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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

CONTENTS

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

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

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

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

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

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

Farinose Exudates in Gymnogrammoid Ferns ECKHARD WOLLENWEBER

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

Review

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

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

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

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

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

Review

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WwW

a
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,

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122

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AMERICAN
FERN
JOURNAL

Volume 68
Number 1

January-March, 1978

QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY

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

Leptogramma pilosa in Alabama

Chromosome Numbers in the Fern Genus Anogramm

UDITH G. opie and GERALD J. GASTONY

Curtis Eugene Delchamps (1925-1977)

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

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

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

On the Distribution of Lycopodium flabelliforme
in Illinois

Review

Missoup; Boranican,

APR 95 1978

GS ;
ARDEN LIBRARY

ERRY M. BASKIN and CAROL C. BASKIN

IRVING W. KNOBLOCH and DONOVAN S. CORRELL 1

‘oO

LUIS D. GOMEZ P.

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

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

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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.

> 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)

LITERATURE CITED
ALT. K. S. and V. GRANT. 1960, Cytotaxonomic observations on the goldback fern. Brittonia 12:

53-170.
BARY, A. de 1877. Vergleichende Anatomie der Vegetationsorgane der Phanerogamen und Farne. /n
W B. Hofmeister, ed. Handbuch der physiologischen Botanik, vol. 3. W. Engelmann,

Leipzig.
BERTI, G. and F. BOTTARI. 1968. Constituents of ferns. Jn L. Reinhold and Y. Liwschitz, eds.
Progress in Phytochemistry, vol. 1. Interscience, London
BLASDALE, W. C. 1893. On certain leaf hair structures. Brethes 1: 252-258.
. 1903. On ceroptene, a new organic compound. J. Amer. Chem. Soc. 25: 1141-1152.
BOHM, B. A. 1968. Phenolic compounds in ferns. II. Phytochem. 7: 1687-1689.
—__——. 1975. Chalcones, aurones and dihydrochalcones. Jn J. B. Harbone, et al., eds. The
Laroisien Academic Press, New Yor
BOTTARI, F., A. MARSILI, I. MORELLI, ane M. PACCHIANI. 1972. Aliphatic and triter-
noid hydrocarbons from ferns. Phytochem. 11: 2519-2523.
BOWER, F. O. 1923. The Ferns, part 1. Cambridge Univ. Press, Cambridge.
CHRIST, H. 1897. Die Farnkrauter der Erde. G. Fischer, Jena.
DIETZ, V. 1978. Vorkommen und Verbreitung von Flavonoid-Aglykonen bei Farnen der Gattung
ret gg Part of a Diplomarbeit, Institut fiir Botanik, Technische Hochschule,

Darmstadt.
DOMIN, K. 1929. The hybrids and wei forms of the genus Pityrogramma (Link). Rozpr. Ceské
Akad., Tfida II, 38(4): 1-80, t. /-XI.
DOUS, a Pigs Uber Wachsdusscheidungen bei Pflanzen; ein Studium mit dem Oberflachenmikro-
p. Bot. Arch. 19; 461-473.

ERDIMAN, H., L. NOVOTNY, and M. ROMANUK. 1966. Flavonols from the fern Cheilanthes
farinosa. Tetrahedron Suppl. 8(1): 71-74.

FAUX, A., M.N. GALBRAITH, D. H. S. HORN, and E. J. MIDDLETON. 1970. The structures
of two ecdysone analogues, cheilanthon A and B, from the fern Cheilanthes tenuifolia. Chem.
Comm, 1970: 243-244

GUERVIN, C., J. LAROCHE, and C. Le COQ. 1971. Stéréostructure des produits d’excrétion sur
les feuilles du Gymnogramme sulphurea. Rév. Gén. Bot. 78: 53-55.

GUNNING, B. E. S. and M. W. STEER. 1977. Plant Cell Biology—an Ultrastructural Approach.
Edward Amold, London.

HABERLANDT, G. F. J. 1918. Physiologische Pflanzenanatomie, ed. 5. W. Engelmann, Leipzig.

HARBORNE, J. B., T. J. MABRY, and H. MABRY, eds. 1975. The Flavonoids. Academic Press,

Lon
HOHLKE, F. 1902. Uber die Harzbehiilter = die Harzbildung bei den Polypodiaceen und einigen
Phanerogamen. Beih. Bot. Centralbl. 11: 8-45.
IMAL 5. W. J. and J. G. BAKER. 1868. Sai Filicum. R. Hardwicke, London
Al, S., T. TOYOSATO, M. SAKAI, Y. SATO, S. FUJIOKA, E. MURATA, Heal eS GOTO.
WER 969. Screening results of plants for phytoecdysons. Chem. Pharm. Bull., Tokyo 17: 335-339.

. T., K. N. N. AYENGAR, and S. RANGASWAMY. 1972. Structure of cheilarinosin, a

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

——, K. N. N. AYENGAR, and S. RANGASWAMY. 1973. Occurrence of cheilanthone-B in
eaeon anthes mysurensis. Indian J. Chem. 11: 1336-1338.

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

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

Central American Species. Flora 165: 507-522,

E. WOLLENWEBER: FARINOSE EXUDATES OF GYMNOGRAMMOID FERNS 27

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

LELLINGER, D. B. 1967. Pterozonium (Filicales: Polypodiaceae). Jn B. Maguire collaborators.
The botany of the Guayana Highland- VII. Mem. New York Bot. Gaed 7: 2-

LINK, H. F. 1842. Icones Selectae Anatomico-botanicae, fasc. 4. Haude & eh Be de.

| aes ai K. 1930. Die Epidermis. Jn K. Linsbauer, ed. Handbuch der Pflanzenanatomie, div.

, vol. 4, part 2. Gebriider Borntraeger, Berlin.

Secrets. U. and E. SCHNEPF. 1976. Elimination Processes by Glands. 3. Organic Substances. Jn
U. Liittge and M. G. Pitman, eds. Encyclopedia of Plant Physiology, New Series. Springer,
Berlin

LYE E tT. F., J. S. LYTTLE, and A. CARUSO. 1976. Hydrocarbons and fatty acids of ferns.
Phytochem. 15: 965-970.

MABRY, T. J. 1973. The chemistry of geographical races. Pure Appl. Chem. 34: 377-

MELCHIOR, H. and E. WERDERMANN. 1954. A. Engler’s Syllabus der Pies ed: 12;

vol 1. Gebriider Borntraeger, Berlin

MOBIUS, M. 1927. Die Farbstoffe der Pflanzen. In K. Linsbauer, ed. Handbuch der Pflan-
zenanatomie, vol. 3, part 1. Gedbriider Borntraeger, Berlin.

MOLISCH, H. 1923. Mikrochemie der Pflanzen, 3rd ed. G. Fischer, Jena.

NAYAR, B. K. 1962. Ferns of India. VI. Cheilanthes. Bull. Natl. Bot. Gard. Lucknow 68: 1-36.
964. Some aspects of the morphology of Pityrogramma calomelanos and Pityrogramma

chrysophylla. J. Indian Bot. Soc. 43: 203-213.

NILSSON, M. 1959. The structure of ceroptene. Acta Chem. Scand. 13: 750-757

—. 196la. Dihydrochalcones from the fronds of Pityrogramma chrysophylla var. marginata.
Acta Chem. Scand. 15: 154-158

. 1961b. Chalcones from the fronds of Pityrogramma chrysophylla var. heyderi. Acta Chem.
Sex nd. 15: 211-212.

OGURA, Y. 1972. Comparative Anatomy of Vegetative Organs of the Pteridophytes. /n W. Z
mermann, et al., eds. Handbuch der Pflanzenanatomie, 2nd ed., vol. 7, part 3. Cochictihen
Borntraeger, Berlin.

RAMAKRISHNAN, G., A. BANERJI, and M. S. CHADHA. 1974. Chalcones from Onychium

auratum. Phytochem. 13: 2317-2318.
RANGASWAMY. S. and R. T. IYER. 1969. Flavonoids of Cheilanthes farinosa. Indian J. Chem. 7:
526.

SCHKUHR, C. 1804-09. Kryptogamische Gewachse. Published by the author, Wittenberg

SCHNEPF, E. 1969. Sekretion und Exkretion bei Pflanzen. Jn L. V. Heilbrunn, ed. Piceeatans
matologia: Handbuch der Protoplasmaforschung, Band 8. Springer, Wie

,and A. KLASOVA. 1972. Zur Feinstruktur von Ol- und dence uals Ber. Deut. Bot.
Gesell. 85: 249-258.

