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American Fern Journal 
A QUARTERLY DEVOTED TO FERNS 


Published by the 


AMERICAN FERN SOCIETY 


9 
EpITors 
Cc. V. MORTON 
R. C. BENEDICT IRA L. WIGGINS 


A. C. SMITH 


VOLUME 50 — > 
1960—!4 


MONUMENTAL PRINTING COMPANY, BALTIMORE, MARYLAND 


MISsouR! BOTANICAL 
GARDEN LIBRAF 


RY 


Contents 


Votume 50, NuMBER 1, Paces 1-160, Issurp AprRIL 25, 1960 
The Genesis of the American Fern Journal —.... gee C. be be 
What is the Role of Spores in Fern Taxonomy? — __. Clai . Bro 
The Endemic Pteridophytes of the California Floral Pas nee 
John Thomas Howell 


First Illustrations of Ferns from Peru and Chile Joseph n 

Evergreer wl aganate erns and the Meanings of oe img Categories 

rth American oo. Warren H. Wagner, Jr. 

The ok he Peruvian Fern Rolla Tryon 
Survival of Hart’s-tongue Fern @ Central New York 

Mildred E. Faust 

Ferns and Allies in Kansas Ronald L. McGregor 


A Mule-train Trip to Sierra Mohinora, Chihuahua __ Donovan S. Correll 
Multicellular and Branched Hairs on the Fern Gametophyte 
Al 


ma G. Stokey 
io a in the Study of Dryopteris Hybrids Edgar T. Wherry 
Abnormal Nuclear Division in Fern Prothallia William J. Crotty 


On a Species of Dryopteris, Subgenus Pyenopteris 
otozi Tagawa and Kunio Iwatsuki 
The Correct Name for the Hybrid Dryopteris cristata ert ‘spinu ulosa 


s Ballard 

C eew FQUInehuI fcc ss — ct Boivin 
Two Problem-species: Schizoloma cordatum Gaud, and Syngramma 
Pree SAU es R. E, aise: 
peevopreris porreti m Bavaria: = oe Pe as oe Poelt 


Notes on Pteridophytes from Australasia and New Caledonia, 2 


Mary D. Tindale 
The Nomenclature of a Japanese Ophioglossum Hans P, Fuchs 
A Synopsis of Sceptridium in Japan oe Makoto Nishida 


The Identification of Aspidium distans Viv. 


Rodolfo E. G. sie Plane 
a, in the Genus Asplenium Found in Northwestern and Cen 
_E 


ORO Os eee 6 its a ae : ve 
Dicayitiok: on Cultivated Ferns, VI. The Ferns Curtently Known 

me tomolre oo . Morton 
Recent Fern Literature: Ferns of Alberta; Ferns of California eo 
American Fern Society 


bo 


<piediie ck come 


VotuMmeE 50, NuMBER 2, Pages 161-224, IssuED JUNE 28, 1960 


Hunting Ferns in the Barrancas of Chihuahua, Mexico 
Irving Knobloch 161 
Ferns in Cultivation, IL Sylvia Leatherman 168 
Some Fern Books for Different Regions of the United Siadey 
. V. Morton 169 
The Genesis of the American Fern Journal: Supplementary 
h C. Sade 179 
Tsoétes melanopoda in Southern Hlinois Robert H. pie Jr, 181 
The Smooth Scouring Rush and Its Complexities — — Richard L. Hauke 185 
The Gametophyte and Young Bpegnyte of Athyrium ior 
. K. Nayar 194 
Recent Fern Titerature 22003) 203 
Notes and News: Ophioglossaceae Wanted 
American Fern Society: Report of President; Report of Treasurer; 


Votume 50, NuMBER 3, PAGES 225-256, IssuED Oct. 18, 1960 


w I Became epee in Ferns —. Edgar T. Wherry 225 


ene in Oubangi, Con G. J. De Joncheere 228 
Ferns Cultivated in Sait caal Sadleria ....-.----—--—-+--- Barbara Joe 232 
Ferns in Cultivation, III. Growing ae Balls .. Sylvia Leatherman 235 
A New Native Hardy Plant Area at the New York Botanical Garden 

R. C. Benedict 236 


Two New Georgia Stations for Lycopodium eomplanatum var. 
fubelliforme 6 _ Thomas A. Hutto 240 
Vein Patterns in 1 Mierosorium seandens and Its Allies 
Mary T. Tindale 241 


A New Station for Trichomanes Petersii in Georgia —... Henry Bookout chi 
Recent Fern Literature 2. ----------——--—— ne 28 
Notes and News: Los Angeles Sons Society; Batters dein R. C. Ching 253 
» 
ee Porn Society 2 265 
Votume 50, NUMBER 4, PAGES 957-304, IssuED DEC. 23, 1960 
On the Lava Fields of Rangitoto —-----—----—--~ Marguerite Crookes bit 
Feeding Hardy Ferns: Wise, rae 2 Risky Methods 4. J. Maeself ai 
Azolla caroliniana Willd. in Georgia ----------- re Wilbur H. Duncan 26 


The Lyeopodiaceae and cu pinaluaaes of Towa _... Tom S. Cooperrider 267 


Patan i cuisiane 9 oe William D. Reese 269 
Ke. peraee Dryopteris Species Based on Characters of the 
MPRISHOTO ce ke ee ee Fern Ward Crane 270 

ie Notes on Ferns, a eg C. V. Morton 275 

Cytological Observations on the Himalayan oa of Athyrium and 

Comments on the Ev mgt Status of the Genus 
ea Mehra and 8, 8. Bir 276 
Notes and News: his PRGtOR TA pine Se ee a en 295 
. 296 


Vol. 50 January-March, 1960 No. 1 


American Fern Journal 


A QUARTERLY DEVOTED TO FERNS 


Published by the 


AMERICAN FERN SOCIETY 


ts 


Eprrors 
Cc. V. MORTON 
RC. BENEDICT IRA L. WIGGINS 


Siftieth COnniversa 


MONUMENTAL PRINTING COMPANY, BALTIMORE, MD. 


Entered as second-class matter at the post office at Baltimore, Md., under the Act 
of March 8.1870, Acceptance for mailing at special rate of postage provided for © 
Sits of February 28, 1925, embodied in Paragraph (d-2) Section 34,40 P. b. 


2 


The American Fern Society 


Council for 1960 
OFFICERS FOR THE YEAR 
Ouam A. Brown, Department of Botany, Louisiana State University, Baton 
Rouge, Louisiana President 
MaxceL RayMonp, Montreal Botanical Garden, 4101 East ~~. Street, 
Montreal, Canada ‘residew 
DowaLp Hurrieston, Longwood Gardens, Kennett Square, Pines 


Seoretary 

Water S. Puiiirs, Department of Botany, University of ssanpge too Tucson, 
Arizona 

C. V. Morton, Smithsonian Institution, Washington 25, D. C. 

Editor-in-Chief 

OFFICIAL ORGAN 
American Fern Journal 
EDITORS 

OC. V. Morron._____._._.....-.. Smithsonian Institutio: roby ee. m 25, D. C. 
R. C. BENEDICT. 185 Hall Street, Brooklyn 5, N. Y. 
Ira L. Wiaeins____. a ——— Stanford University, Stanford, Calif. 
A. 0. Suira_____.. ..Smithsonian Institution, Washington 25, D. C. 


An illustrated quarterly devoted to the general study of ferns. 

Matter for publication — be addressed to C. V. Morton, Smithsonian 
Institution, Washington 2 5, D. C. 

Subseription $2.50, —- of agency handling fees; sent free to — 
bers of the American Fern ae gprnkiores dues, $2.50; sustaining m 
bership, $5.00; - mooaberikte, $50.00). tye xe reprints, if peo aay in 
pao will be f urnished authors at cost. They should be ordered when 
proo: 

ack volumes $2.00 each; single back numbers 50 ach ; Cumula: ative 
Index to vols. 1-25, 25 cents. Ten per cent discount on pte of six volumes 
or more. 


ston, Longwood Gardens, Kennett Square, Pennsylvania. Mem bers and sub. 
aren should allow two months for chan; to take effect. 

cations for membership, subscriptions, orders for back numbers, and 
ne oe imess communications should be addressed to the Treasurer: Dr. 
pareasiy 8. ie. Department of Botany, University of Arizona, Tucson, 


LIBRARIAN AND CURATOR OF THE HERBARIUM 
Dr. H. W. Wagner, University of Michigan, Ann Arbor, Mich. 


A regular loan department is maintained in connection with the library 
and herbarium. Members may F eee books and specimens at any time, 
the wer paying all postal or ogy ote charges. The pages of the “| , Sones 
also are open to members who wish exchanges; a membership = 


to arrange 
published at intervals, to assist thos terested 
rt different localities @ interested in obtaining specim 


American Fern Journal 


Vou. 50 January-Marcu, 1960 No. 


if 


PLETED Se ANNIVERSARY 


Invited Papers Contributed in Commemoration of the 
Fiftieth Volume of the American Fern Journal? 


CONTENTS 
The Genesis of the American Fern Journal Ratpu C. BENEDICT 
What is the Role of Spores in Fern Taxon nomy?_.CLAIR A. BROWN 


The Endemic Pravidepkyiss of the California Floral Provine 
Joun T 


First Illustrations of Ferns from Peru and Chile — os 
wisi taeT Grapeferns and the Meanings of Infraspecific Categories 
s Used in North America Pterido: ofa 


The Ecology of Peruvian Ferns ——-———-——~——- 


ARREN H. WAGNER, JR. 32 
Routa TRYON 46 ° 


Survival of Hart’s-tongue Fern in Central New York : 
5 . 


Miuprep E. Faust 


Ferns and Allies in Kansas.__-—-------_-— 
A Mule-train Trip to Sierra Mohinora, Chihuahua 


Ronatp L. McGregor 62 + 


Donovan S. CORRELL 66 * 


Multicellular and Branched Hairs on the Fern ——. 
A G, STOKEY 


rogress in the Study of Dryopteris Hybrids ean T, WHERRY 


rere Nuclear Division in Fern Prothallia 
cota J. CRorry 


On the Species of Drop | —— nus Pyeno 
1 TAGAWA ret ge TWATSUKI 


The Correct Name for the Si Wieseuils marnten x spinulosa 
FRA 


98-9 


cIs BALLARD 105 * 


fees Borvin 107° 


New. Equisetun.._ 
Two Problemapesen Schizoloma cordatum Gaud. and peepee 
pinnate J. Smith R. E. Horrrum 109 + 


10ther invited papers will appear in oth 


reports of the officers and other Society matters will appear in number 
pC. V, Mj 


er issues during the year The 
b 


9 


a 


2 AMERICAN FERN JOURNAL 


Dryopteris borreri in Bavaria... J. PornT 114 * 
Notes on Pteridophytes from Australasia and New Caledonia, I 
Mary D. TINDALE 117 * 
The Nomenclature of a Japanese Ophioglossum___ Hans P. Fucus 124 * 
A Synopsis of Sceptridium in Japan Makoto NISHIDA 127 *¢ 
The Identification of Aspidium distans Viv. 
Ropotro E, G. Picnt-SerMoiur 133 ¢ 
Hybrids in the Genus Asplenium Found in Northwestern and 
Central Wncone a D. E. Meyer 138 « 


The Genesis of the American Fern Journal 


Rauteu C. Benepicr 


official publication? At the beginning of the Fern Society’s his- 
tory, in 1893, the infant society had for a time exercised such 
control but the numbers of members was so small that mainte- 
* nance of a journal was impractical and the Society arranged with 
Willard N. Clute to take over what had been called the ‘‘Linnaean 
Fern Bulletin’”’ as a personal venture. A year or so afterward, 
Mr, Clute shortened the name to “‘The Fern Bulletin’ as it re- 
mained through the rest of its twenty year existence. Part of the 
arrangement relating to the transfer of ownership called for the 


Genesis or FrrN JOURNAL 3 


designation of the Bulletin as the ‘official’? organ of the Fern 
Society. Each year the officers of the Society arranged with Mr. 

Clute for an agreed upon sum per member which the Society 
paid from dues received. 

Willard Clute’s contributions to the Fern Society were out- 
standing, both as the chief factor in its founding and for his 
successful development of the Fern Bulletin. For approximately 
the first ten years of its existence, there seems to have been an 
abundance of articles that were real contributions, many of 
which are well worth reading now. In fact there was so much 
original work being done by members that two substantial issues 
of papers read at two Society meetings in 1899 and 1900 were 
separately published as ‘‘Fernwort Papers.’”’ 

During the first decade of the twentieth century, contributors 
of articles for the Fern Bulletin became fewer. Older members 
who had been chief contributors in earlier years were passing 
out of the picture. While the Fern Society continued a steady 
increase in numbers, there was a growing feeling that as a well- 
established, botanical society, it had become substantial enough 
to be represented by a publication owned and controlled by its 
members. Fern articles which might well have gone to the 
Fern Bulletin were published in other, more general botanical 
pournals. 

Professor Clute’s interpretation of the decline of contribu- 
tors to the Fern Bulletin was that the Fern Society and the 
Fern Bulletin were coming to the end of a period of real need 
in the field of fern study. He announced that he would continue 
the Fern Bulletin to the end of a twenty year existence, even 
if he had to contribute all its pages himself. To a suggestion that 
he re-transfer the Bulletin back to the Fern Society, he returned 
a definite negative answer. If he had acceded we should now be 
entering upon the sixty-eighth year and volume of the Fern 
Bulletin instead of the fiftieth volume of the American Fern 
Journal. 

Such in brief was the background of developments which led 
in 1910 to the birth of the American Fern Journal. During that 


4 AMERICAN FERN JOURNAL 


year, a group of members set in motion two projects designed to 
determine whether the Fern Society should institute and publish 
its own official bulletin. Plans were made and carried out for the 
nomination of an independent slate of officers to be offered in 
competition with those nominated by the official nominating 
agency of the Society. The Constitution provided for such in- 
dependent nominations then as now, and a single such nomina- 
tion had been made for the secretaryship the preceding year. 
The independent nominee had received a majority of votes but 
the Society Council, as then constituted, had ruled that the 
official candidate should be declared elected. The second step 
undertaken was the preparation and printing of two issues of a 
fern publication to be supported by private contributions. 

The first issue of the new publication made its appearance 
during the summer of 1910 and a second issue appeared in the 
fall of 1910.1 The name chosen was the ‘‘ American Fern Jour- 
nal,’’ the size and format followed that of the Fern Bulletin. 
The seven sponsors signed the Foreword in the first issue. These 
Seven sponsors were: L. S. Hopkins, H. E. Ransier, E. J. Wins- 
low, R. C. Benedict, Philip Dowell, W. C. Barbour, and Nellie 
Mirick. It was my privilege and responsibility to assemble the 
articles and to see to the printing of the first 16-page issue. Mr. 

velyn J. Winslow, of Auburndale, Mass., took charge of the 
distribution and continued to act as the business editor for over 
twenty years more, storing the increasing number of back issues 
in private facilities. Those first two issues were sent to all mem- 
bers of the Fern Society. Others among the seven made their 
contributions in other ways. 

The following autumn, the other step in the program was 
carried out—the presentation of an independent slate of can- 


1 Since this is an account of the birth of a new publication, it is ap- 
propriate to note the place as well as the date of the happy event. The New 
ork Botanical Garden was ti e scene of the event and the venture was 
further supposted by the accession of several Garden staff members as new 
members of the Fern Society during 1911 


GENESIS OF FERN JOURNAL 5 


didates for the four offices of members known to be favorable 
to a Society-owned and controlled publication. All four were 
elected by a good majority and in January, 1911, the follow- 
ing officers became the new Executive Council: President, Dr. 
Philip Dowell; Vice-President, Miss Nellie Mirick; Secretary, 
M . Hopkins; Treasurer, Mr. H. G. Rugg. One of their 
first acts was the adoption of the American Fern Journal as the 
Fern Society’s official publication, with the incorporation of the 
two 1910 issues as parts of a six-number volume. Incidentally, 
1911 proved to be a banner year in the number of new members 
received and in the growth of the membership of the Society. 

Philip Dowell served as President for only one year, and was 
succeeded by C. H. Bissell, of Connecticut, for the two succeeding 
years. In those three years the basic policies by which the Society 
has been run ever since were established. The several accounts 
for ‘‘reserve funds’’ were established. Editors have been ap- 
pointed by the elected Executive Council. Naturally, successive 
Councils have been very happy to maintain in office editors like 
C. A. Weatherby (1915-1940), William R. Maxon (1941-1947), 
and Conrad V. Morton (1948-1960), whose services as the ‘‘ex- 
ecutive’? and responsible editors-in-chief have meant so much to 
the Fern Journal and to the Fern Society. The Journal has never 
had an ‘‘editorial’’ page as did the Fern Bulletin. When the 
Editor writes for the Journal, he does so in his individual, not 
his official capacity. One desirable change was made some years 
back in the Editor-in-chief’s position—he was made a non-elected 
member of the Executive Council. 

185 Haut Srreer, BRooKuyn 5, NEw YorK. 


6 AMERICAN FERN JOURNAL 


What Is the Role of Spores in Fern Taxonomy? 
Cuair A. BRowN 


Ferns are a group of plants that illustrate many changes in 
taxonomic concepts which are useful in establishing systems of 
classification and in the delimiting of genera and species. The 
Linnaean genera were distinguished on such gross morphological 
features as the shape, size, and placement of the sori. Later, 
stelar anatomy became an important criterion. Then such items 
as venation, origin of the sorus, indusium, development of the 
sporangium, scales, hairs, glands on the indusium, chromosome 
number, and, to a lesser extent, spore morphology were used to 
separate species. This is not a definitive paper on the use of 
spore morphology in fern taxonomy but rather an assessment of 
the role that may be attained by this much neglected fiel 

The concept of the structure and the terminology opuel to 
different portions of the spore is quite varied. Douglas H. 
Campbell (1905) described spore structure as follows: ‘‘The 
young spores are thin-walled, but later the wall becomes thicker 
and shows a division into two parts, an inner layer, which gen- 
erally shows a cellulose reaction and is called the endospore 
(intine), and an outer more cuticularized coat, the exospore 
(exine). In addition, a third outer coat (perenium, epispore) 
is very generally present.’’ Structurally, the spore consists of 
a protoplast, the living portion encased in a thin layer, the 
intine, which is surrounded by the exine. This in turn is gen- 
erally enclosed by one or more layers which have been termed 
perispore, epispore, perine, or sclerine by various authors. There 
is no unanimity of usage among those interested in spore mor- 
phology. 

It was recognized early that differences in the number and 
orientation of cleavage planes gave rise to two types of spores 
from the spore mother cell, namely, the tetrahedral and bilateral 
spores. These were so named because, in the case of the former, 
three planes or faces were common to the proximal side of the 
spores tetrahedral or bilateral, and Coniogramme with spores 


Spores IN FERN TAXONOMY 7 
four spores of a tetrad and at their junction developed a tetrad 
scar or Y-mark which was responsible for the term ‘‘trilete 
spore.’’ The second type of spore, the bilateral spore, was formed 
from the division of the spore mother cell by two divisions so 
that the proximal edges touched each other along one line of 
contact. Visually this is equivalent to quartering a sphere with 
two longitudinal cuts at right angles to each other. The contact 
scar of this bilateral spore was a single line, slit, or opening 
which was called a monolete scar or aperture. The terms tet- 
rahedral versus trilete and bilateral versus monolete are used 
more or less synonymously. 

It is surprising that, in all the detailed studies on the ontogeny 
of the sporangium, the number of cells of the annulus, the 
number of spore mother cells, and the number of spores in a 
sporangium, a definitive answer has not been given to the follow- 
ing questions: Can both monolete and trilete spores be produced 
in one sporangium, in one sorus on one frond or in one species? 

Selling (1944) compiled a list of species in the genera Ophi- 
oglossum, Botrychium, Angiopteris, and Polybotrya in which 
the leading type was the trilete spore and the occasional type 
was the monolete spore whereas this was reversed in species in 
the genera Psilotum, Marattia, Schizaea, Dryopteris, and Meso- 
chlaena. He observed spores devoid of any dehiscence mark in 
some species of Lycopodium, Psilotum, Ophioglossum, Crbotium, 
and Schizaea, and mentioned that this condition has been re- 
ported in certain hybrids. Selling (1946) cited Goebel and 
Tammes as observing both monolete and trilete spores in the 
same sporangium of Psilotum. In his 1944 paper for the same 
observation, however, there is no specific mention for the species 
listed that both spore types came from the same sporangium. Also 
he apparently added mention of the trilete form for Psilotum 
nudum from the literature as he wrote specifically “I have seen 
no such cases.”’ 

Copeland (1947) characterized the spore types of monotypic 
genera such as Hemipteris, Ochropteris, and Anopteris with 


8 AMERICAN FERN JOURNAL 


bilateral or tetrahedral. He characterized genera with several 
species in them, such as Pteris, with ‘‘spores tetrahedral or less 
commonly bilateral, smooth, tuberculate or sculptured.’ 

The writer is inclined to question some of the observations that 
a species can possess both spore types normally for the following 
reasons : 

First, the fundamental difference in methods of spore forma- 
tion seems too distinctive to occur in the same species, although 
it is true that bilateral microspores and trilete megaspores are 
well known in [soétes. 

Second, all the species the writer has examined (limited in 
number in relation to the world flora) had either trilete or 
monolete spores. 

Third, contamination is so easy. Fresh, shedding sporophylls 
of Osmunda regalis were collected for spores. A microscopic 
examination of the spore mass showed it to be predominately 

smunda spores but there was a considerable quantity of pine 
and oak pollen. The same observation was repeated on Osmunda 
cimnamomea. Occasional spores of a dryopteroid type were also 
found. The writer has found more pine pollen in the sori of a 


China, than fern spores. It is not unusual to find herbarium 
Specimens with a copious quantity of sporangia and spores that 
came from another sheet in the folder. 

MeVaugh (1935) published a study on fern spores with a key 


indicating that he was cognizant of possible contamination. 
Erdtman (1958) illustrated the proximal face of a Haplosoria 
spore found in a slide made from Serpyllopsis caespitosa var. 
densifolia. 

Palynologists such as Knox (1950), Selling (1944, 1946), 
Harris (1955), and Erdtman (1958) are much better acquainted 
with fern spores than are most fern taxonomists. Harris men- 
tioned one clear-cut occurrence of both spore types in Marattia 


Spores IN FeRN Taxonomy 9 


salicina in his detailed studies on the spores of New Zealand 
pteridophytes. Erdtman (1958) figured a general view of a 
monolete grain of Phanerosorus major whereas a. avaned 
palynogram has a trilete aperture. 

How many spores should be measured to give the size and 
range for distinguishing species? The philosophy of the investi- 
gator has a bearing. Some examine spores in several fields, pick 
out an average-sized spore visually, and then measure several. 
Then they look for smaller or larger spores in order to get the 
range of size variation. Others may measure a given number at 
random and average the spore sizes. Mrs. Knox measured 10 
grains. Selling regularly measured 25 spores when he could 
find that number. Harris reported that he measured 10 spores _ 
but, when he wished a critical comparison between two species, 
he measured 50 of each. The larger the number of spore meas- 
urements, the greater the degree of accuracy in determining 
average size and amount of variation. 

What effect does different processing methods and mounting 
media have upon spore size? Palynologists process peat by either 
of two methods. They boil the peat in 10% potassium hydroxide 
solution or in Erdtman’s acetolysis mixture (9 parts acetic 
anhydride plus 1 part sulphuric acid). Reference slides are 
also prepared by the same reagent used in disintegrating the 
peat. This produces a ‘‘fossilized’’ grain. The protoplast and 
intine are removed and the resulting structure is more com- 
parable to the grains found in processed peat. These reagents 
often destroy the perispore and leave a denuded exine. Many 
have interpreted these denuded exines as immature spores. 
Spores mounted in water tend to swell, and the perispores of 
the sensitive fern, Onoclea sensibilis, and of the ostrich fern, 
Matteuccia pensylvanica, are quickly shed. Potassium hydroxide 
and the acetolysis mixture often over-expand pollen grains with 
increases from 2 to 17% and, in some instances, up to 43%. 
B. B. Christensen (1948) found considerable variation between 
measurements at high dry versus oil immersion and between 


10 AMERICAN FERN JOURNAL 


different observers of the same slides, as great as the expansion 
of the processing reagents. It is reported that pollen mounted 
in glycerin or glycerin jelly tends to swell. Freshly shed to dry 
spores have been mounted in Clarite (no longer available), 
Euparol, Permount, or Hoyers Mounting medium without dehy- 
dration. Unfortunately such mounts are rather opaque. The size 
and degree of wrinkling of the crests in Dryopteris spores de- 
pends upon the amount of moisture they absorb. 

The writer prepares temporary slides from herbarium speci- 
mens as follows: The frond is searched under a 25-power bin- 
ocular dissecting microscope and individual sori which contain 
spores are chosen. It should be mentioned that many herbarium 
specimens are deficient in mature spores and unopened sporangia 
are often filled with immature spores. Also, in the case of ferns 
like Dryopteris serrata, it is almost impossible to pick individual 
sori or sporangia because they form such a dense mass. Here 
scrapings must be used with caution. The sori or sporangia are 
picked with a moistened needle and placed on a slide in a drop of 
lactophenol or pure glycerin and then heated over an alcohol 
lamp until the liquid boils or steams in the case of glycerin. 
This drives out the air from the spores, expands them and makes 
them translucent. Lactophenol was used for many years but 
recently partial destruction of the perispore was observed in a 
few species. Pure glycerin was tried and found satisfactory. 
Slides so prepared last for years, if kept flat. A lactophenol 
slide dries out in a few weeks’ time. It has been the practice to 
remove the cover glass and add a drop of glycerin jelly to make 
more permanent slides. If the cover glass is sealed with paraffin, 
the slides will keep even longer. 

Erdtman’s acetolysis mixture has become one of the chief 
reagents used in spore and pollen studies. It is used where- 
ever there is ample spore material. Acetolysis can be accom- 
plished on the slide by gently heating a few sporangia in a drop 
of the fluid. If the slide is heated too long, the spores turn dark. 
The advantage of this process is that it removes the internal 


Spores IN FERN TAXONOMY am 


contents and renders the grain very translucent; it stains the 
perispore and exine a clear, brownish color. Fern spores which 
have been processed in potassium hydroxide or acetolysis and, 
to a lesser extent in lactophenol, show the exine distinct from 
the perispore. The perispore in some cases is very thin and 
adheres tightly to the exine, giving rise to the belief that spores 
lack a perispore. 

Some measure the exine only; others measure the perispore 
as well. The reason for measuring the exine is that the perispore 
is often greatly distended by the processing. 

There is no information on the relationship between spore size 
and the nutrition of the plant as there is for certain flowering 
plants. Thus there are many reasons for variation in spore meas- 
urements of the same species by different people and a small 
difference of two or three microns may be of little importance. 

