L. crassum Humb. & Bonpl. €x Willd., does not show the typical growth habit of the species, with short Prostrate branches from which the erect elongated branches originate. ire

The present results do not suggest a rational way to divide subg. Urostachya. On the basis of leaf habit, chromosome number, and gametophyte alee 'Wo or three major groups usually have been recognized (Boivin, 1950; He ‘ 1949-1950; Léve, Love & Pichi Sermolli, 1977; Rothmaler, leap a ca < Alston, 1938). The two spore morphologies (Wilce, 1972) do not me ene these major groups. Heterophylly and homophylly per se are not BISUBELS roups ters for Separation, since many species are intermediate. In addition, some Sie Which | interpret as natural ones contain both leaf types. Accurate eng ya numbers and gametophytes are known for approximately ten species of a 8

60 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

totalling perhaps 400 species. Although these characters suggest the existence of more than one group, much more detailed knowledge is necessary before formal recognition at any level is advisable.

Subgenus Lycopodiella can be divided into four groups that are based on differ- ences of growth habit, sporophyll and sporangium shape, mucilage canal distribu- tion, spore morphology and chromosome number. Branching patterns indicate three related groups, but the pattern in L. laterale does not conform with the basic pattern that is otherwise characteristic of the subgenus. The L. inundatum and L. carolinianum groups have a very similar branching pattern, but seem distinct on other grounds (Bruce, 1976c). Chromosome numbers are not known for the L. laterale group, but otherwise point to distinct numbers for the L. inundatum and L. carolinianum groups, whereas in the L. cernuum group, several odd counts make it impossible to deduce the base number unambiguously. Despite the aber- rant branching of L. laterale , there is hardly reason to doubt that it is more closely related to the species of subg. Lycopodiella than to any other group.

In subg. Lycopodium, Wilce (1972) informally recognized seven groups. Som of these are treated as genera by Holub (1975) and Pichi Sermolli (1977). pre the branching patterns of this group are very uniform, other characters are not. Six spore types, five leaf habit types, and four doubtfully distinct types of sporan- gium wall structure are known from the seven groups. Two gametophyte types are represented in the five groups that have been studied. Different chromosome numbers are recorded from four of the groups. These discontinuities coincide to a great extent with the group limits and contribute to the distinction of separate evolutionary units in the subgenus. On the other hand, the same characters are mainly variations of common basic characters which are distinct from those of the other subgenera. It is therefore doubtful that these groups are sufficiently distinct to be given higher than sectional rank.

LITERATURE CITED

BIERHORST, D. W. 1971. Morphology of Vascular Plants. Macmillan, New York. :

BOCK, W. 1962. Systematics of Dichotomy and Evolution. Geol. Center Res. Ser. 2:1-300. Geolog'- cal Center, North Wales, Pennsylvania

BOIVIN. se pion The problem of generic segregates in the form-genus Lycopodium. Amer. Fern J.

PRUE. Fc G. a Development and distribution of mucilag Is in Lycopodium. Amer. J. Bot. 481-49]. - 1976b. Gametophytes and subgeneric concepts in Lycopodium. Amer. J. Bot. 63: 919-924. i Be pe studies in the biology of Lycopodium carolinianum. Amer. Fern J. CHU, M. C.-Y. 1974. A comparative study of the foliar anatomy of Lycopodium species. Amer. J. Bot. 61:681-692. CUTTER, E. G. 1966. Pattems of organogenesis in the shoot. Pp. 220-234 in E. G. Cutter (ed.). as Trends in Plant Morphogenesis. Longmans, Green, London ae oe 1949-1950. Systema Lycopodiorum. Rev. Sudamer. Bot. 8:67-86, 93-116. AY, J. E. 1916. Studies in the New Zealand species of the genus Lycopodium: Part I. Trans. New Zeal. Inst. 48:253—30 . 1919. Studies in the New Zealand species of the genus Lycopodium: Part I1I—The plastic- ity of the species. Trans. New Zeal. Inst. 51:161-216.

B. OLLGAARD: STUDIES IN LYCOPODIACEAE, II. 61

HOLM-NIELSEN, L. B., S. JEPPSEN, B. LOJTNANT, and B. OLLGAARD. 1975. Preliminary report on the 2nd Danish Botanical Expedition to Ecuador. Botanical Institute, University of

HOLUB, J. 1964. aay eine neue Gattung der Ordnung peerage Preslia 36: 1622. ——.. 1975. Diphasiastrum, a new genus in Lycopodiaceae. Preslia 47: 0. LOVE, D. LOVE, and R E.G. PIC Hi SERMOLLIAIO7: Sakae Atlas of the eridophyta. J. Cram duz «elves B...1975.. Salient 1 ceded eae I. Observations on the structure of the sporangium wall. Amer. Fern. J. 65:19-27. ges H. BALSLEV. 1979. ape on the 3rd Danish Botanical Expedition to Ecuador. Rep. . Inst. Univ. Aarhus 4: (in press). PICHI seuwouit Reb. GaAs 7. reas pteridophytorum genera in taxonomico ordine redi- gendi. Webbia 31:313—512. PRIMACK, R. B. 1973. Growth patterns of five species of Lycopodium. Amer. Fern PRITZEL, E. 1900. Lycopodiaceae. Pp. 563-606 in A. Engler and K. Prantl (eds.). Die Natiirlichen Pflanzenfamilien, Teil 1, Abt. 4. W. Engelmann, ie ROTHMALER, W. 1944. Pteridophyten-studien I. Repert. Sp. Nov. sere 54:55-82. STOKEY, A. G. 1907. The roots of Lycopodium pithyoides. Bot. Gaz. 44:57-63. : TROLL, W. 1937. Vergleichende Morphologie der hoheren Pflanzen, me 1, Teil 1. Gebruder entecet Berlin. Reprinted by O. Koeltz, K6nigstein-Taunus, 1967. WALTON, J. and A. H. G. ALSTON. 1938. Lycopodiinae. Pp. 599-606 in F. Verdoorn (ed.). Manua lf Pteridology. M. Nijhoff, The Hague WILCE, i H. saps Section Complanata of the genus " Lycopodium. Nova Hedw. Beih. 19: x + 7233, t. I-XL Bs paleo spores, I. General spore patterns and the generic segregates of Lycopodium. Amer. Fern J. 62:65-79

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62 AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 2 (1979)

SHORTER NOTES

ISOETES BUTLERI IN GEORGIA. — Recently, Baskin and Baskin (Amer. Fern J. 68:7—8. 1978) reported stations for /soétes butleri Englem. from Ken- tucky, Tennessee, and Alabama as representing the known distribution of the species east of the Mississippi River. Since then, we have collected it on 20 Apr 1978 in a cedar glade drained by West Chickamauga Creek in Catoosa County, Georgia, 34°54'20’N and 85°14'48’W (Boom 63, TENN). According to J. M. Baskin (pers. comm.), C. L. Boynton (Biltmore Bot. Stud. 1:147. 1902) reported finding /. butleri in 1899 in Chickamauga Park, which he incorrectly ascribed to Dade Co., Georgia. It is not clear whether this is a sight report or if specimens were collected, but it is likely that this and our localities are in the same area. Dr. Wilber Duncan (pers. comm.) does not record its existence in the state. We believe our collection represents a southeastern extention of the species’ range of approximately 120 miles from the cedar glades of the Central Basin of Tennessee, and constitutes the first documented report of its occurrence in Georgia.

The mature megaspores of our specimens lack the distinct or confluent tubercu- late ornamentation which is typical of most J. butleri, according to Taylor, Mohlenbrock, and Murphy (Amer. Fern J. 65:33-38. 1975), and instead appear rather smooth even when examined at 1300 with the SEM. The taxonomic significance of this character-state appears doubtful, but future collections of additional populations of J. butleri from Georgia should be examined in this con- text. The possibility that smooth megaspores are somehow environmentally in- duced or geographically related should be tested experimentally.

The plants we collected were from a population restricted to a moist mud depression along a small intermittent stream which drains the limestone glade. Within a month after our collection, the visible population had shrunk consid- erably in size as the glade began to enter its annual dry season. Therefore, this species should be sought in early spring in the northwestern Georgia counties. Geologically, Georgia is probably best known to botanists for the sediments of the Coastal Plain and the igneous and metamorphic rocks of the Piedmont. The northwestern portion of the state, however, is characterized by Paleozoic rocks of which a considerable portion is limestone. Typical cedar glades tend to develop in areas where this limestone outcrops, creating ideal conditions for the calciphilous ib butleri.—Brian M. Boom and A. Murray Evans, Department of Botany, Uni- versity of Tennessee, Knoxville, TN 37916.

AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 2 (1979) 63

JUVENILE LEAVES OF THE APOGAMOUS FERN NOTHOLAENA COCHISENSIS.—Cochise’s Cloak Fern is a desert or semi-desert dweller which ranges from central and western Texas to California and south into Mexico at least to the state of Aguascalientes. Material of this species from McKelligan Canyon, El Paso, Texas (Knobloch 2523, MSC) recently has been proven to be apogamous since no archegonia are present and the sporophyte arises from a single cell layer in back of the sinus (D. P. Whittier, pers. comm.).

In sexual species of ferns, the first leaf arises from an upper part of the embryo, but in N. cochisensis, the stem apex arises from a mound of vegetative cells of the prothallus, and all the leaves come from this stem apex.

es ¢* «@ ft ®&

4a & t£éAa & 3 |

FIG. 1, Heteroblastic leaf sequence in N. cochisensis (Knobloch 2523, MSC), x 1.3.

As in most species of ferns, the first leaves of N. cochisensis do not sage the adult ones (Fig. 1). Wagner (Amer. J. Bot. 39:578-592. 1952) crane . heteroblastic development in several other species. In N. a ei leaves are tiny and invariably simple. They vary greatly in shape, much pat - than do adult leaves. The first leaves gradually die as they are segues eal ones. Depending upon growing conditions perhaps, eventually ae ge appear. Why the genes for pinnateness are repressed in early os : de- known, but it seems evident that the extent of photosynthetic leaf The first veloped is sufficient for the requirements of the plant at each stage. } é me leaves are for the most part devoid of the dense indument that a ioe adult leaves.—Irving W. Knobloch, Department of Botany and Plant Pathology,

Michigan State University, East Lansing, MI 48823.

64 AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 2 (1979)

A THELYPTERIS NEW TO FLORIDA.—Over the past two years field studies by Nauman and Austin have turned up two additional species for the flora of the United States, Pleopeltis revoluta (Spreng. ex Willd.) A. R. Smith and Tectaria incisa Cav., both from Broward County, Florida.

A third species was found in November of 1978, Thelypteris grandis A. R. Smith var. grandis. Previously this variety was known from the Greater Antilles and St. Kitts in the Lesser Antilles (Smith, Univ. Calif. Publ. Bot. 59:1—143. 1971) and is here recorded as new for the United States. Other varieties of T. grandis occur in Central and South America. Voucher specimens have been deposited at the following herbaria: FAU, FLAS, UC, and US.

Thelypteris grandis was discovered by Daniel Austin and me while studying the vegetation of the Fakahatchee Strand portion of the Big Cypress National Pre- serve. The plants were found growing in a mixed swamp dominated by Taxodium distichum. About a dozen individuals formed a single colony on one of a series of tram roads. Essentially limestone spoil mounds, the trams are abandoned access roads from a logging operation in the 1940’s and 1950's.

This is the most distinctive species of Thelypteris occurring in Florida. It is easily recognized by its large size, with fronds more than 1.5 m long and 0.5 m wide. Other distinctive features include the creeping rhizome, deeply incised pinnae mostly 2-3.5 cm wide, and the costa being uniformly puberulous below. The type variety is distinguished from the other varieties by the greatly reduced basal segments of the pinnae, the glabrous or sparsely hairy indusia, and the nearly medial sori (Smith, loc. cit.).

Whether 7. grandis is a casual occurrence from the Antilles, native, or an escaped cultigen is at present uncertain; further study may shed more light on this’ question.— Clifton E. Nauman, Department of Biological Sciences, Florida At- lantic University, Boca Raton, FL 33431.

NOTEWORTHY PUBLICATIONS OF THE CHRONICA BOTANICA

BEDDOME, R. H. The Ferns of British India. Includes Bangladesh, Burma, Ceylon, Malaya, Nepal, Pakistan, and Tibet. Set of 2 vols. Reprint ed. 1973. Available CB (cloth bound) or LB (half-leather with gold lettering). CB Rs. 285: LB US $55.

. Supplement to the Ferns of Southern India and British India.

Seas Coe ache CB Rs. 85; LB US $18.50.

———.. Handbook to the Ferns of British India. Reprint ed. 1970. CB Rs. 100; LB US $21.

NAYAR, B. K. Companion to R. H. Beddome’s Handbook. 1974. CB Rs. 75; LB US $14.50 Annales Cryptogamici et Phytopathologici, vol. 8 . Fern Flora of India. The first taxonomic account of the fern flora since Beddome’s classical work ‘‘Handbook to the Ferns of British India’’ published in 1883. An illustrated floristic account containing keys to identifications as well as lucid descriptions of species, genera, and families of Filicopsida. Prepared by the foremost pteridologist of India. About 1000 pages. November

1979.

CB Rs. 550; LB US $115.

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Plant Classification Second Edition

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This classic text has been revised to include recent research: new examples for major manuals on floras in the Pacific Northwest, California, Northeast, and South; reassignment of the members of the Amentiferae to new positions in the Thalamiflorae and Calyciflorae; revisions in the tribes of the grasses; recent findings in evolution; and classification systems of flowering plants. Plant Classification, Second Editionis a superbly illustrated text/reference book written to help readers: acquire essential vocabulary for describing characteristics of plant groups; identify plants by applying the use of keys and descriptions; gain knowledge of plant.taxa through preparation and preservation of specimens to form-an ordered collection;:develop an under- standing of the basis for classification of plant groups; and gain an appreciation of the association of species in natural vegetation.

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Flowering Plants Gymnosperms Ferns Psilophytes and Horsetails Club Mosses Association of Species in Natural Floras Appendix Glossary

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AMERICAN aie FERN ea J O U R N A E July-September, 1979

QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY

A New Nephrolepis Hybrid From Florida CLIFTON E. NAUMAN Spore Morphology of Anemia subgenus Anemia STEVEN R. HILL

The Development of Plantlets from Strobilus Branches in Lycopodium phlegmaria VC, WEE

Incidence of Epiphytism in the Lycopsids JOSEPH M. BEITEL

Cyrtomium fortunei in Louisiana and Mississippi GARRIE P. LANDRY, MICHAEL ISRAEL, ROBERT SCHWARZWALDER, JR, and R. DALE THOMAS

The Fine Structure of the Pre-meiotic Stages

of Sporogenesis in Onoclea sensibilis NORMAN P. MARENGO

Api ; : Pical Dominance in Anemia phyllitidis Gametophytes THOMAS L. REYNOLDS

— — A New Combination in Asplenosorus; — , hes alabamensis New to Kentucky, xuality in Asplenium resiliens

Reviews

MISSOURI BOTANICAL

OCT 16 1979

GARDEN LIBRARY

70, 84,

nN a

| _

The American Fern Society Council for 1979

RICHARD L. HAUKE, Dept. of Botany, University of Rhode Island, Kingston, R.I. 02881. President ROBERT M. LLOYD, Dept. of Botany, Ohio University, Athens, Ohio 45701. Vice-President LESLIE G. HICKOK, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916. Secretary JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916. Treasurer JUDITH E. SKOG, Dept. of Biology, George Mason University, Fairfax, Va. 22030. Records Treasurer DAVID B. LELLINGER, Smithsonian Institution, Washington, D.C. 20560. Editor-in-Chief

JOHN T. MICKEL, New York Botanical Garden, Bronx, N.Y. 10458. Newsletter Editor American Fern Journal EDITOR

DAVID B. LELLINGER Smithsonian Institution, Washington, D. C. 20560

ASSOCIATE EDITORS DAVID W. BIERHORST ..Dept. of Botany, University of Massachusetts, Amherst, Mass. 01002 GERALD J. GASTONY +++r+eeseeee-Dept. of Biology, Indiana University, Bloomington, Ind. 47401 JOHN T. MICKEL. New York Botanical Garden, Bronx, New York 10458

The **American Fem Journal’ (ISSN 0002-8444) is an illustrated quarterly devoted to the general study of fems. It is owned by the American Fern Society, and published at the Smithsonian Institu- tion, Washington, DC 20560. Second-class postage paid at Washington.

Matter for publication and claims for missing issues (made within six months of the date of issue) should be addressed to the Editor.

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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 Fem Society (annual dues, $8.00; life membership, $160.00). Extracted offprints, if ordered in advance, will be furnished to authors at cost, plus postage.

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Newsletter Dr. John T. Mickel, New York Botanical Garden, Bronx, New York 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, hor- ticultural notes, and reviews of non-technical books on fems.

Spore Exchange

Mr. Neill D. Hall, 1230 Northeast 88th Street, Seattle, Washington 98115, is Director. Spores exchanged and collection lists sent on request.

—— Gifts and Bequests | tsand beq the Society enable i ‘ é bers and to others interested in ferns. Botanical books, back issues of the Joumal, and cash or other gifts are always welcomed, and are tax-deductible. Inquiries should be addressed to the Secretary.

AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 3 (1979) 65

A New Nephrolepis Hybrid From Florida CLIFTON E. NAUMAN*

Nephrolepis is a genus of probably 30 species mostly pantropical in distribution. Several species have been problematical in their identification, notably N. biser- rata (Swartz) Schott, N. exaltata (L.) Schott, and N. hirsutula (Forst.) Presl (Christensen, 1932, p. 74; Copeland, 1947, p. 91). Two of these species, N. exal- tata andN. biserrata, occur in Florida and have been a source of difficulty, in part because of plants intermediate in a number of characteristics. The intermediates appear to represent hybrid populations between these two species.

Distributional data show that intermediates occur only in the overlapping por- tions of the parent species’ ranges and not where only one parent occurs. Inter- mediates are found wherever N. exaltata and N. biserrata occur together, usually in disturbed portions of the habitat. The intermediates are more abundant and luxuriant as would be expected of hybrids.

2

20 ym

FIG. 1. Spherical spore of the proposed hybrid. FIG. 2. Aborted tetrad of the proposed hybrid.

Cytology does not rule out the possibility of hybrids between N. biserrata and N. exaltata since both species have been reported to have the same gametic number, n=41, (Abraham et al., 1962; Chiarugi, 1960; Fabbri, 1965; Walker, 1964-65, 1973). ee 89

Spore morphology provides further evidence of hybridity in the intermediates. Nephrolepis is characterized by free, anisopolar, bilateral, monolete ih Dah int Erdtman & Sorsa, 1971). Spores of the intermediate are irregular, unsee : sae alete (Fig. 1) and demonstrate features similar to those described wl ad (1962) for Polystichum hybrids. A large number of aborted spores ce : ee tetrads (Fig. 2) also are characteristic of these plants. Frequency distri : ss - spore size are platykurtic when compared to those of the putative pt _ (Fig. 3). This distribution appears to be a significant feature of Term y (Daigobo, 1967; Kanamori, 1969, 1971; Wagner & Chen, 1965).