SEIGLER, D. S., D. M. SMITH, and T. J. MABRY. 1975. n-Alkans from diploid, triploid and
tetraploid plants of Pityrogramma triangularis flavonoid chemotypes. Biochem. Syst. Ecol. 3:

SERIZAWA, S. and E. WOLLENWEBER. fl Cheilanthes kuhnii var. brandtii and the composi-
tion of its farina. Amer. Fern. J. 67: 107-108.

SMITH, D. M., S. P. CRAIG, and J. cane one 1971. Cytological and chemical variation in
Pityrogramma triangularis. Amer. J. Bot. 58: 292-299.

STAR, A. E. 1977. Intracellular and Exudate Flavonoids of Pityrogramma Gold- and Silver-back
Ferns. 1. Joint Symp. Phytochemical Societies of Europe and North Amer., Ghent. asad
session).

————., and T. J. MABRY. 1971. Flavonoid frond exudates from two Jamaican ferns, Pityro-
gramma tartarea and P. calomelanos. Phytochem. 10: 2817-2818

,D. S. SEIGLER, T. J. MABRY, and D. M. SMITH. 1975a. Internal flavonoid patterns of
diploids and tetraploids of two exudate chemotypes of Pityrogramma triangularis. Biochem.
Syst. Ecol. 2: 109-112.

“7+, Hl. ROSLER. T. J. MABRY, and D. M. SMITH. 1975b. Flavonoid and ceroptin pigments
from frond exudates of Pityrogramma triangularis. Phytochem. 14: 2275- 2278.

28 AMERICAN FERN JOURNAL: VOLUME 68 (1978)

STRASBURGER, E. 1905. Lehrbuch der Botanik, 7th ed. G. Fischer, Jena.

SUNDER, R., K. N.N. AYENGAR, and S. RANGASWAMY. 1974. Crystalline chemical compo-
nents of Challanihes longissima. Phytochem. 13: 1610- 1611.

SWAIN, T. 1973 itive Tracheophytes. In G. snp and J.
Fanaa eds. Chemistry in Botanical Classification. Academic Press, New :

ibn - M., Jr. 1956. A revision of the American species of Notholaena. Contr. Gray "Herb. 179:

a Taxonomic — notes. II. Pityrogramma (including Trismeria) and Anogramma.
Goat. Gray Herb. 1 -76.

———.. 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

The American Fern Society
Council for 1978

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

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

The ‘‘American Fern Journal’ (ISSN 0002-8444) is an illustrated quarterly devoted to the general
study of ferns. It is owned by the American Fern Society, and published at the Smithsonian Institu-
tion, Washington, DC 20560. Second-class postage paid at Washington.

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

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

Orders for back issues should be addressed to the Treasurer.

General inquiries concerning ferns should be addressed to the Secretary.

Back volumes $5.00 to $6.25 each; single back numbers of 64 pages or less, $1.25; 65-80 pages, $2.00
each; over 80 pages, $2.50 each, plus shipping. Ten percent discount on orders of six volumes or more;
postage additional.

Library

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

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

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exchanged and collection lists sent on request.

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“i Bie mes See - ee i’ Lo}
in ferns. Botanical books, back issues of the Journal, and cash or other gifts are always welcomed, and
are tax-deductible. Inquiries should be addressed to the Secretary.

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

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.

| 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.

TRIARCH
Over 5@ Years

of slide manufacture
and service to botanists.

TRIARCH INCORPORATED
P.O. Box 98
Ripon, Wisconsin 54971

———_—__—

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.

-_ 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.

Back volumes $5.00 to $6.25 each: single back numbers of 64 pages or less, $1.25; 65-80 pages, $2.00

each; over 80 pages, $2.50 each, plus shipping. Ten percent discount on orders of six volumes or more;
postage additional.

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

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

Spore Exchange

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

Dr. John T. Mickel,
may borrow books at

Gifts and Bequests

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

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

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

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

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

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.

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
Number 4

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
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 J.-GASTONY ..:4.:...:.. 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.

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Subscriptions $9.00 gross, $8.50 net if paid through an agency (agency fee $0.50); sent free to
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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)

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

of slide manufacture
and service to botanists.

TRIARCH INCORPORATED
P.O. Box 98
Ripon, Wisconsin 54971

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

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

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

The ‘‘American Fern Journal’ (ISSN 0002-8444) is an illustrated quarterly devoted to the general
study of fems. It is owned by the American Fern Society, and published at the Smithsonian Institu-
tion, si niphinaens DC 20560. Second-class postage paid at Washington.

Matter for publication and ~— for missing issues (made within six months of the date of issue)
ace be addressed to the Edito

Changes of address, dues, and se for membership should be sent to Dr. J. E. Skog, Dept.
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General inquiries concerning ferns should be addressed to the Secreta

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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

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.

pH ANALYZER pH controls: |
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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

ee
eiciral

= a

ae Panes
°

Lee

nt
ak
on

2
=
a
>

— @
2s
8

is. FIG. 6. Physiogra

FIGS. 1-5. Distribution maps of Arkansas Thelypter

28 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

We wish to thank Dr. Alan R. Smith for checking the identifications of T.
kunthii and T. torresiana.

REPRESENTATIVE COLLECTIONS:

Thelypteris kunthii: Ashley Co.: Along timber access road SE of Lake Georgia-Pacific; T17S, ROW,
sect. 35, 25 Oct 1975, Johnson 281 (Hendrix College).

Thelypteris torresiana: Ashley Co.: Along timber across road W of Crossett; T18S, ROW, sect. 9, 25
Oct 1975, Johnson 283 (Hendrix College).

Thelypteris noveboracensis: Cleburne Co.: Johnson 393 (Hendrix College). Garland Co.: Moore
6278 (APCR). Montgomery Co.: Taylor 1084 (SIU). Pike Co.: Taylor 2903 (SIU). Polk Co.: Moore
410261. (UARK). Pulaski Co.: Johnson 112 (Hendrix College). Saline Co: Moore 480/54 (UARK).

Thelypteris palustris var. pubescens: Ashley Co.: Johnson 276 (Hendrix College). Bradley Co.:
Demaree 19445 (MO). Drew Co.: Moore 420048 (UARK). Greene Co.: Moore 480685 (UARK).
Hempstead Co.: Moore 480353 (UARK). Izard Co.: Johnson 422 (Hendrix College). Lawrence Co.:
Taylor 1794 (S1U). Little River Co.: Palmer 8359 (MO). Polk Co.: Moore & McWillliam s.n. (U ARK).
Sharp Co.: Wade 167 (UARK). Washington Co.: Henbest 16 (UARK).

Thelypteris palustris var. haleana: Bradley Co.: Cypress swamp near Warren, Demaree 19445
(SMU). Drew Co.: Swamp near Wilmar, Demaree 24624 (SMU).

LITERATURE CITED

DEMAREE, D. 1943. A catalogue of the vascular plants of Arkansas. Taxodium 1:1-88.

LEONARD, S. W. 1972. The distribution of Thelypteris siana in the southeastern United States.
Amer. Fern J. 62:97-99.

SMITH, A. R. 1971. Systematics of the neotropical species of Thelypteris section Cyclosorus. Univ.
Calif. Publ. Bot. 59:1- 143.

ROBERT J. RODIN (1922-1978)

Robert Joseph Rodin was born in Sacramento, California, on 15 July 1922. He
lived in Eureka and in Turlock in his youth. During World War II service in the
lS. Marine Corps, he was stationed in Guam and China. In 1948 he married
Elva Bain, who, with three daughters, survives him. He received a Ph.D. degree
in botany from the University of California at Berkeley in 1951. After graduation,
he accepted a position as a professor of biology at Forman Christian College,
Lahore, West Pakistan. He collected ferns in the Himalayas extensively during
this time. From 1953 until his retirement in 1976, Rodin was a professor of biology
at California Polytechnic University in San Luis Obispo. His research specialties
included the anatomy of the Gnetales, ethnobotany of Ovamboland, and the
taxonomy of the pteridophytes of California, an interest which led to his publish-
ing the *‘Fems of the Sierra’ for the Yosemite Natural History Association. In
recent years he was active in the conservation movement in California. In 1966—
67 he was a Fulbright Professor at the University of Delhi. He also took part in
botanical expeditions to South Africa in 1947 and to southwestern Africa in 1973.
He died in San Luis Obispo on 27 June 1978.—D.B.L.

AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 1 (1979) 29

SHORTER NOTES

VITTARIA LINEATA REDISCOVERED IN GEORGIA. — Vittaria lineata (L.)
J. E. Smith, the Shoestring Fern, is primarily restricted to peninsular Florida
reaching as far north as Citrus Co. on the Gulf coast and Duval Co. on the
Atlantic coast (E. T. Wherry, The Southem Fem Guide, Doubleday, 1964).
However, a disjunct population had been known since 1938 in Lincoln Co., Geor-
gia. This piedmont population grew in pits in south-facing sandstone cliffs. Unfor-
tunately, this area was quarried and the original locality destroyed. Thus, as far as
known, Vittaria has been extinct in Georgia for some years.

This note reports the rediscovery of V. lineata in Georgia on Cumberland
Island, Camden Co. The plants were observed at three localities on the island
some distance apart. Two sites were bark fissures of Live Oak trees (Quercus
virginiana Miller); the third was in a bark fissure of Sweet Bay (Magnolia vir-
giniana L.) All sites were approximately one meter above the ground on the east
and southeast exposures of the tree boles. One specimen (Worthington 336) is on
deposit at the University of Georgia herbarium, and another (Stoneburner s.n., 19
Sept 1978) is in cultivation at the University of Georgia Plant Growth
Facilities.—James G. Bruce, Department of Botany, and D. L. Stoneburner,
James I. Richardson, and Joanne Worthington, Institute of Ecology, University
of Georgia, Athens, GA 30602.

THE DISTRIBUTION OF DRYOPTERIS GOLDIANA AND D. MARGINALIS
IN MISSOURI. — Dryopteris goldiana (Hooker) Gray is one of the rarest ferns in
Missouri. The only known occurrences have been early collections made in two
isolated locations along the eastern border of the state. These were noted by
Steyermark in ‘* Flora of Missouri” (1963, p. 40). In July, 1976, I discovered the
first locality for this species on the western side of the state. Plants were found
growing in rich loess deposits underlain by local limestone formations In a ravine
tributary to an unnamed intermittent drainage north of the Missouri River in
south-central Clay County, in a city park within the city limits of Kansas City.
This particular site is 200 miles from either of the two previously recorded Mis-
souri locations, and suggests that perhaps D. goldiana also occurs in the interior
sections of Missouri, although it has not been recorded from there. Habitat infor-
mation listed in Gleason and Cronquist’s ‘* Manual of Vascular Plants of North-
eastern United States and Adjacent Canada’”’ (1963, p. 24) noted a preference for
moist woods in circumneutral soil. The Kansas City site is Knox Silt Loam oe
PH 5.6-7.3. Other fern species found in association with D. goldiana show the
richness of the site: Adiantum pedatum var. pedatum, Athyr um PyYEOCATP a
Botrychium virginianum, Cystopteris fragilis, Dryopteris mar ginalis, Onoclea
Sensibilis, and Woodsia obtusa, as well as liverworts. The richness of the fern
flora at this site is unique in northwestern Missouri. bois + tox thie

The presence of Dryopteris marginalis (L.) Gray at this site 1s unusual, see
Species is commonly found in shaded crevices of rock ledges and bluff sites

30 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

Missouri where soils are generally derived from sandstone, chert, or granite.
While in the same vicinity of D. goldiana, the plants of D. marginalis were found
at a slightly higher elevation on a north-facing slope. Dryopteris marginalis also
was located on the thick loess deposit, an unusual substrate for this species in
Missouri. This particular location is the northwesternmost occurrence of D. mar-
ginalis in Missouri; Steyermark’s ** Flora of Missouri’ (1963, p. 40) recorded it
from Saline County, 75 miles east of this latest discovery. The species is fairly
common south of the Missouri River in most counties of the Ozarks, but has not
been found in the northwestern part of the state.

The site was nominated as one of Missouri’s State Natural Areas, representing
the rich river hills community type as well as preserving the habitat for a rare
Missouri species, D. goldiana. Through a cooperative agreement between the
Kansas City Parks Department and the Missouri Department of Conservation,
State Natural Area status was approved in February 1978.

Documentary specimens have been filed in the herbaria of the University of
Missouri at Columbia (UMO) and at Kansas City. — Greg F. Iffrig, Missouri
Cooperative Wildlife Research Unit, Stephens Hall, University of Missouri, Co-
lumbia, MO 65211.

GYMNOGRAMMA VS. GYMNOGRAMME. — Confusion has existed over the
proper spelling and gender of the generic name Gymnogramma. Although this
name is illegitimate (see Underwood, Bull. Torrey Bot. Club 29:617-634. 1902),
its wide use in the past has made it often quoted in current literature. The name
has been considered to be either feminine or neuter and has been alternatively
spelled Gymnogramme.

In originally naming the genus Gymnogramma, Desvaux (Berl. Mag. 5:304-
1811) cited the derivation of gramma as from the Greek meaning ‘“‘line.’’ In this
sense, the word gramma (ypauud) is the Doric form of the more usual Attic
Greek gramme (ypaupr) (Pichi-Sermolli, Webbia 21:487-505. 1966). Both words
have the Same meaning and both are feminine in gender. The problem arises from
the similar-looking Attic Greek word, gramma (ypappa), which is neuter and
means “that which is drawn’’ or ‘‘letter.’’ Some authors, unaware of the feminine
Doric form gramma and/or confused by the identical spelling of the different
neuter Attic word gramma, have decided that Desvaux’s use of gramma as
feminine was improper and that the word should either be considered as neuter oF
should be changed to gramme. However, since Desvaux specified that gramma
means *‘line,”’ and treated it as feminine, he clearly was intentionally and cor
rectly employing the Doric form of gramme. Modifications of the generic name to
Gymnogramme or specific epithets to a neuter ending are not correct.

This clarification has a rather wide application since there are names of many
fern genera based on gramma or gramme (e.g. Anogramma, Coniogramme, etc.)
In all cases when these are derived from the Greek word meaning ‘‘a line,”’ the
original spelling, whether as gramme or gramma, should be maintained and
epithets should have a feminine termination. If there are any derived from the

AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 1 (1979) 31

Greek word meaning “that which is drawn”’ or ‘‘a letter,”’ these also should retain
their original spelling but with epithets of a neuter termination. Where a derivation
is not included, gender of the name should follow that published with the
name.—Christopher H. Haufler and Rolla M. Tryon, Gray Herbarium, Harvard
University, 22 Divinity Avenue, Cambridge, MA 02138.

REVIEW

“FERNS,” by Philip Perl and the Editors of TIME-LIFE Books, Alexandria,
Va. 1977. 159 pp. + many photographs; numerous watercolors by Richard Crist.
$8.95.—To say that this volume is beautifully illustrated and contains a wealth of
information for the amateur fern grower is true, but it also is an oversimplification
that does not do justice to the term ‘‘review.”’ In fact, this book is a good example
of a modern, mass-produced encyclopedia volume. It is an attractively packaged
hodge-podge of enticing tidbits of information which are priceless for beginning
fern growers. Unfortunately, the editors were too intent on interjecting poetic and
artistic touches, rather than on fulfilling their implied purpose of giving com-
prehensive information on fern growing, including recent and advanced knowl-
edge of the subject.

The introductory chapter is a delightfully written historical sketch, but is bro-
ken up by the inappropriate interposition of photographic essays having little orno
relationship to the text. The remaining chapters fare a bit better in this regard.
There is much to be gleaned from the horticultural information in chapter four,
although I doubt that many readers will find a ready supply of builder S sand.
Perlite, which is available nearly everywhere and which is included in most soil
Preparations based on the Cornell mix, is omitted here, although many of the
cultural suggestions in chapter five include it.

The photographic illustrations are superb, especially those in the essay on
identifying ferns by family. On the other hand, I found the delicately attractive
watercolors in chapter five to be little more than decorative. They have ye
ethereal quality not always representative of the stark greens and bold forms -
the species they pretend to illustrate. For example, the excellent photographs 0
the Hacksaw fern on pages 70 and 72 bear little resemblance to the illustration on
Page 110. A number of the watercolors appear so similar as to be of little value in
distinguishing between superficially similar but distinct genera.

32 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

The fern encyclopedia comprising chapter five is the meat of the volume, and
occupies the second half of the book. It contains some identifying characteristics
of a variety of ferns, mostly cultivated, their provenance, and hints on their
culture, with recommended temperature, foot-candles of light, and pH readings ad
infinitum, as if the latter two were all that important. I am not convinced that the
average fern grower will ever determine his foot candles or pH, or even care.
Elsewhere in this work the ubiquitous Asparagus-fern is defrocked, but in chapter
five, Selaginella and Equisetum are treated as if they were legitimate ferns. An
extensive and useful bibliography is given on pages 152-153.

Considered as a whole, even with its idiosyncracies, this is a handy and attrac-
tively priced volume for fern growers.—Robert Read, Department of Botany,
Smithsonian Institution, Washington, DC 20560.