No one has studied the differences in spores from apogamous 
ferns as compared to those in which meiosis has oceurred, Man- 
ton (1950) has shown that certain ferns have a polyploid chro- 
mosome series. She reported that the spores of the royal fern 
with a diploid chromosome compliment had a diameter of 80 
microns whereas the tetraploid spore had a diameter of 90 mi- 
erons. Although the chromosome compliment of ‘spores of hybrid 
ferns has been investigated, their spore sizes are seldom men- 
tioned, Manton reported abortive spores in the hybrid of Dry- 
opteris abbreviata X Filix-mas. Erdtman (1958) illustrated ab- 
normalities in Aspleniopsis decipiens and Dipteris chinensis. The 
first showed interesting modifications of the trilete aperture and 
the second had what can be interpreted as a monolete scar with 
a tiny fork near one end. Erdtman and Praglowski (1959) re- 
ported upon an anomolous condition in Pityrogramma hybrida 
var. maxima (= P. calomelanos X chrysophylla) in which the 
normal trilete aperture assumed a variety of shapes. Two of 
the trilete arms were irregularly forked; another spore had a 
monolete aperture; another a monoporate aperture, and the 
last was a grain without any dehiscence mark. Wagner and 


12 AMERICAN FERN JOURNAL 


Boydston (1958) illustrated the variation of spores of Asplenium 
ebenoides, A. rhizophyllum, and those of a hybrid between them. 
They showed several abnormally small grains with a perispore. 

The above remarks show that there is a lack of detailed infor- 
mation and some pitfalls that must be taken into consideration 
in the use of spores in fern taxonomy. The present role can be 
shown by the following selected citations. 

Christensen in Verdoorn (1938) outlined a classification of 
ferns with very little attention to spores. He mentioned spore 
types for certain orders, families, or subfamilies, along with the 
presence or absence of the perispore. 

Selling (1946) presented keys for the identification of Ha- 
waiian ferns by means of the spores. 

Copeland (1947) gave the spore type for nearly every genus 
and occasionally mentioned the surface condition as smooth or 
tuberculate. 

Makino (1949) included spore illustrations for about one-third 
f the Japanese ferns. Unfortunately, they are too small to be 
of diagnostic value. 

Lawalrée (1950) briefly described the spores for most of the 
Belgian species and genera. 

Reed (1953) was perhaps the first to include brief charac- 
terizations and photographs of non-cleared spores in a state fern 
book. Unfortunately, the internal contents obscured certain diag- 
nostic features and the quality of the illustration suffered by the 
method of reproduction. 

Madalski (1954) made ne spore illustrations for every 
fern in the atlas of the Polish flor 

Holttum (1954) omitted the spore characteristics for many 
genera of the Ferns of Malaya. However, he used spore charac- 
teristics as a means to separate Asplenium phyllitidis and sug- 
gested that the spores of Asplenium borneense indicated prob- 
able relationships with Aspleniwm scandens. Mme. Tardieu-Blot 
(1954) considered that spores had good generic and specific diag- 
nostic value and gave brief descriptions and good illustrations. 


Spores IN FERN TAXONOMY 13 


Harris (1955) published a manual of the New Zealand ferns 
based only upon spore characteristics. He presented keys to 
the families, genera, and for the species of many genera; for 
example, a key to the species of Polystichum. He reported that 
six species of Adiantum had similar spores with overlapping 
sizes and it was not possible to determine the species of Adiantum 
by their spores. He called attention to Lindsaca viridis with 
monolete spores in contrast to L. cuneata and L. linearis, which 
have trilete spores. He stated that the spore characteristics 
supported the removal of Lorsoma from the family Hymen- 
ophyllaceae. 

Holttum (1957) indicated Nakai’s proposal to place ferns 
with monolete spores in a distinct subfamily of the Gleicheni- 
aceae apart from those with trilete spores was an unnatural ar- 
rangement because Dicranopteris linearis has trilete spores but 
the closely allied species D. pubigera and D. currant have 
monolete spores 

The foregoing citations are sufficient to show that fern spores 
are becoming increasingly important in fern taxonomy. They 
have been used successfully to distinguish species in some genera, 
to differentiate genera and to characterize families. As studies 
progress and procedures are stabilized, spore data will be used 
also to distinguish polypoid races or strains and to indicate 
hybridity. Spores have as much diagnostic value as scales and 
hairs; the data should be considered as supplemental informa- 
tion, not the sole criterion. It is admitted that there are groups 
where spore data cannot be used to distinguish species. 


BIBLIOGRAPHIC REFERENCES 


CAMPBELL, — H., The spuced ne sgpenige of Mosses and 
Ferns. ed. 2, vii, 1-657. tae a n Co., New York. 

o inantonts RB. B., Measurement as a means of jaeniatying fossil pollen. 
Danmarks psa et carpet Vol. 4 (2): 5-22. 19 

CHRISTENSEN, C., in Verdoorn, Manual of pease: 529-550. 1938. 
Martinus Nijhoff, The esos The Netherlands. 

CopeLANp, E. B., Genera Filicum, Ann. Cr. yptogamici et Phytopathologici. 
5: xvi, = 247. 1947. Chronica Solahies, Waltham, Mass, 


14 AMERICAN FERN JOURNAL 


ErpTMAN, G., Pollen i. Medi Morphology/Plant Taxonomy Gymnospermae, 
eestor Bryop » (An Ca ne to Palynology II). 1-151. 
Almaqvist & Bee Stockh 
ican eg . and J. R. PRAGLOWSK ot x notes on pollen eee Wee 
pollen morphological techniques. Pine an hee er 112 .(2).2-L75-1 
59 


1959. 
Harris, W. F., A manual on the spores of New Zealand Pteridophyta. Bull. 
116: 1-186. fon N. Z. Dept. Sci. & Ind. Res 
Hoirtum, R. , A Revised Flora of Mileve Vol. 2, Ferns of Malaya. 
Si 


. O. Singap: 

—, Morph eloay: Grow: habit, and Clasitaton in the Family Gleicheni- 
aceae. Phytomorphology 7 (2): 168-18 (a; 

Kwox, Euizasetu, M., The spores of teeta Phylloglossum, Selagi- 
nella, and Isoétes and their value in the study of microfossils of Paleozoic 

ge. Trans. and Proc. Bot. Soc. Edinburgh. 35 (3): 209-357. 1950. 

LAWALREE, ANDRE, Flore Générale de Belgique, Ptéridophytes. iv, 1-195. 
1950. Ministér de’agriculture, Jardin Botanique de 1’Etat, Bruxelles. 

Maparsxl, Jo6ser, Atlas Flory Pols Re IT Ziem nee nati LCL), 87 A1G0., 
(2) 33 tab. 1954. Polska Akademia Nauk, Warszaw: 

Makino, Tomiraro, An Illustrated Tora of Japan. Le Ed. 1-1070. 1949, 
indexes. 

Manton, I., ee of Eeilay and Evolution in the i asl xi; 

6. 195 


1-31 0. Cambridge University Press, Cambridge, Engla 
McVaveu, Rogers, Studies on the Spores of some Wocthuestara Ferns. 
ern. Journ CG 73-85. 1935. 


mer, F : 
Reep, Ciype F., The Ferns and Fern-allies of Maryland and Delaware in- 
ea abies of Columbia, xviii, 1-286. 1953. Reed Herbarium, Balti- 


mo 

pecs Gok H., Studies in the Recent and Fossil Species of Schizaea, 

with particular vétereare to their spore characters. Med. Gotebergs ab t. 
—112. 1944, 


—, Studies in Hawaiian Pollen Statistics, Pt. 1. The spores of the 
Hawaiian Pteridophytes. B. P. Bishop Rawat Spec. Pub. 37: 1-8 
1946. 


TARDIEU-BLot, MME., Cryptogamia. Pteridophytes. 1-107. 1954. Par 
BG 


WaGner, W. H., ia i oypston, A New Hybrid Spleenwort pins 
oe ceva! at Fernwood and its cea ies to a Peculiar Plant 
from West Vir, a. Amer. Fern. Jour 8 (4): 146-159. (058 


Louisiana Le UNIVERSITY, iin Rovce 3, Lovistana. 


ENDEMIC PrEeRIDOPHYTES OF CALIFORNIA 15 


The Endemic Pteridophytes of the California 
Floral Province 


JouHn THomas HowELL 


Several years ago I advanced the idea that there is a natural 
California flora and proceeded to outline a California Floral 
Province, using as distinguishing criteria ‘the high endemism 
which is found in both genera and species and the distinctive 
associations in which these genera and species occur’’ (1957, pp. 
133-138). Geographically, the province would extend from the 
vicinity of Coos Bay on the coast of Oregon southward along 
the California coast to northern Baja California south of En- 
senada and to the islands off the coast, while the eastern bound- 
ary would be marked by the mountain ranges that extend north- 
ward from the Sierra San Pedro Martir to the vicinity of Mount 
Shasta in California and thence northwestward to the Oregon 
coast, separating the Siskiyou-Klamath montane area from the 
Caseadian. 

Although this proposal was based entirely on seed plants, I 
took occasion several months later to extend the idea to pteri- 
dophytes and outlined the results in remarks before the Ameri- 
can Fern Society, at its meeting at Stanford University in Au- 
gust, 1957. At that time, I indicated that endemism among spe- 
cies of pteridophytes in the California Floral Province amounted 
to about 27%, using as a basis Maxon’s and Pfeiffer’s treat- 
ments in Abrams’ Illustrated Flora of the Pacific States (1923). 
The impressiveness of this figure was shown by a comparison 
with the percentage of endemism indicated by Morton’s account 
of the pteridophytes in Gleason ’s New Illustrated Flora (1952), 
in which endemic species constitute only about 7% of the entire 
fern population. Endemism as a mark of a California flora is 
thus not restricted to seed plants. 

The present account of pteridophytes endemic to the Califor- 
nia Floral Province is based chiefly on the above-mentioned treat- 
ments by Maxon and Pfeiffer (1923), as they have been accepted 
or somewhat modified by Munz in A California Flora (1959). I 


16 AMERICAN FERN JOURNAL 


have altered Munz’ total slightly (at points apparent in the 
following notes) and have of course applied the concept of 
endemism to a provincial, not a political, area. As accepted here 
the pteridophytes number 98 entities (86 species and 12 sub- 
species and upladamel of which 63 are ‘‘ferns’’ and 35 are 
_“‘fern-allies.’’ Of this total number, 27 entities (21 ferns and 6 
fern-allies) are endemic, which by percentage amounts to 27.55% 
(ferns, 33.33% ; fern-allies, 17.14% ).1 
he following notes give the name, distribution, habitat, ete. 
of the endemic pteridophytes in the California Floral Province 
as I now understand them. The phytogeographic affinity of two- 
thirds of the entities is strongly austral; only eight or nine show 
closer relations to plants of northern floras. 
SELAGINELLACEAE 
SELAGINELLA ASPRELLA Maxon. On exposed dry slopes and 
in rock crevices, most common in the mountains of southern 
California from the San Gabriel Mountains, Los Angeles County, 
to the Laguna Mountains, San Diego County, at elevations of 
5000-8000 feet. An outlying station reported by R. M. Tryon 
(1955, p. 76 and map 51, p. 73) is in the southern Sierra Bt re 
in the Kern River Canyon, Tulare Coun ty, where the plan 
grows at an elevation as low as 2800 feet Pavel 5031, a. 
SELAGINELLA BicELovit Underwood. In gravelly or rocky soil 
or in shallow soil overlying rock surfaces; widespread and fre- 
quently common at elevations from sea level up to 6000 feet, 
from northern Baja California northward through southern 
California to Tulare County in the southern Sierra Nevada and 
to Sonoma County in the central Coast Ranges. R. M. Tryon 
(1955, map 8, p. 21), in mapping the distribution of the Bigelow 
1Tt is interesting to compare these figures with the percentage of 
endemism in several other rll ties in the California Floral Province. Thus 
endemism in Carex amounts 7.66%, in Eriogonum to 52.08%, in Phacelia 
be gel a and in the “arweed Tribe’’ of the Compositae, the Madiinae, 


aie eos in pteridophytes in California in a strictly political sense is 
considerably reduced: Only 15 entities, or 15.30%, are restricted to the 


ENpEMIC PrerRIDOPHYTES OF CALIFORNIA 17 


mossfern, indicates two rather widely separated stations on the 
east side of Baja California that lie beyond the boundary of the 
California Floral Province as I have outlined it. It is to be ex- 
pected that this species will be found to be rather common in 
central Baja California as the mountains there are more inten- 
sively explored. In 1956, George Lindsay and I found depauper- 
ate plants growing in rock crevices in the desert mountains 32.5 
miles southeast of Rosario on the road to El Marmol (Howell 
31012). 

SELAGINELLA CINERASCENS A. A. Eaton. S. bryoides Under- 
wood. On packed clayey soil of open or brushy slopes in the 
coastal fog belt near sea-level, from San Diego County, Cali- 
fornia, southward to the vicinity of Ensenada, Baja California. 
Although this species is not uncommon within its highly re- 
stricted range, it is not frequently collected, perhaps because its 
delicate moss-like filagree over clay is overlooked by collectors 
of vascular plants. 

SeLAGINELLA Hansentt Hieron. S. Bolanderi Hieron. S. 
rupestris (L.) Spring var. Hansenii (Hieron.) Jepson and 
var. Bolanderi (Hieron.) Jepson. Crevices and shelves of 
igneous and metamorphic rocks, 200-6000 feet elevation on 
west slope of the Sierra Nevada from Tulare County north to 
Butte County, with outlying stations on the Marysville Buttes 
(Heller 11802) and on Mount Shasta. I do not know of the 
Hansen moss-fern in the Coast Ranges, but R. M. Tryon (1955, 
map 30, p. 51) indicates two stations for the plant in the Inner 
Coast Range north and south of San Francisco Bay,? and Munz 
(1959, p. 24) eredits it to the Santa Lucia Mountains. 

Exciupep species. Although Selaginella leucobryoides Maxon 
is found only in California, it is an endemic restricted to the 
higher mountains of the Mohave Desert that lie quite beyond the 
limits of the California Floral Province. 


2 The localities north of San Francisco Bay are Knights Valley, Napa 
County, W. R. Dudley, Feb. 23, 1895 (US, and Stanford 50231), and 
‘<Ganyon northwest of Napa, July 30, 1913, Suksdorf 753 (US); both 
collections seem to be S. Wallacei Hieron., which is common in the region.— 
C. V. Morton. 


18 AMERICAN FERN JOURNAL 


TSOETACEAE 

Isorres Orcurtu A. A. Eaton. J. Nuttalla A. Braun var. 
Orcuttii (A. A. Eaton) Clute. Amphibious plants maturing on 
the drying beds of former rain pools (‘‘vernal pools’’) ; coastal 
mesas in northern Baja California and in California from San 
Diego County north through the South Coast Ranges and south- 
ern Sierra Nevada foothills to the Sacramento Valley in Sacra- 
mento County, 100-2000 ft. elevation. 


OPHIOGLOSSACEAE 
OPHIOGLOSSUM CALIFORNICUM Prantl. O. lusitanicum L. subsp. 
californicum (Prantl) Clausen. Moist flats and slopes at widely 
separated stations at elevations below 1000 ft. from central Cali- 
ornia to northern Baja California. The plant is rare or rarely 
detected, being known only from a few collections from Ione, 
Monterey, San Diego, and Ensenada. Published records from 
central Mexico are dubious. Munz (1959, p. 29) gives the 
habitat of the plant as ‘‘vernal pools.’’ A collection I made in 
Balboa Park in San Diego in March, 1932 (No. 8232) came from 
a ‘‘moist grassy slope, growing up through a mat of Selaginella 
bryoides’’; and Miss Eastwood told of making a collection at 
Tone (her No. 12413) on a grassy slope near a blue oak (Quercus 
Douglasn). 
POLYPODIACEAE 
ApIANTUM JorDANII C. Mueller. A. emarginatum D. C. Eaton. 
On shaded rocks or in roeky soil beneath trees or shrubs at ele- 
vations up to 4000 feet; northern Baja California north to the 
California Channel Islands and through the California Coast 
Ranges and Sierra Nevada to southwestern Oregon (lower Rogue 
River Canyon, acc. to Peck, 1941, p. 51). In the North Coast 
Ranges of California a hybrid between A. Jordanii and A. 
pedatum Li. has been described: A. & Tracyi C. C. Hall ex 
Wagener (1956). 
SPLENIUM VESPERTINUM Maxon. A. Trichomanes L. var. 
vespertinum (Maxon) Jepson. Moist crevices of shaded rocks 
below 3000 feet elevation; San Gabriel Mountains, Los Angeles 


ENDEMIC PTERIDOPHYTES OF CALIFORNIA 19 


County, south to the mountains of San Diego County in Cali- 
fornia and (ace. Maxon, 1923, p. 18) to Baja California.* 

CHEILANTHES CALIFORNICA (Hooker) Mett. Hypolepis cali- 
fornica Hooker. C. amoena A. A. Eaton. Aspidotis californica 
Nuttall ex Copeland. Dry slopes in shallow or rockey soil below 
4000 feet elevation, from southern California north through the 
Coast Ranges to Humboldt County and through the Sierra 
Nevada to Butte County, south to northern Baja California. 

In treating the California lace-fern as a Cheilanthes, 1 am ac- 
cepting what I believe to be the correct conservative interpreta- 
tion of Maxon and of Weatherby, and more recently of Wagner 
and Gilbert (1957, pp. 741-743), rather than that of Copeland 
(1947, p. 68) or Munz (1959, p. 35). 

CHEILANTHES CARLOTTA-HALLIAE Wagner & Gilbert. Shaded 
or sunny places around rocks in areas of serpentine ; known only 
in the California Coast Ranges from Marin, San Benito, and San 
Luis Obispo counties. Mrs. Hall’s lip-fern is intermediate be- 
tween C. californica (Hooker) Mett. and ©. siliquosa Maxon 
(Pellaea densa Hooker ; Onychium densum Brack.), and perhaps 
it should have been named as indubitably a hybrid. Concerning 
this fern as I know it on Mount Tamalpais, Marin County, I 
recently remarked: “‘. . in the vicinity of Bootjack (which is 
the type locality), there is to be found such a variety of plants 
intermediate between C. californica and C. siliquosa that it is 
not always possible to tell where one leaves off and the other 
begins, and it is well nigh impossible to delimit the proposed 
species [7.¢., C. Carlotta-halliae| from the numerous variations’’ 
(Howell, 1959, p. 53, 54). It would be enlightening to know if 
this lip-fern originated independently through hybridization in 
the three widely separated localities where it has been reported, 
or if those occurrences represent dispersal from a single cross. 
The field occurrence of the plant on Mount Tamalpais strongly 
suggests hybridization at that place. 


3 San Rafael, Apr. 13, 1882, M. H. Jones 3749. [C.V.M.] 


20 AMERICAN FERN JOURNAL 


CHEILANTHES CLEVELANDII D. C. Eaton. Around rocks in dry 
brushy places below 5000 feet elevation; northern Baja Cali- 
fornia north in California to the northwestern borders of the 
Colorado Desert in Riverside County and northwest to Santa 
Barbara, Santa Barbara County, and to Santa Cruz and Santa 
Rosa islands. 

CHEILANTHES CooperaE D. C. Eaton. In moist shaded rock- 
crevices in summer-dry hills at elevations below 2000 feet, fre- 
quently on limestone (marble) or other metamorphic rocks, with 
an interrupted distribution from Shasta County, California, 


County and in southern California from Santa Barbara County 
southeastward to southwestern San Bernardino County. 
CHEILANTHES FIBRILLOSA (Davenport) Davenport ex Under- 
wood. C. lanuginosa Nuttall var. Brae Davenport. Cali- 
fornia’s rarest endemic fern, known only from the original col- 
lection made ‘‘by the brothers Parish well down in one of the 
passes that open out on the south side of the San Jacinto Moun- 
tains, in June 1882.’’* Actually the type locality is on the west 
side of the mountains and concerning it Munz and Johnston 
(1922, p. 113) wrote as follows: Cheilanthes fibrillosa is ‘known 
only from the type collection by Parish! made ... in the San 
Jacinto Canyon at Oak Cliff at the point where the road to 
Strawberry Valley leaves the canyon bed.5 The plant was found 
among rocks on a gravelly bench. An unsuccessful search for 
4Bull. Torrey Club 12: 21. 1885. 


I know of only three other ferns whose yor pig records rest on single 
collections. Two are holaretic species of Aspleniwm of wide distribution. 
son in California i 


a 
granitic rock between Columbine Lake and Sawtooth Pass, at an elevation 
of about 11,000 fy Howell 17803, saree: on Aug. 7, 1942 (ef. Howell, 
Base Camp Botany, 1942, p. 3; Ewan, Amer. Fern Journ. 33: 29. 1943). 
The fourth fern piles in California Prot a S aiele collection is the wide- 

: ot ; 


.) & 
Holeomb Valley, San Bernardino Mowntainej in 1882 (cf. Munz, 1959, p 


ENDEMIC PTERIDOPHYTES OF CALIFORNIA 21 


this fern was made at the type locality by Parish and Johnston 
in 1918; a similar one by the authors occurred in 1922 when the 
type locality and the water-shed above were examined.’’ 

CHEILANTHES INTERTEXTA (Maxon) Maxon. €. Covillei Maxon 
var. intertexta Maxon. Shaded or partially shaded rock crevices 
and rocky slopes in the Coast Ranges and Sierra Nevada, and, 
according to M. E. Peck (1941, p. 51), to the ‘‘Siskiyou Moun- 
tains of Oregon,’’ from near sea-level to elevations of 9000 ft. 
Two stations just beyond the limits of the California Floral 
Province are: Dixey Mountains, Lassen County, California, and 
near Virginia City, Storey County, Nevada. Both on Mount 
Tamalpais and Mount Diablo in the central California Coast 
Ranges I have collected lip-ferns that I have determined as C. 
Covillei Maxon because of the scaly fronds and on both peaks 
this fern grew at a distance of less than a mile from C. inter- 
texta (ef. Howell, 1959, p. 54, as to Mount Tamalpais plant). 
In both places, the scales of C. Covillei are somewhat more 
ciliate-fimbriate than are the scales of more typical C. Covillei 
from southern California coast or desert, but on both Tamalpais 
and Diablo I had no doubt but that I was dealing with two dif- 
ferent lip-ferns. Although the two species grew rather close 
together in areas where habitats are uniformly alike, each was 
found restricted to its own particular rock-outcrops and no in- 
tergrades were observed on rocks situated between the two. 

CHEILANTHES PartsHm Davenport. Known only from the re- 
gion of the type locality on the rocky desert slopes of Andreas 
Canyon, on the east side of the San Jacinto Mountains near Palm 
Springs, Riverside County, California. Although included here 
as an endemic of the California Floral Province, the Parish lip- 
fern is found in the zone between the California flora and the 
Sonoran Desert flora and it could be considered a borderline 
endemic of the latter flora. However, from both morphological 
and phytogeographie relations to other California lip-ferns it 
seems preferable to regard the Parish lip-fern as Californian 
rather than Sonoran. 


22 AMERICAN FERN JOURNAL 


NorTHOLAENA CALIFORNICA D. C. Eaton subsp. NIGRESCENS 
Ewan. Alewritopteris cretacea (Liebmann) Fournier subsp. 
nigrescens (Ewan unz. Rocky places in the chaparral, south 
side of the San Gabriel Mountains, Los Angeles County, Cali- 
fornia. This subspecies is not very distinct from variants in the 
typical subspecies and was placed in synonymy under N. cali- 
fornica by R. M. Tryon in his revision of the genus (1956, p. 73). 
Munz (1959, p. 35), after reducing N. californica to synonymy 
under Aleuritopteris cretacea, retains for his inclusive species 
the relatively restricted range of N. californica (southern Cali- 
fornia to Arizona and Baja California) rather than the truly 
extended range it should have had [at least to Puebla, Mexico, 
where the type of N. cretacea was collected, if not to Chile 
whither it ranges as N. sulphurea (Cav.) J. Smith (ace. R. M. 
Tryon, 1956, p. 71, map 40)|. Notholaena candida Hooker var. 
accessita Jepson (1923, p. 27) [N. californica f. accessita (Jep- 
son) Ewan], described as a local California endemic of eastern 
San Diego County, is referred to the synonymy of N. californica 
D. C.-Eaton by R. M. Tryon (1956, p. 73), a course that is fol- 
lowed here. 

NorHOLAENA Newserry! D. C. Eaton. Cheilanthes Newberryt 
(D. C. Eaton) Domin. Dry rocky places at lower elevations from 
Ventura County to San Bernardino and San Diego counties and 
San Clemente Island in California and to northern Baja Cali- 
fornia and Guadalupe Island in Baja California. 

PELLAEA ANDROMEDIFOLIA (Kaulfuss) Fée. Pteris andro- 
medifolia Kaulfuss. Pellaea rafaelensis Moxley. (For extended 
synonymy see A. F. Tryon, 1957, p. 179.) Seasonally dry slopes, 
more or less shaded by brush or trees, on rocky ledges or in 
rocky soil, at elevations up to 3500 (or 4000) feet; southern 
California north through the Coast Ranges to Mendocino and 
Tehama counties and through the Sierra Nevada foothills to 
Butte County, west to the Channel Islands, and south to north- 
ern Baja California and Cedros Island (A. F. Tryon, 1957, map 
13). A Thomas Jefferson Howell collection from Roseberg, Ore- 


ENpremic PTERIDOPHYTES OF CALIFORNIA 23 


gon, is regarded as doubtfully authentic by A. F. Tryon (1957, 
p. 182). Pellaea andromedifolia (Kaulfuss) Fée var. pubescens 
D. C. Eaton is a name applied to pubescent plants from the 
southern part of the range of the species. 

PELLAEA BRACHYPTERA (Moore) Baker. Platyloma brachyptera 
Moore. Pellaea Ornithopus Hooker var. brachyptera (Moore) 
D. C. Eaton. Allosorus brachypterus (Moore) O. Kuntze. Dry 
rocky slopes, generally in areas of metamorphic rocks, 2500-8000 
feet elevation; southwestern Oregon south in California to Lake 
County in the Coast Ranges and to Placer County in the Sierra 
Nevada. 

EXCLUDED SPECIES AND VARIETY. Although Pellaca mucronata 
(D. C. Eaton) D. C. Baton including its variety, var. californica 
(Lemmon) Munz & Johnston, is called a ‘“California species”’ 
by A. F. Tryon (1957, p. 157), both var. mucronata and var. 
californica range too far and too commonly into the desert moun- 
tains of southeastern California for either to be regarded here 
as endemic to the California flora in the provincial sense of this 
paper. Var. californica is entered twice by Munz: Once as a 
synonym of P. compacta (Davenport) Maxon (1959, p. 35) and 
again (p. 36) as an accepted variety. 