Pe aaa : : . 431. *Dept. of Biological Sciences, Florida Atlantic University, Boca Raton, eh

Volume 69, number 2, of the JOURNAL was issued June 26, 1979.

66 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

_ N. biserrata s aa REEL GREE | T T T T

Oo. © N. x averyi = 2) a ©. 5 v 7 ee w ~ 6) o ON ¥ T T T T je Ree i 20 30 40 50

Maximum Length (pm)

N. exaltata

| :

ce aR ' T 7 Pa so '

FIG. 3. Frequency distributions of maximum spore length in three of the Florida taxa of Nephrolepis.

C. E. NAUMAN: NEW NEPHROLEPIS HYBRID 67

Morphological intermediacy occurs in several characters, and is demonstrated by polygonal graphs of mean values of eight characters (Fig. 4). Scatter diagrams suggest some degree of backcrossing in addition to hybrid intermediacy (Fig. 5). In Florida, the adaxial surfaces of the costae of N. biserrata typically are tomen- tose; those of N. exaltata are glabrous. The intermediates are slightly pubescent.

N. exaltata N. x averyi

N. biserrata

FIG. 4. Polygonal graphs of mean values of eight morphological characters wd node eg as re of Nephrolepis. A = Pinna length (cm). B = Interpinna distance (mm). C = Fron _ Pe ie Pinna tip shape (1-5 are coded states from obtuse to narrowly attenuate). E = eainceie states from (1-4 are coded states from no lobe to a linear lobe). F = Pinna attachment (1-3 are ¢ %

sessile to subpetiolulate). G = Lower pinna length/width ratio. H = Pinna width (cm).

t Indusium width also is intermediate: in N. eee cia anes 4 ee . t 1.26 mm. bla about 1.35 mm, and in the intermediates abou 08 (3.04-7.79) in N. biserrata,

average 9.53 (6.86-17.93) in the intermediates, 5: and 8.40 (4.30-16.98) in N. exaltata. ne

Mixtures of character states also occur in the intermediates. en - - indusium shape and attachment are examined in typical nih oe ates biserrata and N. exaltata, the two species are distinct. Nephrolep!

i indusi d by a point orbicular, peltate indusia, and N. exaltata has reniform indusia attached by a po

68 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

©) 30- 0 a re) re) fe) O O ° Oo 26- .@) @) re) ©) .@) Oo Oo ©) Oo Oo 2 ~~ _ Oo E O £— e) = e) : © sd be pe * ® o © O ed Oo a Bs oO @ Co 4 © € 18 - a0 & * 56 - Lg ee © c+) 2S code 8° - <° te Ge a e @ & e °@ . ® 0 $33 & O-:N. biserrata e e, Se e @=:N. x averyi a % @,00 Ps @-N. exaltata oe ~ 10 + e ® ~ 2 T t t T LJ a T J 0.8 1.2 1.6 2.0

Pinna Width (cm)

FIG. 5. Scatter diagram of interpinna distance versus pinna width of three of the Florida taxa of Nephrolepis.

C. E. NAUMAN: NEW NEPHROLEPIS HYBRID 69

in the sinus. Occasionally a mixture of indusium types is found on a single plant, apparently always on the intermediates. Pinna shape of the intermediates is falcate like that of N. exaltata. Frond length approaches that of N. biserrata.

At least two characters suggest hybrid vigor. The stipe length of N. biserrata and N. exaltata is 2.3 and 1.7 dm, respectively, but in the intermediates it is 2.7 dm. Trichome length inN. biserrata is consistently 0.3 mm; in the intermediates it is consistently 0.4 mm.

The morphological intermediacy and spore morphology of the hybrids fulfill two of the three criteria for hybridity given by Wagner and Chen (1965). Distribution and habitat ecology also make hybrid origin of the intermediates likely. The hy- brids may be known as:

Nephrolepis < averyi Nauman, hybr. nov. Bt tin

Planta inter N. biserratam et N. exaltatam interposita et verosimiliter per hy- bridionem harum specierum orto. Costae abaxiales leviter pubescentes. Indusia orbicularia vel reniformia, peltata vel in sinu affixa. Sporae abnormales et abor- tivae, sphericae, diametro 32.6 um. :

Epiphytic, epipetric, or terrestrial plants with fronds up to 3.0 m long. Stipes 1.0-5.25 dm long, glabrous, paleate with lanceolate-attenuate, brown scales. Rachises 0.5-2.6 m long, paleate, otherwise glabrous. Pinnae falcate, 3.6-8.0 cm long and 1.0-1.75 cm wide, truncate to truncate-auriculate at the base, narrowly acute at the tip, with serrulate margins, costae abaxially lightly pubescent, lamina adaxially and abaxially glabrous to occasionally pubescent and lightly apie Sori intramarginal to supramedial. Indusia orbicular to reniform, peltate to A tached in the sinus. Annulus with 13 or 14 indurated cells. Spores abnormal an abortive, spherical, 32.6 pm in diameter.

TYPE: Fakahatchee Strand off West Grade, 50 ft E of Indian Mound Slough Bridge, Collier Co., Florida, 29 Jan 1979, Nauman et al. 635 (US; isotypes FAU, FLAS, GH, MSC, NY). : ks and

Associated with its parents N. biserrata and N. exaltata in hammocks on swamps. Distributed in southern Florida (Broward, Collier, Dade, Palm Beac and Pinellas Counties, and probably others). Named after George N. pie

The hybrid is best distinguished from N. exaltata by its larger size ane’ hig x pubescent upper costal surfaces, and from N. biserrata by its falcate pinnae, a narrower fronds.

PARATYPES (all from Broward Co., Florida): Cypress Creek serene vt 1976, Durand 65 (UC), Durand 71 (FSU), 12 Oct 1978, Nauman 4 Nauman 436 (USF).

LITERATURE CITED

ABRAHAM, A., C. A. NINAN and P. M. MATHEW. 1962. Studies on the ¢ the pteridophytes. VII. Observations on one hundred species of so’ Bot. 41:339-421. ; CHIARUGI, A. a) hone chromosomiche delle pteridophyta. Caryologia ee ao CHRISTENSEN, C. 1932. The pteridophytes of Madagascar. Dansk Bot. Ark. 7:1-253, ¢. COPELAND, E. B. 1947. Genera Filicum. Chronica Botanica, Waltham, Mass. — nae DAIGOBO, S. 1967. Variation of spore size in the Jaf Pol : : ey Bot. 42:207-210.

ytology and phylogeny of uth Indian ferns. J. Ind.

70 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

ERDTMAN, G. and P. SORSA. 1971. Pollen and Spore Morphology/Plant Taxonomy. Almquist and Wiksell, Stockholm.

FABBRI, F. 1965. Secundo supplemento alle tavolle chromosomiche delle pteridophyta di Alberto Chiarugi. Caryologia 18:675-728.

KANAMORI, K. 1969. Studies on the sterility and size variation of spores in some Japanese Dryo- pteris. J. Jap. Bot. 44:207-217, t. 14-17. . 1971. Studies on the sterility and size variation of spores in some apogamous ferns. J. Jap. Bot. 46:146-151.

MORZENTI, V. M. 1962. A first report on pseudomeiotic sporogenesis, a type of spore reproduction by which “‘sterile’’ ferns produce gametophytes. Amer. Fern. J. 52:69-79.

WAGNER, W. H., Jr. and K. L. CHEN. 1965. Abortion of spores and sporangia as a tool in detection of Dryopteris hybrids. Amer. Fern J. 55:9-29.

WALKER, T.G. 1964-65. A cytotaxonomic survey of the pteridophytes of Jamaica. Trans. Roy. Soc. Edinb. 66:169-237. . 1973. Additional cytotaxonomic notes on the pteridophytes of Jamaica. Trans. Roy. Soc. Edinb. 69:109-135.

REVIEW

**THE PTERIDOPHYTES OF SURINAME,” by K. U. Kramer. Natuurw. Stud. Suriname Ned. Antill. 93:1-198. 1978.—In format this work is essentially a sec- ond edition of Posthumus’ ‘Fern Flora of Suriname”’ (1928). But it is much revised, corrected, and expanded over its predecessor. Considerable collections have been made in Suriname over the past fifty years, especially in the mountain- ous central portion that is underlain by Roraima sandstones. This area has floristic affinities with Guayana and southern Venezuela, and so contains many species not recorded by Posthumus.

Kramer’s treatment includes 334 taxa of ferns and fern-allies in 63 genera. For the most part, the genera are conservatively and broadly delimited, a course chosen by the author because of the lack of definitive knowledge in splitting such genera. The work contains a very novel and useful multiple-entry key to most genera (a few are treated in a conventional key). By scoring an unknown specimen for up to eight characters, a unique or nearly unique profile results that can be matched against a list of characters for each genus. Those who have worked with incomplete material will appreciate this approach over a conventional key.

The keys to species appear to be accurate and well constructed. Although the Flora lacks descriptions and illustrations, there are few enough taxa, even in the largest genera (Trichomanes, 37; Polypodium, 30: Adiantum, 21), that the keys distinguish them adequately and the chance of arriving at a wrong name is slight. Important literature is given for the Flora as a whole and for each genus. An index showing accepted names and synonyms concludes the volume, which may be purchased for D. fl. 48 (ca. $23.50) from the Foundation for Scientific Research in Suriname and the Netherlands Antilles, Plompetorengracht 9-11, NL-3512 CA Utrecht, Netherlands.—D.B..

AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 3 (1979) 71

Spore Morphology of Anemia subgenus Anemia STEVEN R. HILL*

As a continuation of earlier work on the schizaeaceous genus Anemia Swartz (Hill, 1977), I have examined spores of nine species of subg. Anemia by means of the scanning electron microscope. This subgenus, unlike subg. Coptophyllum, which was revised by Mickel (1962), has not been critically treated since the revision of the Schizaeaceae by Prantl (1881). The present work was initiated to help clarify the relationships of some of the species in the subgenus which have vegetative specializations by examining the spores, which are assumed here to have a more conservative rate of change than the vegetative characters.

Spore samples were collected either in the field or from selected herbarium specimens. Twenty-three specimens of the nine species were mounted on aluminum stubs with double-adhesive tape and coated with a thin layer (200-400 A) of gold- -palladium. The prepared samples were then examined at 15 kV and photographed using the JEOL JSM-U3 Scanning Electron Microscope of the Electron Microscopy Center, Texas A&M University. Proximal, distal, and or- namentation detail micrographs have been included in most cases to allow com- parison at each orientation. Line scales indicate size based upon instrument mag- nification readings and are best used to indicate relative size. In the following list of vouchers, collections cited with an asterisk are illustrated in this paper and known herbarium locations for each collection are cited.

Anemia hirsuta (L.) Swartz. MEXICO: Jalisco: Ayo el Chico, Mc Vaugh 17167 (US). Temascaltepec: Ixtapan, Hinton 1640 (US). Oaxaca: Juchatengo, Hill 1643* (NCU, NY, VT); Villa Alta, Vera-Santos 3517 (US

ce munchii Christ. MEXICO: Oaxaca: Dist. Juchitan, 6 km S of Matias Romero, Hill s.n.* (spore collection only, living material at

Anemia oblongifolia (Cav.) Swartz. Group I (see text). BRAZIL: Goias: Serra dos Pyrenaos, Ander- son, Hill et al. 10296* (NY, UB); Serra Dourada, Inwin et al. 11793* (NY, UB,

Anemia oblongifolia (Cav.) Swartz. Group II (see text). BRAZIL: Goias: Serra dos Cristais, Irwin et al. 13216a* (NY, UB, US); Serra dos Pyrenaos, Inwin et al. 10977 (NY, UB, US); Alto Paraiso, Hill 1036 (VT).

Anemia ouropretana Christ. BRAZIL: Minas Gerais: Serra do Espinhago, [nvin et al. 27976* UB, US).

Anemia pastinacaria Moritz. MEXICO: Guerrero: Plan del Carrizo, Hinton 14650 (US). BRAZIL: Goids: Chapada dos Veadeiros, Invin et al. 24260 (NY , UB, US); Serra do Caiapo, Anderson, Hill et al. 9466* (NY, UB).

Anemia phyllitidis (L.) Swartz. MEXICO: Oaxaca: San Gabriel, Hill 1 Zapotal, — 2563* (US). BRAZIL: Distrito Federal: Corrego Landim, UB, US). Goids: Serra dos Pyrenaos, Irwin et al. 18947 (NY, UB, US); S (VT). Minas Geteis, Serra do Espinhago, Anderson, Hill, et - 9467* (NY, UB).

Anemia pohliana Sturm. BRAZIL: Goias: Jatai, Chase s.n S ete radicans Raddi. BRAZIL: Espirito Santo: Cachoeiro de Itap

(NY,

661 (VT). Chiapas: El Inwin et al. 11333 (NY, erra Dourada, Hill 1089

emirim, Foster & Foster 971*

; * (NY, US). Anemia rotundifolia Schrad. BRAZIL: Bahia: Toca de Onca, Rose & Russell 20099* (N

*Departn address: De- raateriment of Biology, Texas A&M University, College Station, oo 77843. Present of Botany, University of Maryland, College Park, MD

7? AMERICAN FERN JOURNAL: VOLUME 69 (1979)

+

FIGS. and 2. Anemia phyllitidis, proximal and distal. Anderson, Hill et al. 9467. FIGS. 3 and 4. 4- munchu, proximal and distal. Hill s.n. FIGS. 5 and 6. A. pastinacaria, proximal and distal. Anderson, Hill, et al. 9466. Line scale = 20 um,

S. R. HILL: SPORE MORPHOLOGY OF ANEMIA 73

4 27976. FIG. 9. A. bare ans, FIGS. 7 and 8. Anemia ouropretana, proximal and distal. Jrw : 7 sper vpre eiageae Proximal. Foster 971. FIG. 10. A. pohliana, proximo- -lateral. Chase a

difolia, proximal and distal. Rose & Russell 20099. Line scale = 20 u

AMERICAN FERN JOURNAL: VOLUME 69 (1979) 74

FIGS. 13 and FIGS. 15 anc A, hir.

14. Anemia oblongifolia, Group I, proximal 116. A. oblongifolia, Group II, proximal Sula, proximal and distal, Hill 1643.

. > : and distal, Anderson, Hill, et al. ai and distal, Invin et al. 13216a. FIGS. 17 and 18. Line scale = 20 wm.

§. R. HILL: SPORE MORPHOLOGY OF ANEMIA 75

RESULTS

With important exceptions to be noted below, the spores of taxa in Anemia subg. Anemia are tetrahedral with several widely-spaced, narrow ridges on each face separated by smooth unornamented zones. The ridges usually possess large, terete, rod-shaped appendages called baculae.

Spores of Anemia phyllitidis, the type species of the genus and of subg. Anemia and one of the most widespread species, are illustrated in Figs. /, 2, and /9. The variable spores are 50-60 «m in diameter exclusive of the baculae. The ridges are typically very narrow and are separated by unornamented zones or furrows (fos- sulae) 5-10 wm wide. The baculae are 6-10 wm long and truncate, and on the ridges between them are smaller microspines 1-2 um long.

The spores of A. munchii, a narrowly distributed species, differ primarily in their more highly ornamented ridges (Figs. 3, 4, and 20). The trilete scar has at least three rows of spines, compared with the single row in A. phyllitidis, the spores are 10-20 um larger (60-70 «m) on the average, and tend to be more spherical. The baculae are narrower and the ridges tend to lack the smaller micro- spines.

Anemia pastinacaria, another widely distributed species, has variable spores (Figs. 5 and 6) which are intermediate in size between the two described above. There is a tendency towards shorter, more clavate baculae. The trilete scar is more ornamented than that of A. phyllitidis but less so than that of A. munchii, having two rows of baculae.

Several members of subg. Anemia endemic to Brazi tive structures but conservative spore morphology. Four of these taxa were examined, but only one specimen of each was available for study due to scarcity. The first of these was A. ouropretana. This species differed from A. phyllitidis in its small spores (Figs. 7, 8, and 22), which were 40-50 wm in diameter. In addi- tion, the baculae were distinctly rounded and slightly expanded towards the apex. This tendency was shared by spores of A. radicans, but this species IS eg that it relies upon vegetative as well as or instead of gametic een ; sample examined (Fig. 9) possessed generally abortive or abnormally shape apparently non-functional spores as well as several elongated fronds bearing ap cal meristems. Since A. radicans is thought to be a sexually “es species (Lellinger, pers. comm.), the sample may have been atypical. :

The spores of A. pohliana (Figs. 10 and 21), a species also endemic e igre resembled those of the other members already described, but picid - si spores examined in the single specimen available were either abnorma “ ie the else were small and abortive. The scarcity of specimens, bai — , veral populations, and the intermediate nature of the fronds between agi sa other members of the subgenus in addition to the abnormal ogee se ag that this species or this particular specimen is of hybrid origin. ’ requires further study. ere

The fourth Brazilian endemic examined, A. rotundifolia had well formed, distinctly spherical spores. These were s

| have specialized vegeta-

(Figs. 11, 12, and 23), mall for the subgenus,

AMERICAN FERN JOURNAL: VOLUME 69 (1979)

V

FIG. 19. Anemia spore ornamentation. A. phyllitidis. Laughlin 2563. F1G. 20. A. munchii, Hill s.n. FIG. 21. A. pohliana. Chase s.n. FIG. 22. A. ouropretana. Irwin et al. 27976. Line scale = 10 »m. measuring 40 «wm in diameter. The ridges lack microspines, and the baculae ap- proach a clavate shape as in A. ouropretana and A. radicans.

The last group examined included two taxa, A. oblongifolia and A. hirsuta, both wide-ranging species. The spores of A. oblongifolia were seen to fall into two groups which were correlated with two vegetatively distinct groups of specimens. Group I may be exclusively Brazilian and can be characterized by semi-erect fertile fronds which are long-stalked (stipes up to 10 cm long) and which bear narrow pinnae 3-5 mm wide. The spores of this group (Figs. 13, 14, and 24) differ from all of the other spore samples examined by the presence of wide ridges, narrow fossulae, and the complete absence of conspicuous baculae. In addition,

S. R. HILL: SPORE MORPHOLOGY OF ANEMIA

FIG, 23. Anemia spore ornamentation. A. rotundifolia. Rose & Russell 20099. FIG. 24. A. oblon- gifolia, Group 1. Irwin et al. 11793. FIG. 25. A. oblongifolia, Group II. Irwin et al. 13216a. FIG. 26. A. hirsuta, Hil 1643. Line scale = 10 um.

the spore angles are somewhat prolonged as seen in proximal view and the or- namentation of the ridges consists of small (2 «m), branched microspines. These characteristics are also typical of spores of members of subg. Coptophyllum (Hill, 1977). Plants in morphological Group II have low, short- stalked fronds (stipes up to 2-3 cm long) with comparatively broader pinnae 6-8 mm wide. The spores (Figs. 15, 16 and 25) have baculae typical of A. phyllitidis and its allies present in reduced form (3-5 um long), and the occasionally branched microspines of the Coptophyllum g group are also found among them. The fossulae and ridges are intermediate in width between typical examples of each subgenus. The spores

AMERICAN FERN JOURNAL: VOLUME 69 (1979) 78

A.oblongifolia 2

A. hirsuta

A.oblongifolia 1

A. pohliana A. pastinacaria

A. radicans

A. rotundifolia A. munchii

FIG. 27. Possible morphological rel

Bone nemia. ationships among selected spore types of subg. A Hypothetical connection with

subg. Coptophyllum at upper right.