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Plant Classification
Second Edition

Lyman Benson, Pomona College, Emeritus
1979 Casebound 736 pages

This classic text has been revised to include recent research:
new examples for major manuals on floras in the Pacific
Northwest, California, Northeast, and South; reassignment of
the members of the Amentiferae to new positions in the
Thalamiflorae and Calyciflorae; revisions in the tribes of the
grasses; recent findings in evolution; and classification systems of
flowering plants. Plant Classification, Second Edition is a superbly
illustrated text/reference book written to help readers: acquire
essential vocabulary for describing characteristics of plant groups;
identify plants by applying the use of keys and descriptions; gain
knowledge of plant.taxa through preparation and preservation of
specimens to form an ordered collection; develop an under-
standing of the basis for classification of plant groups; and gain
an appreciation of the association of species in natural vegetation.

Section Heads:

Flowering Plants
Gymnosperms

erns

Psilophytes and Horsetails
Club Mosses

Association of Species in

Natural Floras
HEATH

Appendix
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.
Treasurer
JUDITH E. SKOG, Dept. of Biology, George Mason University, Fairfax, Va. 22030.
Records Treasurer
DAVID B. LELLINGER, Smithsonian Institution, Washington, D.C. 20560. Journal Editor
ALAN R. SMITH, Dept. of Botany, University of California, Berkeley, Calif. 94720

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

ASSOCIATE EDITORS
DAVID W. BIERHORST ..Dept. of Botany, University of Massachusetts, Amherst, Mass. 01002
GERALD J. GASTONY ...........,, Dept. of Biology, Indiana University, Bloomington, Ind. 47401
JOHN T. MICKEL New York Botanical Garden, Bronx, New York 10458

The ‘American Ferm 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.

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.

Memoir
Dr. Alan R. Smith, Dept. of Botany, Univ. of California, Berkeley, California 94720, is Editor of
the memoir series “ Pteridologia.”’
Newsletter
= eateries Mickel, New York Botanical Garden, Bronx, New York 10458, is editor of the
sien omega Forum.”” The editor welcomes contributions from members and oe
hembers, including miscellaneous notes, offers to exchange or purchase materials, personalia, hor-
ticultural notes, and reviews of non-technical books on ee. :
Library
New York Botanical Garden, Bronx, New York 10458, is Librarian. Meare
any time, the borrower paying all shipping costs.
Spore Exchange

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

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

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

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)

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

of slide manufacture
and service to botanists.

We welcome samples of your preserved research
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tinued need for quality prepared slides, address
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O:

TRIARCH INCORPORATED
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.
E/2 Jhandewala Ext., New Delhi-110055, India

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

a details or sample copies, call us toll free: mag ad 1388.
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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

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

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

ASSOCIATE EDITORS
DAVID W. BIERHORST ..Dept. of Botany, University of Massachusetts, Amherst, Mass. 01002
GERALD J. GASTONY +++r+eeseeee-Dept. of Biology, Indiana University, Bloomington, Ind. 47401
JOHN T. MICKEL. New York Botanical Garden, Bronx, New York 10458

The **American Fem Journal’ (ISSN 0002-8444) is an illustrated quarterly devoted to the general
study of fems. It is owned by the American Fern Society, and published at the Smithsonian Institu-
tion, Washington, DC 20560. Second-class postage paid at Washington.

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

Changes of address, dues, and applications for bership 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 Fem Society (annual dues, $8.00; life membership, $160.00). Extracted
offprints, if ordered in advance, will be furnished to authors at cost, plus postage.

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

Spore Exchange

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

—— Gifts and Bequests
| tsand beq the Society enable i ‘ é bers and to others interested
in ferns. Botanical books, back issues of the Joumal, and cash or other gifts are always welcomed, and
are tax-deductible. Inquiries should be addressed to the Secretary.

AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 3 (1979) 65

A New Nephrolepis Hybrid From Florida
CLIFTON E. NAUMAN*

Nephrolepis is a genus of probably 30 species mostly pantropical in distribution.
Several species have been problematical in their identification, notably N. biser-
rata (Swartz) Schott, N. exaltata (L.) Schott, and N. hirsutula (Forst.) Presl
(Christensen, 1932, p. 74; Copeland, 1947, p. 91). Two of these species, N. exal-
tata andN. biserrata, occur in Florida and have been a source of difficulty, in part
because of plants intermediate in a number of characteristics. The intermediates
appear to represent hybrid populations between these two species.

Distributional data show that intermediates occur only in the overlapping por-
tions of the parent species’ ranges and not where only one parent occurs. Inter-
mediates are found wherever N. exaltata and N. biserrata occur together, usually
in disturbed portions of the habitat. The intermediates are more abundant and
luxuriant as would be expected of hybrids.

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

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)

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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)

FIG. 19. Anemia spore ornamentation. A. phyllitidis. Laughlin 2563. F1G. 20. A. munchii, Hill s.n.
FIG. 21. A. pohliana. Chase s.n. FIG. 22. A. ouropretana. Irwin et al. 27976. Line scale = 10 »m.
measuring 40 «wm in diameter. The ridges lack microspines, and the baculae ap-
proach a clavate shape as in A. ouropretana and A. radicans.

The last group examined included two taxa, A. oblongifolia and A. hirsuta, both
wide-ranging species. The spores of A. oblongifolia were seen to fall into two
groups which were correlated with two vegetatively distinct groups of specimens.
Group I may be exclusively Brazilian and can be characterized by semi-erect
fertile fronds which are long-stalked (stipes up to 10 cm long) and which bear
narrow pinnae 3-5 mm wide. The spores of this group (Figs. 13, 14, and 24) differ
from all of the other spore samples examined by the presence of wide ridges,
narrow fossulae, and the complete absence of conspicuous baculae. In addition,

S. R. HILL: SPORE MORPHOLOGY OF ANEMIA

FIG, 23. Anemia spore ornamentation. A. rotundifolia. Rose & Russell 20099. FIG. 24. A. oblon-
gifolia, Group 1. Irwin et al. 11793. FIG. 25. A. oblongifolia, Group II. Irwin et al. 13216a. FIG. 26.
A. hirsuta, Hil 1643. Line scale = 10 um.

the spore angles are somewhat prolonged as seen in proximal view and the or-
namentation of the ridges consists of small (2 «m), branched microspines. These
characteristics are also typical of spores of members of subg. Coptophyllum (Hill,
1977). Plants in morphological Group II have low, short- stalked fronds (stipes up
to 2-3 cm long) with comparatively broader pinnae 6-8 mm wide. The spores
(Figs. 15, 16 and 25) have baculae typical of A. phyllitidis and its allies present in
reduced form (3-5 um long), and the occasionally branched microspines of the
Coptophyllum g group are also found among them. The fossulae and ridges are
intermediate in width between typical examples of each subgenus. The spores

AMERICAN FERN JOURNAL: VOLUME 69 (1979)
78

A.oblongifolia 2

A. hirsuta

A.oblongifolia 1

A. pohliana A. pastinacaria

A. radicans

A. rotundifolia
A. munchii

FIG. 27. Possible morphological rel

Bone nemia.
ationships among selected spore types of subg. A
Hypothetical connection with

subg. Coptophyllum at upper right.

S. R. HILL: SPORE MORPHOLOGY OF ANEMIA 79

seem to provide a morphological link between the A. phyllitidis spore type and the
Group I A. oblongifolium spore type described above, which is more typical of
spores of subg. Coptophyllum.

The spores of A. hirsuta (Figs. 17, 18, and 26) pose an interesting problem in
subg. Anemia. The shape of the spores, the widely spaced, narrow ridges, and the
unbranched microspines ornamenting those ridges are typical of those of A. phyl-
litidis, but the large baculae of that group are entirely absent, and the spore angles
are slightly prominent.

DISCUSSION

Figure 27 illustrates a tentative scheme of morphologically related spore types
to summarize the observations presented above. This scheme is not intended to
represent a phylogeny, but is intended to express morphological similarities in the
specimens examined. Basic to this arrangement are the variable spores of Anemia
phyllitidis, to which several other spore types are connected. Anemia radicans, A.
ouropretana, A. munchii, and A. rotundifolia, which are all restricted, specialized
species, have spore types apparently related to and possibly even derived from the
pool of variation seen in A. phyllitidis. Anemia pohliana may be a hybrid derived
from members of this complex, but at any rate its spore type seems closely allied
to that of A. pastinacaria, whose spores suggest a trend either towards baculae
reduction as seen in spores such as those of A. oblongifolia, or in the opposite
direction towards baculae elaboration as seen in A. phyllitidis. The A. hirsuta
spore may be derived from that of A. phyllitidis directly through loss of the large
baculae while retaining the microspines, or may be separately derived from that of
A. oblongifolia by modification of the number of ridges and by simplification of
the complexity of the microspines.