PITyYROGRAMMA TRIANGULARIS (Kaulfuss) Maxon var. PALLIDA 
Weatherby. In California on dry rocky banks or in rocky soil, 
more or less shaded, Sierran foothills below 4000 feet, from Butte 
County to Kern County, and according to Munz (1959, p. 38) to 
Santa Clara County in the Coast Ranges. 

PrryROGRAMMA TRIANGULARIS (Kaulfuss) Maxon var. VISCOSA 
(Nuttall ex D. C. Eaton) Weatherby. Gymnogramme triangu- 
laris Kaulfuss var. viscosa Nuttall ex D. C. Eaton. Ceropteris 
viscosa (Nuttall) Underwood. Pityrogramma viscosa (Nuttall) 


Riverside, Orange, and San Diego counties and on the Channel 
Islands, south to northern Baja California. 
PoLYPODIUM CALIFORNICUM Kaulfuss. P. intermedium Hooker 


24 AMERICAN FrerRN JourNAL 


Island) north in California to the Channel Islands and through 
southern California to Humboldt County in the Coast Ranges 
and to Butte County in the Sierra Nevada. 


of P. californicum associated with the evergreen P. Scouleri 
Hooker & Greville. 


probably of hybrid origin ( Howell, 1949, p. 51). 

PotysticHum Dupieyr Maxon. P. aculeatum (Swartz) Roth 
var. Dudleyi (Maxon) Jepson. In deep shade on moist rocky 
slopes near the coast; central California Coast Ranges from San 
Luis Obispo County (ace. to Munz, 1959, p. 41) to Marin 
County. 

PoLYstICcHUM MUNITUM (Kaulfuss) Pres] subsp. CURTUM 


PoLYSTICHUM MUNITUM (Kaulfuss) Presl subsp. NupATumM 
(D. C. Eaton) Ewan. Aspidium munitum Kaulfuss var. nuda- 
tum D. C. Eaton. P. munitum (Kaulfuss) Presl var. nudatum 


ENDEMIC PTERIDOPHYTES OF CALIFORNIA 25 


(D. C. Eaton) Gilbert. Rocky slopes and cliffs in the California 
mountains at elevations of 4000-7000 feet; Sierra Nevada from 
Tulare County to Plumas County. This variant and the preced- 
ing are not too readily distinguished from the wide-ranging P. 
munitum var. imbricans (D. C. Eaton) Maxon and may repre- 
sent forms in a variable species rather than geographically dis- 
tinet subspecies. 
BIBLIOGRAPHIC REFERENCES 

CopELAND, E. B. 1947. on Filicum, pp. I-XVI, 1-247. 
sedi: J.T. 1949. Marin Flora, Pt teridophyta, pp. 49-55. 

9 The California iota and its Province. Leafl. West. Bot. 8: 133- 


—. 1959. After Ten Years: Names and Notes for the Marin Flora. Leafl. 
t. 9: 53-66. 


JEPson, W. L. belt A Manual of the Flowering Plants of California, 
yee pp. 44, 
Maxon, W. R. 1 3. Pteridophyta (except Isoétaceae) in L. R. Abrams, 


92 id 
ae Titustrated Flora of the Pacific States, pp. 1-35, 38-50. 
Morton, C. V. 1952. Pteridophyta, in H. A. Gleason, New Illustrated Flora 
of the Northeastern geste States and Adjacent Canada, pp. 1- 
Munz, P. A., and I. M. Jonnston. 1922, 1923. The Distribution of South- 
ern Catton Bigeiorhy tes. Amer. Fern Journ. 12: 69-77, 101-122; 
Loe 


MuNz, 8 A., and D. D, Keck. 1959. A California Flora, Lepidophyta, 
Calonlophyta: Pterophyta, pp. 21— 


Peck, M. E. 1941. A Manual of the Higher Plants of Oregon, Pteridophyta, 
pp. 43-57. 
Preirrer, N. 1923. Isoétaceae in L. R. Abrams, An Illustrated Flora 


923 
of the Pacific ‘Binted: pp. 35-38. 
Tryon, A. F. 1957. A apeeg of the Fern Genus Pellaea, Section Pellaea. 
Mo. Bot. Gard. Ann. 44: 
Tryon, R. M. 1955. paces oan and Its Allies. Mo. Bot. Gard. 


WaGn 1956. ‘ Natural Hybrid, X Adiantum Tracyi C. C. Hall. 


Maareie is: "195-26 
Waener, W. H., an ae | F. Girpert. 1957. An Unusual New Cheilanthoid 
: 738-743. 


Fern Fea California. Amer. Journ. Bot. 44 
CALIFORNIA ACADEMY OF SCIENCES, ‘San FRrANcIsco, CALI- 
FORNIA. 


26 AMERICAN FERN JOURNAL 


First Illustrations of Ferns from Peru and Chile 
JosEPH KwAN 


Linnaeus recognized twelve genera of ferns in his Species 
Plantarum of 1753. The largest genus was Polypodium with 
fifty-eight species of which over sixty percent are American. In- 
cluded in his genus ‘‘Polypodium’’ were species now known to 
be species of Polystichum, Dryopteris, Cystopteris, Cyathea, and 
other genera. Some eight authors were cited by Linnaeus in pro- 
viding the bibliographies for the New World species. He cited 
his own works first, as elsewhere in his Species Plantarum, fol- 
lowed by references to the works of Plumier, Petiver, Gronovius, 
Plukenet, Morison, Hans Sloane, and John Ray. Though Lin- 
naeus lists ‘‘Fewillaei peruviana’’ as one of his sources in the 
introduction to his classic, he does not refer to Feuillée in the 
pages devoted to ‘‘Cryptogamia Filices,’’ evidently overlooking 
the fern descriptions contained in his work when writing the 
account of the several American species. This is more unusual 
since Feuillée illustrated two species based on Chilean observa- 
tions not previously pictured. Father Luis Feuillée (1660-1732), 
explorer, astronomer, and botanist, was the author of the first 
herbal for Peru and Chile in which these fern drawings ap- 
peared. 

Luis Econches Feuillée was born at Mane, near Forealquier, in 
Provence, in the year 1660, of humble parents. In his twentieth 
year Luis joined the Order of the Minimi. At this time Europe 
was awakening to a lively interest in the sciences—the Royal 
Society was founded in London when Luis was two years old, 
and the Academie des Sciences was chartered by Louis XIV in 
1666. Feuillée’s taste for the sciences attracted attention in 
official circles and in 1699 he was sent to the Levant as an aide to 
Jacques Cassini on a hydrographic mission to determine the 
character of ports, to map offshore currents, and so forth. His 
mission to the Antilles in 1703 was an extension of this assign- 
ment; on this expedition he went ashore at Martinique, Cara- 
cas, and elsewhere, and returned to Brest in 1706. On Decem- 


AMERICAN FERN JOURNAL VotumE 50, PLATE 1 


JOURNAL 
DES OBSERVATIONS 
PHYSIQUES: 
MATHEMATIQUES 


ET BOTANIQUES. 


Faites par l'ordre du Roy fur les Cotes Orientales 
de lAmerique Meridionale, & dans Ics Indes 
Occidentales, depuis l'année 1707. jufques cn 1712. 

Par leR.P. LOUIS FEUI LLEE, Religicax Minime , 

Mathemasicien , Bocanifie de Ss MAJEST Ey 
© Correfpondant de Académie Royale des Sciencei» 


TOME SECOND. 


A PARIS, AUE S. JACQUES, 
Chez PIERRE GIFFA%T, Livave Co ow du Roy, 
& de l'Academic Roya‘e de Peinture & de Sculpture , 
a Vimage Saace b beri. 


“ pce. XY. 
AVEC APPROBATIONS ET PR IVILCGE DU KOT. 
Tipe Page oF FEUILLSE’S Rake JouRNAL (1714) OF His TRAVELS ON THE 
Coasts OF PERU AND CHILE 


AMERICAN FERN JOURNAL VoLUME 50, PLATE 2 


Planche XL. 


: \ 
LEQ 


Polypodiun radice 
Jguamosa . anlgo 
Pilllabilenm 


» bafore Sony 


FEUILLEE’S ORIGINAL ILLUSTRATION OF POLYPODIUM FEUILLEI BERT. OF 
CHILE 


PL.FemBee Bor. Reg. del. 


ILLUSTRATIONS OF FERNS 29 


ber 14, 1707, Feuillée again set sail for America, but was forced 
to return to Teneriffe by contrary winds in May, 1708; he suc- 
ceeded in rounding the Horn in January and he reached Peru in 
April, 1709. For the next nine months he mapped the port of 
Callao and other smaller roadsteads, directed the artist Pierre 
Giffart! in the sketching of the city’s skyline, described several 
of the animals he encountered, the distinctive balsas of Peru, 
and computed astronomical data before proceeding to Chile on 
the return voyage to France. 

It was upon Feuillée’s visit to Concepcion in January, 1711, 
that he observed, described, and illustrated the fern named Poly- 
podium feuille: in 1829 by Bertero. Feuillée had identified the 
fern he found ‘‘au Nord de la Ville de Pinco’’ as ‘‘Polypodium 
radice squamosa. vulgo Pillabileum,’’ following the current prac- 
tice of applying polynomials which were indeed short to long 
descriptive names for the plants in question. Elsewhere in his 
diary Feuillée more closely locates the habitat as the mountains 
to the east of the little valley called Pinco (1714: 545). Looser 
(1951: 41) corrected Pinco to ‘‘Penco’’ and located it on the 
Bay of Taleahuano. As Looser remarked this drawing (Pl. 2), 
which was Plate 40 in Feuillée’s work of 1714, is an excellent 
illustration of a sterile frond. Looser (1948: 85) says that Poly- 
podium feuillet is ‘‘a very common epiphyte; also on walls’’ in 
the region of Corral, Niebla, and in the vicinity, of Valdivia, 
where it ranges from 400 m. above sea level down to the shores 
of the Pacific. Kunkel (1959) recently published a paper on 
the nine infraspecifie taxa of this fern. Feuillée’s plate stands 
about intermediate between the var. feuillei, that is, the typical 
variety as interpreted by Kunkel, and var. ibanezii Loos. (cf. 
Kunkel’s plate 42), but it will be noted that Looser’s illustration 
of his var. ibanezii (fig. 9) shows a terminal lobe of the frond 
which is broad and irregularly toothed, being rather different 
from Kunkel’s interpretation of that variety. 


1 Giffart was also engraver for Plumier’s great work Nova Plantarum 
Americanarum Genera (1703). 


AMERICAN FERN JOURNAL VoutuME 50, PLATE 3 


[xV. 


Bibs aunor non ramos ponmelis wtentates . page ‘ | 


FrEvUILLEE’S ORIGINAL ILLUSTRATION OF NOTHOLAENA MOLLIS KUNZE OF 
CHILE 


ILLUSTRATIONS OF FERNS 31 


A second fern more briefly described by Feuillée and less accu- 
rately illustrated as well was Notholaena mollis Kunze. This 
second Chilean fern to be illustrated in botanical literature has 
a more obscure origin, for Feuillée fails to record where he ob- 
served this species. This fern is known from Coquimbo? and may 
have been observed there when he visited the port in April, 1710. 
The third volume of Feuillée’s work, published in 1725, was 
illustrated by an unknown artist, Giffart having died in 1723. 
The xerophytic character of this Notholaena (cf. Pl. 3) doubt- 
less engaged Feuillée’s attention for he likely noticed the ‘‘resur- 
rection habit’’ of alternately drying and reviving with wet and 
rainless periods. This fern was evidently not distinctive enough 
to be involved in early species descriptions as was Polypodium 
fewillei. 

Feuillée’s important contribution to the early knowledge of 
flowering plants has been sketched elsewhere by me (1959). 
Among the more notable of these were the genus Fuchsia and 
what has come to be called the ‘‘California Pepper Tree’’ 
(Schinus molle). Feuillée’s later years included his election to 
the French Academy, his investigation of the island of Hierro in 
the Canaries in 1724, and the publication of the third volume 
of his Journal, already mentioned, in 1725. The only English 
translation of Feuillée’s works was a much abridged treatise 
edited by Petiver entitled ‘‘The South Sea Herbal of Feuillée’s 
Medicinal Plants,’’ published in 1715 and including five plates. 
This seems to have been designed for sea captains. Father 
Feuillée died in Marseilles on April 18, 1732, the year of George 
Washington’s birth. 

BIBLIOGRAPHIC REFERENCES 
Ewan, J. Feuillée’s Two Herbals of 1714 and 1725. Herbarist (Herb Soe. of 


Amer.) 25: 60-64. 1959. : 

Feuillée, L. Journal des Observations Physiques Mathematiques et Bota- 
niques. 1714. : 
2W. T. Stearn remarks on the unusual opportunity among pre-Linnaean 


authors for locating the source of Feuillée ’s plants. Cf. his valuable Intro- 
duction to Linnaeus, Sp. Pl. (Ray Soe. reprint, London, 1957), p. 145, 


32 AMERICAN FERN JOURNAL 


Kunkel, G. Uber oe pot Bertero und neue varietiten dieser Art. 
Nova Hedwigia 1(2): 202. 2 pls. 1959. 
Looser, G. Ferns of goat apa This JOURNAL 38: 33-44, 71-87. 1948. 
—, El Genero Polypodium L. y sus representantes chilenos. Rey. Uni- 
Ms ag [Santiago] 36(1): 13-82. 1951 
—, Helechos de Chile Central. Moliniana 1: ie 95. 1955. 
ACKNOWLEDGMENT 
It is a pleasure to acknowledge the assistance of Dr. R. M. Tryon in the 
identification of these ferns. 
TULANE University, New ORLEANS, LOUISIANA. 


Evergreen Grapeferns and the Meanings of Infraspecific 
Categories as Used in North American Pteridophytes 
Warren H. Wacner, Jr.! 

In late years, work on the North American evergreen grape- 
ferns (Botrychium, subg. Sceptridium) has led me to a taxo- 
nomic interpretation that deviates quite strongly from that of 
my predecessors. The plant I designate as Botrychiwm oneidense 
has been construed in the past mostly as a variety or form of 
B. dissectum or of B. multifidum or as their natural hybrid. 
The one now called B. ternatum (or its American representa- 
tive) has been interpreted, if at all, as a form of B. multifidwm. 
The occasion for this paper is to state briefly the philosophy on 
which these interpretations are founded, and, at the same time, 
to call attention to some of the usages of infraspecifie categories 
to be found among other North American pteridophytes. 

Estimates of the total of pteridophytic species that grow in 
North America north of Mexico range from roughly 340 to 380. 
We may never decide on the exact number of species, however, 
but not because of lack of exploration and research. In fact, I 
suspect that the next half-century will yield a tremendous ex- 
pansion of our knowledge. The problem in estimating the num- 
ber involves the taxonomic definitions themselves. There are no 

1 ree pigeon by the Horace H. Rackham School, University of 


Michigan. I a indebted to Dale J. Hagenah for introduci on dig e to the 
i ol and to award G. Vous for advice on historical matter: 


EVERGREEN GRAPEFERNS 33 


absolute, hard and fast applications of taxonomic categories in 
expressing plant relationships. But within certain boundaries 
they are meaningful. 

The subject of infraspecific categories has been much dis- 
cussed, and I have selected a few references (Emerson, 1955; 
Fuchs, 1954; Lewis, 1955; Mayr, Linsley, and Usinger, 1953; Rol- 
lins, 1952; Shinners, 1958; Simpson, 1958; and Weatherby, 
1942) as a introduction to the vast literature on the subject. 
In the following, however, I wish to emphasize that the opinions 
are largely my own, even though the majority of them tend to 
agree with general practice. The major infraspecifie categories 
applied to North American pteridophytes are form, variety, and 
subspecies. 

I think that most of our forms are extremely minor categories 
at best—-some even results of injuries. Forms are trivial vari- 
ants, in my opinion, of no known consequence in the broad 
biology of the species, and they exist as scattered individuals. In 
Plate 4, I have tried to make a graphic representation of the 
category of form in comparison to subspecies and species. Forms 
may be induced by abnormalities or extremes of environment and 
development, or by single- or few-gene mutations. A good ex- 
ample of an environmentally induced form might be what has 
been designated as Woodwardia virginica f. fertilis—just an in- 
completely fertile leaf, with only a fraction of its sori developed. 
It tends to be found in deep, shady swamps. The same rhizome 
growing into sunny areas of the swamp will be normal, with 
fully fertile leaves. Abnormalities of development or ontogeny 
come usually from injuries or disturbance in growth. Trans- 
planting rhizomes of Onoclea sensibilis into the greenhouse often 
results in the leaf form hemiphyllodes, one side sterile, the other 
fertile. The same transplanted rhizomes produce, in my experi- 
ence, also imperfectly fertile leaves, f. obtusilobata. Both of 
these kinds of bizarre leaves are occasionally found in the wild, 
where they are produced by unknown factors (insects, injury 
by mammals, fire?). Taylor Steeves (1959) recently discussed 
Osmunda cinnamomea f. frondosa and f. latipinnula, giving a 


34 AMERICAN FERN JOURNAL 


good, modern example of the causal approach to the study of 
forms. 

In the category of genetic forms are probably most of the 
established crested forms of ferns, the skeletonized leaves, varie- 
gated forms, more or less sterile and divided variants (e.g., 
Polypodium virginianum f. cambricoides, Asplenium platy- 
neuron £. hortonae), and so on. Some years age (1931), Anders- 
son-Kott6 established that certain of these are of simple genetic 
nature, but so far as I know such work has not been carried out 
in recent years. Many of the North American genetic forms de- 
serve experimental study. In my opinion, however, no trivial 
“‘form’’ deserves a formal botanical name, no matter how 
peculiar-looking. In 99 out of 100 cases, I guess that their 
taxonomic significance is likely to be nil. Most are morpho- 
genetic monstrosities, the naming of which seems to be a hold- 
over from an earlier time and tradition, when the biology of 
plant species was poorly if at all understood. 

n one recent manual of the northeastern American flora 
(Fernald, 1950) I estimate that there are 110 unnecessary Latin 
or Greek names of trivial forms of pteridophytes. If we persist 
in naming such sports the sky is the limit: We could name 150 
more right now, for the eastern United States alone, and this 
would be just a beginning. (There must be at least 75 species, 
for example, for which no crested forms have been named; and 
many more for which we have no dwarfed forms named.) 

When classification proceeds to the level of variety, subspecies, 
and species, it now has experimental and statistical methods of 
far more profound significance than were available a half-cen- 
tury ago. Such methods cannot be used, of course, in unexplored 
and remote areas of the tropics that are still in the alpha stage 
of knowledge, but they can surely be adopted for our native 
pteridophytes. Criteria of detailed anatomy (using clearing and 
sectioning techniques, as well as observation under high-powered 
microscopes), gametophyte morphology and biology, chromo- 
somes, ecology and substratum, tests of characters in experimen- 
tal cultures, and—above all—the nature of the breeding popula- 


EVERGREEN GRAPEFERNS 35 


tion under natural conditions—these provide us a much more 
concrete evaluation of diversity and its meaning. 

The term variety, however, has been much confounded by 
different usages. Some botanists are inclined to drop it entirely. 
Some have used it to mean what others call ‘‘form.’’ Others 
have used it for ‘‘subspecies.’’ Still others have been incon- 
sistent. To make matters worse, the term ‘‘variety’’ has been 


minor, category. An example of a zoological definition of sub- 
species is appropriate: “Geographically defined aggregates of 
local populations which differ taxonomically from other such 
subdivisions of a species’? (Mayr et al., 1953). The italics are 
mine. The botanist Fuchs recently (1954) defined a subspecies 
as follows, emphasizing another aspect (and I quote the original 
French) : 

““Soug-mspice: Unité qui groupe les individus dont les 
différences morphologiques et anatomiques son bien établies et 
qui sont liés entre eux par des formes transitoires continues et 
non par des hybrides. C’est-a-dire que des ‘espéces’ liés entre 
elles d’une facon continue par des formes transitoires ne sont 
pas des espéces, mais des sous-espéces. ”’ ‘ 

To Fuchs’ definition, I know of only one exception so far in 
the study of ferns, namely the remarkable hybridization that 
occurs between species of Pteris in Ceylon (Walker, 1958)? 
producing complete intergradation between plants that are 


2 Reviewed, this JourNaL 49: 43, 44. 


FORMS (''VARIETIES, '' ONE SENSE) 


INDIVIDUALS SCATTERED 


THREE IDEALIZED TAXONOMIC SITUATIONS 


SUBSPECIES (''VARIETIES, '' ANOTHER SENSE) 


COMPLETE INTERGRADATION 


SPECIES @ 


FEW HYBRIDS OR NONE 


— GLVIg ‘0G ANNIOA 


IVNGNOf NYG NVOMAINVy 


EVERGREEN GRAPEFERNS 37 


with little doubt good species. This single exception is not suffi- 
cient, I believe, to prevent us from combining three ideas in a 
definition of subspecies or variety that will apply to the vast 
majority of pteridophytes, viz. that, in general, subspecies or 
varieties (a) are morphologically distinct; (b) geographically 
defined, i.e., have distinctive ranges; and (c) if their ranges 
are in contact or overlap, freely and continuously intergrade, 
without sterility barriers. The middle picture in Plate 4 en- 
deavors to show this graphically. 

Two subspecies or varieties, so defined, cannot have the same 
or overlapping ranges and at the same time co-exist in the same 
habits, for if they were shown to do so and still remained dis- 
tinet from each other without intergradation their interpreta- 
tion would then be questionable. If they did not intergrade and 
maintained their distinctions, there would have to be strong 
barrierg to interbreeding. The taxa would therefore not be sub- 
species but species. I thus believe firmly that if two substan- 
tially different taxa do co-exist and extend together over large 
areas without interbreeding then they must be regarded as bio- 
logical species. It would be a misuse of the concept of subspecies 
or variety to apply such categories to them. It would also be a 
misuse, in my opinion, if two undoubtedly distinct species of 
pteridophytes should form hybrids, sterile or fertile (by doubling 
of chromosome complements), and those hybrids are treated as 
subspecies or varieties of one of the parents; well marked inter- 
specific hybrids that lack intergradation with their parental 
species may be expressed taxonomically by means, to be described 
below, other than the use of infraspecific categories. 

Regarding the idea of breeding capacity within species, I do 
not mean, of course, to imply that there may not be intraspecific 
sectors which are incompatible, even in the same population, just 
as there are in animals. I shall not be surprised if there are all 
sorts of undetected biochemical mutants which are limited in 
their ability to breed with at least certain other members of the 
same species. In pteridophytes, the very little understood 
phenomenon of autopolyploidy—sectors of a species with differ- 


38 AMERICAN FERN JOURNAL 


ent multiples of the chromosome number (e.g., 42, 84, 126, 168, 
etc.) —probably divides a species into more or less saucaane 
reeding components (e.g., Polypodium vulgare, Aspleniw 
trichomanes, Cystopteris fragilis s.s.). Irene Manton and fas 
group at the University of Leeds are studying such complexes. 
Autopolyploid isolation might lead over great spans of geologi- 
cal time to varieties, subspecies, and species. We do not really 
yet know, however, to what extent the so-called polyploid forms 
at their origins may be able to exchange genes anyway with 
their diploid progenitors, in spite of the chromosome number 
barrier. The various chromosome races may be, so far as we can 
tell, identical morphologically and ecologically, indicating that 
except for genome differences they are genetically alike and 
therefore sectors of the same taxon. Chromosome number by it- 
self—at least at this stage of our knowledge—should not be used 
as the criterion of subspecies or varieties. These, instead, are 
more reliably defined upon suites of morphological characters 

d upon differences of range 
Current studies of the evergreen species of Botrychium in- 
volve such problems of interpretation. For a number of years I 
have been making mass collections of populations of these plants, 
and recently various members of this Society have very kindly 
made local population samples for me. All the evidence points 
to four distinct taxa in northern North America—B. dissectum 
(ineluding many ‘‘forms’’), B. multifidum, B. oneidense, an 
B. ternatum (or an American representative of it). All four 
taxa co-exist in what must be an unlimited number of localities 
across the large area running from Minnesota to Quebee and 
New England and downward to the southern Great Lakes region. 
I have attempted to obtain all my collections of the species grow- 
ing side by side with one, two, or three of the others. In this 
study I have taken 4,627 specimens of B. dissectum in 82 locali- 
ties; 640 of B. multifidum in 54 localities; 2,173 of B. oneidense 
in 66 localities; and 993 of B. ternatwm in 35 localities. The 
chromosomes of all four taxa have been counted; and all show 
n = 45, so there is no chromosome number barrier to hybridiza- 


Segments small to large Tips undivided to divided Range: N. to 8. 


1 { ternatum 1 dissectum 1 multifidum 
1 multifidum 2  oneidense 2 ternatum 
2 dissectum 3 { multifidum 3 oneidense 
3 oneidense l ternatum 4 dissectum 
Vernation early to late Segments angular to round Young leaves green to red 


1 multifidum 1 ternatum 1 oneidense 
2  oneidense 2 disseetum 2 multifidum 
3 ternatum 3 multifidum 3 dissectum 
4 dissectum 4  oneidense 1 ternatum 
Spores smooth to rugose Margins smooth to dentate Mid-winter leaves green to bronze 
1. multifidum 1 { multifidum 1 § oneidense 
2 ternatum _t pare 1) multifidam 
3 peer on tern 2 ternatum 


3) one ; oes 3 dissectum 


SNUGIAdVUt) NAAADAAAGT 


40 AMERICAN FERN JOURNAL 


tion. There is no obvious correlation of the character complex 
of one species with any of the others. The major variables are 
enumerated below, the species being arranged appropriately : 

hese character comparisons, some still incomplete, were made 
for the most part in mixed populations in the same habitats, 
and will be reported on in detail later.* There is overlap in prac- 
tically all the characters, so that the arrangement is based on 
averages. Where two species are bracketed, the character is es- 
sentially identical in both. There is no question that all four 
species are very closely related. However, the point to be stressed 
regarding these grapeferns is that they do co-exist over a large 
range, yet maintain their distinctness. No one species is similar 
enough to any other in all the variables to be merged with it 
into one species. Botrychium oneidense and B. ternatum could 
be made varieties as well of B. multifidum as of B. dissectum. 
If we maintain two species, B. dissectum on8 B. multifidum, 
then we must maintain the two others by any ““snecies standard”’ 
(cf. Rollins, 1952; Wagner, 1959). 