S. R. HILL: SPORE MORPHOLOGY OF ANEMIA 79

seem to provide a morphological link between the A. phyllitidis spore type and the Group I A. oblongifolium spore type described above, which is more typical of spores of subg. Coptophyllum.

The spores of A. hirsuta (Figs. 17, 18, and 26) pose an interesting problem in subg. Anemia. The shape of the spores, the widely spaced, narrow ridges, and the unbranched microspines ornamenting those ridges are typical of those of A. phyl- litidis, but the large baculae of that group are entirely absent, and the spore angles are slightly prominent.

DISCUSSION

Figure 27 illustrates a tentative scheme of morphologically related spore types to summarize the observations presented above. This scheme is not intended to represent a phylogeny, but is intended to express morphological similarities in the specimens examined. Basic to this arrangement are the variable spores of Anemia phyllitidis, to which several other spore types are connected. Anemia radicans, A. ouropretana, A. munchii, and A. rotundifolia, which are all restricted, specialized species, have spore types apparently related to and possibly even derived from the pool of variation seen in A. phyllitidis. Anemia pohliana may be a hybrid derived from members of this complex, but at any rate its spore type seems closely allied to that of A. pastinacaria, whose spores suggest a trend either towards baculae reduction as seen in spores such as those of A. oblongifolia, or in the opposite direction towards baculae elaboration as seen in A. phyllitidis. The A. hirsuta spore may be derived from that of A. phyllitidis directly through loss of the large baculae while retaining the microspines, or may be separately derived from that of A. oblongifolia by modification of the number of ridges and by simplification of the complexity of the microspines.

The problem requiring the most study is the morphological assessment of the A. oblongifolia complex. The spore analysis presented here suggests that more than one species actually is involved. In addition, the great resemblance of the spores of form 2 in Fig. 27 (referred to as Group I in the text above) with those of subg. Coptophyllum may provide a link between the two subgenera. Since subg. Cop- tophyllum is considered to be the most primitive of the three subgenera (Mickel, 1962), this taxon may aid in our knowledge of the origin of subg. Anemia.

I would like to thank Dr. Elenor Cox and the Electron Microscopy Center at Texas A&M University for aid in obtaining access to the scanning electron micro- scope, Dr. John T. Mickel of the New York Botanical Garden for support while collecting spore samples in Oaxaca, Mexico, and Dr. David B. Lellinger of the U.S. National Herbarium for kindly lending specimens used in this study.

LITERATURE CITED

optophyllum. Amer. Fern. J. Anemia, subgenus C optophyl

HILL, S. R. 1977. Spore morphology of Anemia subgenus C oa et

MICKEL, J. T. 1962. A monographic study of the fern genus oe State J. Sci. 36:349-482.

PRANTL, K. 1881. Untersuchungen zur Morphologie der ene

ceen. Wilhelm Engelmann, Leipzig.

skryptogamen. II. Die Schizaea-

80 AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 3 (1979)

The Development of Plantlets from Strobilus Branches in Lycopodium phlegmaria +: C. WEE

Lycopodium phlegmaria L. is an epiphyte commonly found growing on the trunks and branches of forest trees in Singapore. It is a widely distributed species in the Old World tropics, ranging from Africa to Polynesia. Its pendulous stems are covered with spirally arranged, stiff, ovate-lanceolate leaves (Fig. 1). Both the sterile portions of the stems and the strobili branch dichotomously (Figs. / and 2). The sporophylls are smaller than the foliage leaves and are arranged more com- pactly along the branches. In most cases, the sporophylls are grouped terminally On determinate stems. However, instances of foliage leaves appearing along the strobilus branches have been observed (Fig. 2). This is reminiscent of the condi- tion seen in L. selago L., where the fertile zones alternate with the sterile and the foliage leaves are not much different from the sporophylls (Bower, 1935, p. 200).

Recently, a portion of L. phlegmaria detached from the parent plant was found lying on damp ground on the secondary forest at the MacRitchie Catchment area in Singapore. It was still green, and the ends of the fertile branches showed growth following detachment. Under the humid conditions of the forest floor, the ends of the branches had reverted to vegetative growth and had produced one or two

with long internodes (Fig. 3). At the nodes were leafy structures similar to the sporophylls, In the axils of these structures were obviously vegetative buds, some of which had developed into vegetative shoots (Fig. 3).

Longitudinal sections of the strobilus branches revealed the gradual transforma- tion of the sporangia from fertile to sterile in the acropetal direction. The fertile,

vermiculite similarly stimulated development of vegetative shoots after about four months. The new growths had excessively elongate internodes were smaller until foliage leaves were produced when the axis turned upwards. Roots formed at the points of turning after a further period of two to four months. These shoots could then be removed as plantlets.

These observations give support to Bower’s (1908, p. 164) theory that foliage leaves essentially are sporophylls with the sporangia completely suppressed. In this respect, it should be noted that the strobili of L. obscurum L. and L. flabel-

*Department of

Singapore Botany, University of Singapore, Bukit Timah Road, Singapore 10, Republic of

Y. C. WEE: PLANTLET DEVELOPMENT IN LYCOPODIUM PHLEGMARIA

PU Ag ted

they,

ca)

_ 1. Potted plant showing the pendulous habit and

‘ FIG. 2. Strobilus branches showing groups of foliage owing branch tips developing

new growth at the end of a

FIGS

> eta l-4. Lycopodium phlegmaria. FIG

ics, ge branched strobili (scale in cm)

into oo, Detached portion of plant found on the forest floor sh (scale in mm). FIG. 4. Close-up of portion of Fig. 3 showing

Strobilus br: bilus branch (scale in mm).

82 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

liforme (Fern.) Blanch. also have been observed to produce vegetative exten- sions, but not plantlets (Bierhorst, 1971, p. 15). In Selaginella, however, Goebel (1930, p. 657) managed to induce pieces of strobili to develop foliage leaves by placing them on a moist medium

I wish to record my thanks to Prof. A. N. Rao for his assistance in the anatomi- cal study and for critically reviewing this paper; to Mr. J. Wee for bringing the piece of L. phlegmaria to my attention and for technical assistance; and to Mr. D. Teow for the photographs.

LITERATURE CITED

BIERHORST, D. W. 1971. Morphology of Vascular Plants. Macmillan, New York. BOWER, . a 1908. The Origin of a Land Flora. Macmillan, London

—__—.. . Primitive Land Plants. Macmillan, London

commie. <e 1930. Organographie der Pflanzen. Fischer, fet:

TRIARCH Over 50 Years

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AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 3 (1979) 83

Incidence of Epiphytism in the Lycopsids JOSEPH M. BEITEL*

Epiphytism in pteridophytes is well known, with the families Hymenophyl- laceae, Polypodiaceae, and Aspleniaceae each containing a large number of epiphytic species (Copeland, 1947). In a recent article on epiphytism (Madison, 1977), the systematic composition and major features of vascular epiphytes were examined, based on observations of living plants and a survey of specimens in the Gray Herbarium of Harvard University. Sixty-five families of vascular plants were listed, including the Psilotaceae, Ophioglossaceae, Schizaeaceae, Hy- menophyllaceae, Polypodiaceae, Aspleniaceae, Aspidiaceae, and Davalliaceae. However, two major pteridophyte families, the Lycopodiaceae and the Selaginel- laceae, were not included. These omissions in Madison’s enumeration seriously distort the true incidence of pteridophytic epiphytes.

A survey of the herbaria at the University of Michigan, the New York Botani- cal Garden, the Missouri Botanical Garden, the Field Museum, and the Smithsonian Institution revealed a very substantial number of epiphytic species in Lycopodium s.l. In fact, of the 374 species of Lycopodium s.l. examined, 19 species (51%) were epiphytic. All of the epiphytic species are in the segregate genus Huperzia s.l. (commonly treated as Lycopodium subg. Urostachys). Huperzia, including its 93 terrestrial members, accounted for 76% of all the species of Lycopodium s.1. examined. Huperzia species are far more common and diverse in the tropics than in the temperate zone. Only a few species occur In North America north of Mexico, of which the best known are the two terrestrial species L. selago and L. lucidulum, the epipetric species L. porophilum, and the epiphytic species L. dichotomum. ae :

The epiphytic members of Huperzia fall into two classes, with intermediates between the classes grading from one extreme to the other. At one extreme there is little or no differentiation of the sporophylls, the sporangia being bome on Photosynthetic leaves on normal shoots. At the other extreme, there is htt differentiation of the sporophylls, which are also aggregated into dangling tassels. With few exceptions, the branch systems are pendent, the base of the plant being attached by a mass of roots to boughs and crotches of the supporting trees. Gametophytes of epiphytic species are found buried in leaf mold and mosses in the same habitat as the sporophytes (Bierhorst, 1971, P- 22). 4 wih

The large number of epiphytic Huperzia (192 species), when ao cE sulak Madison’s enumeration, ranks this group as the 13th largest family - oe das epiphytes (out of a total of 65 families) and the 12th largest genus © pereen epiphytes (exclusive of the genera of the Orchidaceae, which was no beyond the family level in Madison’s list).

A comprehensive survey of the spike-mosses (Se Sult in the discovery of some epiphytes, although the great terrestrial or epipetric. At least five epiphytes are know

laginella) would probably re- majority of species are n in Selaginella.

* Departs 48109 “Department of Botany, University of Michigan, Ann Arbor, MI .

84 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

Selaginella oregana D. C. Eaton is an isophyllous spike-moss of the Pacific Northwest rainforest, where it typically hangs from deciduous trees such as Acer macrophyllum (Hitchcock et al., 1969, p. 32), and S. zahnii Hieron. is a heterophyllous spike-moss from New Guinea and the Admiralty Islands, where it grows among mosses and filmy ferns on trees (Wagner & Grether, 1948, p. 65). Three species of Selaginella are known to be facultative epiphytes, although they more commonly are epipetric: §. involvens (Swartz) Spring from China, Japan, Taiwan, and the Ryukyu Islands (Walker, 1976, p. 38), S. molliceps Spring from Africa and Madagascar (Tardieu-Blot, 1964, p. 25), and S. versicolor Spring from West Africa (Alston, 1959, p. 16).

LITERATURE CITED ALSTON, A. H. G. 1959. The Ferns and Fern-allies of West Tropical Africa. Crown Agents,

London. BIERHORST, D. W. 1971. Morphology of Vascular Plants. Macmillan, New York. COPELAND, E. B. 1947. Genera Filicum. Chronica Botanica, Waltham, Mass. HITCHCOCK, Cob. CRONQUIST, and M. OWNBEY. 1969. Vascular Plants of the North- west, vol. 1. University of Washington Press, Seattle, Washington. MADISON, M. 1977. Vascular epiphytes: Their systematic occurrence and salient features. Sel-

byana 2:1-13. TARDIEU-BLOT, M.-L. 1964. Flore du Gabon, 8. Ptéridophytes. Muséum National d’ Histoire Naturelle, Paris.

WAGNER, W. H., Jr. and D. F. GRETHER. 1948. The pteridophytes of the Admiralty Islands. Univ. Calif. Publ. Bot. 2417-110,

WALKER, E. H. 1976. Flora of Okinawa and the Southern Ryukyu Islands. Smithsonian Institution, Washington, DC.

REVIEW

THE NORTH DAKOTA, U.S.A.,”” by Aleta Jo Petrik-Ott, Nova Hedwigia Beiheft 61:1-332. 1979. Dm. 100 (ca. $50.00).—Of all the United States regional fern Floras, this 1s probably the most complete in its treatment. Sixty-five pteridophyte taxa are included. Each is dealt with at length by means of a synonymy, a descrip- tion including chromosome data when known, the time of sporulation, the habitat,

annotated list of excluded names and the like will help lay to rest taxa erroneously included in the flora in the past. A glossary, indices to Latin and common names, and an addendum conclude the volume. This work surely is indispensible for critical taxonomic and floristic work in the upper great plains. —D.B.L.

AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 3 (1979) 85

Cyrtomium fortunei in Louisiana and Mississippi GARRIE P. LANDRY,* MICHAEL ISRAEL,** ROBERT SCHWARZWALDER, JR.,* and R. DALE THOMAS***

On September 2, 1978, we observed a few plants of Cyrtomium fortunei J, Smith in the labyrinth of ravines of the Tunica Hills region 2 miles northwest of Weyanoke, in West Feliciana Parish, Louisiana (Israel in 1979, LSU). A second population of approximately 80 plants was discovered on September 10, about three miles from the first location in deep ravines on Ouida Irondale Road 4 miles east from La-66 (Landry 78574, LSU, MICH). During the exploration of ravines on September 17 in Wilkinson County, Mississippi, which is due north of West Feliciana Parish, a single plant of C. fortunei was observed growing at a waterfall on Clark Creek, north of the settlement of Pond (Landry 78593, LSU). Following these discoveries, we found that C. fortunei had been collected by Dr. R. Dale Thomas, 3 miles North of Tunica, Louisiana inOctober, 1974 (Thomas 42096, NLU, LSU). The collection was identified by Dr. A. Murray Evans. The discov- ery of this fern represents a new addition to the presently known floras of Louisiana (Thieret, 1972) and Mississippi (Evans, 1978).

Cyrtomium fortunei is characterized by a short, erect rhizome covered with dark, lanceolate to ovate, acuminate scales. The fronds are pinnate with 23 or more pairs of lanceolate, dull green pinnae having a prominent, darkened midvein. Cyrtomium fortunei is easily recognized by its papyraceous fronds and minutely serrate pinnae, as opposed to the commonly cultivated C. falcatum Presl, which has coriaceous fronds with entire, glossy pinnae.

Nearly all stages of reproduction were observed in the populations of C. for- tunei. The largest plants were 75-90 cm high and had all fertile fronds. But plants 30 cm or less high also had some fertile fronds. Profuse reproduction was evident from the large number of sporelings having two or three fronds. The largest plants were predominantly on the upper sides of the ravines, whereas the smallest were confined to the ravine bottoms. :

The Tunica Hills region is characterized by hills with loess deposits pera from I to 60 m thick. Elevations in the area vary from 60 to more than 90 m a ie sea level. The hills are highly eroded by streams, and form narrow ridge tops aie correspondingly steep to vertical slopes ending in ravines. The beg ask sified as a mixed mesophytic forest association modified by Magnolia a Sas subtropical broadleaf species. In all locations, the associated gee oot aed nolia grandiflora, Fagus grandifolia, Ilex opaca, C ornus florida, an se cceareris gigantea. Other pteridophytes observed include Adiantum aig ee fragilis, Athyrium pycnocarpon, A. thelypteroides, Polystichum acr and Equisetum hyemale.

ens 3

“Department of Botany, Louisiana State University, Baton Rouge, LA ig 70803 ““Department of Entomology, Louisiana State University, Baton senate =e . *“**Department of Biology, Northeast Louisiana University, Monroe, iets

86 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

Cyrtomium fortunei was introduced into cultivation as early as 1866 (Christen- sen, 1930). Although once popular horticulturally, it has been superseded by more attractive species such as C. falcatum. Cyrtomium fortunei in the Tunica Hills region may have originated as an escape from the numerous, old plantation gar- dens of the area. Naturalization of cultivated pteridophytes in Louisiana is not without precedent; Selaginella braunii Baker and S. uncinata Spring both have become established (Landry, unpubl.). Although MacDougal (1976) found C. fortunei on brick walls in Charleston, South Carolina, in our opinion these plants qualify as escapes or as adventives, but are scarcely naturalized. The occurrence of C. fortunei in this region may represent the first report of its naturalization in the United States (W. H. Wagner, Jr., pers. comm). We would like to express our sincere appreciation to Dr. Warren H. Wagner, Jr. for his assistance in preparing this article.

LITERATURE CITED

CHRISTENSEN, C. 1930. The genus Cyrtomium. Amer. Fern J. 20:41-51. EVANS, A. M. 1978. Mississippi Flora: A guide to the ferns and fern allies, Sida 7:282-297. MacDOUGAL, J. M. 1976. Naturalization of Cyrtomium fortunei in North America. Amer. Fern J.

66:25-26. THIERET, J. W. 1972. Checklist of the vascular flora of Louisiana, Part 1. Lafayette Nat. Hist. Mus. Tech. Bull. 2. Lafayette, Louisiana.

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AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 3 (1979) 87

The Fine Structure of the Pre-meiotic Stages of Sporogenesis in Onoclea sensibilis NORMAN P. MARENGO*

Ultrastructural features of the dividing meiocyte of Onoclea sensibilis L. were described by the author (1977), and those of the young spores arising from this cell by Marengo and Badalamente (1978). Since a light microscopic study of this species by the author (1949) showed an apparent decrease in size of cytoplasmic inclusions from the sporangium initial to the spore mother cell, it is of interest to elucidate these changes on the ultrastructural level.

Individual sporangia dissected from young fertile fronds were routinely fixed in glutaraldehyde followed by osmium tetroxide and embedded in Epon (Spurr, 1969). Sections 0.5um thick were cut from individual sporangia and were examined by phase contrast microscopy without staining. From sporangia iden- tified as containing pre-meiotic stages in sporogenesis, thin sections were cut with a diamond knife, stained with uranyl acetate and lead citrate, and examined with an Hitachi HU-11A electron microscope.

The sporangial initials (Fig. /) appear to contain large, osmiophilic bodies (Fig. !,t) which resemble closely the tannin accumulations described by Ledbetter and Porter (1970). The appearance of vacuoles (v) within these bodies is followed by their disappearance in the later stages of sporogenesis, including newly formed Spores. Mitochondria (m) appear hardly larger than those in later stages and in newly formed spores. Their internal structure, however, differs from that of mitochondria in newly formed spores. Cristae appear not to be shelf-like, but rather mitochondrial material separated by internal cavities. This peculiar and distinctive internal structure persists through at least the early prophase of aaa (Fig. 6,m), following which their cristae extend as plates, continuous across the mitochondrion (Marengo, 1977). Fie. 2.1)

The osmiophilic bodies are still present in early sporogenous cells ( = has With an occasional vacuole apparently replacing the tannin mass (Fig. 2,v). : y time the sporogenous cells approach the rounded-up meiocyte part : Fi miophilic bodies have disappeared and lipid droplets appear in the cytoplasm (11g. 31);

In the dividing sporogenous cell in anaphase (Fig. 4), the Se iecara concentrated in the polar regions of the spindle. The distribution : = ie ana- at mitotic anaphase appears to contrast markedly with that of the af indle phase I, where the mitochondria are concentrated at the uate poet of Preparatory to invading it, and form the mitochondrial plate c _ etic interphase following the first division as well as all phases of the sec division (Marengo, 1949, 1977). =

In the early prophase of the first meiotic division (Fig. Polarity is reflected in a concentration of mitochondria at the ba

5), spore mother cell se of a barely

Scr arses : . le, NY 11548. “Department of Biology, C. W. Post College, Long Island University, Greenva

88 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

FIG. 1. Part of a sporangial initial showing dense accumulations of tannin (t), an incipient vacuole (v), an

amyloplast (a), and several mitochondria (m). Base of the initial is at the left, x 11,516. FIG. 2. - Pp htkeneenss cell showing persistent tannin bodies (t), mitochondria (m), and an amyloplast (a), *

N. P. MARENGO: SPOROGENESIS IN ONOCLEA SENSIBILIS 89

FIG. 3. An older sporogenous cell just prior to ‘‘rounding-up”™ into the spore os ccies tannin bodies are absent and lipid droplets (L) occur, * 4,480. FIG. 4. Sporogenous anaphase : mitochondria concentrated in polar regions of the spindle, x 7,484.