The problem requiring the most study is the morphological assessment of the A.
oblongifolia complex. The spore analysis presented here suggests that more than
one species actually is involved. In addition, the great resemblance of the spores
of form 2 in Fig. 27 (referred to as Group I in the text above) with those of subg.
Coptophyllum may provide a link between the two subgenera. Since subg. Cop-
tophyllum is considered to be the most primitive of the three subgenera (Mickel,
1962), this taxon may aid in our knowledge of the origin of subg. Anemia.

I would like to thank Dr. Elenor Cox and the Electron Microscopy Center at
Texas A&M University for aid in obtaining access to the scanning electron micro-
scope, Dr. John T. Mickel of the New York Botanical Garden for support while
collecting spore samples in Oaxaca, Mexico, and Dr. David B. Lellinger of the
U.S. National Herbarium for kindly lending specimens used in this study.

LITERATURE CITED

optophyllum. Amer. Fern. J.
Anemia, subgenus C optophyl

HILL, S. R. 1977. Spore morphology of Anemia subgenus C oa et

MICKEL, J. T. 1962. A monographic study of the fern genus oe
State J. Sci. 36:349-482.

PRANTL, K. 1881. Untersuchungen zur Morphologie der ene

ceen. Wilhelm Engelmann, Leipzig.

skryptogamen. II. Die Schizaea-

80 AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 3 (1979)

The Development of Plantlets from Strobilus Branches
in Lycopodium phlegmaria
+: C. WEE

Lycopodium phlegmaria L. is an epiphyte commonly found growing on the
trunks and branches of forest trees in Singapore. It is a widely distributed species
in the Old World tropics, ranging from Africa to Polynesia. Its pendulous stems
are covered with spirally arranged, stiff, ovate-lanceolate leaves (Fig. 1). Both the
sterile portions of the stems and the strobili branch dichotomously (Figs. / and 2).
The sporophylls are smaller than the foliage leaves and are arranged more com-
pactly along the branches. In most cases, the sporophylls are grouped terminally
On determinate stems. However, instances of foliage leaves appearing along the
strobilus branches have been observed (Fig. 2). This is reminiscent of the condi-
tion seen in L. selago L., where the fertile zones alternate with the sterile and the
foliage leaves are not much different from the sporophylls (Bower, 1935, p. 200).

Recently, a portion of L. phlegmaria detached from the parent plant was found
lying on damp ground on the secondary forest at the MacRitchie Catchment area
in Singapore. It was still green, and the ends of the fertile branches showed growth
following detachment. Under the humid conditions of the forest floor, the ends of
the branches had reverted to vegetative growth and had produced one or two

with long internodes (Fig. 3). At the nodes were leafy structures similar to the
sporophylls, In the axils of these structures were obviously vegetative buds, some
of which had developed into vegetative shoots (Fig. 3).

Longitudinal sections of the strobilus branches revealed the gradual transforma-
tion of the sporangia from fertile to sterile in the acropetal direction. The fertile,

vermiculite similarly stimulated development of vegetative shoots after about four
months. The new growths had excessively elongate internodes
were smaller until foliage leaves were produced when the axis turned upwards.
Roots formed at the points of turning after a further period of two to four months.
These shoots could then be removed as plantlets.

These observations give support to Bower’s (1908, p. 164) theory that foliage
leaves essentially are sporophylls with the sporangia completely suppressed. In
this respect, it should be noted that the strobili of L. obscurum L. and L. flabel-

*Department of

Singapore Botany, University of Singapore, Bukit Timah Road, Singapore 10, Republic of

Y. C. WEE: PLANTLET DEVELOPMENT IN LYCOPODIUM PHLEGMARIA

PU Ag ted

they,

ca)

_ 1. Potted plant showing the pendulous habit and

‘ FIG. 2. Strobilus branches showing groups of foliage
owing branch tips developing

new growth at the end of a

FIGS

> eta l-4. Lycopodium phlegmaria. FIG

ics, ge branched strobili (scale in cm)

into oo, Detached portion of plant found on the forest floor sh
(scale in mm). FIG. 4. Close-up of portion of Fig. 3 showing

Strobilus br:
bilus branch (scale in mm).

82 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

liforme (Fern.) Blanch. also have been observed to produce vegetative exten-
sions, but not plantlets (Bierhorst, 1971, p. 15). In Selaginella, however, Goebel
(1930, p. 657) managed to induce pieces of strobili to develop foliage leaves by
placing them on a moist medium

I wish to record my thanks to Prof. A. N. Rao for his assistance in the anatomi-
cal study and for critically reviewing this paper; to Mr. J. Wee for bringing the
piece of L. phlegmaria to my attention and for technical assistance; and to Mr. D.
Teow for the photographs.

LITERATURE CITED

BIERHORST, D. W. 1971. Morphology of Vascular Plants. Macmillan, New York.
BOWER, . a 1908. The Origin of a Land Flora. Macmillan, London

—__—.. . Primitive Land Plants. Macmillan, London

commie. <e 1930. Organographie der Pflanzen. Fischer, fet:

TRIARCH
Over 50 Years

of slide manufacture
and service to botanists.

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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

“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

. 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

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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

ea,

,
3,
CYS

8,
cs
pe
:
“ts

oo ey
SH rae egueme
SES re
Stee See
re QPEL
eae 2

&
a)
Be
e

S. CHANDRA: GAMETOPHYTE MORPHOLOGY OF DRYNARIOPSIS 113

initial cell enlarges and divides (Fig. 2) by a wall perpendicular to the first wall,
one that is parallel to the polar axis of the spore. The germ filament elongates
along the equatorial plane of the spore (Figs. 2 and 3) perpendicular to the first
rhizoid. Due to the physical obstruction provided by the spore coat, the germ
filament is deflected and the rhizoid is pushed away to one side. The rhizoid,
which usually emerges first, is dark brown and devoid of chloroplasts. This is
termed Vittaria-type spore germination by Nayar and Kaur (1968, 1971). The germ
filament elongates, becomes 4-8 cells long (Fig. 4), and is composed of short,
stout, barrel-shaped, densely chlorophyllous cells. The basal cell is short, often
rather bulbous, and may bear two or more rhizoids. The germ filaments often are
branched (Figs. 5, 9, 10, 12, 19, and 30), sometimes profusely so, with the
branches developing into separate gametophytes, in contrast to Pseudodrynaria
(Nayar, 1954) and Drynaria (Nayar & Kachroo, 1953), in which branching has
been reported to be rare. Development of the gametophytic plate is initiated when
the germ filament is 4-8 cells long, and may be either Drynaria-type or Aspidium-
type.

Drynaria-type development.—The terminal cell of the germ filament and two or
three cells behind it divide transversely and longitudinally to produce a spatulate
gametophytic plate (Figs. 5 and 6). The plate often becomes 5-10 cells wide and
broadly ovate, but is devoid of any organized meristem (Fig. 7). Only rarely is an
apical cell established early (Figs. 16-18), in which case the young gametophyte
becomes cordate. For the most part, an obconical apical meristem cell is differ-
entiated later by two oblique divisions of one of the marginal cells at the anterior
end of the gametophytic plate (Fig. 8). The apical cell undergoes a considerable
number of divisions and the gametophyte becomes cordate (Fig. 12) before the
apical cell is replaced by a multicellular meristem (Figs. 9-11 ). In Drynaria-type
development, the establishment of an apical meristematic cell usually is much
delayed (Figs. 16-18), and the gametophytes usually develop hairs on the margin
when the gametophytic plate becomes cordate; the younger gametophytes are
naked. In contrast, in Drynaria rigidula, an apical meristematic cell is established
early, at the second division of the terminal cell, often initiating it (Nayar, 1965).

Young gametophytes with slightly asymmetrical meristemati
gametophytes. FIG. 36. Superficial, stalked hair. FIG. 37.
shaped gametophyte.

Marginal hair. FIG. 38. Young, strap-

114 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

As in Aglaomorpha (Nayar, 1965), the young gametophytes of Drynariopsis often
are broader than long, whereas those of Drynaria and Pseudodrynaria are
elongate-oblong at this stage (Nayar, 1965). Soon the apical meristematic cell
becomes positioned at the base of the apical notch. It is then replaced by a
multicellular meristem (Fig. 13) in the usual way, by a transverse wall in the
meristematic cell followed by vertical walls in the outer cell.