The following lvctaatbes the correlation of resemblances in 
nine variables: 

B, multifidum B. dissectum 
B. oneidense 5 4 
B, tern 4 5 

I believe that the taxonomic situation of these plants corresponds 
to the diagram on the right-hand side of Figure 1. Whether or 
not my treatment represents an ultimate taxonomic solution to 
the problem of inter-relationships of these evergreen grapeferns 
remains to be seen, but I think that this situation is a good illus- 
tration of a complex in which the infraspecific categories of sub- 
species or variety do not apply, in spite of the close resemblances 
that exist. 


3 The southern EE hag eS . bye Seagate does not occur to- 
gether with B. multifidum or B. ternatum, so far as I know; in fact their 
ranges do not nate ap. Howe ce Dale M. ‘Smith ie recently discovered 

: 0 i 


n 
ana, though not yet in the same habitat. Our fitting of B. tenuifolium into 
the character spectra for these plants then will have ag ag based probably 
on comparison with B, dissectum which does grow with 


EVERGREEN GRAPEFERNS 41 


Before going further, I should like to give some examples of 
taxa that I would regard as likely to be valid subspecies or varie- 
ties; the following list will be ample 
BoTRYCHIUM LANCEOLATUM var. LANCEOLATUM (Eurasia, western No rth 

America) and B. LANCEOLATUM var. ANGUSTISEGMENTUM (eastern North 

America). 

PTERIDIUM AQUILINUM var. LATIUSCULUM (North America, northern Europe, 
eastern Asia), and P. AQUILINUM var. PUBESCENS (western North America, 
Mexico). 

OSMUNDA REGALIS var. REGALIS (Eurasia) and O. REGALIS var. SPECTABILIS 


orth Am mae 
PHYLLITIS SCOLOPENDRIUM vV: ¢ BOOLOPENDRTUE (Europe) and P. SCOLOPEN- 


DRIUM var. AMERICANA (ster North America 
CRYPTOGRAMMA CRISPA var. A (Europe, pia tharenthtn Asia) and C. 
CRISPA var. ACROSTICHOIDES Vues Asia, North America) 


The example of bracken in the United States and Canada is a 
particularly good one because the two varieties grade into each 
other nicely where their ranges overlap (Tryon, 1941; Weather- 
by, 1942), and the picture is approximately like that of sub- 
species (or variety) in Figure 1. Most of the others are so well 
separated that their intergradation is unknown or absent. 

In contrast to these there are some designated subspecies or 
varieties that should be considered quite skeptically and deserve 
research, because there is a real doubt as to whether these are 


vironmental modifications. It goes almost without saying that a 
given fern genotype when grown in a deep swamp will produce 
plants of different aspect (larger, more divided leaves ; petioles 
and petiolules attenuate ; segments longer, more remote; texture 
thinner; sporangia fewer, if any) than in an open, dry field. 
By the same token, a mesophytie plant with certain characters 
in average habitats in Indiana and Michigan may be expected 
to have a different aspect if growing at very high latitudes or 
altitudes, e.g., Alaska (the plant smaller, simpler; petioles con- 
tracted; segments shorter, overlapping; texture firmer or more 
leathery ; sori or sporangia abundant and confluent). These 


2 


42 AMERICAN FERN JOURNAL 


changes are so familiar to a field worker that he usually makes 
his taxonomic treatment accord, although it is true that ulti- 
mately the field observations should be confirmed experimentally, 
if there is any doubt, by uniform culture techniques. However, 
even without experiments, the probability that such changes are 
merely environmental is extremely high. Such taxonomic segre- 
gates as the following are questionable : Botrychium simplex var. 
simplex and B. simplex var. tenebrosum, B. virginianum subsp 
virginianum and B. virginianum subsp. europaeum, B. multi. 
fidum subsp. multifidum and B. multifidum subsp. intermedium, 
Adiantum pedatum var. pedatum and A. pedatum var. aleuti- 
cum, Sea luec munitum var. munitum and P. munitum var. 
imbric 

A revision of the Ophioglossaceae by Robert T. Clausen (1938) 
has numerous examples of such subspecies and varieties, since 
the author depended so strongly on characters that students of 
these plants more interested in natural, biological populations 
recognize to be readily modified by environment. The examples 
cited above may not stand experimental tests and are probably 
mere forms. It is almost certain that individuals tinct as 
different subspecies or varieties growing in the same regions 
belong to one basic genotype and do not wa seta subspecifi¢ 
separation. For example, specimens of B. virginianum from 
Michigan identified as subsp. ewropacum are with little doubt 
only dwarfed forms of the typical subspecies 

To set up subspecies or varieties for distinct taxa that grow 
side by side in abundance and maintain their distinctions defies 
any definition of species, in my opinion. If two taxa co-exist over 
a large range and maintain their characters, they should be in- 
terpreted as species. This means that not only do the taxa have 
diagnostic features of sufficient number and nature to be readily 
separated, but they do not interbreed to form fertile popula- 
tions, or, if they do interbreed, it is to produce only occasional 
sterile hybrids or hybrids with decidedly diminished fertility. 
The varieties designated below would seem to represent, in 
actuality distinct species: Lycopodium alopecuroides var. alo- 


EVERGREEN GRAPEFERNS 43 


pecuroides and L. alopecuroides var. adpressum, Botrychium dis- 
sectum var. dissectum an . dissectum var. oneidense, 
lunaria var. lunaria and B. lunaria var. minganense, Gymno- 
carpium dryopteris var. dryopteris and G. dryopteris var. ro- 
bertianum, Dryopteris spinulosa var. spinulosa and D. spinulosa 
var. intermedia. : 

The first of these pairs, involving the Lycopodium inundatum 
complex in North America, greatly needs research. C. V. Morton 
recently called to my attention that the Athyrium filix-femina 
complex is another that deserves careful investigation. The case 
of Botrychium lunaria vs. B. minganense was revived several 
years ago when we wrote a report (Wagner and Lord, 1956) to 
show that when these plants grow together in the same habitats, 
there may be detected at least 14 differences, including even those 
of juvenile plants and leaf primordia. They do not interbreed, 
and we therefore considered them as sympatric species, the range 
of B. lunaria completely overlapping that of B. minganense 
(the ‘‘species’’ of Figure 1). I believe that the same interpreta- 
tion should be given to the other examples listed. With all the 
obvious differences between Dryopteris spinulosa and D. inter- 
media, species which flourish together in the same habitats inti- 
mately associated, it seems to me illogical to treat them as varie- 
ties of one species. Their common hybrid, D. X fructuosa, has 
abortive spores, indicating that, along with morphological dif- 
ferences, there is a clear-cut breeding barrier. 

The mention of a hybrid brings us to the other use of the 
eategories of subspecies or variety, which I would hold to be a 
misuse. It has been common in pteridophyte taxonomy to desig- 
nate plants which are more or less obviously interspecific hybrids 
as varieties or subspecies of one of the two parents, as follows: 
A < B = AB, but AB is treated as A var. AB or B var. AB. 
None of the following are varieties, in my opinion: Equisetum 
hyemale X kansanum = E. hyemale var. intermedium, Woodsia 
glabella X ilvensis = W. ilvensis var. alpina, Cystopteris bulbi- 
fera X fragilis = C. fragilis var. laurentiana, Dryopteris eris- 
tata X goldiana = D. cristata var. clintoniana, D. intermedia X 


44 AMERICAN FERN JOURNAL 


spinulosa = D. intermedia var. fructuosa, Asplenium montanum 
< pinnatifidum = A. pinnatifidum var. trudellu. 

To be true, most of these were interpreted as subspecies or 
varieties before evidence leading to concepts of hybridity was 
adduced; the error, it seems to me, is to persist in treating plants 
as ats of one of the parents, after the indications are pre- 
sented that they are interspecific hybrids. In each of the cases, as 
has been shown with at least some assurance (in the work of 
R. L. Hauke, D. F. M. Brown, R. F. Blasdell, E. T. Wherry, 
S. Walker, and the writer, respectively), the ‘‘varieties’’ in 
question are actually not members of one species but are crosses, 
containing the heredities of two species. If they are to be made 
subspecies, or varieties, it could be of one parent just as well as 
the other. Some of these hybrids, of course, are evlcied plants, 
well established in the biotic community, largely as ‘‘sexual 
oe through doubling of chromosomes (ampuldioioia or 
as ‘‘asexual species’’ through some form of vegetative, non- 
sexual propagation such as apogamy (cf. Emerson, 1955). Other 
interspecific hybrids among pteridophytes are entirely sterile, 
and do not reproduce significantly by vegetative means. Hybrid 
taxa are best expressed either as a formula (e.g., Dryopteris 
intermedia *K marginalis, the species arranged iiphabationlle) 
or as a binomial (D. & pittsfordensis; or, if a successful, sexual 
or asexual species, D. pittsfordensis, without the ‘‘X’’). I do 
not believe that it is ever appropriate to designate them as varie- 
ties of one of the parents. 

Having described the difficulties in the taxonomic interpreta- 
tion of the evergreen grapeferns and having briefly reviewed in 
the light of my own bias the usages of the various infraspecific 
categories, I do not believe by any means that the conclusions of 
this short review are necessarily definitive for either problem. 
Mostly, I think, it should be re-emphasized that there are avail- 
able today tools (such as techniques for studying anatomy, life- 
cycles, chromosomes, constancy of characters, and breeding 
ability) for making more objective analyses of taxonomic rela- 
tionships than we have had previously. Under these conditions, 


EVERGREEN GRAPEFERNS 45 


perhaps, a certain conceptual solidarity among North American 
pteridologists in how they apply the infraspecific categories is 
more appropriate than ever before. Perhaps the notes presented 
here will call to the forefront some of the principles and prob- 
lems we need to face. 

LITERATURE CITED 
ANDERSSON-KortTé, IrMA, 1931. The genetics of ferns. Bibliographica 


ica 8. 
CLAUSEN, R. T. 1938. A monograph of the Ophioglossaceae. Torrey Bot. 
: 1-177. 
Emerson, ALFRED E. 1955. Biological species. Encyclopedia Brittanica. 
0. go. 
FERNALD, M. L. 1950. Gray’s Manual of Botany, 8th Ed. American Book 


Co. 

Fucus, Hans Perer, 1954. Essai d’une nouvelle sega yee taxa 
‘¢genre,’’ ‘‘espéce,’’ ‘‘sous-espéce,’’ ‘‘varieté,’’ et ‘‘forme. gme 
Congr. Intern. Bot., Compt. Rend. des Seances & Rapp. & Communic. 

i, os 


EWIS, HARLAN, 1955. Specific nee Infraspecifie Categories in Plants. In 
Biological Systematics. Sixtee Ann. Biol. Colloquium Proc. 13-20. 
Oregon State College, Suen ore 

Mayr, E., Linsey, E. G., and R. Sam, 1953. Methods and prin- 
ciples of Systematic Zoology gonna Hill. New Yor 

Rouuins, Reep C. 1952. a today and eeeiberow. Rhodora 654: 
1-19 


SuHinners, Luoyp, 1958. New varietal names in Argemone (Papaveraceae). 
Southwestern Naturalist 3: 212-237. 

Smpson, G. G. 1958. The species problem (review). Science 127: 245, 246. 

STEEVES, TAYLOR 1 An ai agedatte of two forms of Osmunda 


cinnamomea. Rhodora 61: 223-23 
R . JR. 1941. Revision - the genus Pteridium. Gray Herb. 


TRYON, 
Contr. 134: 1-7 

WALKER, Evor G. 1958. Hybridization in some species of Pteris L. 
Evolution 12 “ 1): 82-92. 

WAGNER, W. s oe grapeferns qgpelcany Botryehium 
ternatum: a ae NT report. This JOURNAL 49: 103. 


Waaner, W. H., Jr. and Lois P. Lorn, 1956. The Satis and ecyto- 
logical distinctness of poten ong minganense and B. lunaria in Michigan. 
Bull. Torrey Club 83: 261-280. 

WEATHERBY, C. A. 1942. oe Rhodora 44: 157-167. 


University or MicnigaAN, ANN Arbor, MICHIGAN 


46 AMERICAN FERN JOURNAL 


The Ecology of Peruvian Ferns 
Rouua TRYON 


The Andes dominate Peru. They traverse its length and are 
responsible for the principal types of vegetation, their distribu- 
tion patterns, and for the richness and the diversity of the flora 
as well. The Altiplano is the high central land from 10,000 to 
14,000 feet elevation. It may be rather flat although usually 
it is broadly rolling; where it is penetrated by valleys there is 
considerable relief. It is bordered, except in the north, by a 

roken chain of high mountains many of which rise to 18,000 feet 
or more and support permanent snow fields and glaciers. These 
highlands are relatively cool and dry and the vegetation consists 
of grasslands, sedgelands, semi-desert shrubs, and cacti. Locally 
small woods, especially of Polylepis, may be found. Above the 
limit of agriculture, at about 13,000 feet up to the limit of 
vascular plant life at about 17,000 feet, is the Puna where the 
cold prevails and low cushion plants are the principal botanical 
feature. 

Along the eastern slopes of the Andes there is a transitional 
climatic and vegetational band from about 11,000 down to 6,000 
feet, and below this the tropical forest begins. This is the west- 
ern edge of the largest forest in the world for it extends un- 
broken for some 1,700 miles through the Amazon Basin to the 
Atlantic Ocean. On the Pacific side, the slopes of the Andes 
become progressively drier, below 10,000 or 8,000 feet to the 
coast, and are usually barren or with a sparse desert vegetation. 
In the north of Peru, these desert conditions become less pro- 
nounced and at the northernmost tip there is a small forested 
area. This otherwise barren coastal zone is relieved only at 
intervals by green irrigated river valleys and by the naturally 
verdant lomas. 

This paper is based primarily on the notes, collections, and 
observations made in Peru from July to November of 1956." This 


grSupported by a grant from the National Science Foundation (NSF 


Ecouogy or Peruvian FrErns 47 
trip was made for the purpose of providing a background for a 
taxonomic study of the ferns of Peru; I was accompanied by 
my wife Alice Tryon, to whom I am indebted for many of the 
results obtained. The principal places where we studied ferns 
are the following: The Montafa fern group at Iquitos, Tingo 
Maria, La Merced, and Potrero; the Ceja Thicket fern group at 
Carpish and Machu Picchu (the ruins); the Ceja Serub fern 
group near Huacapistana; the Sierra fern group at Cuzco, 
Huaneayo, and Tarma; and the Loma fern group at Amancaes 
and Lachay, in the Department of Lima. Our own material has 
been supplemented by that of others, largely taken from the 
publications listed in the bibliography. I am under particular 
obligation to Dr. Ramén Ferreyra, to Dr. César Vargas, and to 
Dr. Pedro Coronado for aid to my studies while in Peru. 

In the ecological classification of the ferns of Peru, two fern 
vegetation types, the Forest Fern Vegetation and the Steppe and 
Serub Fern Vegetation, are recognized on the basis of various 
characters of the plants. Each of these is subdivided into two 
ecological fern groups, primarily on the basis of environment 
and geography and secondarily on the basis of floristics. An 
intermediate between two of these groups is recognized because 
of its prominence. This classification is necessarily subject to 
local exceptions and transitions. However, it may be expected to 
portray the broad correlations between the plants and their 
environment. 

I have included only the native Pteridophyta in their natural 
habitats. Such introduced species as Adiantum Capillus-Veneris 
and Pteris vittata, on the sea cliffs near Lima, and such native 
species as Trismeria trifoliata, Equisetum bogotense, and E. 
zylochactum, along irrigation ditches and in similar habitats in 
the naturally dry coast and coastal valleys, will not receive 
further mention. 

I. Tue Forest Fern VEGETATION 

The forest fern vegetation is characterised by the presence of 
more or less obligate epiphytes and, among the terrestrial ferns, 
by continuous growth throughout the year (or at least the leaves 


48 AMERICAN FERN JOURNAL 


remain in a fresh condition), by sterile plants, by species with 
large leaves (5 feet long or more), some of which, having widely 
creeping rhizomes, form conspicuous colonies, and by species 
with some modification of the leaf for vegetative reproduction. 
In a given locality there is a relatively large number of species 
(about 50 to 100 within a mile radius) many of which are locally 
very rare. The area of this fern vegetation is that of the forest 
(Montafia) and moist Ceja de la Montana (Ceja Thicket). 

The following are some of the epiphytic ferns: Polypodium 
duale, P: filicula, P. percussum, P. plumula, P. angustifolium, 
P. polypodioides (var. Burchellii), P. phyllitidis, P. pectinatum, 
Eschatogramme panamensis, Asplenium serratum, A. auri- 
culatum, Ophioglossum palmatum, and a number of species of 
Elaphoglossum. Some of these species may also grow on bare 
rocks, or may survive, at least for a time, on fallen branches. 
These species are biologically similar to the Sierra ferns. They 
may become dormant during the relatively mild dry season, 
especially if growing on the more exposed branches, and they 
have small leaves; the plants are usually fertile and none of the 
species have leaves modified for vegetative reproduction. When 
the plant is dormant, the leaves of most of the species curl and 
probably can revive after a brief dormancy; those of Ophioglos- 
sum. die. 

The following are some of the species with large leaves: 
Nephrolepis biserrata (to 12 feet), Lygodium volubile (to 40 
feet), Hemidictyum marginatum (to 1 feet), Hypolepis paral- 
lelogramma (to 22 feet), Adiantum pectinatum (to 8 feet), and 
the tree ferns such as Alsophila microdonta and A. elongata 
(both to 10 feet). Others with large leaves frequently form con- 
spicuous colonies: Dennstaedtia cicutaria, Hypolepis hostilis, 
Pteris grandifolia, Gleichenia bifida, G. Bancroftii, Dicranopteris 
pectinata, and Pteridium aquilinum (var. arachnoideum). The 
last species is the most aggressive and may oecupy whole hillsides 
after they have been cleared for agriculture. 

There are two kinds of modifications of the leaf for vegetative 


Ecouoay or PERUVIAN FERNS 49 


reproduction. The following species have buds on the lamina: 
Polystichum  platyphyllum, Thelypteris macrotis, Bolbitis 
crenata, and Diplazium cristatum (all with buds along the rachis 
toward the apex of the lamina), Doryopteris pedata var. palmata 
(buds at the base of the lamina) and Tectaria incisa (buds at 
the base of the pinnae and along the pinna-rachises). In these 
species the buds are persistent and develop into plantlets, espe- 
cially on the old leaves, while still attached to the leaf. In 
Dennstaedtia arborescens there are deciduous buds in the axils 
of the pinnae. Other species have a rachis-tip that roots to 
produce a new plant. In Adiantum deflectens, Trichomanes 
diversifrons, and Asplenium radicans the rachis-tip is elongate. 
In Trichomanes Hostmannianum it is greatly elongate and may 
produce roots at intervals before developing a new plant at the 
tip; while in Danaea Moritziana the tip is elongated but not 
otherwise modified. Asplenium radicans has all, or nearly all, 
of the leaves rooting, whereas all of the other species mentioned 
also have leaves with normal apices. 

The reason for the considerable number of sterile plants in 
this fern vegetation needs investigation. The fertile leaves may 
be fugacious as in Bolbitis crenata, B. Lindigti, Polybotrya 
caudata, and P. osmundacea, and present only during a brief 
period of the year; or plants of some species may grow where the 
environment is not conducive to the formation of fertile leaves; 
or perhaps some species are not freely fertile, even in suitable 
habitats. 

Ta. Tue MontaNa Fern Group 

This is the fern group of the forest which occupies the large 
area east of the Andes and a very small area in the Department 
of Tumbes that is an extension of the Gulf of Guayaquil vegeta- 
tion. It is warmer than the Ceja Thicket and fog is not typical. 
There are some 60 to 140 inches of rain annually, most of it from 
October to April. 

It is difficult to distinguish the group, by its species, from the 
Ceja Thicket ferns. Many of the species of the Montafia are not 


AMERICAN FERN JOURNAL VoLUME 50, PLATE 5 


1000 FT. 
(300 M.) 


¢ 

Ee | es 

ig ee tS 
Fe} & gg 


DISTRIBUTION OF ECOLOGICAL FERN Groups In Peru. Map anp D1 
MATIC PROFILE 


AGRAM- 


Ecotocy or PERUVIAN FERNS 51 


Ba iat 


sufficiently common to be useful in defining the group and those : 
that are may also grow in the Ceja Thicket. The best floristic 
distinction, perhaps, is the presence, only in the Montana, of 
numerous and often rather common species of Adiantum with 
broad or dimidiate segments such as: A. petiolatum, A. obliquum, 
A. latifolium, A. macrophyllum, A. anceps, A. peruvianum, A. 
platyphyllum, A. pectinatum, A. tetraphyllum, A. fruetuosum, 
A. macrocladum, A. tomentosum, A. terminatum, A. pulveru- 
lentum, and A. villoswm. 

Furthermore it is difficult to distinguish floristic elements 
among the ferns of the Montafia, for the reasons mentioned 
above, although among its several hundred species there are 
some that are confined to the higher elevations and others to the 
lower ones. 

Is. Tue Cesa THICKET FERN GROUP 

The Ceja Thicket occupies locally favorable situations along 
the higher eastern slopes or low summits of the Andes at eleva- 
tions of about 6,000 to 11,000 feet. These are moist, cool areas 
where clouds and fogs are present most of the year. Tree ferns 
and the bamboo Chusquea are the dominant elements in the 
landscape, both extending well above the dense shrubby growth. 
The best floristic distinction from the Montafia ferns is the ab- 
sence of the Adiantums mentioned above and the presence of such 
species as Hymenophyllum Ruizianum, Gymnogramma fleruosa, 
Lycopodium complanatum, L. clavatum, L. Jussiaei and L. 
pendulinum. 

INTERMEDIATE BETWEEN Ia AND IIa: THE Cesa ScrvB FERN GROUP 

This fern group occurs in the same zone as the Ceja Thicket 
ferns but in drier situations (dry Ceja de la Montafia) where 
foggy conditions do not regularly occur. It is intermediate 
between the Sierra fern group and the Montafa fern group and 
forms a pronounced transition zone between them. Floristically, 
there is a mixture of species from the Sierra fern group and 
the Montafia group, especially of the more widely distributed 
ones. For example, Pellaea ovata, P. sagittata, and Woodsia 


HABITATS 
LACHAY) ; 


or E 
a. C 


oo 
COLOGICAL 
EJA 


FERN Groups IN Peru. 1, Sierra Group (Near TarMA); 2, Loma Group (LoMA 
Scrusp Group (NEAR HvuAcapisTana); 4, MonraNa Group (Iquitos) 


IVNUNOL NYA NVOIUIIVy 


9 ULVIg ‘0G ANATION 


Ecouogy or PERUVIAN FERNS 585) 


montevidensis, eee Sierra ferns, may pe in the company 
of Polypodium crasst folium, Pity artarea, Nephroleptis 
ae and N. pists tea are naiatet Montana species. 
of the ferns have small leaves, are terrestrial, are usually 
ae and do not have leaves modified for vegetative reproduc- 
tion. In these characters, the group resembles the Steppe and 
Serub fern vegetation. The plants grow more or less throughout 
e year or at least maintain leaves in fresh condition and in 
this respect it resembles the Forest fern vegetation. A few 
species such as Pteridium aquilinum and Pteris muricata have 
large leaves. 


Il. Tur Srepre aND ScruB FERN VEGETATION 


The Steppe and Scrub fern vegetation is characterised by the 
absence of obligate epiphytes and by terrestrial plants with a 
definite dormancy during which the leaves either die or become 
curled, with fertile and small leaves (less than 2 feet long), and 
by the absence of species that form conspicuous colonies or have 
leaves modified for vegetative reproduction. In one locality there 
are relatively few species (about 5-15 within a mile radius) but 
most of these are frequent. This fern vegetation occupies the 
rather dry western portions of Peru. It has a mosaic distribu- 
tion, for most of the coastal and adjacent Andean slopes are too 
dry for ferns, and in the Altiplano and the mountains it is 
mostly confined to rocky hillsides, cliffs, and similar habitats. 

TIA. THE SIERRA FERN GROUP 

The Sierra ferns have a seasonal dormancy, but the dormancy 
may be intermittent due to infrequent rains during the dry 
season. The available moisture is largely in the soil or in rock 
crevices and there is a greater number of species where there is 
local seepage or perhaps conditions suitable for the condensation 
of water from the atmosphere. There is little information con- 
cerning this latter factor although it may be an important one. 
The rainfall, some 20 to 44 inches annually, is by no means 
deficient in itself but the very dry air, the winds and the strong 


54 AMERICAN FERN JOURNAL 
insolation at the high altitudes combine to reduce its effective- 


ess. 

Most of the Sierra ferns may be divided into two kinds. The 
xeric element is typical of the drier sites and these species curl 
and retain their leaves during the dry season; they may revive 
and continue activity during brief moist periods. Such species 
are: eilanthes incarum, C. pruinata, C. myriophylla, C. 
iiirtoo: Polypodium pycnocarpum, Pellaea ternifolia, Sclagi- 
nella peruviana, Notholaena nivea, and N. aurea. The mesic ele- 
ment is typical of the locally moister habitats and in these species 
the leaves die during the dry season, unless in an unusually 
favorable place. Adiantum Poiretii, A. digitatum, Cystopteris 
fragilis, Asplenium fragile, Thelypteris Rosei, and oodsia 
montevidensis are typical of this element. Two species of this 
element are exceptional in having proliferous buds on the leaf. 
These are Asplenium fragile and the closely related A. Gilliesii. 
A bud develops on the petiole which may become greatly elongate 
and act as a stolon. The petiole below the bud is persistent after 
the lamina has withered and the plantlet develops at what ap- 
pears to be its tip. 


IIs. THe Loma Fern Group 

The loma fern group is part of the unique vegetation occurring 
at intervals along the coast of Peru (and Chile) north to about 
8°S. Latitude. This vegetation develops in response to local 
physiographic conditions that, in winter, lead to more or less 
constant fog and guara (heavy mist) at certain hills and valleys. 
The summer months are continuously dry. Although a rather 
lush vegetation may be developed by the unusual moisture con- 
ditions, the flora, on the other hand, is evidently restricted by 
the long continuously dry season. 

The loma ferns represent, almost wholly, a selection of species 
from the Sierra ferns. To my knowledge, only three of the 
species are not also among the Sierra ferns: Ophioglossum 
nudicaule, a Polypodium near P. lasiopus, and a Dryopteris near 
D. patula, but it is not certain that they do not also occur there. 


Harr’s-ronaue iN New York DE 


i) 


Further study must be made before it will be known if the last 
two are perhaps endemic to the lomas. The most common species 
are: Polypodium pycnocarpum, Adiantum subvolubile, and A. 
digitatum; others are: Ophioglossum petiolatum, Anogramma 
leptophylla, Woodsia montevidensis, Notholaena peruviana, and 
Adiantum Poiretii (vars. hirsutum and sulphureum) . 