90 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

i - ed

tis %,

*

bite 5. Early prophase of the first meiotic division. The polarized chromosomes have a mitochondrial neentration at their base (barely discernible in the marked area), x 12,272. FIG. 6. Enlargement of

ma : . . é : a ; os Fig. 5, showing mitochondria (m) with atypical internal structure and nuclear mem- € (nm) undergoing the disorganization characterizing the onset of meiosis, x 33,750.

N. P. MARENGO: SPOROGENESIS IN ONOCLEA SENSIBILIS 9]

discernible (by electron microscopy) polarized ‘‘bouquet’’ of chromosomes (Marengo, 1949). This pachytene “‘bouquet’’ of the meiotic prophase has been described in animal meiosis (Cicada) by Shaffer (1920) and by Plough (1917) in Rhomaleum.

An enlargement (Fig. 6) of the marked area in Fig. 5 shows mitochondria with still peculiar and distinctive internal structure and a disorganized nuclear mem- brane (nm) reflecting the onset of meiosis.

It appears from the sequence of events described that the large, **plastid-like”’ inclusions of the author’s paper of 1949 were in all probability the osmiophilic accumulations of tannin described herein, since the mitochondria, although un- dergoing an internal reorganization during the pre-meiotic stages of sporogenesis, show no appreciable decrease in size.

LITERATURE CITED

LEDBETTER, M. C. and K. R. PORTER. 1970. Introduction to the Fine Structure of Plant Cells. Springer- Verlag, Berlin, New York. ae MARENGO, N. P. 1949. A study of the cytoplasmic inclusions during sporogenesis in Onoclea

sensibilis. Amer. J. Bot. 36:603-613. am . 1977. Ultrastructural features of the dividing meiocyte of Onoclea sensibilis. Amer. J. Bot. 64:600-601.

———., and M. A. BADALAMENTE. 1978. The fine structure of the newly formed spore of Onoclea sensibilis. Amer. Ferm J. 68:52-54 : PLOUGH, H. H. 1917. Cytoplasmic structures in the male germ cells of Rhomaleum micropterum

Beauv. Biol. Bull. 32:1-12, t./. ‘ol. Bull SHAFFER, E. L. 1920. The germ-cells of Cicada (Tibicen) septemdecim. (Homoptera). Biol. Bull.

38:404—475 ; SPURR, A. R. 1969. A low viscosity epoxy resin embedding medium for electron microscopy. /. Ultrastruct. Res. 26:31-43.

REVIEW

if

“ATLAS OF FERNS OF THE BRITISH ISLES,” edited by A. C. Jermy, et al. 101 pp. Botanical Society of the British Isles and the British Pteridological a ety, London, 1978. ISBN 0-901158-01-1. & 3.50.—This atlas updates Perring an Walters’ ‘‘ Atlas of the British Flora’’ (1962). The nomenclature follows ae Europaea,”’ with a few recent and necessary changes. The 95 maps ne ae for each 10 km square, and will be of interest mostly to students of the se flora. Accompanying each map, however, is a brief paragraph aa. phytogeographical and ecological notes of great value to taxonomists an oe eho turalists. Usually a reference to a specific, detailed paper peareruir ee also is given. The notes on differentiating difficult species and hybri : se ee very welcome. Problems of ecology and taxonomy still awaiting eo ae pointed out. This volume is more than an atlas, and is a welcome addition pteridophyte literature for Britain.—D. B. L.

92 AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 3 (1979)

Apical Dominance in Anemia phyllitidis Gametophytes

THOMAS L. REYNOLDS*

The phenomenon of apical dominance (generally defined as the inhibition of lateral buds or meristems by the apical bud or meristem) occurs generally throughout the plant kingdom and does not appear to be limited to higher vascular plants. Albaum (1938a) demonstrated that the central growth zone, located in the indentation of the roughly heart-shaped prothallus of Preris longifolia, was analo- gous to an apical meristem. If this apical meristem was excised from the prothal- lus, numerous adventitious gametophytes, each with a central growth zone, would arise from the previously inhibited cells. The meristematic zone was considered responsible for the maintenance of apical dominance as long as there was a con- tinuum of living cells between this meristem and the rest of the prothallus. As far as | am aware, the present report is the first to document meristem-induced apical dominance in isolated prothallial tissues where there was a total lack of cellular continuity between the meristem and the affected cells.

MATERIALS AND METHODS

Ripe sporangia of Anemia phyllitidis (L.) Swartz were collected from green- house-grown plants. Prior to the experimental manipulations, the spores were surface sterilized by suspending them for 1 minute in a freshly made solution of 1% sodium hypochlorite (Clorox), followed by several rinses in sterile, distilled water. The final spore suspension (10 ml) was transferred to a sterile culture tube. Subsequent dilutions of this suspension were made until a spore density of 8.0 x 10° spores/ml was obtained. One milliliter aliquots of this suspension were asepti- cally transferred to 125 ml Erlenmeyer flasks containing 67 ml of Miller and Miller’s (1961) sterile, modified Knop’s medium. The spore cultures were placed on a shaker under 400 ft-c of white light for 6 to 8 weeks at 25 + 2°C. The gametophytes which formed served as stock material for the experiments.

Nearly mature, cordate gametophytes were harvested in a sterile transfer room. They were graded for uniformity in size and were cut into equal halves. Ten apical halves each possessing a meristematic notch and ten basal halves each lacking a meristematic notch were placed in a single Petri plate containing filter paper moistened with 10 ml of the sterile medium. These cultures were sealed with Parafilm and returned to 400 ft-c white light for 21 days. During this period, the halved gametophytes were provided fresh medium weekly. Basal halves cultured independently of the apical halves served as a control. Counts of the number of adventitious gametophytes formed were made at the end of three weeks. Each experiment was performed in duplicate, and the entire series of experiments was repeated once.

“Department of Botany, Ohio State University, Columbus, OH 43210.

T. L. REYNOLDS: APICAL DOMINANCE IN ANEMIA GAMETOPHYTES 93

OBSERVATIONS AND DISCUSSION

When excised apical and basal halves were cultured together (Figs. /-3), the formation of heart-shaped, bisexual gametophytes by the basal halves was almost completely suppressed; only one out of 200 explants produced a notched gametophyte in 21 days. The apical halves continued their normal differential growth to form cordate prothallia with both antheridia and archegonia (Fig. 2). Each basal half produced numerous antheridia but no archegonia, became spatu- late in form, and completely lacked a central meristematic zone (Fig. 3). These plants retained their distinct morphology as long as they remained in culture with the meristematic plants. When basal fragments were cultured alone (for 14 days as a control) or were removed for 14 days from the influence of the apical explants, there was a rapid proliferation of lobes with notched meristems and archegonia in 200 out of 200 explants (Fig. 4).

div ae oe @ HP a oh ‘) ee

@

FIG. 1. Division of a nearly

FIGS. 1-4. Apical dominance in the gametophyte of Anemia ice mature, cordate gametophyte into apical and basal portions.

of an apical half — a peat rnd Eventually a single, whole gametophyte forms. bye mg half cultured in association with apical halves forms an amorphic male gametophyte. The ac ce represent antheridia. FIG. 4. Basal half cultured alone produces numerous, adventitious ee phytes (stippled outgrowths), each with a notched meristem. Abbreviations: div = division, m = ap meristem.

These results indicate that, contrary to the well documented reports on ud plants (Miller, 1968), the apical meristem of Anemia can exert BS epttenas the the mature basal cells of the gametophyte without tissue —- sone h two regions. This example of apical dominance is probably cenmunmeanhe amie the production of some diffusible chemical substance by the meristem, PN halves readily form adventitious gametophytes when removed from sep err of of the apex. However, the regulatory mechanism does not require a jun living tissue to be effective.

94 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

The exact nature of the controlling chemical produced by the apex is, as yet, unknown. Albaum (1938b) proposed that the meristem acts as a source of auxin which is transported basipetally in intact gametophytes to inhibit the division of mature cells. Voeller (1966) established that Anemia possesses an antheridogen system and, at least initially, produces two types of gametophytes. The male type is spatulate in outline, lacks a notch meristem, and bears only antheridia. The hermaphroditic type is cordate, possesses a notch meristem, and produces both antheridia and archegonia. In multispore culture, both morphological forms exist at the same time, but the male is induced only by the presence of an antheridogen produced by the meristem of the hermaphrodite. Due to the limits of information provided by this study, the possible role of antheridogen in suppressing or modify- ing the expression of totipotency by differentiated cells cannot be ascertained. It does seem plausible, however, that antheridogen and possibly other substances, such as auxin, produced by the apex may be intricately involved in apical domi- nance and the maintenance of the normal form of intact gametophytes.

LITERATURE CITED

ALBAUM, H. G. 1938a. Pees growth, regeneration, and adventitious outgrowth formation in fern prothallia. Amer. J. 137-44 aan Inhibitions pias - growth hormones in fern prothallia and sporophytes. Amer. J. Bot. 33:

tee . H. 1968. Fern ee iophyics a as eApermental, Patera, et wih 34:361-440.

,and P. M. MILLER. 1961. on the growth and development of the ed of the fern, Onoclea sensibilis. Amer. J. Bot. 48:154—- 159.

VOELLER, B. R. 1966. Gibberellins and growth in ferns. pp. 247-258. In Proceedings of the Interna- tional Symposium on Plant Stimulation, Bulgarian Academy of Science Press, Sofia.

SHORTER NOTES

A NEW COMBINATION IN ASPLENOSORUS.—Recently, Taylor and Moh- lenbrock (Amer. Fern J. 67:66. 1977) published the name Asplenium x herb- wagneri for the hybrid between Asplenium pinnatifidum and A. trichomanes. This is appropriate as long as the Walking Fern is considered as a species of Asplenium. However, when the Walking Fern is considered as the separate genus Camptosorus, as it commonly is, all taxa containing genomes of both Asplenium and the Walking Fern must be considered as part of the hybrid genus Asplenosorus. In the above-mentioned case, the hybrid between Asplenium (Asplenosorus) pinnatifidum—itself the fertile hybrid between Camptosorus rhizophyllus and Asplenium montanum—and Asplenium trichomanes has one set of chromosomes from each of Camptosorus rhizophyllus, Asplenium montanum, and A splenium trichomanes. The name for this hybrid thus becomes Asplenosorus x herb-wagneri (Taylor & Mohlenbrock) Mickel.—John T. Mickel, New York Botanical Garden, Bronx, NY 10458

AMERICAN FERN JOUNAL: VOLUME 69 NUMBER 3 95

CHEILANTHES ALABAMENSIS NEW TO KENTUCKY.—In the past, Cheilanthes has been represented in the flora of Kentucky by C. lanosa (Michx.) D. C. Eaton andC. feei Moore. Both are xerophytes restricted to various types of dry, rocky outcrops. Cheilanthes lanosa is widespread in the southern and west- ern counties of Kentucky. on both calcareous and non-calcareous substrates: C. feei is known only from crevices in blocky Silurian dolomite cliffs in Bullit County, where it reaches the southeastern limit of its range. Now a third species of Cheilanthes, C. alabamensis (Buckl.) Kunze, can be added to the Kentucky flora. The Alabama Lip Fern is frequent on limestone cliffs and ledges from Tennessee, Missouri, and Kansas south to northern Florida, Louisiana, Texas, and Mexico. We suspected that it might occur in Kentucky due to its presence in Montgomery County, Tennessee on bluffs along the Red River.

On 15 April 1979, we searched for likely sites on the Dot topographic sheet covering south-central Logan County. We chose to examine cliffs along a sharp bend of the Red River where it loops into Kentucky. The high, limestone cliffs were predominantly sheer and very dry, not the typical habitat of C. alabamensis. After a short search, we found an outcrop of well weathered limestone with an abundance of small cavities and recesses. Soon we located a small clump of juvenile C. alabamensis plants. About a hundred yards further, we discovered a large specimen from which we made collections (Cranfill 3761, APSC, KY). Plants associated with the Lip Fern include Aquilegia canadensis, Arabis laevigata, Heuchera pubescens, and Taenidia integerrima, as well as the ferns Asplenium platyneuron, A. resiliens, Cystopteris bulbifera, C. tennesseensis, Pel- laea atropurpurea, and P. glabella. In addition, there was an abundance of both epiphytic (on Ulmus americana) and epipetric Polypodium polypodioides. ;

No additional plants of C. alabamensis were discovered during an intensive, two-hour search. The apparent rarity of this fern probably stems from two related factors: the northern locality with its harsh winters and the abundance of Heuch- era and Polypodium competing at the site. Winter die-back, which is not normal in C. alabamensis, was evident in both clumps, especially in the juveniles. The winter climate may weaken the gametophytes and young sporophytes sufficiently so that they can barely compete with the Heuchera and Polypodium. Although abundant on the cliff face, these plants are most robust in the niche that the fern finds most suitable. These factors could account in part for the frequency of all three plants. An intensive investigation of similar cliffs in Todd and Christian counties would very likely reveal additional stations for the Alabama Lip Fern.—&. Cranfill, Division of Biological Sciences, University of Michigan, Ann Arbor, MI 48109 and Laurina Lyle, Department of Botany, Austin Peay State University, Clarksville, TN 37040.

96 AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 3 (1979)

SEXUALITY IN ASPLENIUM RESILIENS.—The widespread species A. resi- liens Kunze ranges from eastern, central and southwestern United States to southern South America. In the southeastern United States, cytological investiga- tion by Wagner (Amer. Fern J. 56:3-17. 1966) has shown it to be an apogamous triploid producing 32 spores per sporangium. This condition is strongly correlated with an apogamous life cycle, whereas the 64-spored condition indicates normal sexual reproduction.

In preparing a treatment of Asplenium of the Chihuahuan Desert region, I counted spores from the sporangia of 50 different collections of A. resiliens depos- ited at TEX/LL. These ranged from central Texas and southern Arizona to south- em Mexico, with a few from South America, but most were from the Chihuahuan Desert region of Texas and Mexico. Several unopened sporangia per specimen, from separate individuals when possible, were counted. Almost all specimens yielded counts of ca. 32 spores per sporangium (rarely ca. 64 aborted spores). However, in two specimens from the Chisos Mountains of west Texas, ca. 64 spores per sporangium were counted in each of five sporangia from each sheet. The spores were smaller than in the other populations, but otherwise were normal.

Apparently this is the first report of sexuality in this species. The putative sexual plants are morphologically indistinguishable from the apogamous ones. Thus, both sexual and apogamous forms appear to occur in Asplenium resiliens , a situation similar to that described by Wagner, Farrar and Chen (Amer. Fern J. 55:171-178. 1965) for Pellaea glabella var. glabella.

The 64-spored specimens, both from Brewster County, Texas, are: Green Gulch, Chisos Mountains, 28 Mar 1937, Warnock 608; and South River [Rim], 26 Aug 1937, Warnock 608.

Although both bear the same collection number, the former bears the collector’s signature on the label, and the latter label is typed and includes at least one clear error, It is likely that the latter label is in error and that both sheets represent a single collection from Green Gulch. Collections of A. resiliens from Green Gulch made during other years (Correll 13666: Warnock & Tharp 6837; Wendt & Lott 2144, 2144A) and numerous other Chisos Mountains specimens examined uni- formly display 32 spores per sporangium.—Tom Wendt, Rama de Botanica, Col- egio de Postgraduados, Chapingo, Edo. México, México.

NOTEWORTHY PUBLICATIONS OF THE CHRONICA BOTANICA

BEDDOME, R. H. The Ferns of British India. Includes Bangladesh, Burma, Ceylon, Malaya, Nepal, Pakistan, and Tibet. Set of 2 vols. Reprint ed. 1973. Available CB (cloth bound) or LB (half-leather with gold lettering). CB Rs. 285: LB US $55.

. Supplement to the Ferns of Souther India and British India.

se ees tht in CB Rs. 85; LB US $18.50. ————.. Handbook to the Ferns of British India. Reprint ed. 1970. CB Rs. 100; LB US $21. NAYAR, B. K. Companion to R. H. Beddome’s Handbook. 1974. CB Rs. 75; LB US $14.50 Annales Cryptogamici et Phytopathologici, vol. 8 . Fern Flora of India. The first taxonomic account of the fern flora since Beddome’s classical work ‘‘Handbook to the Ferns of British India’ published in 1883. An illustrated floristic account containing keys to identifications as well as lucid descriptions of species, genera, and families of Filicopsida. Prepared by the foremost pteridologist of India. About 1000 pages. November 1979,

CB Rs. 550; LB US $115.

The Chronica Botanica Co. E/2 Jhandewala Ext., New Delhi-110055, India

AMICRIGAN Volume 69 FERN a JOURNAL October-December, 1979

QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY

Disjunct netnasiptent of Isoetes macrospora in Southern Tennessee W. MICHAEL DENNIS, A. MURRAY EVANS, and B. EUGENE WOFFORD 97

Three New Elaphoglossums from Guatemala JOHN T. MICKEL 100

The Role of Temperature in the Vegetative Life Cycle of Isoétes butleri JERRY M. BASKIN and CAROL C. BASKIN 103

Gametophyte Morphology of the Fern Genus

Drynariopsis ( Polypodiaceae) SUBHASH C ore

: B. GENA, T. ae BHARDWAJA, A New Species of Selaginella from India C. 4A. K. YADAV. 119 Shorter Note: A pomp nami Medium 122

for Growing Fern Pr

iRI ROTANICAC mrss 102

Review Jan 16 1980 122 American Fern Journal GARDEN LBSARY 123 Index to Volume 68 Z

124 Erratum

The American Fern Society Council for 1979

RICHARD L. HAUKE, Dept. of Botany, University of Rhode Island, Kingston, R.1. 02881. President ROBERT M. LLOYD, Dept. of Botany, Ohio University, Athens, Ohio 45701. Vice-President LESLIE G. HICKOK, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916. Secretary JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916. Treasurer JUDITH E. SKOG, Dept. of Biology, George Mason University, Fairfax, Va. 22030. ecords Treasurer DAVID B. LELLINGER, Smithsonian Institution, Washington, D.C. 20560. Editor-in-Chief

JOHN T. MICKEL, New York Botanical Garden, Bronx, N.Y. 10458. Newsletter Editor American Fern Journal EDIT

OR DAVID B. LELLINGER Smithsonian Institution, Washington, D. C. 20560

ASSOCIATE EDITORS DAVID W. BIERHORST .-Dept. of Botany, University of Massachusetts, Amherst, Mass. 01002 GERALD J. GASTONY ‘ss1+s+eee+Dept. of Biology, Indiana University, Bloomington, Ind. 47401 JOHN T. MICKEL New York Botanical Garden, Bronx, New York 10458

The **American Fer Journal’’ (ISSN 0002-8444) is an illustrated quarterly devoted to the general study of fems. It is owned by the American Fern Society, and published at the Smithsonian Institu- tion, Washington, DC 20560. Second-class postage paid at Washington.