Under cultural conditions, the gametophytes become distinctly cordate with a
multicellular meristem and cluster of rhizoids on the lower surfaces (Figs. 14 and
15) about 10-12 weeks after spore germination. The midrib is initiated when the
gametophytes are about four months old. The midrib is thin and very conspicuous.
The mature gametophytes are cordate, broader than long, grow flat on the sub-
strate, and have a deeply notched apex, as in the other species of drynarioid ferns.
The meristem consists of 3-5 elongate cells perpendicular to the surface. The
lobes are composed of uniformly thin-walled, densely chlorophyllous, polygonal
cells arranged in more or less radiating rows. In contrast to Merinthosorus (Baj-
pai, 1964), the walls of the wings do not possess collenchyma-like thickenings at
the corners. As in other drynarioid ferns (Nayar & Kachroo, 1953; Nayar, 1954,
1961, 1965; Bajpai, 1964), the marginal cells of young gametophytes develop uni-
cellular, papillate hairs, which begin to form only after the gametophytes have
become distinctly spatulate. The hairs are thin-walled, cylindrical, with a rounded
anterior end, and are chlorophyllous. They originate as a protuberance of one of
the superficial cells, which elongates and is cut off as a hair. When young, the
hairs have a swollen tip with a prominent nucleus surrounded by a few vacuoles
on all sides (Fig. 37). Simple, club-shaped hairs like those found in Aglaomorpha,
Merinthosorus, Photinopteris, and Pseudodrynaria rarely are found on the sur-
face of Drynariopsis gametophytes (Fig. 36). Branched, club-shaped, polypodioid
hairs as found in other drynarioid ferns (Nayar, 1965) and most other
poly podiaceous genera, are absent in Drynariopsis.

Drynariopsis gametophytes are attached to the substrate by unicellular, elon-
gate rhizoids. On adult thalloid gametophytes, the rhizoids generally are restricted
to the lower surface of the midrib. Rhizoids generally are absent on the wing and
margins of cordate-thalloid gametophytes, but in strap-shaped gametophytes,
marginal rhizoids are often very frequent and in some cases are characteristically
tec as on the gametophytes of the Grammitidaceae and some

olypodiaceae (Nayar & Raza, 1970). The rhizoids are soft and nearly hyaline; the
rhizoid wall is markedly colored, and often is reddish brown, even at an early
developmental stage.

Aspidium-type development.—In contrast to Drynaria-type gametophyte on-
pr ; the Aspidium-type is variable in Drynariopsis with regard to the sequence
ne po . All variations characteristic of Aspidium-type development have
t ot served in Drynariopsis. The most common is that the filament terminates
“ae unicellular hair (Fig. 20). The terminal cell becomes beaked, and the protrud-
ics Pipes is cut off by a transverse wall. Plate formation is initiated by cells

ind the terminal cell dividing longitudinally (Figs. 2/-23). Sometimes the
penultimate cell of the germ filament also is sluggish. A broad, usually lopsided

S. CHANDRA: GAMETOPHYTE MORPHOLOGY OF DRYNARIOPSIS 115

plate is formed by differential cell division behind the sluggish anterior region
(Figs. 23 and 26). The larger side often bulges out, pushing the terminal hair to one
side. All cells except the basal cell may divide longitudinally and take part in plate
formation. An obconical meristematic cell is differentiated laterally in the thallus
by two oblique divisions in one of the marginal cells on the more expanded side of
the plate (Figs. 23 and 26). It may be formed either early during plate formation or
only after the plate becomes several cells wide. This type of gametophyte de-
velopment has been reported in Christiopteris (Nayar, 1967), Merinthosorus (Baj-
pai, 1964; Nayar, 1965), and Platycerium (Bauke, 1878; Orth, 1936; Stokey &
Atkinson, 1954). Sometimes the terminal cell of the germ filament, after producing
a hair, divides longitudinally. One of the daughter cells then produces a hair (Fig.
27). The hair-bearing cell is sluggish, but the other daughter cell actively divides
by an oblique division to form a meristematic cell (Fig. 28, M). The resulting plate
may be slightly asymmetrical (Figs. 29 and 3/). Marginal, unicellular hairs are
produced continuously, whether a meristematic cell is differentiated early or not.
In some gametophytes, a plate is initiated prior to hair formation. The terminal cell
divides longitudinally (Fig. 24), with one daughter cell initiating plate formation. A
lateral meristematic cell forms in the plate (Fig. 25). The other daughter cell of the
terminal cell remains sluggish, and may bear a hair.

The gametophyte grows by the activity of the meristematic cell. Since this cell is
lateral, the young gametophyte is lop-sided. The asymmetry is more marked when
meristematic cell formation is delayed. Soon the meristematic region becomes
notched and apical by unilateral growth (Fig. 7). As the gametophytes grow,
however, the asymmetry is lost, and the adult thallus is symmetrically cordate
(Fig. 15). Gametophyte development beyond the establishment of an apical meri-
stematic cell is similar to that in Drynaria-type.

Ameristic gametophytes.—Occasionally ameristic gametophytes occur in cul-
ture (Figs. 32-35). In such cases, a single, obconical meristematic cell may not be
developed at all, and the marginal cells on one side develop a multicellular meri-
stem directly. Ameristic gametophytes also have been reported in Christiopteris
(Nayar, 1967) and in Merinthosorus (Bajpai, 1964).

In some cases, the gametophytes elongate by diffuse growth, become irregular
and strap-like with a rounded apex, and often bear profuse marginal rhizoids (F : é
38). No organized meristem is ever differentiated, but the cells constituting the
anterior region of the strap-like gametophyte are smaller in size and gps more
actively than do the cells in the posterior half. No midrib is produced. T € rise
stem is neither very active nor well defined. All cells of the anterior region o : oi
meristematic activity. This type of gametophyte development, —
Kaulinia-type by Nayar and Kaur (1969, 1971), has been reported in many g
of the Polypodiaceae (Nayar & Kaur, 1969).

Mature gametophytes.—The ee i , ae
14-16 weeks after spore germination. The mature g cae
bear sex organs aie pesca on the lower surface of the meen seo ti
are the common type found in the Polypodiaceae (Nayar, ste a sheeple
superficial, but on gametophytes growing under crowded conditions,

heraclea reach maturity about
tophytes are cordate and

116 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

may develop along the gametophyte margins. The antheridia are small, sub-
globose, and have an undivided opercular cell and a funnel-shaped basal cell.
Schlumberger (1911) and Schmelzeisen (1933) have reported the occasional occur-
rence of a divided cap cell of the antheridium in Drynariopsis, but these were not
observed in the present study. Archegonia are produced after midrib formation
and usually are found mixed with antheridia. The archegonia are the usual type
reported in advanced leptosporangiates (Stokey 1951; Nayar, 1962), with a short,
slender neck composed of four rows of cells having three or four short cells in
each row. The neck is curved away from the apex of the gametophyte, and the
neck canal cell is binucleate and swollen toward the tip at maturity.

DISCUSSION

The drynarioid ferns are phyletically a problematic group, and commonly are
ascribed a microsorioid ancestry. Among the drynarioid ferns, Drynariopsis usu-
ally is regarded as the most primitive genus and is considered to be nearest the
group’s probable ancestor (Copeland, 1947; Holttum, 1947, 1954; Nayar & Devi,
1964; Nayar, 1965).

Drynariopsis exhibits several peculiarities in its gametophyte morphology
which are otherwise unknown in the Polypodiaceae. The gametophytes are naked
in early stages, and the obconical meristematic cell may be differentiated quite
early in development, as in Microsorium, or it may be differentiated only after the
formation of a spatulate plate, as in Drynaria and Pseudodrynaria (Nayar, 1954,
1959, 1961, 1965; Nayar & Kachroo, 1953). Besides the common Drynaria-type
gametophyte development, which is characteristic of the drynarioid ferns and
most of the Polypodiaceae, Drynariopsis gametophytes exhibit all the variations
characteristic of Aspidium-type development. This is uncommon in the Poly-
podiaceae, but has been reported in Merinthosorus drynarioides (Bajpai, 1964;
Nayar, 1965), Christiopteris (Nayar, 1967), and Platycerium (Bauke, 1878; Orth,
1936, Stokey & Atkinson, 1954). Among ferns of other phyletic lines, this pattern
is characteristic of Blechnaceae, Davalliaceae, Dryopteridaceae, and Olean-
draceae (Nayar & Kaur, 1969, 1971). However, a strap-shaped gametophyte simi-
lar to that of Drynariopsis is found in C hristiopteris and in several microsorioid
genera of the Polypodiaceae, e.g., Colysis, Pleopeltis normalis (Nayar, 1962),
Dendroglossa (Nayar, 1963a), Leptochilus, Paraleptochilus (Nayar, 1963b), and
Lepisorus (Nayar & Raza, 1970).