About 80 percent of the species of flowering plants on the 
lomas are endemic and the few, or no, endemics among the ferns 
is in striking contrast. This may be due to a slower rate of 
evolution among the fern species but I think that it is more 
likely that it reflects their superior means of dispersal. Ttas 
quite possible that, from the Andes, spores of these species reach 
the lomas with sufficient frequency to negate the effects of their 
geographic isolation. 

BIBLIOGRAPHY 
CrerRATE, E. Notas sobre la vegetacion del Valle de Chiquian. Folia Bi- 
ologica Andina-Bot. 1: 9-389. : 
Frerreyra, R. Comunidades vegetales de algunas lomas costaneros del Peru. 
Est. Exper. Agric. ‘La Molina’. Bol. 53. 1953 
and 


WEB nape A. Phytogeography of the Peruvian Andes, (in Maebride, 
Flora of Peru). Field Mus. Nat. Hist. Bot. Publ. 13: 13-81. 1936. 
El mundo vegetal de los Andes Peruanos. Lima, 1945. 
Gray Hersartum or Harvarp UNIVERSITY, CamBRIDGE, Massa- 
CHUSETTS. 


Survival of Hart’s-tongue Fern in Central New York 
Miuprep E. Faust 


One hundred and fifty years after the hart’s-tongue, Phy/litis 
scolopendrium (L.) Newm. var. americana Fern., was first re- 
ported in America, at Split Rock, New York, July 20, 1807, by 
Frederick Pursh, it is still persisting at least in the immediate 
vicinity. Trees which shaded it have been cut, a TNT explosion 
took place near by, and quarrying has destroyed many of the 


56 AMERICAN FERN JOURNAL 


sheltering rocks. To be sure, the plants are ie luxuriant but 
they will probably remain until man completely changes the 
habitat. When this station was rediscovered jae er 30, 1879, 
by the Syracuse Botanical Club, Charles Peck, who had been 
convinced that this station had long been destroyed, wrote then: 
**Pursh has been vindicated and botanists everywhere will re- 
joice. I regard this rediscovery of the Scolopendrium in its origi- 
nal locality after a lapse of more than seventy years as scarcely 
less important than the discovery of the Epipactis. It shows con- 
clusively the persistence of species when left alone.’’! 

Reports of the numbers, or more often of the thriftiness of 
the plants in the stations, have been given from time to time but 
not until 1916 was an organized census taken. Mabel Hunter 
(1922) fortunately chose the Jamesville Woods substation in 
Clark Reservation. In 1920 she repeated the census, found the 
plants increasing, and suggested that the census be a yearly one. 
Not until 1936 was another organized census taken in the Reser- 
vation by Lillian R. Sedgwick, David Caldwell, and myself. 
Since then this has been continued every five years with the 
assistance of botany students at Syracuse University and a num- 
ber of members of the Syracuse Botanical Club—Lillian Sedg- 
wick, Nettie M. Sadler, Mrs. Ellis Hinman, and Eleanore Porter. 
Because of the growth on talus we felt it would hinder survival 
to go into the area each year. 

DisTRIBUTION AND History 

In central New York the native localities are still restricted to 
Onondaga and Madison Counties, where the ferns grow on talus 
slopes of glacial ravines and plunge basins. Coolness is the out- 
standing characteristic of all the ravines. The cool air pockets 
with the associated northern plants of the White Lake areas are 
well described by Petry (1918). The plants grow in the black 
humus of the beech, maple, hemlock, and yellow birch forests 
well below the overhanging ledges (largely of Onondaga lime- 


1Letter (in Syracuse University) to Mrs. Myers, Oct. 3, 1879, Albany. 


Tiart’s-TONGUE IN New YorK ‘ 57 


stone) on east- and north-facing slopes or in similar shaded 
areas. 

A summary of the history as given by Maxon (1900) and 
Hunter (1922) is outlined below. The Roman numerals are used 
for Hunters’ stations and colonies and capital letters for sub- 
stations. 

Geppes (Spiir Rock) (Pursh, 1807) 
II. Currrenaneo Fanus (Cooper, 1830?) 
III. Prerryvitte Fauis (Ledyard, 1898) 
IV. JAMESVILLE 

Howrerr’s Gorge (Foote, 1866) 


i 


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JAMESVILLE Woops (Maxon, 1900) 
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S 
. Munnsvitin (Stebbins, 1934) (Page?) 
VI. BatpwinsvIttE (Larsen, 1959) 

Stations I, IV and VI are in Onondaga and II, III and V in 
Madison County. 

About 1900, H. D. House transplanted several of the plants 
from Chittenango Falls to a ravine near Munnsville. These per- 
sisted until at least 1920 (Hunter 1924) but probably disap- 
peared soon afterwards and were not found in 1946. 

By 1924, West White Lake was “eovered with the blastings 
of quarrying operations’ (Hunter 1924) and Green Pond, often 
called ‘‘Scolopendrium Lake’’ because of the abundance of the 
fern, was in the direct line of quarrying. In December united 
efforts were made to have this unusual White Lake area pre- 
served and included in the Clark Reservation but when these 
failed the owners were persuaded to hold off operations until 
the hart’s-toneue could be saved and many transplanted in the 
Reservation (Hunter 1924) (House 1926). Over 1000 plants 


<4 


58 AMERICAN FERN JOURNAL 


were transplanted by the summer of 1925. Some 350 placed 
in a ravine east of the steps were washed out the spring of 1925; 
around 360 were planted in Colony VI of the Jamesville Woods 
substation and others in original or other ravines throughout the 
park. The Green Pond was destroyed in 1925. East White Lake 
was in the line of quarrying and was half filled in in the late 
thirties, and so no more hart’s-tongues. 

Through the efforts of the Syracuse Botanical Club an 
amendment to the State Law was passed in 1930 to protect the 
hart’s-tongue in these two counties. During the thirties the trees 
around the Rock Gorge substation were cut, so that by 1942 
there was only one mature and two young plants and these were 
gone before 1945. In 1934 Stebbins (1935) found two well- 
established plants in a ravine near Munnsville. For a while this 
was believed to be a native area and thus the region was extended 
to the east. Drs. House and Maxon had searched for the fern in 
this locality years before with no results. It was suggested by 
r. Robert Crockett that it was probably planted by the late 
Mr. Page. We consulted Dr. House and he consulted the son of 
Mr. Page, who said that his father did plant some in that vicini- 
ty. On November 15, 1946, Eleanore Porter, Nettie Sadler, and 
I found a very characteristic ravine, not with two, but with two 
groups of the ferns, one with nine and the other eleven mature 
plants. This is some distance west of the ravine where Dr. House 
planted some much earlier. 

The most recently reported station VI was found by Michael 
J. Larsen in the vicinity of Baldwinsville, in October, 1959. 
There was one large plant with about twelve large leaves with 
many sori. It is on a north-facing bank of a cool ravine where 
many of us have botanized without seeing it. It is about ten 
miles northwest of the original Split Rock station. 


i} 


METHODS OF THE CENSUS 
The census now taken every five years from 1936 through 
1956 follows that of Hunter, in which the counts included three 
groups, mature plants which had at least one leaf with sori, 


TABLE 1 


IV. JAMESVILLE STATION 


H. JAMESVILLE Woops SUBSTATION 


Colony 1916 1920 1928 1936 1941 1946 1951 1956 
(After Hunter) 
x 159 197 100 159 125 164 249" 
I 109 115 
53 159 130 253 783* 
IIt a 8 
Ivi 119 206 
129 258 307 583* 272, 
Vv 29 20 
VI 136 92 4182* 108 91 73 110 89 
Totals 569 638 390 667 635 1110 1393* 
B. Lirr.e LAKE (GREEN LAKE) SUBSTATION 
I 40+ 3 4 19 52* 9 
Il 50 93 132 175 438* 255 
Totals 90 96 136 194 490* 264 
Totals for Clark © 
reservation 728 486 803 829 1600 1657* 


* Highest count. 


1 Colonies IT & IIT and IV & V combined in 1936 because of fluctuation of areas. 


2 About 360 transplanted to colony VI in 1925. 


WYOK MAN NI GTADNOL-S, LUV]T 


60 AMERICAN FERN JOURNAL 


young plants (we consider those one inch and more in length 
and without sori, whereas Hunter considered leaf-shape), and 
sporelings and prothallia. Whenever possible the counts were 
made from the bottom toward the top of the ravines, as the 
young ones can be seen more readily and there is not as great a 
hazard for the plants. Counts were made during the late fall 
or early spring. The former is better, after the early frosts when 
the tall herbs, like Impatiens, are killed and before the snowfall. 


RESULTS AND DISCUSSION 

Table I shows the census of mature plants in the ravines of 
Clark Reservation from 1916 until 1956. The fluctuation in 
counts is to be expected in a plant of such restricted range and 
the counts, having been made at five year intervals instead of 
yearly, show an incomplete picture of survival. Miss Hunter 
found an growth of 12.1% in the Jamesville woods substation in 
the years between 1916 and 1920. This shows a trend toward 
conservation since before this people were free to pick and dis- 
tribute plants and areas were being cleared and burned. It is 
encouraging to see the gradual but fairly constant increase from 
1936 through 1956. The history of the area will help to explain 
the 16 years between 1920 and 1936 with its general reduction in 
numbers. Certainly the destruction of half of the Jamesville 
stations would lower the count in general but the transplants 
should have aided the increase in the park. However, plants 
become established very slowly ; the early Thirties were very dry, 
and many of those which had apparently become established 
could not endure this drought. For example, colony VI seemed 
to maintain its transplants at least until 1928. The very evident 
rise in numbers throughout the preserve from 1941 until the 
present can be due in large part to the passage of the 1930 law 
protecting the ferns and to the constant alertness of the park 
ranger, Mr. Ryan, during many years. Most of these canyons are 
out of the direct line of any of the paths in the park. With the 
moving of the steps farther away from colony I of Green Lake 
these plants seem to be increasing. We have noted, as did Miss 


Harr’s-TONGUE IN NEw YORK 61 


Hunter, that sometimes during dry seasons the prothaffia will be 
more abundant when the mature plants, which are more exposed, 
are drying. Also there is a decided shift in the location from 
year to year or between counts. That is the reason we had to 
combine some of the colonies. They are in anastomosing canyons 
and it is almost impossible at times to separate the colonies. Oc- 
casional yearly counts have helped to show a slight correlation 
between the number of prothallia and mature plants the follow- 
ing year. In 1946, we had a cool rather dry summer with many 
cool nights and there were a large number of prothallia in most 
of the ravines. The highest number of plants for all the stations 
was in 1951. Not only are the numbers increasing but the plants 

e ‘‘thriving.’’ In 1945, one plant in the park had 133 old 
leaves and with 127 new leaves unrolling in the spring. These 
were 3.5 inches wide and some 26 inches long. These ravines will 
probably never have the moisture and coolness accompanied by 
the rare northern plants of the White Lake area but they will 
thrive under the present protection of the park. 

Survival outside of Clark Reservation is difficult to predict. 
In 1951, there were 216 plants in the stations outside as com- 
pared with 1600 in the park. Chittenango Falls is the only 
other station in a park. Unfortunately, a path was cut through 
one colony, which was destroyed by 1956. The other area is 
decreasing since a clearing was made near it. Perryville Falls 
has never had many plants and they are small with narrow 
leaves. It has improved since many of the raspberries are being 
replaced by trees. Munnsville remains the same. Howlett’s 
which had 87 plants in 1925 had 3 in 1956 and 9 the following 
year. It is in one of the most exposed areas and near a path. 
Evergreen Lake, the only one left in the destroyed White Lake 
Area, is owned by people who are interested in preserving it. 
With a count of more than 50 when discovered by Dr. Petry in 
1920 it had 73 in 1951. The original area at Split Rock continues 
to maintain itself, but the plants are small. However in 1951 
there were 11 mature, 60 young, and 105 prothallia. 


62 AMERICAN FERN JOURNAL 


SUMMARY 
1. Two new stations for hart’s-tongue have been discovered in central 
New York since 1922. One is apparently a planted one and the origin of the 


e ntibes of plants in the stations in Onondaga and Madison 
devittiee have been counted at five year intervals from 1936 until 1956. 

3. The number of mature plants in Clark Shier as mane from 
1916 until 1956 shows a decided increase. 


REFERENCES 
House, Homer D. 1926. Saving the Hart’s Tongue. This JOURNAL 16: 
3 


—37. 
Hunter, Maser R. 1922. The ees Status of Scolopendrium in New 
York State. This JouRNAL 9: 28-36. 
192: otes on Soljendiam vulgare Sm. This JourNAL 14: 102-04. 
Micon, W. R. 1900. the Occurrence of the Hart’s Tongue in America. 
Fernwort Papers 30- 46 
Petry, LOREN C. 1918. Studies on the Vegetation of New York State-II. 
The Vegetation of a Glacial Plunge Basin and its Relation to Tempera- 
ture. Bull. Torrey Club 45: 203-10. 
STEBBINS, J. LEDYARD, Jr. 1935. Further Notes on the Hart’s Tongue. This 
JOURNAL 35: 


Ferns and Allies in Kansas 
Ronaup L. McGreeor 

Kansas is a state characterized by cycles of drought and good 
moisture years. Summers result in the drying of habitats to a 
point where only a few localized spots remain moist. In periods 
of drought, often lasting for a period of years, even the most 
protected habitats dry completely. Thus ferns have a very local 
distribution and only a few of the most common species are of 
somewhat general occurrence; a protected habitat insuring 
a moisture supply is a necessity. All habitats, whether limestone 
or sandstone, are neutral or above in ph. with the result that 
rock inhabiting species are found on both limestone and sand- 
stone cliffs, except for Cheilanthes feei and Pellaea dealbata, 
which occur only on strictly calcareous sites. 

In the eastern half of Kansas rocky wooded hillsides, particu- 
larly those with at least small rocky cliffs, have a fern flora. 


FERNS IN KANSAS 63 


Often only one or two species are encountered and these seldom 
in abundance. An average undisturbed site will have Cystopteris 
fragilis var. protrusa, Botrychium virginianum, and Woodsia 
obtusa. Rarely Adiantum pedatum and Botrychium dissectum 
var. obliquum will be found. Sandy oak-hickory areas, particu- 
larly if sandstone cliffs are present, generally have a greater 
diversity of species and a larger number of each species. 

In the state 20 genera of ferns and fern allies are found, con- 
sisting of 32 species, 7 varieties and subspecies, and 5 forms. 
Most of these are species familiar to any student of ferns. In the 
following described locations will be found most of the species 
known for the state 

One interesting locality, and the only place in Kansas where 
such a place is found, isin NEY NEY, Sec. 1, T. 358., R. 25 E., 
Cherokee County. This is in the small Ozark area in extreme 
southeastern Kansas where a 100 foot high steep north-facing 
bluff occurs for a short distance along Shoal Creek. The upland, 
above the bluff, is an open wooded area with Quercus marilandica, 
Q. stellata, Q. Shumardii, and Carya tomentosa as the dominants. 
The soil is shallow over surfacing cherty Boone limestone of 
Mississippian Age. In April and May scattered colonies of 
Ophioglossum Engelmannii occur in open places and Isoétes 
Butleri is rare on shallow soil in depressions of surfacing lime- 
stone. 

At the top of the bluff an abrupt 8 4 of limestone aver- 
aging 15 feet in thickness occurs. A carpet of Woodsia obtusa 
is found just above this outcrop and is conspicuous until June 
at which time it dries and is not evident again until fall. Along 
the ledge of limestone are found a few species erowing from 
erevices and pockets. These are: Asplenium platyneuron, A. 
resilicns, Cheilanthes lanosa, Cystopteris fragilis var. simulans, 
Pellaea atropurpurea, and Pellaea dealbata. 

Below the ledge of limestone is found a moist, steep, talus 
slope on which grow Quercus alba, Cornus florida, Lindera 
benzoin, Ilex decidua, Physocarpus opulifolius var. intermedius, 
Viburnum rufidulum, and Forestiera acuminata, with Betula 


64 AMERICAN FERN JOURNAL 


nigra along the stream at the bottom. Occurring on this slope 
as scattered individuals and colonies are: Adiantum pedatum, 
Asplenium platyneuron, Botrychium virginianum, Camptosorus 
rhizophyllus, Cystopteris fragilis var. protrusa, Polystichum 
acrostichoides, Thelypteris hexagonoptera, and Woodsia obtusa. 
On the flood-plain terrace at the foot of the slope is found 
Equisetum hyemale var. pseudohyemale. 

A mile west of this site is another similar location but with 
less diversity. Here, however, occur two species known nowhere 
else in Kansas. These consist of a few plants of Cheilanthes 
alabamensis and Pteridium aquilinum var. pseudocaudatum. 
Botrychium dissectum var. dissectum was collected once at this 
location. 

A second fern site in Kansas is located in the SW14 NE! Sec. 
36, T. 33 8., R. 11 E., Chautauqua County. This is an area 
where sandstone of the Douglas Group, Pennsylvanian Age out- 
crops or surfaces. A forest of Quercus stellata and Q. marilandica 
dominates the scene; but along sandstone cliffs are found Carya 
exana, deicesces arborea, Viburnum rufidulum, Opuntia 
humifusa, Saxifraga terana, and Bouteloua hirsuta. This site is 
characterized by a 10 to 20 foot sandstone cliff along the east 
side of a small creek. Above the cliff are a series of four barren 
sandstone flats 5 to 20 yards wide which merge with a gently 
sloping, sandy, oak-wooded upland area. On shallow soil around 
these bare areas is an abundance of Selaginella rupestris and our 
only known location for Ophioglossum Engelmannii on sandy 
soils. On shaded sandstone rocks are found Cheilanthes lanosa, 
Woodsia obtusa, and Asplenium platyneuron. On the upper 
edge of the cliff are masses of Cheilanthes lanosa, Woodsia 
obtusa, and Selaginella rupestris. On the vertical side of the 
partially shaded west-facing cliff are found: Asplenium platy- 
neuron, A. Trichomanes, Cheilanthes lanosa, Cystopteris fragilis 
var. tennesseensis, Dryopteris marginalis, Pellaea glabella, and 
Woodsia obtusa. One small canyon cuts through this cliff and 
where it joins the creek are to be found Onoclea sensibilis. 
Polystichum acrostichoides, and Thelypteris palustris. On a 


Ferns In KAnsAs 65 


large block of sagas which has broken loose from the cliff 
is the only known Kansas colony of Polypodium polypodioides 
var. Miskovtihiis 
A little known habitat . ferns is to be found in the NEY 
NEY, See. 5, T. 34 5., 5 W., Barber County. A few canyons 
in the area ave mae ee outcrops of argillaceous, cal- 
careous, dolomite associated with gypsum. On a few of these 
small rock outcrops occur colonies of Cheilanthes Feei and Pellaea 
tropurpurea. Small sandy rivulets in these canyons have 
colonies of Equisetum laevigatum subsp. Funstonu and Marsilea 
mucronata. The only trees present are Populus Sargentti, Celtis 
reticulata, and Sapindus Drummondu. At this location Pellaea 
atropurpurea has been found growing in a clump of Opuntia. 
The High Plains topography of western Kansas is one of 
general monotony. The upland, however, is marked by thou- 
sands of depressions of various sizes. Most average but three to 
ten meters in diameter and a foot in depth. These depressions 
are known as buffalo wallows and their origin attributed to the 
wallowing of buffalo. Probably a number of factors account for 
their presence including wind scour, differential eolian deposi- 
tion, differential compaction, solution- subsidence, and the wal- 
lowing of buffalo. These depressions in the buffalo grass prairies 
hold water in the spring until late June. In about 15% of these 
wallows Marsilea mucronata occurs as small plants only two or 
three inches high. As hot summers arrive the wallows slowly dry 
and sporocarps are produced in abundance. In such areas road- 
side ditches often are carpeted by Marsilea for miles along county 
roads. In early fall the numerous sporocarps appear as though 
thousands of small beans had been spread in the dried ditches. 
In SEY, SWI, See. 14, T. 11 S., R. 4 W., Ottawa County is 
found an area known locally as Rock City. It consists of several 
clusters of round sandstone concretions four to twelve feet in 
diameter formed by erosion of Dakota Sandstone. These con- 
ceretions are located at the lower slope of a hill and adjacent to 
a river valley plain. Several of these concretions have fissures 
from which grow Cheilanthes Feei and Pellaea glabella. Under 


66 AMERICAN FERN JOURNAL 


the shaded side is found Woodsia obtusa. Though the small area 
is frequented by picnickers, these ferns have persisted for over 
90 years. 

The sand hill areas of central and western Kansas are not 
without a few interesting fern allies. In section 18, T. 22 8., R. 
5 W., Reno County, are several small sand dune enclosed ponds. 
These invariably contain colonies of Marsilea mucronata but 
rarely are sporocarps produced unless the ponds dry completely 
by mid-summer. This is the only known site for Pilularia 
americana in Kansas, but it is relatively common around the 
margins of these Bonde and regularly produces sporocarps. 
Seattered colonies of Equisetum laevigatum subsp. laevigatum 
occur in the area. 

A few other species are found rarely in the state and may be 
briefly listed. Azolla mexicana is found in old oxbow swamps in 
he Kansas River Valley near Lawrence. Isoétes melanopoda 
occurs in a moist swale in a nearly pure stand of Eleocharis three 
miles southwest of Neodesha, Wilson County. Equisetum arvense 
is known from a few moist sandy areas in eastern Kansas but is 
very local in occurrence. 

Several species have been listed for Kansas but must be ex- 
cluded. They were distributed from a fern garden to various 
herbaria and have been listed in manuals. These are: Athyriwm 
Filiz-femina (L.) Roth, Athyrium lien ae (Spreng. ) 
Tidest., Athyrium thelypterioides (Michx.) Desv., Cystopteris 
bilbifete (L.) Bernh., Dryopteris ean (Hook.) Gray, and 
Osmunda regalis L. 

THe UNIVERSITY of KANSAS, LAWRENCE. 


A Mule-train Trip to Sierra Mohinora, Chihuahua 
Donovan 8. CorRELL 
Botanists of our day, similarly to botanists of yore, strive to 
use the most modern ways of transportation within their means 
to carry on their field work. Richard Spruce and others in South 
America, during the nineteenth century, used waterways wher- 


Trip To SrerRA MOHINORA 67 


ever possible; Charles Wright and his contemporaries, in the 
early exploration of our own Southwest, used wagons, often 
belonging to the army, as did J. K. Small and his colleagues in 
exploring southern Florida at the turn of the twentieth century. 
Cyrus Pringle traveled mostly by train in his great Mexican 
exploratory work. Very few, with the exception of such ec- 
centrics as Rafinesque, resorted primarily to walking. 

Today, in the same tradition, we use some form of automotive 
transportation to take us within comparatively easy striking 
distance of most of our goals. Very seldom, and then only to 
reach otherwise inaccessible regions, is it necessary for one to 
have recourse to primitive and time-consuming means of travel, 
such as by mule- or pack-train. 

In October of last year, Dr. Howard Scott Gentry and T de- 
cided to leave momentarily the comforts of modern conveniences 
and venture out into the remote areas of southwestern Chihua- 
hua. Dr. Gentry hoped to satisfy a twenty year desire to attain 
the summit of Sierra Mohinora. He and his assistant, Juan 
Areuelles, were primarily interested in collecting seeds of wild 
plants for chemical analysis by the United States Department of 
Agriculture. I was hoping to obtain some additional fern collec- 
tions for Dr. Irving W. Knobloch’s and my work on the ferns 
and fern allies of Chihuahua which we expect to send to press 
later this year.t 

After assembling our supplies in Parral, we started west in a 
Chevrolet Carryall for what we thought would be Ciénega 
Prieta. The narrow road we travelled is used primarily for 
hauling logs and raw lumber to Parral from San Juan and other 
points more than 100 miles away. As it winds through the moun- 
tains of northern Durango and southern Chihuahua, the road 
traverses some spectacular, as well as dangerous, and botanically 
fascinating country. The canyons and breaks of the Rio San 

1] wish to acknowledge a Fellowship given me by the John Simon Guggen- 
heim Memorial Foundation in support of this work. TI also wish to acknowl- 
edge the help of Mr. C. V. Morton in identifying some of the more difficult 
collections. 


68 AMERICAN FERN JOURNAL 


Juan and Rio Verde are especially rugged and botanically rich. 
The most hazardous stretch on the entire 120 mile trip is along 
the west side of the canyon formed by the Rio Verde. On 
boulders and ledges of ravines sloping into this canyon were to 
be found such species as Polypodiwn erythrolepis, Woodsia 
mexicana, and Asplenium monanthes. 

Because of the primitive nature of much of the road and our 
frequent botanical stops, we did not arrive in the sawmill town 
of San Juan until the afternoon of the third day. On the way 
over from Parral, however, we made a good haul of ferns, in- 
cluding the uncommon Notholaena Aschenborniana and Pellaea 
allosuroides, as well as Asplenium Palmeri, Cheilanthes mezi- 
cana, Notholaena Grayi, Pellaea sagittata var. cordata, and a 
peculiar Polypodium which appears to be a hybrid of PB; seu ibio- 
lepis and P. thyssanolepis. 

Upon arrival in San Juan we were fortunate to find that a 

-muleteer (‘‘arriero’’) had just come in with a caravan from 
Guadalupe y Calvo. Since our contact in San Juan was at that 
time in Ciudad Chihuahua on business, we quickly engaged this 
muleteer for a trip of indefinite length but with his assurance 
that he would eventually place us on the summit of Sierra 
Mohinora. 

The next day, Friday, October 9, with our supplies loaded on 
pack animals and with Dr. Gentry astride a mule and me settled 
on a small mountain pony, we headed for adventure. We would 
share walking with Juan since our muleteer did not own another 
mount. It was amusing and at the same time chagrininge that 
the muleteer, by his sears, considered me, a mere 170 
pounder, to be a ‘‘fat man.’’ When I mounted his small horse 
he winced. 

The greatest inconvenience in traveling with a mule-train is 
that one must gear one’s life to the needs and progress of the 
animals. Erratic lunches are conditioned on reaching a grazing 
spread, and the same holds for camping, plus the need for water. 
Looking down from the undulating hick of a mule can also be 
frustrating when one sees what one takes to be Cheilanthes pyra- 


Trip To SrerrRA MoHINORA 69 


midalis or, wait, is it C. angustifolia; or, is that Polypodium 
guttatum or P. Hartwegianum on that shaded boulder? Time 
and again this would happen and it would always be a case of 
clambering from the mule to verify, and usually collect, the 
species or of settling the question by mentally deciding that 
there were already abundant collections of that particular 
species. 

A mule-train also has its virtue in that the slow pace gives one 
an opportunity to enjoy the gradual and ever-changing land- 
scape. ach promontory revealed scenes that strikingly re- 
sembled those of woodeuts found in old travel-books. 