Matter for publication and claims for missing issues (made within six months of the date of issue) should be addressed to the Editor.

Changes of address, dues, and applications for membership should be sent to Dr. J. E. Skog, Dept. of Biology, George Mason University, Fairfax, Va. 22030.

Orders for back issues should be addressed to the Treasurer.

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). Extracted offprints, if ordered in advance, will be furnished to authors at cost, plus postage.

Back volumes $5.00 to $6.25 each; single back numbers of 64 .25; 65-80 pages, $2.00 each; over 80 pages, $2.50 each, Plus shipping. Ten percent di f six volumes or more; postage additional.

ages or less, $1 ra

Library Dr. John T. Mickel, New York Botanical Garden, Bronx, New York 10458, is Librarian. Members may borrow books at any time, the borrower paying all shipping costs.

Newsletter stesane Mickel, New York Botanical Garden, Bronx, New York 10458, is editor of the —— Fiddlehead Forum.”’ The editor welcomes contributions from members and non- members, including miscellaneous notes, offers to exchange or purchase materials, personalia, hor- ticultural notes, and reviews of non-technical books on fems.

oo Spore Exchange T. Neil D. Hall, 1230 Northeast 88th Street, Seattl : ae , le, Washington 98115, is Director. res exchanged and collection lists sent on request. vat . oe

Gifts and Bequests

Otte ak r ae

a eq i y I } d to ot interested in ferns. Botanical books, back issues of the Joumal, and cash or other gifts are always welcomed, and are tax-deductible. Inquiries should be addressed to the Secretary.

AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 4 (1979) 97

Disjunct Populations of Isoétes macrospora in Southeastern Tennessee

W. MICHAEL DENNIS*, A. MURRAY EVANS**, and B. EXGENE WOFFORD**

Isoétes macrospora Dur. is a northern species widely distributed throughout eastern Canada and the northeastern United States west to Minnesota (Pfeiffer, 1922; Wherry, 1961). Recently, populations of this species were found in the Little Tennessee and Hiwassee Rivers of southeastern Tennessee. The only other report of this species from the southeastern United States is that of Svenson and Griscom (1935), who cite it from northern Virginia. This report was accepted by Massey (1944, 1960) and by Harvill, Stevens, and Ware (1977) for the Virginia Flora, but was not mentioned by either Fernald (1950) or by Gleason and Cronquist (1963).

There had been some confusion concerning the county of this Virginia collec- tion. The voucher located in the Brooklyn Botanical Garden Herbarium is labeled ‘‘Isoetes macrospora Dur., Shallow Water. Passage Creek. Macrospores Average 650 uw. Frederic Co., VA. L. Griscom and H. K. Svenson 5561, July 2, 1933.” Massey (1944) cited the collection as being from Passage Creek in Shenandoah County. This is verified by C. E. Stevens (pers. comm), who corresponded with Svenson, and indicated that Svenson still recalls the locations along Passage Creek as being in Shenandoah County. Recently, this location was examined by A. M. Evans, and plants of J. macrospora were found to occur frequently in the shallow, cobble-bottomed pools of the Elizabeth Furnace area.

Both the 1935 and 1978 collections from Passage Creek and the recent collec- tions from the Little Tennessee and Hiwassee Rivers represent significant disjunc- tions from the glaciated, northern, contiguous range of J. macrospora to the unglaciated Ridge and Valley province of Virginia and Tennessee. The Passage Creek population is ca. 300 miles from the northern range, and the Tennessee populations are ca. 450 miles from the Virginia population.

In the Little Tennessee and Hiwassee Rivers, the known populations of I. macrospora are limited to sections of river between upstream dams and im- pounded waters of mainstream reservoirs of the Tennessee River. The habitat is essentially riverine and consists of an alternating series of pool and riffle areas. Water depth is generally shallow (<2 m), but varies depending on releases from upstream dams. These releases also help to maintain a cool water temperature throughout the year (15 to 19° C in July, 1978). Individual plants of/. macrospora were found rooted in sand-filled crevices of the rocky, cobble substrate in the moderate- to fast-flowing water of pool areas that are 0.1 to 1.5 m deep. Plants growing in areas of weak currents have slightly erect to recurved leaves which spread from the corm in a symmetrical ring. In areas of strong current, the leaves are prominently recurved and fixed by the downstream current, giving the plants a striking, swept-back aspect.

*Tennessee Valley Authority, Division of Water Resources, Muscle Shoals, AL 35660. **Department of Botany, University of Tennessee, Knoxville, TN 37916. Volume 69, number 3, of the JOURNAL was issued October 5, 1979.

98 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

Other species found growing in association with J. macrospora in the Little Tennessee River include Podostemum ceratophyllum, Callitriche heterophylla, Potamogeton tennesseensis, P. epihydrus, P. nodosus, Nitella sp., and Fontinalis sp. Perhaps the most notable associate is the rare red alga, Boldia erythrosiphon Herndon. This alga was first described in 1964 from Big Walker Creek in Giles County, Virginia (Herndon, 1964) and presently is known from a limited number of localities in the southeastern United States. Interestingly, B. erythrosiphon has been found in the three known southeastern localities for /. macrospora.

Isoétes macrospora in Virginia and Tennessee may represent a recent range extension or it may be a disjunct relic of past floras. The widespread distribution of many aquatics has been attributed to long-range dispersal by waterfowl. Ac- cording to Sculthorpe (1967), Jsoétes rhizomes and leaf bases are often eaten by waterfowl, and the disjunct populations of /. macrospora in Virginia and Tennes- see might have resulted from this dispersal mechanism since migratory waterfowl do frequent the Little Tennessee and Hiwassee Rivers. The distance from the northeastern United States populations of /. macrospora (ca. 700 miles) and the requirement of introduction of both mega- and microspores of these heterospor- ous organisms into a habitat conducive for sporophyte establishment and de- velopment are constraints on the hypothesis of recent introduction by waterfowl. The alternative hypothesis is that these disjunct populations may be relics of a past flora in a region that no longer provides generally favorable environmental conditions for this species. This hypothesis is made less likely by the fact that the Tennessee populations occur in two rivers that in recent history have become controlled by upstream dam releases, which have significantly altered the flow and temperature characteristics of the rivers. Surveys of other eastern Tennessee watercourses that offer seemingly suitable habitat for /. macrospora have failed thus far to uncover additional populations. The explanation for the occurrence of this northern species in the southeast thus remains an intriguing problem.

Specimens cited:

NNESS : Monroe Co.: Little Tennessee River at Jones Ferry, 26 Jul 1978, B. E. Wofford & W. M. Dennis 78-133; Little Tennessee River at Tomatlo Ford, 26 Jul 1978, B. E. Wofford & W. M. Dennis 78-134; Little Tennessee River at SW end of Davis Island, 26 Jul 1978, B. E. Wofford & W. M. Dennis 78-135. Polk Co.: Ca. 1.1 mi from hwy. 411 bridge over Hiwassee River, 27 Jul 1978, B. E. Wofford &

21 Aug 1978, B.E . Wofford & A. M. Evans 78-168; Shallow shoals of Hiwassee River, 1.6 road miles NW of intersection of hwy. 30 and state road 2518, 21 Aug 1978, B. E. Wofford & A. M. Evans 78-169. : GINIA: 2 Co.: Passage Creek, 8 Nov 1978, A. M. Evans s.n.

pecimens are deposited in the University of Tennessee Herbari i ; i will be distributed at a later date. a eee

LITERATURE CITED

shite M. L. 1960. Gray’s Manual of Botany, 8th ed. American Book Company, New York,

alas acc oh M., Jr., C. E. STEVENS, and D. M. E. WARE. 1977. Atlas of the Virginia Flora, Part , Pteridophytes through Monocotyledons. Virginia Botanical Associates, Farmville, VA.

W. M. DENNIS ET AL.: ISOETES MACROSPORA IN TENNESSEE

HERNDON, W. R. 1964. Boldia: A new Rhodophycean genus. Amer. J. Bot. 51:575—581. MASSEY, A. B. 1944. Virginia Ferns and Fern Allies. Va. Poly. Inst. Bull. 37(7).

. 1960. The Ferns and ‘‘Fern Allies”’ of Virginia, 3rd ed. V.P.I. Agric. Exten. Serv. Bull. 256. PFEIFFER, N. E. 1922. Monograph of the Isoetaceae. Ann. Missouri Bot. Gard. 9:79-233. SCULTHORPE, C. D. 1967. The Biology of Aquatic Vascular Plants. William C. Clowes and Sons,

London. SVENSON, H. K. and L. GRISCOM. 1935. Isoetes macrospora in the Shenandoah Valley. Amer. mJ. 25:70-71. WHERRY, E. T. 1961. The Fern Guide; northeastern and midland United States and adjacent Canada. Doubleday, Garden City, NY.

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100 AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 4 (1979)

Three New Elaphoglossums from Guatemala JOHN T. MICKEL*

In preparing a treatment of the Guatemalan species of Elaphoglossum, I have found that three of the 37 taxa recognized apparently are unnamed. This is a fairly modest number of novelties; the percentage will be much greater in Mexico, southern Central America, and the Andes from Venezuela to Bolivia. Because of the large number of species in the genus (over 500), the complexities of their taxonomy, the very few regional treatments, and the lack of a usable monograph, the taxonomy of the genus is in chaos and a modern treatment is sorely needed. Efforts to correct this situation are currently underway.

sof Elaphoglossum apodum var. latum Mickel, var. nov.: °

apart, running at a 60-70° angle; hydathodes lacking; blade scales subulate, orange, especially concentrated on the midvein and margin, some scales on the abaxial blade surface, very sparse on the adaxial surface; blade also with minute, erect, glandular hairs; fertile fronds not seen, but ca. 4 as long as the sterile ones to 13 cm long, 1 cm broad) in var. apodum, with linear, orange scales on the abaxial midvein, lacking among the sporangia; spores not seen, but with low

“ . H TYPE: Along Rte. 5 between Semococh and La Laguna on road to Chajmayic, Depto. Alta Verapaz, Guatemala, alt. 500 m, 10 May 1942, Steyermark 46368 (F). PARATYPES: GUATEMALA: Alta Verapaz: Vicinity of Cubilquitz, 1.5-2 mi S of Cubilquitz, Steyermark 44475 (F, US). Izabal: Along Rio Frio and tributaries, Steyermark 41554 (F, US). COSTA RICA: Puntarenas: Osa Peninsula, Lloyd 459 (NY), Seidenschnur 108 and 109 (both NY). COLOMBIA: Magdalena: Santa

Marta, H. H. Smith 2688 (NY). DOMINICAN REPUBLIC: Azua: S iej pepe sd : San Juan, Loma La Vieja, Ekman H

somewhat the figure for E. backhousianum Moore, which was described from living material from Mexico. Although that species is Clearly allied toE. apodum and has the frond form of var. latum, it has a stouter rhizome (over 2 cm diam.), larger fronds (45-75 cm tall, 7.5-10 cm broad), and darker brown blade scales than does var. latum.

Material of E. apodum from much of South America is distinctly different from any of the above. The fronds are very long with a narrow base and probably represent yet a different variety or species.

*New York Botanical Garden, Bronx, NY 10458.

J. T. MICKEL: NEW ELAPHOGLOSSUMS FROM GUATEMALA 101

a19F Elaphoglossum lanceum Mickel, sp. nov.

49%

_ Rhizomata breviter repentia vel suberecta, 1 mm diametro; rhizomatis paleae lineari-lanceolatae castaneae, 7 mm longae; phyllopodia nulla; frondes fasciculati, (

basi cuneatae, margine integrae vel crenulatae; nervi conspicui liberi, simplices vel 1-furcati, 2-3 mm inter se distantes, angulo 35—40° abeuntes; hydathodi con-

“TYPE: Southeast shoulder of Cerro Zempoaltepetl, Dto. Mixe, Edo. Oaxaca, Mexico, in rich, wet, pine-oak cloud forest below Patio de Arena, alt. 8200 ft, Mickel 4900 (NY; isotypes F, UC, US).

PARATYPES

MEXICO: Oaxaca: Dto. Ixtlan, 65 km N of Ixtlan, 44 km S of Valle Nacional, Mickel 5165 (NY). EL SALVADOR: Chalatenango: Cerro EI Pital, Seiler 16] (NY). GUATEMALA: Quezaltenango: Fuentes Georginas: Volcan de Zunil, Standley 85996 (F); Volcan Zunil opposite Sta. Maria de Jestis, Steyer- mark 35147 (US). HONDURAS: Lempira: Montafia de Celaque, Hazlett 2600 (F).

Epiphytic in wet forests at 1500-2500 m elevation. This species is distinct in the lance-shaped blade, thin texture, scales among the sporangia, and the highly per- forated spore crests. Generally the plants are small, but the one large Guatemalan specimen closely resembles E. buchii and E. smithii of the West Indies. However, the Guatemalan plant has a thinner texture, somewhat more blade scales, and spores with highly fenestrated crests in contrast to the unperforated crests of the West Indian species.

Elaphoglossum stellatum Mickel, sp. nov. BR:

Rhizomata breviter repentia, ca. 2 mm diametro; rhizomatis paleae linear- lanceolatae castaneae integrae, ca. 2 mm longae; phyllopodia conspicua; frondes fasciculati, 6-21 cm longi, 0.4—-0.9 cm lati; stipites laminae dimidio breviores, pilis stellatis sparse vestiti; laminae steriles lineares, apice acuminatae, basi cuneatae; nervi vix conspicui liberi, simplices vel 1-furcati, ca. 1 mm inter se distantes,

angulo 60-70° abeuntes; hydathodi nulli; laminae dorso pilis stellatis sparse, veniralter paleis minimis ciliatis vestitae; frondes fertiles quam yee spe breviores angustiores longiusque stipitati, usque 10 cm longae, 0.5 cm — s i laminae subaequilongus; lamina pilis stellatis tantum secus costam abaxialem pilosula; sporae monoletae, breviter cristatae.

FYPE: Volcan Tecuamburro, along trail to San Francisco Tecuamburro on

summit of volcano, N of Chiquimulilla, Depto. Santa Rosa, Guatemala, Steyer- mark 33155 (F; isotype US).

PARATYPE: Rio Samald, below Zunil, Depto. Quezaltenango, Guatemala, Steyermark 35012 (F).

Terrestrial or epipetric, at 2000-2500 m elevation. Elaphoglossum stellatum closely resembles a small, slender specimen of E. petiolatum, but can be sco guished from it by the lack of glandular dots on the abaxial surface, the lac s broader scales on the adaxial surface and stipe, and the lighter rhizome scale

102 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

color. It is distinguished from E. pilosum by the darker rhizome scales, small blade size, and narrow blade. I have seen specimens from South America that are similar to this (small, narrow, with stellate hairs on both blade surfaces), but the specimens lacked names and had somewhat different rhizome scales.

I gratefully acknowledge the support provided by National Science Foundation grant DEB 77-25582. I also thank Dr. Rupert Barneby for his kind help with the Latin descriptions.

“*THE HOME GARDENER’S BOOK OF FERNS,” by John Mickel with Evelyn Fiore. Ridge Press/Holt, Rinehart & Winston, New York. 256 pp. $7.95.—The science of botany, with its multitude of terms, can be a bit formidable to the uninitiated. John Mickel’s book does an excellent job of demystification. On the strictly botanical side, it discusses fern morphology, taxonomy, reproduction, and ferns found in various natural habitats. On the horticultural side, it treats the general principles of indoor and outdoor cultivation and deals specifically with reproducing ferns from spores and by vegetative means. Useful lists of fern societies and of public fern displays are provided. Although the text is marred by occasional small errors, these are more than compensated for by the delightfully light-hearted style in which the book is written. The author’s enthusiasm about

This book is invaluable for those who are just starting out to grow ferns. Since it touches on so many interesting botanical details which do not usually come to the attention of non-botanists, it also will be useful and interesting to more experi- enced growers.—D.B.L.

AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 4 (1979) 103

The Role of Temperature in the Vegetative Life Cycle of Isoetes butleri JERRY M. BASKIN and CAROL C. BASKIN*

Isoétes butleri Engelm. occurs in Georgia (Boynton, 1902), Tennessee, Mis- souri, Kansas, Arkansas, Oklahoma (Pfeiffer, 1922; Fernald, 1950), Alabama (Reed, 1965; Kral, 1973), and Kentucky (Baskin & Baskin, 1978).

Throughout its range it is restricted to shallow soils over limestone. In Tennes- see, /. butleri is restricted to cedar (limestone) glades of the Central Basin and occurs in very shallow soils that range from 5 to 10 cm deep down to bedrock. In the Isoétes habitat, these shallow soils are waterlogged much of the time from late autumn to mid-spring, but frequently are below the wilting coefficient during the remainder of the year (Freeman, 1933). Common associates of J. butleri in the Tennessee cedar glades include Leavenworthia spp., Nothoscordum bivalve (L.) Britt., Ophioglossum engelmannii Prantl, Sporobolus vaginiflorus (Torr.) Wood, and Scutellaria parvula Michx.

In the middle Tennessee cedar glades, leaves of J. butleri come above the soil surface in early to mid-March, and micro- and megasporangia develop on the adaxial surfaces of the leaf bases in early to mid-April. Senescence of the leaves begins in mid-May, and by mid-June the leaves as well as the roots are dead. By this time, the spores are mature. The in in an inactive (q t) state i the soil until autumn, when growth of new roots and leaves is initiated. Growth of new roots begins in early October and is soon followed by leaf growth. By mid October, roots are 0.5—3 cm long and leaves are 0.5—1.5 cm long. Roots continue to grow until mid November, but there is little additional leaf growth until the following spring, when the leaves emerge above the soil surface. ee

Thus, the timing of the vegetative life cycle in. butleri is one of the physiologi- cal adaptations that allows this drought-intolerant species to persist in its summer-dry, winter-wet habitat. In this study we investigated the role of tempera- ture in regulating the timing of root and leaf growth in the vegetative life cycle of /. butleri.

GENERAL METHODS

Corms of I. butleri were collected from a cedar glade in Rutherford County, Tennessee on 10 June 1975 and 13 June 1977, after all the leaves and roots were dead. Corms were washed gently and the dead leaf bases (containing intact micro- or megasporangia on the adaxial surfaces) and dried, shriveled roots were ue moved. Within two days after collection, the corms were placed ona layer of so cm thick in waxed paper cups (7 cm deep and 8.2 cm in diameter) with drainage holes in the bottoms. Five corms were placed on the layer of soil in each cup. Some of the corms were covered with 2.5 cm of soil; others were not buried. In the latter case, the cups were covered with aluminum foil.