Drynariopsis heraclea is unusual among most of the other drynarioid ferns in its
gametophyte development, but shows similarities with Merinthosorus in having
Aspidium-type development. The possibility of segregating Merinthosorus from
other drynarioid ferns on the basis of gametophyte morphology, as suggested by
Nayar (1965), can be ruled out. Copeland (1947) considered Merinthosorus to
have evolved from Microsorium through Aglaomorpha meyeniana, but the
similarities | in gametophyte development of Merinthosorus with those of

Drynariopsis indicate that Merinthosorus is nota derivative of Aglaomorpha. And
Holttum S contention (1954) that Merinthosorus has been derived from
Drynariopsis is supported.

S. CHANDRA: GAMETOPHYTE MORPHOLOGY OF DRYNARIOPSIS 117

On the basis of comparative gametophyte morphology, it is likely that Drynari-
opsis is more intimately related to Merinthosorus than to any other drynarioid
fern, and possibly the latter is derived directly from Drynariopsis. The similarities
with Aglaomorpha in vegetative structures and in fertile fronds presumably are
due to parallel evolution.

I am greatly indebted to Prof. Prescillano M. Zamora, Department of Botany,
University of the Philippines, Diliman, Quezon City, for constant encouragement
and helpful suggestions.

LITERATURE CITED

ATKINSON, LENETTE R. and ALMA G. STOKEY. 1964. Comparative morphology of the
gametophyte of the homosporous ferns. Phytomorphology 14:51-70.
BAJPAI, N. 1964. Gametophyt phology of Merintt Copel. J. Indian Bot. Soc. 43:549-555.
BAUKE, H. 1878. Zur Kenntniss der sexuellen G tion bei der Gattungen Platycerium, Lygodium
und Gymnogramme. Bot. Zeit. 36:753-760, 769-780.
BOWER, F. O. 1923-28. The Ferns, vols. I-III. Cambridge Univ. Press, Oxford.
COPELAND, E. B. 1947. Genera Filicum. Ronald Press, New York.
HOLTTUM, R. E. 1947. A revised classification of Leptosporangiate ferns. J. Linn. Soc. (Bot.)
53:123-158.
————. 1949. The classification of ferns. Biol. Rev. 24:267-296.
————. 1954. Flora of Malaya, vol. II. Ferns of Malaya. Gov’t. Printing Office, Singapore.
NAYAR, B. K. 1954. Studies in Polypodiaceae-II. Contributions to the morphology of Pseudo-
drynaria coronans (Wall.) C. Chr. Phytomorphology 4:379-390.
. 1959. Studies in Polypodiaceae-VI. Further observations of the morphology of Drynaria
Bory. J. Univ. Gauhati 9:95-103.
————. 1961. Ferns of India II. Drynaria and Pseudodrynaria. Bull. Natl. Bot. Gard. 56:1-30.
—————.. 1962. Morphology of the spores and prothalli of some species of Polypodiaceae. Bot. Gaz.
123:223—232. ; :
. 1963a. Contributions to the morphology of some species of Microsorium Link emend.
Copel. Ann. Bot. n.s. 27:89-100. :
. es. Contributions to the morphology of Leptochilus and Paraleptochilus. Amer. J. Bot.
50:301-308.
. 1965. The gametophyte and juvenile leaves of the drynarioid ferns. Bot. Gaz. raspation
. 1967. Morphology of the spores and prothallus of Christiopteris tricuspis. Amer. Fern. J.
nape 1964. Spore morphology of Indian ferns-III. Polypodiaceae. Grana Palynol.
er ey Phen oe. 1953. Studies in Polypodiaceae-I. Contributions to the Seti of
Drynaria Bory: D. quercifolia (L.) J. Sm. and D. propinqua (Wall.) J. Sm. Phytomorphology
3:411-423.
_ Spore germination in homosporous ferns. J. Palynol. 4:1-14.
a eae pre ee prothallial development in homosporous ferns. Phytomor-
hology 19:179-188. é
: py KAUR. 1971. Gametophytes of homosporous ferns. Bot. ee iy i
————, and F. RAZA. 1970. The prothalli of some Polypodiaceae-II. J. ore eile ae
ORTH, H. 1936. Morphologische und physiologische Untersuchungen an Farnp

25:104—150. i d die
SCHLUMBERGER, 0. 1911. Familienmerkmale der Cyatheaceen und der Polypodiaceen, unt! te

Beziehungen der Gattung Woodsia und verwandter Arten zu

102:383—414.,

118 AMERICAN FERN JOURNAL: VOLUME 69 (1979)

SCHMELZEISEN, W. 1933. Beitrage zur Entwicklungsgeschichte der Prothallien einiger Marattia-
ceen, Cyatheaceen und Polypodiaceen. Flora 127:46—80

STOKEY, ALMA G. 1951. The contribution by the gametophyte to the classification of the homo-
sporous ferns. Phytomorphology 1:39-58.

. 1960. Multicellular and branched hairs on the fern gametophyte. Amer. Fern J. 50:78-87.

, and LENETTE R. ATKINSON, 1954. The gametophyte of five species of Platycerium.

Phytomorphology 4:165-172.

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AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 4 (1979) 119

A New Species of Selaginella from India
C. B. GENA, T. N. BHARDWAJA and A. K. YADAV*

During a recent survey of pteridophytes of southeastern Rajasthan, India car-
ried out in September 1977, we collected a Selaginella which is different from all
the known species of this genus. It is being described now as a new species, and is
an addition to the 59 species of the genus already recorded from India (Alston,
1945; Panigrahi & Dixit, 1966a, p. 11, 1966b, 1967, 1968; Bole & Almeida, 1977).

Selaginella rajasthanensis Gena, Bhardwaja & Yadav, sp. nov.

Selaginella reticulatam affinis. Caulis prostratus, alterne ramificans: caules la-
terales non ramificantes praeter basalem tantum qui semel tantum dichotome
ramificans. Folia et sporophylla dimorpha et serrata. Folia lateralia elliptica,
mediana vero ovata; megasporophylla ventralia, cordata. Megaspora tuberculis
rotundis. Microsporophylla dorsalia, oblique lanceolata, costa cum carina ad-
axiali, margine serrata. Microspora exino laevi et crista sinuata et triradiata.

Stem glabrous, prostrate, bearing rhizophores throughout, 1.5-2.5 cm long, the
posterior part (S—7 mm) unbranched, the anterior part bearing alternate branches
up to 1.0 cm long, the first one from the base invariably dichotomously branched
in mature plants (Fig: 1). Leaves serrate, obtuse, dimorphic, the lateral ones (Fig.
2) elliptical or oval, 1.5 mm long, 1 mm wide, the median ones (Fig. 3) ovate, 1 mm
long, 0.5 mm wide. Strobili single at the end of every lateral branch, up to 7 mm
long, bearing two ventral rows of megasporangia and | to 3 microsporangia or
none at all dorsally. Mega- and microsporophylls dissimilar. Megasporophylls
cordate, 1.5 mm long, 1 mm wide, serrate, mucronate, borne in the same plane as
the lateral leaves. Megaspores light yellow, trilete, spherical, 224-240 um in
diam., with many small, round tubercules on the exine. Microsporophylls ob-
liquely lanceolate, 2 mm long, 1 mm wide, serrate, obtuse with a serrately mar-
gined adaxial keel or flap on the midrib (Fig. 5), borne in the same plane as the
median leaves, with only a few (1 to 3) bearing microsporangia. Microspores
brown, trilete, spherical, 38 4m in diam., smooth, the triradiate ridge wavy (Fig.
6).

TYPE: Kundakhoh, Shahabad, Kota, Rajasthan, India, growing on an isolated
moist rock, Sept 1977, C. B. Gena & A. K. Yadav (PUN 2610) (PUN; isotypes:
Pteridophyte Biology Lab., Govt. College, Ajmer, India, No. PBL/77/S1-6/28/
671, B, BM, CAL, K, LWF, NY, US).

