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 


9 


06 


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 


— 


Ww 


~ 


\o 


— 
WwW 


PS 


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N 


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 


108, 


ON 
an 


nN 
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_ 


97 


99 


109 


119 


122 


122 


123 


124 


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 


we 


~ 


‘oO 


LUIS D. GOMEZ P. 


w 


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. 

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 
wit over st $2.50 each, plus shipping. Ten percent discount on orders of six volumes or more; 


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. 


ore erresen re renars scons cert 


Soe NO MORE!!! after rare ferns, old-world epiphytes, all 18 sp. 
diggs & many cv’s, ferns of Malaysia, Thailand, Phillippines, W. 
here caiy at'te sh Sp. unknown to cultivation, and available 

e ERED SPECIES, 6 Upland Ave., Dorchester, 
MA 02124. $1.00/catalog, i 


pacers Ti tse es 


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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———.. 1964. The Ferns of — Polypodiaceae (Dennstaedtieae to Oleandreae). Contr. Gray Herb. 
194; 1-253. 

VERDOORN, F., ed. 1938. Manual of Pteridology. Martinus Nijhoff, The Hague. 

WEATHERBY, C. A. 1920. Varieties of Pityrogramma triangularis. Rhodora 22: 113-120. 

WIESNER, J. 1876. Uber die Se aaprge Beschaffenheit der geformten Wachsiiberziige 
pflanzlicher Oberhaute. Bot. Zeit. 34: 225-236. 

ene E. 1972. Flavonols from the fronds of Pityrogramma chrysoconia. Phytochem. 


ona Die Verbreitung spezifischer Flavone in der Gattung Primula. Biochem. Physiol. 
Sthsieran 166: 419-424. 
. 1975a. Flavonoid-Muster als systematische Merkmale in der Gattung Populus. Biochem. 
Syst. Ecol. 3: 35-4 
: 1975b. Flavonoid- Masten im Knospen-Exkret der Betulaceen. Biochem. Syst. Ecol. 3: 
4 -52. 
oe Flavonoide Exkrete in Goldfarnen und Silberfarnen. Zeitschr. Pflanzenphys. 78: 
-349, 
. 1976b. Flavonoid exudations in farinose ferns. Phytochem. 15: 2013. 
. 1976c. Die Komponenten des ‘‘Mehls”’ bei Cheilanthes und Notholaena—ein chemotax- 
onomisches Merkmal? Ber. Deut. Bot. Gesell. 89: 243-246. 
1977a. Die Zusammensetzung des Flavonoid-Mehls bei einigen Farnen. Zeitschr. Pflan- 
seuphys, 85: 71-76. 
. 1977b. Chalkone und Dihydrochalkone als Mehl-bestandteile bei Farnen (Gattungen 
Cheilanthes und Notholaena). Zeitschr. Naturforsch. 32C: 1013-1014. 
———, J. FAVRE-BONVIN, and P. LEBRETON. 1978. Ein Butyryl-Flavonol aus dem Mehl von 
Notholaena affinis. 
veel SC HNEPF. 1970. Vergleichende Untersuchungen iiber die flavonoiden Exkrete von 
ehl-” und “Ol-’’ Driisen bei Primeln und die Feinstruktur der Driisenzellen. Zeitschr. 
Pflanzenphys. 62: 216-227. 
ZOPF, W. 1906. Zur Kenntnis der Sekrete der Farne. I. Drisensekrete von Gold- und Silberfarnen. 
Ber. Deut. Bot. Gesell. 24: 264- 272: 


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 


33 


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 
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Matter for publication and claims for missing issues (made within six months of the date of issue) 
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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 
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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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platyceriums & many cv’s, ferns of Malaysia, Thailand, Phillippines, W. 
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teria, CA 93013. $1.00/catalog. 


-_ 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; 
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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 
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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. 


TRIARCH 
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Volume 68 
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October-December, 1978 


QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY 


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? 


Reviews 


American Fern Journal 


Index to Volume 68 


Erratum 


PHILLIP M.RURY 9%9 


MERYL A. MIASEK 109 


119 


wrssour! BOTANICAL 


JAN 10 1079 


GARDEN LIBRARY 


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 
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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 
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GERALD J.-GASTONY ..:4.:...:.. Dept. of Biology, Indiana University, Bloomington, Ind. 47401 
JOHN T. MICKEL New York Botanical Garden, Bronx, New York 10458 


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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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and service to botanists. 


We welcome samples of your preserved research 
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your requests for catalogs or custom preparations 
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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 


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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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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: | 
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e mature development, growth, and yield 


pH ANALYZER $19.95 prepaid 
ows PH status instantly. 


recision instrument sh 
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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 


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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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Section Heads: 


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Psilophytes and Horsetails 
Club Mosses 

Association of Species in 


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Glossary 

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AMERICAN ee 
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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. 
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American Fern Journal 
EDITOR 
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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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Dr. Alan R. Smith, Dept. of Botany, Univ. of California, Berkeley, California 94720, is Editor of 
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= eateries Mickel, New York Botanical Garden, Bronx, New York 10458, is editor of the 
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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 


TRIARCH 
Over 5© Wears 


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and service to botanists. 


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P.O. Box 98 
Ripon, Wisconsin 54971 


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. 


The Chronica Botanica Co. 
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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 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. 


Section Heads: 


Flowering Plants 
Gymnosperms 
Ferns 
Psilophytes and Horsetails 
Club Mosses 
Association of Species in 
Natural Floras 
Appendix 
Glossary 


HEATH 


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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) 
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Changes of address, dues, and applications for bership should be sent to Dr. J. E. Skog, Dept. 
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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- 
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exchanged and collection lists sent on request. 

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| tsand beq the Society enable i ‘ é bers and to others interested 
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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: 


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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 


Gy e.e 


Ny 


ea, 


, 
3, 
CYS 


8, 
cs 
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“ts 


@ 

oo ey 
SH rae egueme 
SES re 
Stee See 
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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.”’ 


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