*School of Biological Sciences, University of Kentucky, Lexington, KY 40506.

104 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

Studies on root and leaf growth were carried out in an unheated greenhouse in Lexington, Kentucky and/or in temperature-controlled incubators in the labora- tory. In the unheated greenhouse, the windows were kept open at all times, and temperatures were near those out-of-doors. Continuous thermograph records of air temperatures in the unheated greenhouse were used to calculate mean daily maximum and minimum temperatures and the number of hours that corms were exposed to “‘chilling’’ temperatures between 0.5 and 10°C. From 1 May to 1 September, the soil in which corms were buried was watered to field capacity once each week; during the remainder of the year, it was watered daily except when frozen in winter. These watering regimes were given to approximate the soil moisture conditions that corms would likely be exposed to in the cedar glade habitat.

In the laboratory, four incubators were programmed on 12/12 hr thermoperiods

to approximate habitat temperatures from early spring to late autumn: March, 15/6° C; April, 20/10° C; May and June 30/15° C; July and August, 35/20° C; Sep- tember, 30/15° C; October, 20/10° C; November, 15/6° C (U.S. Dept. Comm., 1965). For the buried corms, leaves were considered to be emerged when their tips were 5 mm above the soil surface; for the nonburied corms, leaves and roots had to be 5 mm long before they were considered to have initiated growth. Nonburied corms were examined for growth at intervals in a darkroom under a green safe lamp, which consisted of a green fluorescent tube wrapped with a No. 24 dark green sheet of ‘‘Cinemoid.’’ Peak intensity of the light as determined with an ISCO Model SR Spectroradiometer was at 550 nm, and essentially all the energy impinging on the corms was between 500 and 600 nm. There was no radiation above 600 nm.

PROCEDURES AND RESULTS

Leaf and root growth under natural temperature regimes.—Fifteen corms col- lected on 10 June 1975 were kept in the unheated greenhouse until November 1976. During the spring of 1976, each plant went through its normal above-ground annual growth cycle. The leaves emerged in early spring, remained green and healthy until late spring, and then yellowed and died in early summer. On 15 September, 1 and 15 October, and 15 November 1976, the corms were removed from the soil to determine if leaf and root growth had been initiated; after each examination, the corms were reburied. Mean daily maximum and minimum tem- peratures were calculated for the 15-day period prior to the various dates when the corms were examined.

On 15 September, there were no visible signs of leaf or root growth, although the corms had been kept continuously moist since 1 September. Mean daily maximum and minimum temperatures for the previous 15 days were 31.0 and ie C, respectively. On 1 October, none of the corms had initiated leaf growth,

ut four of them had initiated root growth. Temperatures for the last 15 days of September were 25.6 and 16.2° C. By 15 October, 10 corms had leaves that were

BASKIN & BASKIN: LIFE CYCLE OF ISOETES BUTLERI 105

5-17 mm long and all corms had roots that were 6-30 mm long. For the first 15 days of October, temperatures were 24.4 and 13.6° C. By 15 November, all corms had leaves that were 1—1.5 cm long and roots that were 1—4 cm long. Tempera- tures during the first half of November were 14.7 and 6.6° C.

Forty corms collected on 10 June 1975 were kept in the unheated greenhouse until 1 October 1976. On this date, 20 corms were transferred to a greenhouse that was heated in the winter, while the other 20 were retained in the nonheated greenhouse. Temperatures in the heated greenhouse ranged mostly from 20 to 30° C during the day and from 15 to 20° C at night. The cups containing the corms were examined for emergent leaves at weekly intervals in both greenhouses until 1 April 1977, and the soil in all cups was kept continuously moist.

In the unheated greenhouse, there was no leaf emergence prior to 16 February 1977. Between 17 and 23 February 1977 (when mean daily maximum and minimum temperatures were 16.2 and 6.4° C, respectively), leaves on eight corms emerged. Between 24 February and 2 March (21.7 and 8.9° C), leaves from seven corms emerged; between 3 and 9 March (17.8 and 7.8° C), leaves from three corms emerged; and between 10 and 16 March (14.8 and 3.0° C), leaves from the remain- ing two corms emerged. In the heated greenhouse, leaves from only one corm emerged, between 9 and 16 March. When the experiment was terminated on I April 1977, the corms in the heated greenhouse were examined and all were still alive.

Initiation of leaf and root growth.—Rates of initiation of leaf and root growth were compared for corms before and after they were exposed to summer tempera- ture and soil moisture regimes in the unheated greenhouse. Eighty corms (16 cups containing 5 corms each) were covered with soil and transferred to the unheated greenhouse, where they remained until 1 September 1977. A second set of 80 corms was not buried, and cups containing them were covered with aluminum foil. Twenty corms (4 cups of 5 corms each) were placed at each of the four alternating temperature regimes. On 1 September, the 80 buried corms in the unheated greenhouse were removed from the soil and placed on a layer of soil in cups which then were covered with aluminum foil. Twenty corms were placed at each tem- perature. None of the 80 corms kept in the unheated greenhouse during summer showed any signs of root or leaf growth on | September. Beginning with incuba- tion at the four thermoperiods, both sets of corms were examined under the ose safe light at 5 to 10 day intervals for leaf and root growth, at which time the sot

was watered if needed. Observations were made on both groups of corms for 170

days. i There was no root or leaf growth on any corm placed at 30/15 or = al ay either June (June corms) or September Co on tea ga epee ath 20/10 and 15/6° C) in September int lated autumn temperatures ( ture in June Fig. 1). At 20/10° C, only 12 of the June corms initiated root growth within 170 obi : vate von September corms produced roots within 35 days. At 15/6° C, 1 oon cet placed at this temperature in June and September produced baal ef sh Pe took 120 days for the June corms but only 35 days for the September .

106 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

contrast, the rate of initiation of leaf growth was slower for September corms than for June corms (Fig. ]). At 20/10° C, eleven of the June corms but only seven of the September corms initiated leaf growth. At 15/6° C, leaves were produced by all of the corms, but growth began sooner for those placed in the incubator in June than in September.

LEAVES 20 started in June 18 0 15/6°C 16F 4 20/10 14

started in September '2F 0 15/6°C 4 20/10

NUMBER OF CORMS THAT INITIATED GROWTH

0 ev 80" "60 80" "100 "120" 40" 166 TIME (DAYS)

FIG. 1. Growth of leaves and roo = j summer temperature and soil oi al nero tens Mase 3 pica sao oF 3/20? Fey ey Osan = .. rms collected in June.—Corms collected on 10 June alienating Gocsann Ps of soil, and 20 corms were placed at each of the four were Guam ee € regimes. The soil was watered regularly, and the cups leaves emerged fro emergent leaves at 5- to 10-day intervals for 205 days. No oie m corms incubated at 30/15 or 35/20° C, and no leaf emergence

corms at 15/6 and 20/10° C occurred until after 80 days (Fig. 2). Leaves had

BASKIN & BASKIN: LIFE CYCLE OF ISOETES BUTLERI 107

emerged from all 20 corms at 15/6° C after 178 days but from only 13 corms at 20/10° C after 205 days.

Effects of chilling on subsequent leaf emergence.—Corms collected on 10 June 1975 were buried in cups of soil and placed in the unheated greenhouse. On 13 September, 1 October, and 1 December 1975 and on 2 January and | February 1976, 20 corms were placed at each of the four thermoperiods, and the cups were examined for emergent leaves at 5 to 10 day intervals for 95 days. After removal to

20r 15/6°C = ‘ 18r = a 16- ” E = * 14 3} ont I2e i S 1OF Oo W Oo ge ew Brie, le Spr ar = ob 30/15 and 35/20 o-+ jo¢_»-0d-0-4-00-0! 010-00 9-0! see e—o O. 80. 100 (20.140. 160. 0 ...<O00 TIME (DAYS) from buried fIsoétes butleri placed at the four temperatures on 12

FIG. 2. Emergence of June 1975.

the incubators, the soil in the cups was kept continuously moist. From the ther- mograph records, we calculated the number of hours that corms in the nonheated greenhouse were exposed to temperatures between 0.5 and 10° C. These tempera- tures were chosen because they are known to promote vernalization, bud break, and afterripening of seeds (Leopold, 1964; Stokes, 1965; Salisbury & Ross, 1969). Corms of/. butleri transferred to the incubators on 13 September had received 0 h of chilling; on 1 October, 22 h; on 1 December, 486 h; on 2 January, 926 h; and on 1 February, 1,218 h. After 95 days, there was no leaf emergence from corms placed in the incubators in September. However, leaves of corms transferred from the unheated greenhouse to the incubators during autumn and winter asa a increase in ability to grow (Fig. 3). At 30/15 and 35/20° C, only nine or fewer o : re corms had emergent leaves after 95 days, regardless of when they were placed in

108 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

the incubators. At 20/10 and 15/6° C, there was an increase in the number of emergent leaves as well as an increase in the rate of emergence. For example, whereas 6 and 11 of the corms placed at 20/10 and 15/6° C, respectively, on 1 October 1975 had emergent leaves after 90 days, all corms placed at these temper- atures on | February 1976 had emergent leaves after 10 and 30 days, respectively.

30/15

20 igt | Oct. 1975 | Dec. 1975 15/6°C 16F wo 14 = ° 2 12+ 15/6°C - 4 (1OF 3 > 4 a. ek 20/10 5 30/15 oOo a 4+ Se aa fe w 35/20 30/15 e20/10 Ww 2r oo2—0 S02 oe) a 35/20 = fe) o-ct-¢L l l l l l 1 ' f f l l l | l l > . 2 Jan. 1976 & = 20 © 15/6°C ce SP S '8F 20/10 ws & '6F ~ 14h w © 12- Ww «10k oO = > z

TIME (DAYS)

FIG. 3. Emergence of leaves from

buried corms of Isoétes butleri vwinedhaes Hiecware S butleri placed at the four temperatures on 1975.

mergence for corms placed at the four temperatures on 13 September

DISCUSSION

When the vegetative growth cycle of J. butleri ends in June with the senescence of roots and leaves, the corms will produce new leaves and roots if they are incubated at autumn (20/10 and 15/6° C) but not at summer (35/20 and 30/15° C) temperatures (F igs. 1 and2). However, even when corms are incubated at 20/10 or 15/6° C, initiation of leaf and root growth is very slow. At 20/10° C, three corms had roots within 15 days and only one had leaves after 60 days. At 15/6° C, one corm had roots after 25 days and one had leaves after 30 days (Fig. 1). At both

BASKIN & BASKIN: LIFE CYCLE OF ISOETES BUTLERI 109

20/10 and 15/6° C, the first leaves to emerge from buried corms did so only after 80 days (Fig. 2). This low temperature requirement for leaf and root growth in /. butleri prevents vegetative growth during the summer because summer habitat temperatures are too high. Thus, although soils in the cedar glade habitat of /. butleri may be moist for several days following summer rains, the corms do not produce new leaves and roots. If roots and leaves were formed during brief periods of favorable soil moisture in summer, they would no doubt be killed within a few days after the soil again dried to the wilting coefficient. Occasionally a short period of favorable temperatures for leaf and root growth follows a summer rain (Baskin & Baskin, 1970). However, the slow rate of growth initiation in corms of I. butleri in summer prevents the formation of leaves and roots.

Root and leaf growth are delayed until temperatures in the habitat become suitable for growth and completion of the vegetative life cycle. In the nonheated greenhouse, there was no root or leaf growth until after 15 September. From 15 September to 1 October, the mean daily maximum and minimum temperatures were 25.6 and 16.2° C, respectively, and during this time none of the corms pro- duced leaves and only four of them produced roots. Most of the corms initiated root and leaf growth between 1 and 15 October when mean daily maximum and minimum temperatures were 24.4 and 13.6° C, respectively. These temperatures correspond very closely to the mean daily maximum (23° C) and minimum (12° C) air temperatures for October in middle Tennessee (U.S. Dept. Comm.., 1965). This explains why root and leaf growth of J. butleri in middle Tennessee begins in October. Also, in autumn the soil in the cedar glades stays moist for many days following rains; therefore, soil moisture does not limit growth and survival of new leaves and roots. i

Data presented in Fig. / indicate that exposure to summer temperature and soil moisture regimes enhanced the subsequent rate of root, but not leaf, initiation in autumn. In the unheated greenhouse in autumn, corms produced roots earlier than leaves, and roots grew faster than leaves. In the field in autumn, roots grow rapidly and attain much of the potential growth before winter. However, leaf growth is slow and leaves do not emerge from the soil surface until spring.

In plants of /. butleri, chilling resulted in a widening of the temperature range, and an increase in rate, of leaf emergence after corms were transferred from the unheated greenhouse to the incubators. With an increase in the number of hours of chilling, there was an increase in the number of corms with emergent leaves and in the rate of leaf emergence at 15/6°C. However, the number of days from 13 September 1975 to 100% leaf emergence did not decrease (Fig. 3). Whereas none of the corms placed at 15/6° C in September and only 11 of those placed at bar thermoperiod on October had emergent leaves after 90 days, all corms placed at 15/6° C on 1 December, 2 January, and 1 February had emergent leaves after 90, 55, and 30 days, respectively. The number of days (from 13 September) for re leaf emergence for corms placed at 15/6°C on 1 December, 2 January, o Be February was essentially the same, i.e., 168, 165, and 171 days, respective * > 20/10° C, the number of corms with emergent leaves as well as the rate of on emergence increased (Fig. 3) and the number of days from 13 September to 100%

110 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

leaf emergence decreased. Whereas none of the corms placed at this thermoperiod in October and December had emergent leaves after 90 days, leaves emerged on all corms placed at 20/10° C on 2 January and 1 February in 55 days or less (Fig. 3). The number of days (from 13 September) for 100% leaf emergence for corms placed at 20/10° C on 2 January and 1 February was 165 and 141, respectively. At 30/15° C and 35/20° C, leaves emerged on one to nine corms for all dates of transferral except 13 September. This may indicate that chilling widened the tem- perature requirement for initiation of leaf growth in a small number of plants, but another possible explanation is that growth had been initiated at lower tempera- tures in the nonheated greenhouse and merely continued at the high ther- moperiods in the incubators.

The changes in the physiological responses of the plants caused by chilling during winter allow leaves to emerge in the field in spring at temperatures that, considering time-temperature relationships, were unsuitable for their emergence the previous autumn. In the nonheated greenhouse, leaf growth was initiated in early October, but there was no leaf emergence during autumn although daily temperatures ranged from 15—20° C maximum and 5-10° C minimum. However, leaves on some corms emerged in the spring between 17 and 23 February, when mean daily maximum and minimum temperatures were 16.2 and 6.4° C, respec- tively, as well as between 24 February and 2 March, when temperatures were 21.7 and 8.9° C, respectively.

LITERATURE CITED

BASKIN, J. M. and C. C. BASKIN. 1970. Germination of winter annuals in July and survival of the seedlings. Bull. Torrey. Bot. Club 98:272-276.

. 1978. Geographical distribution of Isoétes butleri in the southeastern United States. Amer. Fern J. 68:7-8.

BOYNTON, C. L. 1902. Notes from a collector's field-book. Biltmore Bot. Stud. 1:143-150.

FERNALD, M. L. 1950. Gray’s Manual of Botany, 8th ed. Amer. Book, New York.

FREEMAN, C. P. 1933. Ecology of cedar glade vegetation near Nashville, Tennessee. J. Tenn. Acad. Sci. 8:143-228.

KRAL, R. 1973. Some notes on the flora of the southern states, particularly Alabama and middle Tennessee. Rhodora 75:366-410.

LEOPOLD, A. C. 1964. Plant growth and development. McGraw-Hill, New York.

PFEIFFER, N. E. 1922. Monograph of the Isoetaceae. Ann. Mo. Bot. Gard. 9:79-233, t. 12-19.

REED, C. F. 1965. Isoetes in southeastern United States. Phytologia 12:369-400.

Soe F. B., and C. ROSS. 1969. Plant physiology. Wadsworth Publ., Belmont, California. » P. 1965. Temperature and seed dormancy, pp. 746-803. JnW. Ruhland, ed. Encyclopedia of plant physiology, vol. 15/2. Springer-Verlag, Berlin, Heidelberg, and New York.

UNITED STATES DEPARTMENT OF COMMERCE, WEATHER BUREAU. 1965. Climatography of the United States 86-35. Decennial census of the United States climate. Climatic summary of the United States. Supplement for 1950 through 1960. Washington, D.C.

AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 4 (1979) 111

Gametophyte Morphology of the Fern Genus Drynariopsis (Polypodiaceae)! SUBHASH CHANDRA*

As the gametophyte morphology of various fern groups has become better known during the past several decades, it has become increasingly apparent that gametophytes provide dependable morphological criteria for taxonomic and phyletic studies. Bower (1923-28) and Holttum (1949) pointed out that the com- parative morphology of fern gametophytes can be of significance in understanding different phyletic groups. Stokey (1951, 1960), Atkinson and Stokey (1964), and Nayar and Kaur (1968, 1969, 1971) discussed the role of fern gametophytes in taxonomic and phyletic studies. According to Nayar and Kaur (1968, 1969), the patterns of spore germination and gametophyte development characterize various taxonomic groups, and so provide dependable data for comparative morphology.

Nayar (1954, 1961, 1965), Nayar and Kachroo (1953), and Bajpai (1964) studied gametophyte morphology of some drynarioid ferns. Until now, no detailed studies of the gametophytes of the monotypic genus Drynariopsis have been made, al- though Nayar (1965) reported that Drynariopsis gametophytes show Drynaria- type development. This study of Drynariopsis heraclea gametophyte development was made to compare the gametophytes of this primitive genus with those of other drynarioid ferns.

Spores were collected from the fernery of the Natural Science Research Center, University of the Philippines. Gametophytes were raised on sterilized Knop’s nutrient agar medium in petri dishes (Nayar, 1962). The cultures were maintained at 24+2° C under 600 ft-c of light from four fluorescent lamps placed horizontally above the culture dishes. All observations on morphology and development of the gametophytes are based on these laboratory cultures. To study cellular structure, the gametophytes were mounted in an acetocarmine solution, which induced par- tial plasmolysis of the cells and so rendered the cell outlines clear. Drawings were made using a camera lucida.

OBSERVATIONS

The spores of D. heraclea are densely granulose, monolete, perineless, plano- convex in lateral view, oblong in polar view, and of medium size. This is con- firmed by Nayar and Devi (1964) and by Nayar (1965). Fresh spores contain green plastids and prominent, yellowish oil globules. The spores germinate rather slowly; the germ filaments emerge in 3-4 weeks. At germination, the ot = off a small, lens-shaped rhizoid initial cell at the proximal pole by a wall in parallel to the equatorial plane of the spore (the wall perpendicular to the polar axis of the spore). This cell soon forms the first rhizoid (Fig. 1) and essa the direction of the proximal pole (Fig. 2). The distal, daughter gametophytic

*National Botanical Research Institute, Lucknow 226001, India. Ge ‘Portion of a Ph.D. thesis presented in October 1978 to the Faculty of the Graduate School, University of the Philippines, Diliman, Quezon City, Philippines.