The species is confined to Rajasthan, India and is fertile from September to
November. Selaginella rajasthanensis resembles most .s reticulata (Hook. &
Grev.) Spring in general appearance, but it is morphologically quite different from
the latter in the following respects: habitat (prostrate vs. erect), branching (un-
common, except for the lowest vs. repeated), rhizophore abundance (common vs.
absent), lateral leaf shape (elliptical or oval, serrate, obtuse vs. ovate-oblong,
aristate, acuminate), median leaf shape (ovate, serrate, obtuse vs. elliptical, aris-
tate, acuminate), strobilus size (long vs. short), disposition and number - ati
gia (megasporangia 12-14, ventral, microsporangia 1-3 or none at - . >
megasporangia 1-3, basal, microsporangia 9-12, distal); megasporophy pe

, Ajmer 305001, India.
*Pteridophyte Biology Laboratory, Department of Botany, Government College, Aj

FIGS

Late
x

AMERICAN FERN JOURNAL: VOLUME 69 (1979)

. 1-6. Details of S. rajasthanensis Gena, Bhardwaja and Yadav. FIG. 1. Habit, x 2.5. FIG. 2.
ral leaf, x 35. FIG. 3. Median leaf, x 85. FIG. 4. Megasporophyll, x 25. FIG. 5. Microsporophyll,

5. FIG. 6. Microspore, x 430.

C. B. GENA, ET AL.: A NEW SELAGINELLA FROM INDIA 121

(cordate, serrate, mucronate vs. deltoid, aristate, cuspidate); microsporophyll
shape (obliquely lanceolate, serrate, obtuse, with an adaxial, serrately margined
keel on the midrib vs. lanceolate, aristate, accuminate); megaspore size and or-
namentation (larger, the exine with many small, round tubercles vs. smaller, the
exine granular); microspore ornamentation (smooth with a wavy triradiate ridge
vs. spiny with a straight triradiate ridge).

This new species also differs conspicuously from the recently described S.
tama-montana Serizawa (1978) from Japan in branching pattern; the —_ rtary
dense mats. The number and disposition of mega- and microsporangi
tama-montana are rather like those of S. reticulata. The strobili of the aaa
species resemble those of S. reticulata in the number and disposition of mega- and
microsporangia, but sporophyll shape and size is characteristically like that of
vegetative leaves in the former. There are significant differences in mega- and
microspore size and ornamentation also. These three species may be identified as

ollows:

. Plants erect, rooting at the base only, lacking rhizophores. Plants repeatedly branched; leaves and
sporophylls aristate; strobilus with 1-3 basal megasporangia and many distal microsporangia;
megaspores granular; microspores spiny, the triradiate ridge straight ..................... S. reticulata

. Plants prostrate or creeping, with rhizophores present throughout
2. Lateral branches not divided, except for the lowermost, which dichotomize only once; leaves and

sporophylls serrate; strobili with many ventral megasporangia and only a few, irregularly scat-
tered microsporangia; megaspores with round tubercles; microsporophylls with a serrately ma
gined keel on the midrib; microspores smooth, the triradiate ridge wa vy ue S. rajasthanensis
2 Plants frequently branched; leaves and sporophylls ll
with 1-4 basal megasporangia and 6-10 distal microsporangia; megaspores with many reticula-
ee microspores with uneven tubercles, the triradiate ridge straight .......... . tama-montana
Thanks are due to Principal A. B. Mathur and Prof. A. N. Parashar, Govern-

ment College, Ajmer, for providing laboratory facilities, and to Prof. K. M.

Matthew of St. Joseph’s College, Tiruchirapalli, India, for rendering the Latin

description. The University Grants Commission, New Delhi, India provided fi-

nancial assistance for survey of Rajasthan Pteridophytes.

LITERATURE CITED

ALSTON, A. H. G. 1945. An enumeration of the Indian species of Selaginella. Proc. Natl. Inst. Sci.
India 11:211-235.
BOLE, ~ oh and M. R. ALMEIDA. 1977. Four new species of pteridophytes from Bombay Presi-
. J. Bombay. Nat. Hist. Soc. 74:320-325.
Stat G. and R. D. DIXIT. 1966a. tes on twelve interesting species of Selaginella from
India. Symp. Indian Sci. Congr., Chandigarh.
. 1966b. Studies in the systematics of inet Selaginella—I

_—

—

II. Proc. Natl. Acad. Sci. 36:102-

; . : :222-233, f.
eee. SGT. Studies in the systematics of Indian Selaginella-II. J. Indian Bot. Soc. 46:222-233,
1

: | te 8.
. 1968. Studies in the systematics of Indian Selaginella-I. Proc. Natl. Inst. Sci. India
09

34:1 : . Bot.
SERIZAWA, S. 1978. A new species of Selaginella from central Honshu of Japan. J. Jap
: 3

122 AMERICAN FERN JOURNAL: VOLUME 69 NUMBER 4 (1979)

A SIMPLIFIED NUTRIENT MEDIUM FOR GROWING FERN PROTHALLIA.
—Germinating fern spores and culturing prothallia for scientific or educational
purposes is usually accomplished by sowing the sterilized spores on Knudson’s
agar (Bot. Gaz. 79:345-379. 1925; Bot. Gaz. 101:721-758. 1940). The preparation of
this medium requires precise weighing of small amounts of essential minerals
which are dissolved in water, to which agar is added to solidify the medium. For
some time, my laboratory has been successfully using a medium consisting of a
solution of 15 gm of agar per liter of water in which is dissolved one commercial
nutrient plant tablet (Black Magic Plant Food Tablets, Black Magic, Inc., Box
578-P, Hermosa Beach, CA 90254). The resulting medium provides mineral nutri-
ent yielding fern prothallia in numbers and size equal to those grown on conven-
tional nutrient media requiring precise quantitative mineral additives. —Norman
P. Marengo, Dept. ef Biology, C. W. Post College, Long Island University,
Greenvale, NY 1154

AMERICAN FERN JOURNAL

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

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

‘ 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

ae nd flora, 5; Atlas

me, 70; Synaptospory: A
speci

Reynolds, T L. Apical dominance in Anemia phyllitidis
gametophytes, 92

Richardson, J. I. (see J. G. Bruce)
bi nigh

e).
Te ae fol pam in the Teen life cycle of [soétes butleri,
103

Salvinia minima, 15; rotundifolia, 15

Schizaeaceae

Shwariihhdie RoI be e G. P. Landry)

opoemnegte! 10. 12, 82, 83, 119; braunii, 86; flabellata, 9; involvens,
84; kra

a, 9-12, var. aurea, 9; oregana, 84; martensii var.
manana: 9 ar. variegata, 9; ee ee rajas-
thanensis "18-1 reticulata, 119, 121; tama—montana, 121; ver-
sicolor

Sexuality in hs ae ste oe
Short, J. W. Pa yilits assert newly discovered in Alabama,

As cael nutrient medium for prothallia, 122
Spore morphology of Anemia subgenus Anecenial 71
<aatidoior Db. (see JOG. )

ae in eos Lint eaigg ja “The — patterns and in-

Studies of the Azolla- Anabaent ahaa using Azolla mexicana,
lL. 1 labora
F punta y: A hypothesis (rev.). Ra
onomic revision of the New Zealand species of Asplenium
po

Taylors W.C.&D. M. pees Srey gate in Arkansas, 26
4,

Tectaria heracleifolia, 1 incisa, 14, 64, f. incisa, 16;

Thelypteris, 26, 27, 64: ee 16; grandis, 64; interrupta, 16;
kunthii, 16, 26-29; normalis, 16; noveboracensis. sik ovata V
ovata, 16; palustris. 26. var. haleana, 26, kak var. pubescens, 7
27; reptans 1 torresiana, 16, 26-28; totta, 16

. Lan
Three new Elaphopossums dics ‘uate 100
omlinson, P. B e D. C. Adams)
Pivieus aaa
roward pop vee Florida, 14

— R.M ec. up peat
Vittaria 29; lin ne.
niaindy lineata Selescos n Geor;
Webster. T. R. rh ata tere che wr vee sagenedi a

nat ocd um heanain S 80
Wendt, T. goes ality in Asplenium resiliens, 96
Wofford. B. E. (see W. M. Dennis)
os aia -: ag

I virginica, 8

hcotecamalg ‘ heels G. ade e)
Yadav, ec: i

pp. a oe (see ap H.

ERRATUM
Page 37, Table 2. For ‘““FLABELLIFORME”’ read ‘‘DIGITATUM.”’

NOTEWORTHY PUBLICATIONS OF THE CHRONICA
BOTANICA

BEDDOME, R. H. The Fems of British India. Includes Bangladesh,
Burma, Ceylon, Malaya, Nepal, Pakistan, and Tibet. Set of 2 vols.
Reprint ed. 1973. Available CB (cloth bound) or LB (half-leather

with gold lettering). CB Rs. 285; LB US $55.
. Supplement to the Ferns of Southern India and British India.

se eer Ss CB Rs. 85; LB US $18.50.

— ——. Handbook to the Ferns of British India. Reprint ed. 1970.
CB Rs. 100; LB US $21.

1979.

CB Rs. 550; LB US $115.

The Chronica Botanica Co.
E/2 Jhandewala Ext., New Delhi-110055, India

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