Our trail (cf. map) left San Juan by way of Chinatt, with 
Ciénega Prieta lying to the south. It was either up or down, 
with very little level country for relief. On the floor of the 


ra 


seal ‘f 


camaneo!| 


rT 
EST CHIHUAHUA 


: MULE-TRAIN ALONG STREAM NEAR TECOLOTE. BELOW: W apo: 
tana “PRiputaRY oF Rio pEL SoLpapo. PHoTograpus By H. 8. GEN 
mixed pine forest were many herbs among which were scattered 
plants of the tiny orchid Malaxis Ehrenbergi, with its incon- 
spicuous spike of purplish brown flowers. 

In spite of having to break into the routine of a mule-train 


70 AMERICAN FERN JOURNAL 


we were able to pick up quite a few ferns while traveling over 
Sierra Chinatii during the day. We found Cheilanthes lendigera 
in the crevices of ledges and Pellaca ternifolia, Polypodiwm 
ole A and Cheilanthes notholaenoides on some large, shaded 
boul 

ace ‘slightly saddle-weary, to put it mildly, I, for one, was 
relieved to settle for a short first-day when our jd bletegr brought 
us to camp at Agua Amarillo, a few miles from Chinatti. During 
a quick foray here we found some of the finest colonies of an- 
other orchid, Triphora mexicana, that we had ever seen, as well 
as beautiful plants of Cheilanthes bib era var. arizonica 
protruding from crevices of sheltered ledge 

The next day we traveled to the Aiden of Rio Loera above 
Nabogame, country of the Tarahumare and Tepehuane Indians. 
At noon we stopped for lunch along a small stream at Tecolote 
(Owl). On these north-facing moist ledges and about the base 
of mammoth boulders were, among other species, luxuriant 
colonies of Polypodiuwm subpetiolatum and Adiantum Poireti, 
while on rather dry oak slopes were colonies of the uncommon 
Cheilanthes angustifolia and C. Kaulfussii. Downstream a little 
way, on a most unlikely ledge, were numerous plants of Aspleni- 
um castaneum, whose fronds were rooting near their tips, an 
desires occurrence in this species. 

ogame, we set up a two-day camp on a small promon- 

tory Shee: Indian corn-fields. Through these fields a small 
stream meandered on its way from high mountains in the east 
to larger rivers in the north which eventually feed into Rio 
Fuerte. About the base of boulders along its bank were extensive 
colonies of Athyrium Filix-femina var. asplenioides. The long, 
forested ravines that ran above camp and the tuffaceous bluffs 
that towered above all were rich in ferns. Twenty-one species 
were collected here, notably Asplenium Adiantum-nigrum, A. 
resiliens, A. exiguum, Phanerophlebia auriculata, Woodsia mol- 
lis, Selaginella pallescens, Dryopteris patula var. Rossii and a 
most unusual and rare Asplenium, A. Pringlei. 

Early in the morning of our first day at Nabogame I was on 


Trip TO SreRRA MOHINORA 71 


the tuffaceous cliffs above camp jumping from ledge to ledge in 
search of good sporulating fronds of Elaphoglossum pilosum 
when, unaccountably, I jerked a ligament in my left knee. The 
remainder of our journey was, for me, accompanied by much 
agony. In riding an animal, the continuous knee-rolling motion 
was not only painful but not at all conducive to healing. After 
staying in the saddle for several hours I would literally fall off 
my mount and wobble about on my bad leg. With soft brown 
eyes looking at me our muleteers would solemnly wag their heads 
and say in doleful tones ‘‘muy malo.’’ They didn’t know the 
half of it! Philosophically, I considered that my condition was 
nothing more than the result of an occupational hazard. 

As we sat around the fire that night several of the local in- 
habitants drifted in with some of their handiwork in the nature 
of thickly woven woolen blankets. Most were black or off-white 
in color, the wool having been taken from the animals and thence 
woven directly into blankets. Some of the ‘‘white’’ blankets 
had several irregular cross-lines of orange-brown wool dyed with 
vegetable dye. Although most were as soiled as the animals that 
contributed the wool, Dr. Gentry told me that the old man from 
whom I had purchased a blanket had assured him that it had 
been washed in a mountain stream, apparently at Agua 
Amarillo! 

On the morning of October 12 we broke camp and followed 
the valleys lying east of Chihuite to the base of some high peaks 
that stood between us and Guadalupe y Calvo. As we struggled 
up along the open-wooded trail we saw a beautiful Lobelia with 
large indigo-blue flowers. Growing on boulders along the trail 
were Polypodium guttatum, P. Hartwegianum, and P. thyssano- 
lepis. It was here, as my youngsters would say, that I made a 
real ‘‘boner.’’ Riding above the edge of a steep ravine I could 
see arched and hanging from the opposite bank some fronds of 
a Thelypteris that were fully three feet long. With the thought 
in mind that I would collect specimens when I would undoubt- 
edly see the plant later on, they were passed by. They were not 


AMERICAN FERN 


OURNAL 


A BOVE v GORGE OF TRIBUTARY OF Rfo 1 

STANDING NEXT TO COLONY OF BLECHN 

OF COLONY OF BLECHNUM STOLONIFERUM 
PHOTOGRAPHS BY H. §., 


JEL SOI 
STOLON 


8 


IFER . BEL CLOSE-UP 
HOWN IN SCENE 
TRY. 


DADO, WITH 
F I 


THE AUTHOR 
yw 
( 


ABOV 


> 50, PLATE 


Pa 


d 


Trip TO SrerRA MOHINORA 73 


seen again. This is an all too common occurrence with field 
botanists. 

After a day’s stay in Guadalupe y Calvo, where Dr. Gentry 
and I enjoyed the luxury of a crude shower-bath, and a shave 
by a somewhat inebriated barber, we reassembled our scattered 
equipment and started on the last leg of our trip to reach our 
main objective—Sierra Mohinora. Climbing out of the rugged 
canyon that held Guadalupe y Calvo, we eventually reached a 
mountain stream in a rich coniferous forest. As we jogged along 
on our mounts I had been noting from a little distance large 
plants beneath the trees of what I took to be the common 
Pteridium aquilinum var. pubescens when, by chance, I saw 
standing above some exceptionally bright green fronds obese 
spikes of a Botrychium. Struggling from my horse, I quickly 
gathered some fine specimens of B. cicutarium. Who would have 
expected to find plants two feet tall! 

Keeping a steady grinding pace, we passed through the Indian 
settlements of Zorrillo, Tahonas, and Osera, and late in the after- 
noon came out into a clearing which held the small colony of La 
Rocha, at an elevation of 7,500 feet on the northeast slope of 
Sierra Mohinora. We set up camp in a pine forest on a small 
tributary of Rio del Soldado. Growing in pine needle duff on a 
large boulder at the edge of our camp was an extensive colony of 
the wild potato Solanum polytrichon. 

The next day, October 15, I was in for one of the thrills of a 
lifetime. Following the small stream that ran by our camp I 
soon found myself quite abruptly in a wonderful gorge (PI. 7). 
Although many rich areas had been and were still to be found, 
none were more fascinating than this gorge on the headwaters 
of Rio del Soldado. It faced southwest and was sheltered by a 
dense forest of balsam and pine. The stream, in eutting through 
the rock substrata, had formed flumes and miniature falls from 
which an imperceptible and meliorative mist arose. Sheltered in 
this fern paradise were found 23 species of ferns, 6 of which 
were new to the known flora of Chihuahua. 

This was the kind of place a fern enthusiast usually sees only 


VotuME 50, PLATE 8 


N Fern JOURNAL 


TRIP 1 {It OF SIERRA 


Route OF MULE-TRAIN r0M SAN JUAN TO SI 
MOHINORA AND RETURN, OCTOBER 9-21, 1959. 


Tre to Srerra MonINora 48 


in his dreams. Ferns, ferns everywhere! Dryopteris cinna- 
momea, in multitudinous forms, draped from every cliff and 
ledge, while several species of Polypodium formed large mats 
over great boulders. Following seams beneath dripping ledges 
were delicately green plants of the filmy fern Trichomanes radi- 
cans, while in mud at the base of dripping precipices were 
carpets of Hymenophyllum tunbridgense. Woodsias, Aspleniums 
and Cystopteris fragilis grew intermingled and entwined over 
mossy rocks, and Plagiogyria semicordata and Thelypteris pilosa 
luxuriated on the face of cliffs. Another rare fern, Dryopteris 
parallelogramma, rose in erect clumps alongside masses of 
Athyriums on the edge of the water, while Blechnum stoloni- 
ferum pete a seepage bank and climbed about the base of tree 
trunks (Pl 

After I retuned to camp completely laden with specimens, 


recessed walls behind the plunging water were glistening fronds 
of Adiantum pedatum and a small colony of Polypodium vulgare 
var. columbianum, both new to the State of Chihuahua. On 
nearby ledges were fine plants of Notholaena incana and Chei- 
lanthes farinosa. Woodsias grew like grass on the steep slopes 
below the falls. 

Ever since our leaving San Juan the weather had been un- 
settled and, at times, downright threatening. We had continu- 
ous difficulty drying blotters to keep ahead of our collecting. 
Consequently, most of our presses were in a state of semi-dryness 
when we left our camp at La Rocha and started for the summit 
of Sierra Mohinora. However, since it was cool and, at night, 
even cold, thus providing us a real natural refrigeration, we de- 
cided to simply bale up our ‘‘hay’’ while on the summit and 
wait until our descent to complete the necessary drying. 

With a native of La Rocha as guide we had little difficulty in 
reaching the top. We found that the long plateau which forms 
the summit of Sierra Mohinora is covered with a fine forest of 
pines and on its upper north-facing slopes are dense stands of 


76 AMERICAN FERN JOURNAL 


not only pines but also Douglas Fir and spruce (Picea chihua- 
huana). The ultimate summit, designated by a geodetic survey 
marker, is a grassy plot bordered by stunted weather-beaten 
pines. Here, our altimeter gave a reading of 10,300 feet altitude. 
Finding the herbaceous vegetation disappointingly sparse, we 
concluded that the herds of goats that have summer-grazed the 
region from time immemorial had doubtless exterminated most 
of these plants. Besides, the region had apparently already been 
heavily frosted. 

rom the very summit a breath-taking panorama spread west- 
ward to the Pacific. Range after range rolled across Sonora to 
engulf great barranca after great barranca in a seemingly end- 
less repetition. In the immediate foreground, towers of stone 
fell precipitously to forested canyons below. Threaded across 
the face of these towers were scragely plants of Selaginella Un- 
derwooduvi and in shaded crevices below were delicately fragile 
plants of Woodsia mexicana. 

We pitched camp just below the summit in the protection of a 
stand of pines. As dusk fell, great billowy clouds rolled in from 
the southwest to blanket our camp in an eerie twilight. The glow 
of our fire against massive trunks of the towering pines created 
an imaginary wall within which we slowly moved about. The 
gentle wheezing and munching of our animals in the shadows 
beyond lent an air of unreality to our surroundings. With what 
might be called ‘‘quiet efficiency,’’ we fell to raising a make- 
shift shelter for the night. This was far from a waste of time 
and energy for we had no sooner begun than a sleety drizzle com- 
menced to fall and continued doing so through most of the night. 
Snuggling down in our bags it was not too long before the color 
of our noses matched the red Mexican bandanas we were using 
for night-caps. 

The next day, after thawing out and having collected every 
plant species in sight, we returned to our camp-site at La Rocha. 
On the way down from the summit one of the pack-mules got 
(fide our muleteer) a branch under his tail and away he went! 
The run-away results totaled up to a smashed “kitchen”? with 


Trip TO SrerrRA MOHINORA 77 


the resultant loss of all our coffee (after that the rest didn’t 
count). We did retrieve a few scattered oats which later turned 
out to be mixed with an assortment of pulverized glass. A later 
incident which compensated somewhat for this tragic loss was 
the finding of a small colony of Botrychium Schaffneri on an 
open brushy slope near our camp at La Rocha 

Unfortunately, our head muleteer, who was unaccustomed to 
the rigorous life of a field botanist, had become increasingly 
more disgruntled and uncooperative. No amount of persuasion, 
even ‘‘mas dinero,’’ could convince him to take us from La Rocha 
to San Rafael, on a lower tributary of Rio Mohinora. We had 
been given glowing accounts of this essentially tropical region 
where bananas, coffee and oranges were said to be grown. e 
had visions of a profusion of tropical ferns draping from trees 
and hanging from the walls of cliffs. Here we were only a few 
hours journey away! Since I, the so-called ‘‘fat man,’’ had 
already worn out a horse and was well on the way to wearing out 
a so-called ‘‘muy fuerte’’ mule, perhaps it was just as well we 
went no further ; besides, after our loss from the run-away mule, 
we were running low on grub and my game leg was getting no 
better ! 

As we left La Rocha we noted that the beautifully clear morn- 
ing had given way to a threatening day. e great ridge that 
formed the summit of Sierra Mohinora was already blanketed in 
clouds. It was a losing race with the elements. Within an hour 
a cold drizzle set in which soon turned into a steady chilling rain. 
We felt betrayed, but our spirits remained high. October in 
Chihuahua is supposed to be a month of crystal clear skies with 
weather feigning early spring. We were definitely not prepared 
for the rainy wintry weather we had been encountering. 
these thoughts in mind, we trudged across mountain trails of 
slippery rock faces and open valleys that had become quagmires. 
As we entered Guadalupe y Calvo after dark, we felt thankful 
for the gregarious nature of man whereby he would congregate 
and build centers of habitation, no matter how humble or 
wretched they may be. A change of clothes along with a good 


78 AMERICAN FERN JOURNAL 


mess of hot beans, washed down with unpalatable, but also hot, 
ersatz coffee put a new light on things. 

The next morning, with little ado, we packed and started on 
the last lap of our trip. We were not only still intact, but we 
noted that our ranks had increased by the addition of a lone 
burro, which soon became the whipping-boy for the caravan. 

After a night’s stop at Tecolote (Pl. 8), we traveled on the 
next day to the upper slopes of Sierra Chinatu, within two 
hours of San Juan. The animals were, to put it mildly, com- 
pletely fagged, so we made camp. Searching our meagre 
‘kitchen’? we came up with a handful of rice, along with a few 
scrawny potatoes and two eggs we had purchased from the 
Tarahumares. The eggs helped to thicken the greaseless stew 
which we gratefully consumed amid our thoughts. 

Dr. Gentry and I got an early start the next day, Wednesday, 
October 21, and rode on into San Juan in search of some break- 
fast while the muleteers and Juan searched for a burro that had 
started back home during the night. With about fifteen pounds 
already shed from my frame and little twitches of hunger with- 
in, it is not at all strange that I recall those last two hours on 
the trail as the most fernless encountered during the entire trip. 

Texas ResEarcH FounpaTIoN, RENNER, TEXAS. 


Multicellular and Branched Hairs on the 
Auma G. Stoxey! 

Hairs are a common feature on the sporophyte of ferns and 
are usually present on young parts if not on the mature. Bower 
1923) made a sharp distinction between simple hairs and multi- 
cellular scales as indicators of phylogeny. As might be expected, 
hairs are not as highly developed or as numerous on the gameto- 
phyte. It remains to be seen if the types and distribution of 

hairs will be of any assistance in questions of phylogeny. 


1 This account is based on studies of cultures hig have been main- 
tained in connection with the investigations carried on in mi gusmaen with 
Dr. Lenette R. Atkinson for the past ten years, ana" my own earlier 
cultures Q 


Hairs or Fern GAMETOPHYTES 79 


There are many families, especially among the primitive ferns, 
in which hairs are entirely lacking on the gametophyte. They 
have not been found in the Marattiaceae, Osmundaceae, Hymeno- 
phyllaceae, Matoniaceae, Dipteridaceae, Pipa aera Cheiro- 

ittar e 


of the gametophyte.) The most primitive families with gameto- 
phyte hairs are the Gleicheniaceae and Schizaeaceae; they are 
present, also, in the somewhat more advanced Cyatheaceae and 
Loxsomaceae. 

In the Gleicheniaceae the hairs are never numerous, but have 
been described for several species (Campbell, 1908, Stokey, 
1952). They are usually two-celled, less frequently three- or 
four-celled, and borne on the dorsal or ventral surface of the 
midrib. In the Gleicheniaceae, and also in the Cyatheaceae and 
Loxsomaceae, the hair arises from a special initial cell, a wedge- 
shaped cell on the anterior face of a young superficial cell near 
the apex of the thallus (Figs. 6,7). Growth at this stage is rapid 
and the young hair soon projects above the surface, forms two 
cells, rarely more, and then curves towards the apex. A section 
of thallus of Hicriopteris glauca (Thunb.) Ching with a young 
hair is shown in Fig. 3, and an older hair of Gleichenia vulcanica 
Blume in Fig. 2. The terminal cell elongates, becomes slightly 
bulbous and filled with a heavily-staining substance, and 
protoplasm disappears; the same process follows later in the 
basal cell 

In the Schizaeaceae the hair arises as a simple papilla on a 
marginal cell, less frequently on the surface (Bauke, 1878). 
hee mature the hair is usually two-celled and curved towards 
the The hair shown in Fig. 1 was found on a thallus of 
Aaonie  phyllitidis (L.) awaits only 46 days old. Hairs are not 
abundant in this family, and on Lygodium palmatum (Bernh.) 
Swartz they were found on only two prothalli (Rogers, 1923) ; 
they agreed in type with those of Anemia and Mohria, 

In the Cyatheaceae and Loxsomaceae, the multicellular hairs 
arise not only from a special initial but are of a special scale-like 


rf x 5 
AMERICAN FERN JOURNAL VoLuME 50, PLATE 9 


eg ere rae HAIRS. Fig. 1, ANEMIA PHYLLITIDIS ; 2, GLEICHENIA eh sce 
3, HICRIOPTERIS GLAUCA; fi LOXSOMOPSIS COSTARICENSIS; 5-7, ALSOPHILA 
EX scat rc ‘ OPTERIS TENELLA; 9, 10, CYCLOPELTIS eeaauas , 
NEPHROLEPIS CORDIFOLIA; 12, N. ACUMINATA ; 13, OLEANDRA hoger 14, 
CYLOPELTIS PRESLIANA, Fias, 1-2, 6-14, & 120; Fas. 4; 5; 0. 


Hairs or FERN GAMETOPHYTES 81 


type (Bauke, 1876; Goebel, 1912; Stokey, 1930; Stokey & Atkin- 
son, 1956). These hairs are one to several cells wide, many cells 
long, and usually have a tapering slender tip, as in Loxsomopsis 
costaricensis Christ (Fig. 4) and Alsophila excelsa R. Brown 
(Cyathea Brown Domin) (Fig. 5), in which the hairs are 
often much longer. Bauke described them in his account of the 
Cyatheaceae as ‘‘Borsten.’’ They arise, usually after arche- 
gonium production has begun, on or near the cushion on both 
dorsal and ventral surfaces, curving towards the notch and 
making a green brush-like growth large enough to be seen with 
the naked eye. 

In the families formed in recent years by the breaking up of 
the old comprehensive family Polypodiaceae, hairs, unicellular, 
multicellular, and branched, are more abundant and varied than 
in the four families discussed. It is convenient to consider them 
on the basis of Copeland’s classification (1947), with the addi- 
tion of the Grammitidaceae which he recognized later (1951). It 
is in these higher families that the familiar type of papillate 
hair, unicellular and usually glandular, has developed in pro- 
fusion. The multicellular hair, even when two-celled, is much 
less common and seldom present in abundance. A two-celled 
hair may properly be considered a multicellular hair; it seldom 
appears as a chance modification of a one-celled hair, and its 
variants are apt to be three- or four-celled. 

In the Davalliaceae multicellular and even branched hairs are 
found occasionally. On the thallus of Davallia denticulata 
(Burm.) Mett. a few two-celled and branched hairs were found. 
On the gametophyte of Oleandra the abundant long unicellular 
hairs are often borne on extensions of marginal cells and sug- 
gest two-celled hairs; examples of true two-celled hairs were 
found in O. wallichii (Hook.) Presl (Fig. 13). In Nephrolepis, 
multicellular hairs were more abundant, especially on the thallus 
of N. cordifolia (L.) Presl, but less numerous and shorter on N. 
acuminata (Houtt.) Kuhn (Fig. 12). On Humata heterophylla 
(J. EB. Smith) Desv., a marginal three-celled branched hair was 
found near the notch. On the thallus of Arthropteris orientalis 


AMERICAN FERN JOURNAL VOLUME 50, PLATE 10 


29 
PROTHALLIAL HAIRS. Fig. 15, PLATYCERIUM ALCIOORNE ; 16, P. GRANDE; 17, 
OGRAMM T 
22, C 523, 2 2 
PYRROSIA CHINENSIS; 25, PESSOPTERIS CRASSIFOLIA; 26, THELYPTERIS BIOL- 
CTE 


LEYI; 27, XIPHOPTERIS DELITESCENS; \ NOPTERIS JUBIFORME; 29. C. 
ASPLENIFOLIA, ALL X 120 


Hatrs or Fern GAMETOPHYTES 83 


(Gmel.) C. Chr. several examples of three-celled simple hairs 
were found on the ventral surface near the archegonia. On that 
of A. macrocarpa (Cordem.) C. Chr. there were three-celled 
simple hairs on the dorsal and ventral surfaces, and branched 
hairs with as many as six cells appressed to the ventral surface. 
Arthropteris tenella (Forst.) J. Smith had more hairs, all ap- 
pressed to the margin and curved towards the apex; some were 
simple and others elaborately branched (Fig. 8). 

n the large and diverse assemblage of the Aspidiaceae, it is 
not unusual to find species in which multicellular hairs are borne 
on the gametophyte. On the prothallus of Cyclopeltis crenata 
(Fée) C. Chr. two-celled hairs were found on the surface but 
more abundantly on the margin, often with a glandular tip 
which gave a reaction for wax with Sudan IV (Figs. 9, 10); a 
few branched hairs were found on the surface. The same type 
of two-celled hair was found in large numbers on the thallus of 
C. presliana (J. Smith) Copel. even two or three on a single 
marginal cell. On the thallus of Bolbitus quoyana when eight 
weeks old there appeared on the margin two-celled hairs curving 
towards the apex (Fig. 18); at three months they were present 
on the margin in a close growth of curved colorless hairs of two 
or three cells with the terminal sometimes inflated; a few of the 
hairs were branched. Later branched hairs appeared on the 
dorsal surface. On the prothallus of Phanerophlebia caryotidea 
(Wall.) Copel. and on that of Rumohra aristata (Forst.) Ching 
a few short two-celled hairs were found. Pteridrys australis 
Ching had a liberal development of branched hairs with four to 
eight cells on both dorsal and ventral surfaces of the midrib. 
Two-celled hairs are not uncommon on the surface and margin 
of the gametophyte of Pleocnemia conjugata (Blume) Presl; 
branched hairs are relatively rare. he five species of Tectaria 
in our cultures showed multicellular or branched hairs or both: 
T. incisa Cav., T. decurrens (Presl) Copel. T. irregularis 
(Presl) Copel., 7. griffithi (Baker) ©. Chr., from Nepal, and 
T. subtriphylla (H. & A.) Copel. The hairs were mostly on the 
surface, rarely on the margin, usually three-celled but sometimes 


84 AMERICAN FERN JOURNAL 


four- or five-celled. On a rather sparse culture of Heterogonium 
pinnatum similar branched hairs were found on several gameto- 
phytes. Cyclosorus is unusual in having acicular hairs on the 
gametophyte of several species usually one-celled but in the case 
of C. parasiticus occasionally two-celled (Fig. 14). Thelypteris 
(Goniopteris) biolleyi (Christ) Proctor has an unusual type of 
hair—branched stellate hairs similar to those found on the 
sporophyte, but there were no indications of apogamy. They 
appeared rather sparingly at three months on the ventral sur- 
face before archegonia had developed; they were present in great 
numbers on both margin and surface of cushion and wings when 
the cultures were nine months old, and archegonia were abundant 
(Fig. 26). 

In the Aspleniaceae there is a considerable range in regard to 
hairs on the gametophyte, as was pointed out by Wagner (1953). 
A type of multicellular hair which ends in a gland was found on 
Diellia gametophytes, and also on those of Ceterach dalhousiae 
(Wagner, 1952); he found the same type on the thallus of 
Asplenium leucostegioides Baker (1953). The same type is pres- 
ent also on the prothallus of C. officinarum with considerable 
variation in length (Figs. 21, 22). It occurs also on the gameto- 
phyte of A. flabelliforme Cav., appearing when the prothalli are 
three to four months old, on the margin near the notch both 
before and after the production of archegonia. On some gameto- 
phytes of the same age and on some slightly older, apogamous 
apices outgrowths associated with clathrate scales ap- 

ed. On the prothallus of A. septentrionale (.) Hoffm 
among the many long unicellular hairs there are occasionally 
two-celled and even branched hairs. The prothallus of a tetra- 
ploid Phyllitis scolopendrium var. americana Fernald (spores 
from a Michigan plant) when five months old had a considerable 
number of two- and occasionally three-celled hairs, straight or 
slightly curved, on both dorsal and ventral surfaces, as well as 
on the margin near the apex. There were only one-celled hairs in 
a culture of Phyllitis scolopendrium, presumably diploid, raised 
from spores collected in Miirren, Switzerland. The gametophyte 


Hairs or Fern GAMETOPHYTES 85 


of Pleurosorus rutifolius (R. Brown) Fée when less than four 
months old bore a considerable number of long two-celled hairs 
among and in front of the archegonia (Figs. 19, 20). 

The description and figures by Klein (1881) of the prothallus 
of Polypodium heracleum Kunze |Drynariopsis heraclea 
(Kunze) Ching] is the classic account of branched hairs on a 
fern gametophyte. Recent work indicates that the Polypodiaceae 
sensu stricto is the group in which branched hairs appear on the 
gametophyte of the largest number of species, although the range 
in type is not known to be as great as in some other groups. 
Multicellular simple hairs are not unusual in young cultures 
which later produce branched hairs. The most common type of 
branched hair is usually three-celled, such as that of Paltonium 
lanceolatum (l.) Presl (Fig. 23), and that of Pyrrosia lingua 
(Thunb.) Farw. (Fig. 24), with the four- or five-celled, as in 
Pessopteris crassifolia (L.) Underw. & Maxon (Fig. 25) less fre- 
quent. This type of hair is usually found on the surface, on or 
near the midrib and rarely on the margin. 