AMERICAN FERN JOURNAL: VOLUME 69 (1979)

112

oo 6 e {7\ a e] aig esenew, eS e Be 7 EEE ERA RLY, wp cr 0) BOOT EY QEON WX SY

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@

oo ey SH rae egueme SES re Stee See re QPEL eae 2

& a) Be e

S. CHANDRA: GAMETOPHYTE MORPHOLOGY OF DRYNARIOPSIS 113

initial cell enlarges and divides (Fig. 2) by a wall perpendicular to the first wall, one that is parallel to the polar axis of the spore. The germ filament elongates along the equatorial plane of the spore (Figs. 2 and 3) perpendicular to the first rhizoid. Due to the physical obstruction provided by the spore coat, the germ filament is deflected and the rhizoid is pushed away to one side. The rhizoid, which usually emerges first, is dark brown and devoid of chloroplasts. This is termed Vittaria-type spore germination by Nayar and Kaur (1968, 1971). The germ filament elongates, becomes 4-8 cells long (Fig. 4), and is composed of short, stout, barrel-shaped, densely chlorophyllous cells. The basal cell is short, often rather bulbous, and may bear two or more rhizoids. The germ filaments often are branched (Figs. 5, 9, 10, 12, 19, and 30), sometimes profusely so, with the branches developing into separate gametophytes, in contrast to Pseudodrynaria (Nayar, 1954) and Drynaria (Nayar & Kachroo, 1953), in which branching has been reported to be rare. Development of the gametophytic plate is initiated when the germ filament is 4-8 cells long, and may be either Drynaria-type or Aspidium- type.

Drynaria-type development.—The terminal cell of the germ filament and two or three cells behind it divide transversely and longitudinally to produce a spatulate gametophytic plate (Figs. 5 and 6). The plate often becomes 5-10 cells wide and broadly ovate, but is devoid of any organized meristem (Fig. 7). Only rarely is an apical cell established early (Figs. 16-18), in which case the young gametophyte becomes cordate. For the most part, an obconical apical meristem cell is differ- entiated later by two oblique divisions of one of the marginal cells at the anterior end of the gametophytic plate (Fig. 8). The apical cell undergoes a considerable number of divisions and the gametophyte becomes cordate (Fig. 12) before the apical cell is replaced by a multicellular meristem (Figs. 9-11 ). In Drynaria-type development, the establishment of an apical meristematic cell usually is much delayed (Figs. 16-18), and the gametophytes usually develop hairs on the margin when the gametophytic plate becomes cordate; the younger gametophytes are naked. In contrast, in Drynaria rigidula, an apical meristematic cell is established early, at the second division of the terminal cell, often initiating it (Nayar, 1965).

Young gametophytes with slightly asymmetrical meristemati gametophytes. FIG. 36. Superficial, stalked hair. FIG. 37. shaped gametophyte.

Marginal hair. FIG. 38. Young, strap-

114 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

As in Aglaomorpha (Nayar, 1965), the young gametophytes of Drynariopsis often are broader than long, whereas those of Drynaria and Pseudodrynaria are elongate-oblong at this stage (Nayar, 1965). Soon the apical meristematic cell becomes positioned at the base of the apical notch. It is then replaced by a multicellular meristem (Fig. 13) in the usual way, by a transverse wall in the meristematic cell followed by vertical walls in the outer cell.

Under cultural conditions, the gametophytes become distinctly cordate with a multicellular meristem and cluster of rhizoids on the lower surfaces (Figs. 14 and 15) about 10-12 weeks after spore germination. The midrib is initiated when the gametophytes are about four months old. The midrib is thin and very conspicuous. The mature gametophytes are cordate, broader than long, grow flat on the sub- strate, and have a deeply notched apex, as in the other species of drynarioid ferns. The meristem consists of 3-5 elongate cells perpendicular to the surface. The lobes are composed of uniformly thin-walled, densely chlorophyllous, polygonal cells arranged in more or less radiating rows. In contrast to Merinthosorus (Baj- pai, 1964), the walls of the wings do not possess collenchyma-like thickenings at the corners. As in other drynarioid ferns (Nayar & Kachroo, 1953; Nayar, 1954, 1961, 1965; Bajpai, 1964), the marginal cells of young gametophytes develop uni- cellular, papillate hairs, which begin to form only after the gametophytes have become distinctly spatulate. The hairs are thin-walled, cylindrical, with a rounded anterior end, and are chlorophyllous. They originate as a protuberance of one of the superficial cells, which elongates and is cut off as a hair. When young, the hairs have a swollen tip with a prominent nucleus surrounded by a few vacuoles on all sides (Fig. 37). Simple, club-shaped hairs like those found in Aglaomorpha, Merinthosorus, Photinopteris, and Pseudodrynaria rarely are found on the sur- face of Drynariopsis gametophytes (Fig. 36). Branched, club-shaped, polypodioid hairs as found in other drynarioid ferns (Nayar, 1965) and most other poly podiaceous genera, are absent in Drynariopsis.

Drynariopsis gametophytes are attached to the substrate by unicellular, elon- gate rhizoids. On adult thalloid gametophytes, the rhizoids generally are restricted to the lower surface of the midrib. Rhizoids generally are absent on the wing and margins of cordate-thalloid gametophytes, but in strap-shaped gametophytes, marginal rhizoids are often very frequent and in some cases are characteristically tec as on the gametophytes of the Grammitidaceae and some

olypodiaceae (Nayar & Raza, 1970). The rhizoids are soft and nearly hyaline; the rhizoid wall is markedly colored, and often is reddish brown, even at an early developmental stage.

Aspidium-type development.—In contrast to Drynaria-type gametophyte on- pr ; the Aspidium-type is variable in Drynariopsis with regard to the sequence ne po . All variations characteristic of Aspidium-type development have t ot served in Drynariopsis. The most common is that the filament terminates “ae unicellular hair (Fig. 20). The terminal cell becomes beaked, and the protrud- ics Pipes is cut off by a transverse wall. Plate formation is initiated by cells

ind the terminal cell dividing longitudinally (Figs. 2/-23). Sometimes the penultimate cell of the germ filament also is sluggish. A broad, usually lopsided

S. CHANDRA: GAMETOPHYTE MORPHOLOGY OF DRYNARIOPSIS 115

plate is formed by differential cell division behind the sluggish anterior region (Figs. 23 and 26). The larger side often bulges out, pushing the terminal hair to one side. All cells except the basal cell may divide longitudinally and take part in plate formation. An obconical meristematic cell is differentiated laterally in the thallus by two oblique divisions in one of the marginal cells on the more expanded side of the plate (Figs. 23 and 26). It may be formed either early during plate formation or only after the plate becomes several cells wide. This type of gametophyte de- velopment has been reported in Christiopteris (Nayar, 1967), Merinthosorus (Baj- pai, 1964; Nayar, 1965), and Platycerium (Bauke, 1878; Orth, 1936; Stokey & Atkinson, 1954). Sometimes the terminal cell of the germ filament, after producing a hair, divides longitudinally. One of the daughter cells then produces a hair (Fig. 27). The hair-bearing cell is sluggish, but the other daughter cell actively divides by an oblique division to form a meristematic cell (Fig. 28, M). The resulting plate may be slightly asymmetrical (Figs. 29 and 3/). Marginal, unicellular hairs are produced continuously, whether a meristematic cell is differentiated early or not. In some gametophytes, a plate is initiated prior to hair formation. The terminal cell divides longitudinally (Fig. 24), with one daughter cell initiating plate formation. A lateral meristematic cell forms in the plate (Fig. 25). The other daughter cell of the terminal cell remains sluggish, and may bear a hair.

The gametophyte grows by the activity of the meristematic cell. Since this cell is lateral, the young gametophyte is lop-sided. The asymmetry is more marked when meristematic cell formation is delayed. Soon the meristematic region becomes notched and apical by unilateral growth (Fig. 7). As the gametophytes grow, however, the asymmetry is lost, and the adult thallus is symmetrically cordate (Fig. 15). Gametophyte development beyond the establishment of an apical meri- stematic cell is similar to that in Drynaria-type.

Ameristic gametophytes.—Occasionally ameristic gametophytes occur in cul- ture (Figs. 32-35). In such cases, a single, obconical meristematic cell may not be developed at all, and the marginal cells on one side develop a multicellular meri- stem directly. Ameristic gametophytes also have been reported in Christiopteris (Nayar, 1967) and in Merinthosorus (Bajpai, 1964).

In some cases, the gametophytes elongate by diffuse growth, become irregular and strap-like with a rounded apex, and often bear profuse marginal rhizoids (F : é 38). No organized meristem is ever differentiated, but the cells constituting the anterior region of the strap-like gametophyte are smaller in size and gps more actively than do the cells in the posterior half. No midrib is produced. T € rise stem is neither very active nor well defined. All cells of the anterior region o : oi meristematic activity. This type of gametophyte development, — Kaulinia-type by Nayar and Kaur (1969, 1971), has been reported in many g of the Polypodiaceae (Nayar & Kaur, 1969).

Mature gametophytes.—The ee i , ae 14-16 weeks after spore germination. The mature g cae bear sex organs aie pesca on the lower surface of the meen seo ti are the common type found in the Polypodiaceae (Nayar, ste a sheeple superficial, but on gametophytes growing under crowded conditions,

heraclea reach maturity about tophytes are cordate and

116 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

may develop along the gametophyte margins. The antheridia are small, sub- globose, and have an undivided opercular cell and a funnel-shaped basal cell. Schlumberger (1911) and Schmelzeisen (1933) have reported the occasional occur- rence of a divided cap cell of the antheridium in Drynariopsis, but these were not observed in the present study. Archegonia are produced after midrib formation and usually are found mixed with antheridia. The archegonia are the usual type reported in advanced leptosporangiates (Stokey 1951; Nayar, 1962), with a short, slender neck composed of four rows of cells having three or four short cells in each row. The neck is curved away from the apex of the gametophyte, and the neck canal cell is binucleate and swollen toward the tip at maturity.

DISCUSSION

The drynarioid ferns are phyletically a problematic group, and commonly are ascribed a microsorioid ancestry. Among the drynarioid ferns, Drynariopsis usu- ally is regarded as the most primitive genus and is considered to be nearest the group’s probable ancestor (Copeland, 1947; Holttum, 1947, 1954; Nayar & Devi, 1964; Nayar, 1965).

Drynariopsis exhibits several peculiarities in its gametophyte morphology which are otherwise unknown in the Polypodiaceae. The gametophytes are naked in early stages, and the obconical meristematic cell may be differentiated quite early in development, as in Microsorium, or it may be differentiated only after the formation of a spatulate plate, as in Drynaria and Pseudodrynaria (Nayar, 1954, 1959, 1961, 1965; Nayar & Kachroo, 1953). Besides the common Drynaria-type gametophyte development, which is characteristic of the drynarioid ferns and most of the Polypodiaceae, Drynariopsis gametophytes exhibit all the variations characteristic of Aspidium-type development. This is uncommon in the Poly- podiaceae, but has been reported in Merinthosorus drynarioides (Bajpai, 1964; Nayar, 1965), Christiopteris (Nayar, 1967), and Platycerium (Bauke, 1878; Orth, 1936, Stokey & Atkinson, 1954). Among ferns of other phyletic lines, this pattern is characteristic of Blechnaceae, Davalliaceae, Dryopteridaceae, and Olean- draceae (Nayar & Kaur, 1969, 1971). However, a strap-shaped gametophyte simi- lar to that of Drynariopsis is found in C hristiopteris and in several microsorioid genera of the Polypodiaceae, e.g., Colysis, Pleopeltis normalis (Nayar, 1962), Dendroglossa (Nayar, 1963a), Leptochilus, Paraleptochilus (Nayar, 1963b), and Lepisorus (Nayar & Raza, 1970).

Drynariopsis heraclea is unusual among most of the other drynarioid ferns in its gametophyte development, but shows similarities with Merinthosorus in having Aspidium-type development. The possibility of segregating Merinthosorus from other drynarioid ferns on the basis of gametophyte morphology, as suggested by Nayar (1965), can be ruled out. Copeland (1947) considered Merinthosorus to have evolved from Microsorium through Aglaomorpha meyeniana, but the similarities | in gametophyte development of Merinthosorus with those of

Drynariopsis indicate that Merinthosorus is nota derivative of Aglaomorpha. And Holttum S contention (1954) that Merinthosorus has been derived from Drynariopsis is supported.

S. CHANDRA: GAMETOPHYTE MORPHOLOGY OF DRYNARIOPSIS 117

On the basis of comparative gametophyte morphology, it is likely that Drynari- opsis is more intimately related to Merinthosorus than to any other drynarioid fern, and possibly the latter is derived directly from Drynariopsis. The similarities with Aglaomorpha in vegetative structures and in fertile fronds presumably are due to parallel evolution.

I am greatly indebted to Prof. Prescillano M. Zamora, Department of Botany, University of the Philippines, Diliman, Quezon City, for constant encouragement and helpful suggestions.

LITERATURE CITED

ATKINSON, LENETTE R. and ALMA G. STOKEY. 1964. Comparative morphology of the gametophyte of the homosporous ferns. Phytomorphology 14:51-70. BAJPAI, N. 1964. Gametophyt phology of Merintt Copel. J. Indian Bot. Soc. 43:549-555. BAUKE, H. 1878. Zur Kenntniss der sexuellen G tion bei der Gattungen Platycerium, Lygodium und Gymnogramme. Bot. Zeit. 36:753-760, 769-780. BOWER, F. O. 1923-28. The Ferns, vols. I-III. Cambridge Univ. Press, Oxford. COPELAND, E. B. 1947. Genera Filicum. Ronald Press, New York. HOLTTUM, R. E. 1947. A revised classification of Leptosporangiate ferns. J. Linn. Soc. (Bot.) 53:123-158. ————. 1949. The classification of ferns. Biol. Rev. 24:267-296. ————. 1954. Flora of Malaya, vol. II. Ferns of Malaya. Gov’t. Printing Office, Singapore. NAYAR, B. K. 1954. Studies in Polypodiaceae-II. Contributions to the morphology of Pseudo- drynaria coronans (Wall.) C. Chr. Phytomorphology 4:379-390. . 1959. Studies in Polypodiaceae-VI. Further observations of the morphology of Drynaria Bory. J. Univ. Gauhati 9:95-103. ————. 1961. Ferns of India II. Drynaria and Pseudodrynaria. Bull. Natl. Bot. Gard. 56:1-30. —————.. 1962. Morphology of the spores and prothalli of some species of Polypodiaceae. Bot. Gaz. 123:223—232. ; : . 1963a. Contributions to the morphology of some species of Microsorium Link emend. Copel. Ann. Bot. n.s. 27:89-100. : . es. Contributions to the morphology of Leptochilus and Paraleptochilus. Amer. J. Bot. 50:301-308. . 1965. The gametophyte and juvenile leaves of the drynarioid ferns. Bot. Gaz. raspation . 1967. Morphology of the spores and prothallus of Christiopteris tricuspis. Amer. Fern. J. nape 1964. Spore morphology of Indian ferns-III. Polypodiaceae. Grana Palynol. er ey Phen oe. 1953. Studies in Polypodiaceae-I. Contributions to the Seti of Drynaria Bory: D. quercifolia (L.) J. Sm. and D. propinqua (Wall.) J. Sm. Phytomorphology 3:411-423. _ Spore germination in homosporous ferns. J. Palynol. 4:1-14. a eae pre ee prothallial development in homosporous ferns. Phytomor- hology 19:179-188. é : py KAUR. 1971. Gametophytes of homosporous ferns. Bot. ee iy i ————, and F. RAZA. 1970. The prothalli of some Polypodiaceae-II. J. ore eile ae ORTH, H. 1936. Morphologische und physiologische Untersuchungen an Farnp

25:104—150. i d die SCHLUMBERGER, 0. 1911. Familienmerkmale der Cyatheaceen und der Polypodiaceen, unt! te

Beziehungen der Gattung Woodsia und verwandter Arten zu

102:383—414.,

118 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

SCHMELZEISEN, W. 1933. Beitrage zur Entwicklungsgeschichte der Prothallien einiger Marattia- ceen, Cyatheaceen und Polypodiaceen. Flora 127:46—80

STOKEY, ALMA G. 1951. The contribution by the gametophyte to the classification of the homo- sporous ferns. Phytomorphology 1:39-58.

. 1960. Multicellular and branched hairs on the fern gametophyte. Amer. Fern J. 50:78-87.

, and LENETTE R. ATKINSON, 1954. The gametophyte of five species of Platycerium.

Phytomorphology 4:165-172.

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AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 4 (1979) 119

A New Species of Selaginella from India C. B. GENA, T. N. BHARDWAJA and A. K. YADAV*

During a recent survey of pteridophytes of southeastern Rajasthan, India car- ried out in September 1977, we collected a Selaginella which is different from all the known species of this genus. It is being described now as a new species, and is an addition to the 59 species of the genus already recorded from India (Alston, 1945; Panigrahi & Dixit, 1966a, p. 11, 1966b, 1967, 1968; Bole & Almeida, 1977).

Selaginella rajasthanensis Gena, Bhardwaja & Yadav, sp. nov.

Selaginella reticulatam affinis. Caulis prostratus, alterne ramificans: caules la- terales non ramificantes praeter basalem tantum qui semel tantum dichotome ramificans. Folia et sporophylla dimorpha et serrata. Folia lateralia elliptica, mediana vero ovata; megasporophylla ventralia, cordata. Megaspora tuberculis rotundis. Microsporophylla dorsalia, oblique lanceolata, costa cum carina ad- axiali, margine serrata. Microspora exino laevi et crista sinuata et triradiata.

Stem glabrous, prostrate, bearing rhizophores throughout, 1.5-2.5 cm long, the posterior part (S—7 mm) unbranched, the anterior part bearing alternate branches up to 1.0 cm long, the first one from the base invariably dichotomously branched in mature plants (Fig: 1). Leaves serrate, obtuse, dimorphic, the lateral ones (Fig. 2) elliptical or oval, 1.5 mm long, 1 mm wide, the median ones (Fig. 3) ovate, 1 mm long, 0.5 mm wide. Strobili single at the end of every lateral branch, up to 7 mm long, bearing two ventral rows of megasporangia and | to 3 microsporangia or none at all dorsally. Mega- and microsporophylls dissimilar. Megasporophylls cordate, 1.5 mm long, 1 mm wide, serrate, mucronate, borne in the same plane as the lateral leaves. Megaspores light yellow, trilete, spherical, 224-240 um in diam., with many small, round tubercules on the exine. Microsporophylls ob- liquely lanceolate, 2 mm long, 1 mm wide, serrate, obtuse with a serrately mar- gined adaxial keel or flap on the midrib (Fig. 5), borne in the same plane as the median leaves, with only a few (1 to 3) bearing microsporangia. Microspores brown, trilete, spherical, 38 4m in diam., smooth, the triradiate ridge wavy (Fig. 6).