There is much variation in the age of thallus at which 
branched hairs appear, but it is regularly later than that for 
simple unicellar hairs. Pleopeltis hastata (Thunb. ) Moore began 
the production of branched hairs at 75 days; Pessopteris had 
branched hairs with seven cells at three months; Pyrrosia lingua 
had branched hairs with three to six cells at 10 weeks. In Phle- 
bodium aureum (L.) J. Smith, Belvisia spicata (L.) Mirb., and 
Campyloneuron phyllitidis (L.) Presl branched hairs were late 
in developing with some variation in different sets of cultures. 
They appeared sparingly in a culture of P. virginianum L. at 
seven to nine months. They were found on the following species 
of Polypodium at varying times: P. chnoodes Spreng., P. pec- 
tinatum L., P. plebejum Schlecht. & Cham., P. repens Aubl., and 
P. vexatum D. C. Eaton. They appeared also on Aglaomorpha 
meyeniana Schott, Microsorium scolopendria (Burm.) Copel., 
and M. punctatum (L.) Copel.; in addition to branched hairs, 
which appeared at six months, they also bore clathrate scales. 
Nayar (1957) has reported branched hairs on the prothallus of 


86 AMERICAN FERN JOURNAL 


Drymoglossum piloselloides (:) Presl. The large branched hairs 
on the thallus of Platycerium (Straszewski, 1915; Stokey & At- 
kinson, 1954) are similar to the branched hairs described for 
other members of the family except that they are unusually large 
and abundant; the stalk cells are green and the tips of the 
branches glandular (Pigs. 15, 1 

In Loxogramme, a genus whose systematic position is under 
question (Copeland, 1947; Holttum, 1949), multicellular hairs 
have appeared in two species. On the prothallus of L. parksti 
Copel. two-celled marginal hairs appeared sparingly at nine 

ths; when two to six years old several two- or three-celled 
hairs could always be found towards the tip of a lobe or branch 
of the thallus usually on the margin (Fig. 17) but also on the 
surface. The same type of two-celled hair appeared sparingly. 
on L. avenia Presl at 13 months. 

In the Grammitidaceae some species have a heavy growth of 
branched hairs with an occasional two-celled hair along the mar- 
gin (Stokey & Atkinson, 1958). Some of the hairs have a slender 
spine-like branch, as in Xiphopteris delitescens (Maxon) Copel. 
(Fig. 27), or even two such branches as in Ctenopteris jubiformis 
(KIf.) J. Smith (Fig. 28) ; or all branches may have a glandular 
cell at the tip as in C. suspensa (L.) Copel. (Fig. 29). The hairs 
of the Grammitidaceae differ from those of the Polypodiaceae s.s. 
in arising regularly on the margin rather than on the surface. 

The significance of multicellular and branched hairs on the 
fern gametophyte can hardly be considered apart from the larger 
question which would include the distribution and type of uni- 
cellular hairs. We need much more information about the occur- 
rence and distribution of all types of hairs on both young and 
old gametophytes. 

LITERATURE CITED 
Bavuke, H. 1876. Entwickelungsgeschichte des Prothalliens bei den Cyathe- 

aceen. Pringsh. Jahrb. f. Wiss. Bot. 10: 49-116. 

—. 1878. Zur Keimungsgeschichte der Schizaeaceen. Prings. Jahrb. f. Wiss. 

Bot. 11: 603-650. 

Bower, F. O. 1923, The Ferns. Vol. I. Cambridge. 


Dryopteris Hyprips 87 


CAMPBELL, D. H. 1908. sa prothallium of Kaulfussia and Gleichenia, Ann. 
Jard. 102 


. 1951. i ‘ (2) 2°97) 
—. 1955. Ctenopteris in America. Phil. Journ. Sei. 84: 475. 
GorBeEL, K. 1912. Archegoniatenstudien XIV. Loxsoma in das System der 
Farne. Flora 105: 33-52. 
Hourrum, R. E. 1949, The Classification of Ferns. Biol. Rev. 24: 
Kuen, L. 1881. Bau und Verzweigung einiger romarantie basalt 
Akad. Naturf. 42: 333-396. 


Nova Acta K. Leop-Carol. Deutsch. Akad. Natu 33- 
Nay B. K. 1957. Studies in the Polypodiaceae. IV. Drymoglossum 
pi Journ. Indian Bot. Soe : 169-179. 
Rogers, L. M. 1923. pipe o the prothallium of Lygodium palmatum. 
75: 75-84 
ecient BS 30. The abe! of the Cyatheaceae. Bot. Gaz. 90: 1-45. 


52. The gametophyte of the Gleicheniaceae. Bull. Torrey Cl. 77: 323— 


Sroxry, A. G. & ATKINSON, L. R. 1954. The gametophyte of five species of 
eid oortine Phytomorphology 4: 165-172 
56. The gametophyte of Loxsoma Cankinghanitt R. Br. and Lox- 
: Lae costaricensis Christ. Phytomorphology 6: 249-261. 
—. 1958. The gametophyte of the Grammitidaceae, Phytomorphology 8: 


391-403. 

STRASZEWSKI, H. von. 1915. Die Farngattung Platycerium. Flora 108: 271- 
310. 

WAGNER, ch H. Jr. 1952. The fern genus Diellia. Univ. Calif. Publ. Bot. 


ae 1053. ie Asplenium prototype of the genus Diellia, Bull. Torrey Cl. 
: 76-94. 
ane Hoiyoke CoLuEcE, SourH HapLey, MASSACHUSETTS. 
Progress in ag eg of eee Hybrids 
AR T. WHER 

n the course of ee for Beene hybrids to send to 
Dr. Stanley Walker for the cytotaxonomic studies recently re- 
ported in this Journau,! I felt a need for a diagram bringing 
out their inter-relationships. Now that data as to chromosome 
numbers can be added, it has seemed worth while to publish such 
a diagram. Before it can be presented, however, some considera- 


s JOURNAL . 1959 
2 Mieonchote oe ele “ eed ‘Nomenclature, Art. Hi 1.1956, 


88 AMERICAN F'ERN JOURNAL 


tion of nomenclatorial matters is necessary. 

For simplicity all parental taxa are here treated as species and 
designated by binomials, Dryopteris being abbreviated to ‘‘D.’’ 
and epithets uniformly decapitalized. According to standard 
procedure? the taxa forming hybrids are placed in alphabetical 
order, with the symbol between; and when a hybrid has re- 
ceived an individual epithet, this is preceded by that symbol. 
Only basionyms—that is, name-bringing synonyms—are cited; 
additional synonyms can be found if desired in indexes. 


mar .2 x spi 4 


spe4 €X lee.2 X uli.4 


gol. 


DIAGRAM OF DRYOPTERIS SPECIES ms Hysrips. 
EXPLANATION IN TExT ABO 


The diagram represents the five most widespread species in 
the northeastern United States—D. cristata, D. goldiana, D. 
intermedia, D. marginalis, and D. spinulosa, and the ten hybrids 
between them. Their epithets are abbreviated to three letters 
each. The ploidy is represented by numerals, placed in paren- 


Dryopreris Hysrips 89 


theses when inferred though not yet confirmed. These hybrids 
are as follows 

D. crisrara X GoLDIANA. Reported by Benedict® in 1909. It 
was inferred to be the same as D. atropalustris Small* by the 
writer® in 1942, but this is now realized to have been a mistake. 
As pointed out by Walker, D. clintoniana presumably arose from 
a clone of this inferentially triploid hybrid through chromosome- 
doubling. 

D. cristata X INTERMEDIA = D. X Boorri. This taxon, pub- 
lished as a species of Aspidium by Tuckerman? in 1843, was sug- 
gested to be a hybrid by several subsequent workers, and formal- 
ly so treated by Dowell’ in 1908 

D. cRISTATA X MARGINALIS = D. X SLOSSONAE. The first hy- 
brid to be recognized as such in this country, by Davenport® in 
1894. It was named as a species of Nephrodium by Hahne® in 
1904, and transferred to Dryopteris by the writer?? in 1942. 

D. cRISTATA X SPINULOSA = D. X ULIGINOSA. Discovered in 
Europe, this was named Aspidium spinulosum var. wliginosum 
A. Braun,!! and later Lastrea uliginosa by Newman n!? in 1849. 
Its hybrid nature was pointed out by Milde!* in 1858, and the 
epithet was transferred to Dryopteris by Druce.'* 

D. GouDIANA X INTERMEDIA. Reported by Dowell!® in 1908. 
Guessed to be D. separabilis Small!® by the writer’ in 1942, but 
as pointed out by Walker such is not the case. 


aoe Torrey Club 36: 47. 1909. 
s 


16 Ferns SE. States 284. 193 
17 Guide East. Ferns, ed. 2, 161. 1942. 


90 AMERICAN FERN JOURNAL 


D. GOLDIANA X MARGINALIS = D. X LEEDSII. This hybrid was 
reported by Dowell!® in 1908 and assigned the epithet leedsu by 
the writer!? in 1942. The colony along the Susquehanna River 
yielding the specimen on which this was based has been reduced 
by road widening, but young plants are still appearing. These 
are more numerous than would be expected to be produced by a 
hybrid, but an explanation has now been found. Study by Mrs. 
C. W. Crane showed that many of the plants there are a normal- 
spored species; and as noted by Dr. Walker, this is a tetraploid, 
evidently resulting from chromosome-doubling in a clone of the 
diploid hybrid. 

D. GotpIANA & sPINULOsA = D. X poyseri, nom. nov. This 
was discovered by Poyser?® and assigned the cumbersome name 
Nephrodium cristatum clintonianum £. silvaticwm in 1908; it was 
indicated to be the present hybrid by Benedict?! the following 
year. It is here assigned an epithet in honor of William Ald- 
worth Poyser (1882-1928), noted Philadelphia fern student. 
The type specimen, from near Swarthmore, Pennsylvania, is pre- 
served in the Academy of Natural Sciences, Philadelphia. 

D. INTERMEDIA X MARGINALIS. Reported by Benedict*? in 
1909. 

Dryopteris triploidea Wherry, hyb. nov. (D. intermedia X 
ae 


e quam parentes majores, aspect i D. spinulosae 
dinties, i.e. pinnis ascendentibus, te inferionibus basalibus 
longioribus quam sequentibus ; rhachis ursum, venae, et indusia 


glandulas capitatas  ferentia ; sierie feiperfaetae planta 
triploi dea 

TYPE: Clayy ille, New York, collected by B. D. Gilbert, in the 
Gilbert Herbarium, Harvard University. 

The tradition that D. intermedia and D. spinulosa grade into 
one another and are therefore only varietally distinct is mani- 
festly based on the frequent occurrence of this hybrid, which 

18 Bull, Torrey oy 85: 139. 1908. 

19 apa a1: 

20 Fern Bull. 16: 13. 1908. 

21 Bull. Tors Club 36: 47, 1909. 

22 Tdem 


Dryopreris Hysrips 91 


has commonly been known as D. spinulosa var. fructuosa. It was 
first noted in 1900 by B. D. Gilbert,?* who identified his speci- 
mens with the European Lastrea dilatata glandulosa T. Moore, 
which he renamed Dryopteris spinulosa glandulosa. The next 
year he renamed the plant Nephrodium spinulosum fructuosum™ ; 
although the plant intended by Gilbert is the American hybrid, 
the name is technically based on Lastrea dilatata glandulosa 
Moore, which is stated unequivocally as a synonym. This English 
plant must presumably be different, since one of the parents of 
the American hybrid, D. intermedia, does not grow in England. 
Therefore, the American plant is in need of a new name. In any 
case, the epithet fructwosum is not available as a specific epithet 
under Dryopteris as the name is preoccupied. The new epithet 
here proposed refers to the interesting finding by Manton and 
Walker2® that this hybrid, as is predictable by the first-named 
parent being diploid and the second tetraploid, is indeed triploid. 
It should be noted that most specimens in herbaria labelled 
‘<fructuosa’’ are merely luxuriant intermedia. 

D. MARGINALIS X SPINULOSA = D. X PITTSFORDENSIS. First 
published as a species by Slosson,?* and four years later inter- 
preted by her? as this hybrid. 

The foregoing diagram shows that order is at last appearing in 
the heretofore rather confused picture of inter-relationships in 
Dryopteris in the northeastern United States. No longer is there 
any excuse for shifting epithets around from one status to an- 
other under a multiplicity of species. Thanks to the observations 
of spores by Mrs. Crane and of chromosomes by Dr. Walker, 
hybrids can be recognized as such and their parentage estab- 
lished. Further progress can be made if members of the Ameri- 
ean Fern Society will keep on the lookout for additional ma- 
terial, especially of those hybrids which, as indicated in the 
above diagram, have not as yet been available for cytologic study. 


23 Fern Bull. 8: 11. 1900. 

24 List N. Amer. Pterid. 37. 1901. 
25 Nature 171: 1116. 1953. 

26 Rhodora 6: 75. 1904. 

27 Fern Bull. 16: 99. 1908. 


92 AMERICAN FERN JOURNAL 


Special thanks are due to Mrs. C. W. Crane, without whose 
generous aid in checking the spores of specimens of the hybrids 
discussed the preparation of this article would not have been 
possible. 


Abnormal Nuclear Division in Fern Prothallia 
m J. Crotty 
The purpose of this paper! is to report the occurrence of ab- 
normal nuclear divisions in the prothallia of three genera of 


L., the ladder-brake fern, Matteuccia Struthiopteris (.) To- 
saeh: the ostrich fern, and Dryopteris sp. Amitosis is usually 
defined as the simple constriction of the nucleus without the 
formation of condensed chromosomes or spindles; the term nu- 
clear fragmentation is often used when more than two nuclei are 
so formed (5). Since amitosis or direct division has been a sub- 
ject of recurring interest since nuclear division was first de- 
scribed, and, since the controversy over the significance of this 
process is still not satisfactorily settled, we feel it worthwhile 
to record these observations and suggestions for the use of the 
fern prothallia in the exploration of this problem. After describ- 
ing our observations, we shall refer briefly to possible environ- 
mental conditions that might have stimulated the abnormal 
divisons. 

The observations reported here were made on both living and 
fixed (in Navashin’s or Carnoy’s fixative) prothallia which were 
in either the early filamentous or later plate-like growth stages. 
They were stained with a pyronin-methyl green mixture. In no 
case was a single nucleus actually followed through the division 
described here; rather, the sequence was pieced together from the 
study of many isolated stages, a few of which are pictured in the 
figures. Since reports of amitosis have been regarded by some 

1 This baad id etagaer was aided by gpeate grants from the National Sci- 

indation. Acknowledgments are gratefully made to Miss Nancy 


ne 8 
Coddington for Neer el assistance, and to Dr. Norman Marengo for the 
ores of the ostrich fer 


ABNORMAL NucLEAR DIVISION 93 


investigators as due to the fusion rather than to the constriction 
and separation of nuclei, anyone employing an indirect method, 
such as used here, must consider the possibility that he is read- 
ing the actual sequence of events backwards. In the discussion 
we shall consider why we favor amitosis as the interpretation 
for these observations. 

A thallus cell is normally uninucleate unless the first nuclear 
division attendant upon differentiation into a rhizoid, antheridi- 
um or archegonium has occurred and the new cell wall has not 
yet formed. However, according to our experience acquired dur- 


1 2 


CO @ 


> 


S 


ww 
> 


Fia. 1, CONSTRICTING NUCLEI SEEN AT ONE OPTICAL PLANE; 2, SAME NUCLEI 
AT DIFFERENT PLANE SHOWING CONTINUITY BETWEEN THEM; 3, INTERPRETA- 
M 


THREE NUCLEI OF UNEQUAL SIZE; 5, NORMAL NUCLEUS; 
ELONGATE NUCLEUS SHOWING INCIPIENT CONSTRICTION LINE; 
NUCLEI; 8, BINUCLEATE CONDITION 


7, SEPARATING 


ing a cytochemical study (soon to be reported), we can detect 
such differentiation even before nuclear division has occurred, 
and certainly very shortly thereafter. Our interest was aroused, 
therefore, when we noticed a binucleate thallus cell for the first 
time several months ago. Thereafter, during a period of about 
five months, we found frequent binucleate and multinucleate 


AMERICAN FERN JOURNAL 


Fias. 9, 10, OSTRICH FERN, CONSTRICTING NUCLEI; Figs. 11-16, Prrris 

VITTATA NUCLEI DIVIDING AMITOTICALLY; Fig. 14, THREE NUCLEI FORMED; 

Fig. 15, FRAGMENTED NUCLEI, SOME VESCICULAR; Fiqa. 16, FRAGMENTED 
NUCLEI 


ABNORMAL NuciEAR Division 95 


cells in the prothallia germinated from several sowings of Pteris 
vittata spores and in one sowing each of Matteuccia and Dryop- 
teris. No abnormal divisions have been seen in prothallia germi- 
nated in the last three or four months. A common abnormality 
resembled that sketched in figure 1. At one optical level (using 
apochromat oil immersion objectives) we saw what looked like 
two nuclei pressing against one another. At a different optical 
plane, however, it was apparent that the two nuclei were con- 
tinuous (figure 2). Figure3 is our interpretation of what figure 
1 would look like when viewed from the side instead of above and 
is similar to figures reported in the literature (4). Prior to these 
observations, we had assumed that the binucleate condition was 
the result of a regular mitosis. The finding of more and more of 
these closely associated nuclei, some of which were of unequal 
size (figure 4), indicated the possibility that amitotie divisions 
might be occurring. We therefore looked carefully for stages 
that might indicate either normal or aberrant mitosis (e.g. abor- 
tive spindle formation) or true amitosis. Even though spindle 
components are sensitive to the pyronin stain, we found no evi- 
dence of spindle formation. On the other hand, the occurrence 
of enlarged (figure 6) and constricted nuclei (figures 1, 9, 10, 
11, 13) were very common, especially in cells adjacent to bi- and 
multinucleate ones. Incipient constrictions or plate-like in- 
growths were fairly common is these elongated nuclei (figure 6) 
but they proved difficult to photograph satisfactorily. In multi- 
nucleate cells (figures 4, 14, 15, 16), the number of nuclei ob- 
served ranged from three to eight. Some of these multiple nuclei 
were little more than empty vesicles (figure 15) containing a 
pyronin-staining body (presumably a nucleolus) and a lightly- 
stained nuclear membrane. The conclusions that we have 
reached, therefore, are that these abnormal divisions are amitotie 
and that the sequence of events leading to the binucleate cell is 
illustrated by the following figures in the order given: Figures 
5, 6,7 (or 3) and 8 and 9, 10, 11 (or 13) and 12. 

These observations had never been made by us before this 
period even though we had looked at literally thousands of 


96 AMERICAN FERN JOURNAL 


prothallia with special attention focused on the nuclei of the 
cells. Nor as indicated above, have we found such abnormal 
divisions in sowings of the past three or four months. Since we 
used the same collection of spores of Pteris before, during, and 
after the period of these observations, it seems pertinent to con- 
sider possible environmental conditions that might have stimu- 
lated such divisions. The spores are sown in standardized 
amounts in 10 or 20 ml. of mineral nutrient media, in 50 ml. 
erlenmeyer flasks; some of the spores were disinfected with a 
chlorox solution, and others washed only with sterile water. The 
flasks are placed in a refrigerator-incubator under known con- 
ditions of light (65-85 footcandles from incandescent lamps) and 
temperature (25°C) for a given length of time (one, two weeks, 
ete.). After using some of the prothallia for specific experimen- 
tal work, the remaining material in flasks was then shifted to 
other uncontrolled and saci light intensities to make room 
for new sowings. During the period of these observations, we 
experienced trouble ai the temperature-regulating equipment 
so that the temperature was noted to vary from 18° to 30°C on 
a few occasions. All the prothallia with abnormal divisions 
showed evidence of some microbial contamination by the time 
the observations were made (or the material was fixed) since 
little attempt was made to maintain aseptic conditions in the 
original flasks beyond the time the prothallia were removed for 
other experimental purposes. Although the possible environ- 
mental variables listed above are numerous, we feel that it would 
be worthwhile to attempt to define the conditions that stimulate 
these abnormal divisions and we hope that others will also try to 
discover such conditions. Changes in temperature and light in- 
tensity seem to be possible factors. If infection is a factor, this 
might prove more difficult to reproduce. The fern prothallium 
strikes us as an ideal organism for studying such abnormal di- 
visions in vivo but the first step will be to learn how to induce 
them. 

There is a striking similarity between what we have observed 
in these fern prothallia and the pictures reported by Cutter, 


ABNORMAL NUCLEAR DIVISION 97 


Wilson, and Freeman (1) as amitotie divisions in the develop- 
ing endosperm of coconut. That amitosis does occur in both ani- 
mal and plant cells has ey been shown but there seems to 
be some controversy about: (1) whether specific reported in- 
stances are actually cases of actos or are really cases of fusing 
nuclei (which have previously been formed by normal mitosis) ; 
(2) whether or not amitosis is to be regarded as a degenerate, 
pathological condition; and (3) whether or not it is an unusual 
but ‘‘normal’’ situation which could be followed by normal 
mitosis (2, 3, 4, 5). Some discussion of our reasons for reading 
the sequence of stages as we do seems appropriate in view of the 
first of these questions. We can say nothing with assurance 
about the second and third of these controversies. 

Essentially, our arguments for reading the sequence as we do 
are as follows: (1) Thallus cells are normally uninucleate. If 
amitosis has not occurred, we would have to postulate that nor- 
mal mitosis (or mitoses in the case of multinucleate cells) had 
oceurred and that the daughter nuclei had, or were in process 
of fusing. This in itself, seems more unusual and abnormal than 
amitosis. (2) In order to explain the appearance of three, un- 
equal-sized nuclei in a cell (figure 4) by normal mitosis, we 
would have to assume either an unequal mitotic distribution of 
nuclear material, or else an unequal swelling of two nuclei with 
respect to the third, smaller one. Both of these alternatives seem 
much less probable than the amitotic explanation. (3) It is 
more difficult to explain the attitude of nuclei such as those 
shown in figures 3, 7 and 13 as figures of fusion than as figures 
of constriction. One does not expe ect to see merging nuclei 
fusing in such a local manner as in figure 13. (4) The occur- 
rence of many apparently empty nuclei in the multinucleate 
cells is more easily explained by amitosis than by normal mitosis. 

BroLocy DEPARTMENT, WASHINGTON SQuaRE COLLEGE, NEw 
York UNIVERSITY. 

REFERENCES CITED 
(1) Currer, V. M., Witson, K. S., and B, FREEMAN. 1955. Nuclear be- 
havior and cell eden in the eats endosperm of Cocos nucifera. 

Amer. Journ. Bot. 42: 109-115. ° 


98 AMERICAN FERN JOURNAL 


(2) Karer, J. M. 1940. Amitosis I. Bot. Rev. 6: 164-180. 
(3) —. 1951. Amitosis II. Bot. Rev. 17: 105-108. 
(4) Suarp, L. W. 1934. Introduction to Cytology. McGraw-Hill. 
(5) Witson, E. B. 1928. The Cell in Development and Heredity. Mac- 
millan Co. 
On the Species of Dryopteris, — ch hes tyaly 
Motozi Tagawa AND Kun WAT 
In 1855, Thomas Moore described is genus Saar 
which was characterized by having a hard coriaceous texture 
and the peculiarity of imparipinnate fronds with a free termi- 
nal pinna like the lateral ones. Pycnopteris was subsequently 
referred to the genera Aspidium, Lastrea, or Nephrodium, and 
finally recognized by Ching as a subgenus of Dryopteris. Ching 
also described a monotypic genus Microchlaena, which seems to 
the writers not particularly different from Pycnopteris. The 
seven eee listed below may be referred to the subg. 
Pycnopt 
Fcdleniia aha Pycnopreris (Moore) Ching, Bull. Fan Mem. 
Inst. Biol. 8: 371. 1938. 
Pycnopteris Moore, Gard. Chron. 1855: 468. 1855. Type: 
Aspidium sieboldii van Houtte. 
py oe Ching, Bull. Fan Mem. Inst. Biol. 8: 325. 1938. 
ype: dium yunnanense Christ. 
if Besa SIEBOLDII (van Houtte) Kuntze, Rev. Gen. Plant. 
2: 813. 1891; Ogata, Icon. Fil. Jap. 3: t. 127. 1930; Ching, 
Bull. Fan ree Inst. Biol. 8: 396. 1988; H. Ito, in Nakai & 
Honda, Nov. Fl. Jap. Polypod. Dryopt. 1: 25, cum t. et fig. 
1939; Ohwi, Fl. Jap. Pterid. 83, t. 41. 1957; Tagawa, Col. 
Ill. Jap. Pterid. 92, 218, fig. 182. 1959. 
Aspidium sieboldii van Houtte ex Mett. Fil. Hort. Lips. 87, t. 
20, fig. 1-4. 1856. 
Pycnopteris sieboldii Moore, Gard. Chron. 1855: 468, eum fig. 


Nephrodium sieboldii Hook. Fil. Exot. t. 31. 1859; Sp. Fil. 4: 
87. 1862. 


This species was described from cultivated plants introduced 


DryopTeris, SUBGENUS PYCNOPTERIS a9 


from Japan to Europe by Siebold. As will be noted further on, 

the identification of the Chinese plants referred to D. sieboldii 

by Ching is dubious. The species is mainly restricted to south- 

western Japan, where it is common in Kyushu; it occurs spo- 

radically also in warm-temperate regions of Honshu and 
hikoku. 

2. DRYOPTERIS TOYAMAE Tagawa, Act. Phytotax. Geobot. 8: 167. 
1939; Ohwi, Fl. Jap. Pterid. 84. 1957; Tagawa, Col. Til. Jap. 
Pterid. 93, 214. 1959. 

Type: A plant cultivated in the Botanical Garden of the Uni- 
versity of Kyoto, transplanted from Mount Kokuzoyama, 
Nagasaki Prefecture, Kyushu, Japan, Tagawa 2552 (KYO!). 

When the senior writer described the present species, he made 
a comparison with Dryopteris enneaphylla only. As will be dis- 
cussed under that species, D. toyamae is distinct by the crenation 
of the pinnae and the presence of teeth below the apex of the 
pinnae. Also the fact that the terminal pinna is more deeply 
dissected than the lateral ones was pointed out. The direct 
affinity of this species is no doubt with D. sieboldii, from which 
it may have been derived by some cytological variation. The 
pinnae are deeply pinnatifid near the base, thus forming one or 
two pairs of free pinnules; the sinuses become progressively shal- 
lower toward the apex of the pinna (figs. 1, 2). 

Tagawa 3435 from Taiwan was referred to D. toyamae by the 
senior writer.! If D. toyamac is really derived from D. sieboldu, 
its occurrence in Taiwan, where genuine D. sieboldii is not found, 
is difficult to explain. It is conceivable that a plant like Tagawa 
3435 was derived from the Taiwan D. enneaphylla in the same 
manner as D. toyamae from D. sieboldii in Japan. If this sup- 
position is correct, the Taiwan plant must be segregated, for its 
alliance with D. toyamae would be unnatural. 

3 Dryopreris BopINtERI (Christ) C. Chr. Ind. Fil. 254. 1905; 
Ching, Bull. Fan. Mem. Inst. Biol. 8: 398. 1938. 

Aspidium bodinieri Christ, Bull. Acad. Géogr. Bot. Mans 1902: 


1Act. Phytotax. Geobot. 9: 90. 1940. 