TYPE: Kundakhoh, Shahabad, Kota, Rajasthan, India, growing on an isolated moist rock, Sept 1977, C. B. Gena & A. K. Yadav (PUN 2610) (PUN; isotypes: Pteridophyte Biology Lab., Govt. College, Ajmer, India, No. PBL/77/S1-6/28/ 671, B, BM, CAL, K, LWF, NY, US).

The species is confined to Rajasthan, India and is fertile from September to November. Selaginella rajasthanensis resembles most .s reticulata (Hook. & Grev.) Spring in general appearance, but it is morphologically quite different from the latter in the following respects: habitat (prostrate vs. erect), branching (un- common, except for the lowest vs. repeated), rhizophore abundance (common vs. absent), lateral leaf shape (elliptical or oval, serrate, obtuse vs. ovate-oblong, aristate, acuminate), median leaf shape (ovate, serrate, obtuse vs. elliptical, aris- tate, acuminate), strobilus size (long vs. short), disposition and number - ati gia (megasporangia 12-14, ventral, microsporangia 1-3 or none at - . > megasporangia 1-3, basal, microsporangia 9-12, distal); megasporophy pe

, Ajmer 305001, India. *Pteridophyte Biology Laboratory, Department of Botany, Government College, Aj

FIGS

Late x

AMERICAN FERN JOURNAL: VOLUME 69 (1979)

. 1-6. Details of S. rajasthanensis Gena, Bhardwaja and Yadav. FIG. 1. Habit, x 2.5. FIG. 2. ral leaf, x 35. FIG. 3. Median leaf, x 85. FIG. 4. Megasporophyll, x 25. FIG. 5. Microsporophyll,

5. FIG. 6. Microspore, x 430.

C. B. GENA, ET AL.: A NEW SELAGINELLA FROM INDIA 121

(cordate, serrate, mucronate vs. deltoid, aristate, cuspidate); microsporophyll shape (obliquely lanceolate, serrate, obtuse, with an adaxial, serrately margined keel on the midrib vs. lanceolate, aristate, accuminate); megaspore size and or- namentation (larger, the exine with many small, round tubercles vs. smaller, the exine granular); microspore ornamentation (smooth with a wavy triradiate ridge vs. spiny with a straight triradiate ridge).

This new species also differs conspicuously from the recently described S. tama-montana Serizawa (1978) from Japan in branching pattern; the —_ rtary dense mats. The number and disposition of mega- and microsporangi tama-montana are rather like those of S. reticulata. The strobili of the aaa species resemble those of S. reticulata in the number and disposition of mega- and microsporangia, but sporophyll shape and size is characteristically like that of vegetative leaves in the former. There are significant differences in mega- and microspore size and ornamentation also. These three species may be identified as

ollows:

. Plants erect, rooting at the base only, lacking rhizophores. Plants repeatedly branched; leaves and sporophylls aristate; strobilus with 1-3 basal megasporangia and many distal microsporangia; megaspores granular; microspores spiny, the triradiate ridge straight ..................... S. reticulata

. Plants prostrate or creeping, with rhizophores present throughout 2. Lateral branches not divided, except for the lowermost, which dichotomize only once; leaves and

sporophylls serrate; strobili with many ventral megasporangia and only a few, irregularly scat- tered microsporangia; megaspores with round tubercles; microsporophylls with a serrately ma gined keel on the midrib; microspores smooth, the triradiate ridge wa vy ue S. rajasthanensis 2 Plants frequently branched; leaves and sporophylls ll with 1-4 basal megasporangia and 6-10 distal microsporangia; megaspores with many reticula- ee microspores with uneven tubercles, the triradiate ridge straight .......... . tama-montana Thanks are due to Principal A. B. Mathur and Prof. A. N. Parashar, Govern-

ment College, Ajmer, for providing laboratory facilities, and to Prof. K. M.

Matthew of St. Joseph’s College, Tiruchirapalli, India, for rendering the Latin

description. The University Grants Commission, New Delhi, India provided fi-

nancial assistance for survey of Rajasthan Pteridophytes.

LITERATURE CITED

ALSTON, A. H. G. 1945. An enumeration of the Indian species of Selaginella. Proc. Natl. Inst. Sci. India 11:211-235. BOLE, ~ oh and M. R. ALMEIDA. 1977. Four new species of pteridophytes from Bombay Presi- . J. Bombay. Nat. Hist. Soc. 74:320-325. Stat G. and R. D. DIXIT. 1966a. tes on twelve interesting species of Selaginella from India. Symp. Indian Sci. Congr., Chandigarh. . 1966b. Studies in the systematics of inet Selaginella—I

_—

—

II. Proc. Natl. Acad. Sci. 36:102-

; . : :222-233, f. eee. SGT. Studies in the systematics of Indian Selaginella-II. J. Indian Bot. Soc. 46:222-233, 1

: | te 8. . 1968. Studies in the systematics of Indian Selaginella-I. Proc. Natl. Inst. Sci. India 09

34:1 : . Bot. SERIZAWA, S. 1978. A new species of Selaginella from central Honshu of Japan. J. Jap : 3

122 AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 4 (1979)

A SIMPLIFIED NUTRIENT MEDIUM FOR GROWING FERN PROTHALLIA. —Germinating fern spores and culturing prothallia for scientific or educational purposes is usually accomplished by sowing the sterilized spores on Knudson’s agar (Bot. Gaz. 79:345-379. 1925; Bot. Gaz. 101:721-758. 1940). The preparation of this medium requires precise weighing of small amounts of essential minerals which are dissolved in water, to which agar is added to solidify the medium. For some time, my laboratory has been successfully using a medium consisting of a solution of 15 gm of agar per liter of water in which is dissolved one commercial nutrient plant tablet (Black Magic Plant Food Tablets, Black Magic, Inc., Box 578-P, Hermosa Beach, CA 90254). The resulting medium provides mineral nutri- ent yielding fern prothallia in numbers and size equal to those grown on conven- tional nutrient media requiring precise quantitative mineral additives. —Norman P. Marengo, Dept. ef Biology, C. W. Post College, Long Island University, Greenvale, NY 1154

AMERICAN FERN JOURNAL

Manuscripts submitted to the JOURNAL are reviewed for scientific content by one or more of the editors and, often, by one or more outside reviewers as well. During the past year we have received the kind assistance of J. G. Bruce, C. Haufler, R. L. Hauke, R. M. Lloyd, J. Montgomery, R. Oliver, G. R. Proctor, J. E. Skog, A. R. Smith, R. G. Stolze, W. C. Taylor, J. Utley, and R. A. White, to whom we are deeply indebted. We welcome suggestions of other reviewers and offers of assistance.—D. B. L.

ANNUAL INDEX

123

INDEX TO VOLUME 69

Acrostichum, 42, 45; aureum, 42, 43, 45; danaeifolium, 15, 42, 44, 45 Acrostichum in Florida,

sh inson. igi abies in Florida, 42 atum, 29

ia, 7 79; adiantifolia, 15; subg aca AS . 7-19: h 1, 74, 76, 78, munchii, 71, 72, 75 8, 79; oblongifolia, 71, 76-79; ouru- tana, 71 76, 78, 79; ABI na. 1: i 3; pohlia

. 76, 28. 79; ER ¥ 4 Bag Se 15,7 77-7

Apic ld i in Anemia phyllitidi

ig technique for Selaginella, 9

Aspidiaceae, 83

Aspleniaceae, 83

‘esecouesiua 5, 94, 96; er rum, 5; flaccidum, 5; herb-wagneri, 94; um lanes yeas 94; platyneuron, 7,

5. Var. teen. - veil . 47, 95, 96; rhizophyllum, 47;

anes 94: Mile dentatum 14, 15

ig arate flora (rev.), 5

neri

Asplenosorus herb-wag'

Athyrium escis ee 1, 8 pendcanon. 29, 47, 85; thely- pteroides, 85

Atlas sal ferns of the British Isles (rev.), 9

Aus tin, D. F.,G. B. Iverson & C. E. ae A tropical fern grotto

"Broward = Florida, | cee 17. 20, 23, 24; sainnipiee se 23; filiculoides, 17, 21, 23; mexicana, 17-24; = nnata MECC,

Baskin, J. in. The ‘ole Saal in the vegeta- tive life cycle of bettas butleri, 103 Baskin, C. C. (see in)

Beitel, J. M. Sewigence of san in the lycopsids, 83 Beitel, J. M. (see J. G. Bruce

Bhardwaja, T. N. (see a B. Gena)

Blechni: |

-M.& “s ve ans. Isoétes butleri in Georgia, 62 Bch, 41; alae, nee ro ee

B sey, Ves | NIT 7 1 iff her . ne

t Stina KI Cy P| 1 . FA 1

{Fev;),'S Bruce, J. . M. Beitel. A community of Lycopodium sametophytes in Michigan, 33 Bruce, J. G oe neburne, sa Richardson & J. Worthington. 29

Vittaria lineata re

Camptosorus hizphyes, 94 Campvlo ;

oom dra, S. Gam yetophyte morpholon of the fern genus Drynari- ig ope ) 11 Che thes bern, bie feei, 95; lanosa, 95 heile ensis new to Kentucky, 95

_ seeceeeiga 115 dy esi of {Lycopodium gametophytes in Michigan, 33 Coniogram

Cranfill, R Fads alabamensis new to Kentucky, 95 — sloanei, 15; subma

a

um fort ei in ima ana and Mississippi, 85 mor bulbifera, 95; fragilis, 29, 85; tennesseensis, 47, 95 Davalliace ath Sauce: R. J. Rodin, 28 ee. 116 W.M.. let

A. Ben ie macrospora in ‘southeastern esa 97

The chenigenl of Pagers from strobilus branches in Lyco podium a, 80

pora i Tennes-

ee, 97 The distribution of Dryopteris goldiana and D. marginalis in Mis- 29

souri, Drynaria, 113, 114, 116; rigidula, nariopsis, 111, 114, 116, 117; heraclea 111, 113, 115, 116

Dryopteris, 6-8; celsa, 6; cristata = cristata, 6, 7; goldiana, 29, _ i ia, 7; marginalis, 29 separabilis, 8; sls E

sre tame 100; apodum var. ¢ m, 100, var. A

. Majus ; backhousi 100; buchii, 101; Sie 101;

ney 101; pilosum, 102; smithii, 101; stellatu

Equisetum, |, 3; x ferrissii, 1-4; giganteum, 1; subg. Hippochaete 1; hyemale, 1, 3, . affine, 1, 3; laevigatum, 1, 3; x moorei, 1; myriochaetum, 1; ramosissimum, 1-5, s subsp. debile, 1, subsp. ramosi m, 1, 3; x sch

Equisetum ramosissimum in North America, |

Evans, A. M. (see B. M. Boo . M. Dennis)

Evolutionary patterns and processes in ferns (rev.), 16

hs s (rev.), 31

he fine structure of the pre-meiotic stages of sporogenesis in peste sibilis, 87 Gametophyte morphology of the fern genus Drynariopsis phe youn ge 111 Gena, C. B., ae & A. K. Yadav. A new species of ~ 7 Ile fi

Gymace ramma, 30 Gymnogramma vs. gymnogramme, Ha pss .C. H. & R. M. Tryon. ern vs. Gymnogramme,

la R. L. Equisetum ramosissimum in North America, | Hill, S. R. Spore morphology of Anemia subgenus pi nemia, 7 Hot, R. Ww. sk a opp. Sti oe Azolla- sncmapien sym-

ae home gardener’s book of ferns (rev.), 102

Huperzia,

Selim anarstig e, 83

Iffrig, G. ae isin of Dryopteris goldiana and D. mar- ginalis in Misso!

Incidence of parce sm in the lycopsids, 83

Isoétes, 98, 103; bi utleri. 6 62, 103, 106-109; macrospora, 97, 98

si

‘ D. F. Austin) rmy, A.C. et Ge eds. Atlas tm of the British Isles (rev.), 91 TESS, - M. _ W. C. Taylor

loch, I. W. Tuvenile leaves of the apogamous fern “caret

f eae ee |

perio ramer, K. he pteridophytes of Suriname (rev.), 70; Syeueanabony: a es aingsgsta v.), 48

Landry, _M. Israel. R. Schwarzwalder, Jr. & R. D. Thomas.

y rtomium fortune in a na and Mississippi, 85

pis Le echane Lov vi ne ah BE olin patterns and processes in ferns (rev.), 16 Lye boars Lycopodiella, . oe 33-35. 37, 39. 40, 49, 50, 52-55, re ici ap grren var. furcatum, 54, 55, 59; annotinum, 33. 41, 58; ap- essum, 33; carolinianum, 56, 58, 60; ae 57; cer-

124

nuum, ey 55, 58, 59; — 93535, 37,29) 8 pie ora

35, 57; crassum, 53, 59; dendroideum, 33,

dichotomum, 83; cine 33,31, 30: 9: flabelliforme, 37, 80, 82: lateral

inundatum, 54, 56, 58-60; le, 49, 55-57, 59, 60; subg dotis, 49; lucidulum, 33-35, 39-41, 51, 83;

, 58, 60; . Lycopodium, 52, 58, 60; num-

ul ium, 49, 51, 52, 58; obscurum, 33-35, 37, 39, 40, 80;

rus, ’ 33,39, 57; die. Veale. 49, SI, 54, 57-59; subg. Urostachys, 49, 83

Macrothelypteris torresiana, 16 agraw, T, W. & L. J. Musselman. Notes on 7 dispersion of Deyopuatle spores in the Great Dismal Swam

Marengo, N. P. The fine structure of the sae isi stages sporogenesis in Onoclea sensibilis, 87; A simplified nutrient medium for growing fern prothallia, |

Merinthosorus, 114-117; slate 116

Mickel, J. T. The home gardener’s book of mr (rev.), 102; Anew

combination in Aprons, 94; Three new Elaphoglossums uate 00

agraw) auman, C. E. A new Nephrol hybrid from Florida, 65; A Thelypteris new to Florid: Nauman, C. E. (see D. F. ead in) 5-68; x averyi, Pes 68, 69; biserrata, 15, 65-69; exal- Posts 15, 65-69; hirsutula, 6: Asplenosorus, en A new Nephrolepis hybrid from la, 65 A new species of Selaginella po pan 119 Notes on the dispersion of Dryopteris spores in the Great Dismal Swamp, 6 Notholaena cochisensis, 6. —— B. Studies i in Lycopodicee, Il. The — patterns

Onoclea sensibilis, 29, 87 Ophioglossaceae, 8:

Ophioglossum engelmannii, 103; — 14,

Osmunda cinnamomea, 7, 8; regalis, 8, var. ok. 15

Petrik-Ott, A. J. The pteridophytes of mgr Nebraska, South ta and North Dakota, U.S.A. (rev.), 84

Pellaea atropurpurea, 47, 95; glabella, 95, var. glabella, 96

Phlebodium aureum, 15

Photinopteris

oo 47; Sakina m, 47, 48, var. americanum, e cue te ne newly discovered in Alabama, 4

Platycerium, 115,

Pleopeltis hay an faa 64

» £13, ‘ie 116

lotaceae, 7) Psilotum nudum, 15 Pteridium aquilinum, 35, var. latiusculum, 15 The pteri es of Kansas, Nebraska, ‘South Dakota and North Dakota, U.S.A. yea ), 84 The pteridophytes of Suriname (rev.), 70

AMERICAN FERN JOURNAL: VOLUME 69 (1979)

Pteris longifolia, 92; — 15; vittat eviews: Asplenium hybrids in the Neo

ye

ae nd flora, 5; Atlas

me, 70; Synaptospory: A speci

Reynolds, T L. Apical dominance in Anemia phyllitidis gametophytes, 92

Richardson, J. I. (see J. G. Bruce) bi nigh

e). Te ae fol pam in the Teen life cycle of [soétes butleri, 103

Salvinia minima, 15; rotundifolia, 15

Schizaeaceae

Shwariihhdie RoI be e G. P. Landry)

opoemnegte! 10. 12, 82, 83, 119; braunii, 86; flabellata, 9; involvens, 84; kra

a, 9-12, var. aurea, 9; oregana, 84; martensii var. manana: 9 ar. variegata, 9; ee ee rajas- thanensis "18-1 reticulata, 119, 121; tama—montana, 121; ver- sicolor

Sexuality in hs ae ste oe Short, J. W. Pa yilits assert newly discovered in Alabama,

As cael nutrient medium for prothallia, 122 Spore morphology of Anemia subgenus Anecenial 71 <aatidoior Db. (see JOG. )

ae in eos Lint eaigg ja “The — patterns and in-

Studies of the Azolla- Anabaent ahaa using Azolla mexicana, lL. 1 labora F punta y: A hypothesis (rev.). Ra onomic revision of the New Zealand species of Asplenium po

Taylors W.C.&D. M. pees Srey gate in Arkansas, 26 4,

Tectaria heracleifolia, 1 incisa, 14, 64, f. incisa, 16;

Thelypteris, 26, 27, 64: ee 16; grandis, 64; interrupta, 16; kunthii, 16, 26-29; normalis, 16; noveboracensis. sik ovata V ovata, 16; palustris. 26. var. haleana, 26, kak var. pubescens, 7 27; reptans 1 torresiana, 16, 26-28; totta, 16

. Lan Three new Elaphopossums dics ‘uate 100 omlinson, P. B e D. C. Adams) Pivieus aaa roward pop vee Florida, 14

— R.M ec. up peat Vittaria 29; lin ne. niaindy lineata Selescos n Geor; Webster. T. R. rh ata tere che wr vee sagenedi a

nat ocd um heanain S 80 Wendt, T. goes ality in Asplenium resiliens, 96 Wofford. B. E. (see W. M. Dennis) os aia -: ag

I virginica, 8

hcotecamalg ‘ heels G. ade e) Yadav, ec: i

pp. a oe (see ap H.

ERRATUM Page 37, Table 2. For ‘““FLABELLIFORME”’ read ‘‘DIGITATUM.”’

NOTEWORTHY PUBLICATIONS OF THE CHRONICA BOTANICA

BEDDOME, R. H. The Fems of British India. Includes Bangladesh, Burma, Ceylon, Malaya, Nepal, Pakistan, and Tibet. Set of 2 vols. Reprint ed. 1973. Available CB (cloth bound) or LB (half-leather

with gold lettering). CB Rs. 285; LB US $55. . Supplement to the Ferns of Southern India and British India.

se eer Ss CB Rs. 85; LB US $18.50.

— ——. Handbook to the Ferns of British India. Reprint ed. 1970. CB Rs. 100; LB US $21.

NAYAR, B. K. Companion to R. H. Beddome’s Handbook. 1974. CB Rs. 75; LB US $14.50 Annales Cryptogamici et Phytopathologici, vol. 8 . Fern Flora of India. The first taxonomic account of the fern flora since Beddome’s classical work ‘‘Handbook to the Ferns of British India’’ published in 1883. An illustrated floristic account containing keys to identifications as well as lucid descriptions of species, genera, and families of Filicopsida. Prepared by the foremost pteridologist of India. About 1000 pages. November

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