100 AMERICAN FERN JOURNAL 


248. 1902; op. cit. 1906: 115. 1906. Type: Tou- hae Kwei- 
chow, Chins: Bodinier. 

A distinct, large species, attaining three meters in height, 
with 7 to 15 pairs of larger lateral pinnae. Known from Kwei- 
chow, Yunnan, and Szechwan, but rare. 

4. DRYOPTERIS ENNEAPHYLLA (Baker) C. Chr. Ind. Fil. 263. 
1905; Hu & Ching, Icon. Fil. Sin. 7: t. 6. 1930; Ching, Bull. 
Fan Mem. Inst. Biol. 8: 398. 1938. 

Nephrodium enneaphyllum Baker, Journ. Bot. TOL: 170. 
Type: Ichang, Hupeh, China, Henry 3217. 

Dryopteris sieboldii var. heteroneura Tagawa, Journ. Jap. Bot. 
12: 487. 1936; H. Ito, in Nakai & Honda, Nov. Fl. Jap. Poly- 
pod. Dryopt. 1: 26. 1939 (pro form.). Type: Between Mi- 
harashi and Miyama, Province of Taito, Taiwan, Tagawa 
1034 (KYO!). 

Dryopteris heteroneura Ching, Bull. Fan Mem. Inst. Biol. 8: 


Dryopteris heteroneura differs from D. sieboldii in venation 
and in the serration of the margins of the pinnae (Figure 4). 
The margins are doubly-serrate, or broadly crenate with sharply 
toothed lobules. The basal posterior veinlets are usually apart 
from the insertion of the costules and arise directly from the 
costae, and between the first and the second basal posterior vein- 
lets two basal anterior veinlets are borne. The other veinlets 
are borne in regular alternate sequence. Each veinlet runs to a 
sharp tooth. Although such a feature could be influenced by 
the breadth of the pinnae, it does not occur in Japanese D. 
sieboldii, even when the pinnae are broader than those of the 
type specimen of D. heteroncura. However, apart from the type 
specimen, this feature is not so typically developed. An addi- 
tional peculiar feature is the doubly-serrate margins, the serra- 
tion extending to the very apex of the pinnae. In D. sieboldii, 
the margins are almost entire at and below the apex. The same 
difference is seen between Cyrtomium fortunei and C. falcatum. 
Although the peculiar mode of venation is not common to all 


DryYoprTeris, SUBGENUS PYCNOPTERIS 101 


Taiwan representatives, these may be considered conspecific. 
n 


and D. pseudosieboldiu, the original description, the description 
and figure given by Ching, and the specimen Henry 7881 (TI 


wy 


1 
2 
a 

SS 
ee 5 
Fias. 1-3, DRAWN FROM TYPE OF DRYOPTERIS TOYAMAE. 1, A 
BasaL LATERAL PINNA OF A FERTILE FROND, 2, A BASAL 
LATERAL PINNA A STERILE FROND, X 3, LOBULES OF A 

RTILE NNA, 2. Ss. 5, DRAWN FROM TYPE 0: 
DRYOPTERIS HETERONEURA (= ENNEAPHYLLA). 4, A PORTION 


F A STERILE Pinna, X 1; 5, Avic AL PART OF A STERILE PINNA, 
xt 


have been taken into account. It appears that D. enneaphylla is 
quite identical with the species mentioned above except in the 
serration of the pinnae. However, in some specimens of D. 
heteroneura from Taiwan the pinnae are sometimes regularly 


102 AMERICAN FERN JOURNAL 


lobate-erenate up to one-fourth down to the costa, t.e. to about 
the same degree as that shown in the figure of D. enneaphylla by 
Hu and Ching; Tagawa 3094, 3170, 3380 (all KYO) afford suffi- 
cient examples. In D. pseudosicboldii such crenations occur 
more typically, as seen in Tagawa 2054 (KYO). Similar crena- 
tions are also seen in some specimens of D. sieboldi from Japan, 
as for instance N. Nakajima (KYO) and 7. Baba (KYO), and 
Hiroshima H. N. School 52 (T1), but they are usually irregular. 
A further study is necessary of the Chinese plants that have been 
identified as D. sieboldii; the materials available to us are inade- 
quate. 

4a. DRYOPTERIS ENNEAPHYLLA var. pseudosieboldii (Hayata) 

Tagawa & Iwatsuki, comb. nov. 

Dryopteris pseudosieboldti Hayata, Icon. Pl. Formos. 4: 171, 
fig. 111. 1914. Type: ope! ee Province of Tainan, 
Taiwan, Hayata & Sasaki 

Dryopteris sieboldii var. Fas dae Tagawa, Journ. Jap. 

ot. 12: 487. 1936. 

Ching considered that D. pseudo-sieboldii was identical with 
D. podophylla (Hook.) Kuntze of southern China, but it may be 
that he did not see any authentic or topotypice specimens, for it 
is entirely different from D. podophylla in scales, in margins of 
the pinnae, and in the position of the sori. In D. pseudosieboldui, 
the scales are brown and lanceolate, being broadest near the 
base; each veinlet runs into a sharp tooth of the lobule except 
the anterior basal one, which stops a little beyond half-way; the 
sori are scattered and restricted to the marginal part of the 
pinnae, thus leaving a rather broad sterile space on each side 
of the costae. In D. podophylla, on the other hand, the scales 
are black and subulate; the margins of the lobules are subentire 
and neither the anterior nor the posterior basal veinlets reach to 
the margin; sterile bands occur in the fertile pinnae both along 
the costae and the margin. Thus, D. pseudosieboldii is clearly to 
be distinguished as a species distinct from D. podophylla by the 
characters noted above, in spite of the opinion of Ching, 


Dryopteris, SuBGENUS PYCNOPTERIS 103 


The distribution of the sori in D. pseudosieboldui is character- 
istic ; they are restricted to a marginal band, thus leaving a sterile 
band along each side of the costae. On the contrary, in D. 
sieboldii the sori are costal, thus leaving sterile marginal bands. 
Dryopteris pseudosieboldii has more than five pairs of pinnae, 
sometimes up to eight pairs, whereas D. sieboldii has usually four 
pairs or fewer (rarely five pairs). The two species may be dis- 
criminated by these characteristics. However, D. pseudosieboldu 
is quite identical with Taiwan plants of D. enneaphylla except in 
distribution of the sori; this difference is apparent but is hardly 
to be considered sufficiently important to segregate two species, 
and therefore D. pseudosieboldii is here regarded as a variety. 
5. neato nih ms (Hook.) Kuntze, Rev. Gen. Plant. 

2: 813. 1891; Ching, Bull. Fan Mem. Inst. Biol. 8: 399. 1938. 

Se Eads maak . ourn. Bot. 5: 236, t. 1. 1853. 

Type: Hongkong, M. Cham 

Nephrodium podophyllum Hook. ‘Sp. Fil. 4: 87. 1862. 

As mentioned above, this is another distinct species, which is 
restricted to southern China: Fukien, Kwangtung, Hongkong, 
and Hainan. 

6. DRYOPTERIS NEOPODOPHYLLA Ching, Bull. Fan Mem. Inst. 
Biol. 8: 401. 1938 
Type: Between Pin-fa and San Chouen, Kweichow, China, 
Cavalerie 2883. 

The writers have not had the opportunity of studying this 
species. 

7. DRYOPTERIS YUNNANENSIS (Christ) Copel. Gen. Fil. 122. 
9477 


2 The first renaming of the illegitimate Polypodiwm ty, He Wall. 
Hook. (non Aiton) was as Phegopteris elongata J. Smith (Hist. Fil. 933, 


1875). By the International Code of Botanical asker Art. 72, Nota, 
this is consider m mith an a sfer. Therefore, 
the ies na or the species appears to be Dryopteris elongata J 
Smith) K ntze, Rev. Gen. Plant. 2: 811. 1891. An en pe nomencla- 


tural synonym is Dryopteris khasiana (a ele ae te rata and a taxonomic 
synonym is D. yunnanensis (Christ) Copel.—c. V. 


104 AMERICAN FERN JOURNAL 


Polypodium elongatum Wall. ex Hook. Sp. Fil. 4: 234. 1862, 
non Aiton 1789 nec Schrad. 1818. 

Aspidium yunnanense Christ, Bull. Herb. Boiss. 6: 965. 1898. 
Type: Mengtze, Yunnan, China, Henry 9038A. 

Dryopteris khasiana C. Chr. Ind. Fil. 272. 1905; Contr. U. S. 
Nat. Herb. 26: 276. 1931. [New name for Polypodium 
elongatum Wall. ex Hook., non Aiton] 

Microchlaena yunnanensis Ching, Bull. Fan Mem. Inst. Biol. 
8.325: 4. VI, fig. 1. 1988. 

Much impressed by the catadromous mode of venation, Ching 
described the new genus Microchlaena, which was based solely on 
this species. Copeland (1947) alluded to Ching’s genus briefly 
and made it a synonym of Dryopteris, since Dryopteris as a 
whole has the catadromic plan of venation. A comparison with 
the type of venation in D. heteroneura is useful. As noted above, 
the sequence of veins is not always completely consistent, even 
within a single species. In an Assam specimen of D. yunnanensis 

‘ collected by Gustav Mann (October, 1888, KYO), some veinlets 
are anadromie in sequence. Thus the venation is not a feature 
sufficient to discriminate the genus Microchlaena. 

Besides the ‘‘peculiar’’ mode of venation, the lack of scales 
above the stipe base and the dark green colour of the frond were 
noted by Christensen (1931) and also by Ching (1938). As a 
matter of fact, the texture of D. yunnanensis resembles that of 
D. dickinsii or of D. atrata. However, small seales are, although 
rarely, found on the costae beneath, especially at the very base. 
Although the lateral pinnae are numerous (up to 20 pairs com- 
pared to 15 pairs at most, in D. bodinieri, otherwise the largest 
species of Pycnopteris), the imparipinnate blades are distinctive 
enough to allow us to consider D. yunnanensis also a member of 
the subgenus Pycnopteris. 


DEPARTMENT OF Botany, UNIversItTy oF Kyoto, Kyoto, JAPAN. 


DRYOPTERIS CRISTATA X SPINULOSA 105 


The Correct Name for the Hybrid Dryopteris 
cristata X spinulosa 


FRANCIS BALLARD 


A hybrid between Dryopteris cristata (L.) A. Gray and D. 
spinulosa (O. F. Muell.) Watt has been known in Europe for 
more than a hundred years. In Clapham, Tutin, and Warburg’s 
Flora of the British Isles (p. 42, 1952) it is assigned the name 
““Dryopteris uliginosa (A. Braun) Kuntze,’’ but since such a 
name is not listed in Christensen’s Index Filicum and Supple- 
ments I have been asked to clarify its status. 

The first use of the epithet wliginosum in this context was the 
description of Aspidium spinulosum b) uliginosum A. Braun in 
Doell’s Rheinische Flora (1843), which reads as follows: ‘‘ Moor- 
liebender, kurzstachlicher Schildfarn. Strunk ziemlich kurz, mit 
wenigen, locker stehenden, breit-eiférmigen, kurz zugespitzten, 
braungelben Spreublittchen; Wedel doppeltgefiedert-fieder- 
spaltig, Fiederchen geniihert, an der Basis auf der unteren Seite 
etwas breiter, die an der Basis stehenden ein wenig kiirzer als 
die folgenden; Ziihne der Fiedertheilchen ziemlich kurz, stachel- 
spitzig, anliegend. Diese noch weiter zu beobachtende Form 
wurder yon A. Braun im Moos bie Freiburg mit der ersten 
Varietiit und mit Aspidium cristatum gefunden.’’ The ‘‘first 
variety’’ mentioned was var. elevatum, which is typical spinulo- 


Newman in proposing Lastrea uliginosa in 1849 made no refer- 
ence to Doell’s Rheinische Flora, but he did in fact adopt the 
epithet wliginosum from that work, as he explains in his “‘A His- 
tory of British Ferns’’ (p. 163. 1854). He stated: ‘‘The first 
notice of this fern which I have seen is in D6ll’s ‘Rhenish Flora,’ 
pp. 17 and 18, but I believe it to be copied from a prior work or 
a MS. of Professor A. Braun... The next notice of this fern, 
and the first of its occurrence in England, is from my own pen; 
and, by a strange omission, subsequently amended by myself, no 
allusion whatever is made to the earlier description I have just 

ited.’ Newman assigned the common name ‘‘Lloyd’s Fern’”’ 


106 AMERICAN FERN JOURNAL 


to this plant, in honour of a Mr. Lloyd, a gardener who had pro- 
vided him with the material of it. 

Druce in his various publications referred this plant to 
Dryopteris as D. uliginosa Ktze. Why Druce should have at- 
tributed the epithet to Kuntze I cannot imagine. It was probably 
just an error, which was copied by Clapham, Tutin, and War- 
burg. I think that we shall have to accept Druce’s combination 
as validly published, since it is quite clear from his reference in 
me Comital Flora that he was referring to Newman’s plant. 

specimen he quotes is one of those quoted by Newman: 
aetaehas Cheshire Mr. George Pinder.’’ Druce’s first use 
of the name was in 1908, although he prefixes the eon with 

a question mark. Perhaps one should take the reference in Hay- 
ward’s Pocket Book (1909). 

In any ease, the combination Dryopteris uliginosa (A. Braun 
ex Doell) Druce antedates D. uliginosa (Kunze) C. Chr. (Ind. 
Fil. Suppl. 3: 100. 1934). The important synonymy is as follows: 
DRYOPTERIS ULIGINOSA (A. Braun ea Doell) Druce, List of Brit. Plants 87. 

; Druce in eee. s Bot. Pocket Book, ed. 13, 259. 1909; Comital 
Fl. Brit. Isles 383. 1932 
Aspidium spinulosum var. oa A. Braun ex Doell, Rhein. Fl. 17. 
1843. 


Lastrea uliginosa Newm. Phytol. 3: 678. 184! 
Lophidium uliginosum Newm. Phytol. 4: 371. tis, Hist. Brit. Ferns 163. 
1854. 


Aspidium uliginosum Nyman, Consp. Fl. Eur. 866. 1884, non Kunze, 1847. 
Nephrodium sia Rouy, Fl. France 14: 414. 

Polystichum uliginosum P. Fourn. Quatre Fl. France 17. 1947. 

Royau eas GaRDENS, Kew, ENGLAND. 


1 This cross, pan ein ace cristata  spinulosa has been reported from the 
age States a paler f times ie Bull, Torrey Club 35: 136, 1908; Fern 
Bull, 77: 35 


D, . Fer 1g Ga | 911, Reed, Ferns and Fern- 
allies ie Muvy bait a peewie as, 1953), but is not well known. The 
name uliginosa for it has not come into the American literature, but it is 


now available. The sorreapoudin oan of cristata glad aan on the 
other hand is well known under the name D. Boottii.—C. V. Mor 


New EquisetuM 107 


A New Equisetum! 
BERNARD BOrviIn 


In the ane Arctie there occurs a plant that could easily 
Ss as a small Equisetum arvense or, if fertile stems are 


raun|, because, as in the latter, the fertile stems turn green 
and produce branches. However, closer examination reveals that 
the production of green branches is normal in this plant and not 
as in £. arvense, just an exceptional individual variation. Fur- 
thermore, there is some overlapping in the time of appearance of 
fertile and sterile stems and the behavior of the plant is in this 
respect, intermediate between that of HZ. arvense and E. pratense. 

In this plant, which I am naming F. calderi, the fertile stems 
appear first but the sterile stems begin to appear while the 
fertile stems are still present and only in sporesis. Most, if not 
all, of the fertile stems soon turn green and produce green sterile 


may be briefly stated as follows: Central cavity very small (one- 
fifth to one-half of the diameter in arvense), about as small as 
the carinal cavities; sheaths from the middle of the stem with 
4 or 5 (6) teeth (8-12 in arvense), these (0.6) 0.8-1.0 (1.3) mm. 
long (1.5-2.5 mm. in arvense); branches trigonous with teeth 
0.4-1.0 mm. long (1-2 mm. in arvense) ; upper sheath of fertile 
stems 4-9 mm. long (16-19 mm. in arvense), with 4 to 5 teeth 
only, these very short, 1.5-3.0 mm. long. This Arctic vicariant 
of Equisetum arvense Li. may be named and described as follows: 
Equisetum calderi, sp. nov. Affine Equiseto arvensi L. et inter 
sectiones Subvernalia hit et Vernalia Braun stans. Surculi 
annui dimorphi, fertiles praecoces vel subcoetanei, persistentes 
et ramulos serotinos proferentes. Surculus sterilis gracilis, 5-2 
em. alt., erectus vel decumbens, caudatus, 4-5 (6)-goniatus, 
vaginis 4-5 (6)-dentatis, dentibus (0.6) 0. 8-1.0 (1.3) mm. longis, 


1Contribution No. 34 from the Plant Research Institute, Research 
Branch, Canada Department of Agriculture, Ottawa, Canada. 


108 AMERICAN FERN JOURNAL 


lacuna eerie minima. Frons simplex, sparsa, verticillis (3) 
n planta, ramis revlon solidis, internodiis (2) 3-5 
ramo, dentibus vaginarum ternis, 0.4—1. 0 n longis. 
Sureulus fertilis 5-12 em. ongus, primo simplex, d aetide yamosus 
et viridescens, internodiis pallide brunneis viridescentibus, 
vaginis pallide viridibus, vagina aap are 4-9 mm. longa, denti- 
bus 4 vel 5 in vagina 
RANKLIN : Senn é Calder 3747, ierolisher Bay, Baffin Island, 
63°45'N, 68°32’W, in stream 0.5 mile north of camp, upright in 
3”-6” of water, both fertile and sterile plants present, July 5 
1948 (DAO type) ; Senn & Calder 3798, from same locality, bed 
of dried up river, July 7, 1948 (DAO); D. Coombs 112, Erik 


Harbour, Baffin Island, 72°40'N, 76°36°W, Aug 4 
( ; D. Coombs 121, Eclipse Sound, Bylot Island, 72 55 N, 
80°15'W, in sand, Au 1 (DA ’. Bruggeman 


July 26, 1951 el a Bruggemann 186, northeaster 
Ellesmere Island, 82°31'N, 62° 45°W, on ey tussocks among 
mosses, grasses and willows, found only i mall area on a 


steep, wet, north-facing slope at foot of ne sacra cliff, 15’ to 
30’ above sea level, July 4, 1951 (DAO). 

EWATIN: W. J. Cody 1077, Southampton Island, Coral Har- 
bor, 64°09’N, 83°18’W, fairly common in gravelly muck in 0.5"- 
2” water, July 1, 1948 (DAO); Cody 1674, from same locality, 


mes str 
shallow slow-flowing water, Aug. 11, 1948 (DAO); Cody 1997, 
from same locality, common in moist limestone gravel, Aug. 11, 
1948 (DAO) ; Cody 1703, from same locality, fairly common in 
Carex meadow, feds bh 1948 (DAO); Cody 1554, from same 


locality, common er 1”, rooted in shallow muck over lime- 
stone gravel, July 5. 1948 (DA 
MAcKE W. I. Findlay 260, Coppermine, 67°50'N, 115° 


10°W, mibiat putes strobili very few, vegetative shoots quite 
common, Aug. 6, 1951 ae 

This species probably occurs also in Greenland, but the only 
collection at hand, Sérensen 202, from Scoresbysund, consists 
mostly of small fragments and is only doubtfully referable here. 


Two PROBLEM-SPECIES 109 


The sterile shoots vary from erect to depressed, but all the 
fertile collections (Bruggemann 186, Senn & Calder 3747, Find- 
lay 260, and Cody 1703 & 1077) show only erect sterile stems, the 
depressed phase being apparently sterile. 

It is a pleasure to attach to this species the name of my friend 
and co-worker J. A. Calder. 

CANADA DEPARTMENT OF AGRICULTURE, OTTAWA, CANADA. 


Two Problem-species: Sige cordatum Gaud. and 
Syngramma pinnata J. Smith 
R. E. aie 

It is clear from Copeland’s Genera Filicum (pp. 55, 57) that 
he considers both Schizoloma cordatum and Syngramma pinnata 
to be nearly related to the genus Tacmitis. I agree with this state- 
ment, and wish to present further evidence for it than that given 
by Copeland. My conclusion will be that ae species should be 
transferred to Taenitis. 

Schizoloma cordatum has a fairly wide distribution in Borneo, 
the Philippine Islands, Celebes, and the Moluccas, but has not 
been very frequently colleeted. I believe the reason to be that it 
is confined to limestone, and of very local occurrence, often in 
places not easily accessible. Certainly, the only locality from 
which it is known in Sarawak is the limestone hill at Bau, near 
Kuching. But ee dealing with morphology, a note on nomen- 
clature is necessa 

The genus Schizoloma was established by Gaudichaud in 1824, 
and in it he placed three very diverse species: S. cordatum, NS. 
guerinianum and 8S. billardieri. Fée later removed S. cordatum 
to a new genus Schizolepton and S. guerinianum to Isoloma, 
leaving S. billardieri in Schizoloma. Copeland states that in so 
doing Fée misconstrued Schizoloma, but he does not explain what 
he means by that statement. Fée correctly construed Schizoloma 
Gaud. as a mixture, and had the right to choose how he would 
resolve that mixture. In so doing, he effectively chose S. bil- 
lardieri as the type species of Schizoloma. 8. billardiert (= 


110 AMERICAN FERN JOURNAL 


Lindsaea ensifolia Swartz) undoubtedly belongs to the genus 

Lindsaea as defined in the recent monograph of Kramer and as 
accepted by Copeland, and thus Schizoloma becomes a synonym 
of Lindsaea. The correct name for Schizoloma cordatwm is thus 
Schizolepton cordatum (Gaud.) Fée; there is no other known 
species of Schizolepton. 


i) 


PARAPHYSES, ALL X , TAENITIS BLECHNOIDES ; 2, SYNGRAMMA 
PINNATA; 3, anne, ‘CORDATUM; 4, SYNGRAMMA ALISMIFOLIA 
Schizolepton differs from Lindsaea in having rigid cylindrical 
bristles, each consisting of a single series of cells, as a covering 
of the young parts of its rhizome. The rhizome of Lindsaea bears 
distinetly flat seales. In its sorus Schizolepton superficially re- 
sembles those species of Lindsaea (e.g. L. ensifolia) which have 
a continuous sorus all along the edge of each leaflet. But in 
Lindsaea the sorus is protected by a thin indusium which is quite 
distinct in substance from the true margin of the frond. In 
Schizolepton the sorus is in an apparently marginal groove, and 
the two edges of the groove are exactly alike in substance. 


1Cf, Kramer, Acta Bot. Neerl. 6: 97-138, 1957. 


Two PROBLEM-SPECIES 111 


think that the conclusion is inescapable that the sori in Schizo- 
lepton and Lindsaea, though superficially similar, have had quite 
different evolutionary origins. Thus, both in protective bristles 
on the rhizome, and in the nature of the protection of the sorus, 
Schizolepton is quite different from Lindsaca. Equally, it is 
distinet from Isoloma (it seems to me that Copeland is wrong in 
suggesting that Isoloma is related to Schizolepton). A third 
striking difference is the presence in Schizolepton of abundant 
rather thick hair-shaped paraphyses (fig. 3), as long as the 
sporangia, each consisting of a row of about eight cells, the lower 
cells gradually more slender, the apical one not of distinctive 
form; paraphyses in Lindsaea are much fewer and shorter, 
usually of two cells. (Fée gave a drawing of a paraphysis of 
Schizolepton, but omitted to show that it consists of a row of 
separate cells.) 

Fée compared Schizolepton to Vittaria, and in my opinion the 
apparently marginal soral groove in the two genera is of exactly 
comparable structure, the two lips of the groove being of equal 
substance. But it is certain that Schizolepton and Vittaria are 
not closely related, and I believe the development of similar sori 
in the two cases is due to parallel evolution. 

In his comments on Schizolepton, Copeland writes ‘‘related to 
Taenitis, in spite of being indusiate.’’ As we have just noted, 
it is not indusiate. But it is certainly related to Taenitis, agree- 
ing in rhizome-bristles, in venation, and in the nature of its 
paraphyses. Copeland states that Taenitis has ‘‘peculiar pa- 
raphyses which resemble abortive sporangia.’’ But in fact the 
paraphyses of Taenitis are similar to those of Schizolepton, con- 
sisting of a single row of thin-walled cells (fig. 1) ; the difference 
is that in Taenitis there are more cells in each paraphysis (about 
16) and the apical 10-12 cells are much shorter than the rest. 
In a dried specimen, the cells of a paraphysis collapse and turn 
brown, but if they are soaked in an alkaline bleaching solution 
they lose the brown colour and expand to their orginal shape. 
The paraphyses in Taenitis blechnoides develop before the spo- 


112 AMERICAN FERN JOURNAL 


rangia, and their swollen ends form a very effective protection 
for the young sporangia. 

It appears to me that Schizolepton differs from Taenitis only 
in having the sorus in an apparently marginal groove instead of 
in a superficial position between midrib and edge. If we take the 
parallel case of Vittaria, we find some species with sori in shallow 
superficial grooves lying between midrib and edge, and others 
with sori in apparently marginal grooves. Comparing Vittaria 
with the related genus Antrophyum, it seems probable that the 
elongate superficial sorus is primitive in Vittaria, and that the 
apparently marginal sorus is due to the upgrowth of the inner 
edge of a groove which at first is superficial. The sorus of Schizo- 
lepton bears the same relation to that of Taenitis blechnoides as 
the marginal type of sorus bears to the superficial type in Vit- 
taria. As both kinds of sorus can occur in the one genus Vittaria, 
they could also occur in Taenitis (though here intermediate con- 
ditions are lacking) ; in fact, it would be quite a natural arrange- 
ment to transfer the species Schizolepton cordatum to Taenitis, 
and this I now do. The new combination is Taenitis cordatum 
(Gaud.) Holttum.? 

Regarding Syngramma pinnata J. Smith, Copeland remarks 
‘distinguished from Taenitis only by the sori and paraphyses.”’ 
Here again therefore we have to look at paraphyses (fig. 2), and 
find that Copeland’s statement is incorrect, because he did not 
know the true form of the paraphyses of Taenitis (though they 
were figured by Beddome in the Ferns of British India, ¢.54, in 
1866). There is in fact close agreement between the paraphyses 
of Syngramma pinnata and Taenitis blechnoides; in S. pinnata 
the distal swollen cells are longer and fewer than in Taenitis, 
being about midway between those of Taenitis blechnoides and 
Schizolepton cordatum. But in other species of Syngramma the 
paraphyses are different; they have a glandular terminal cell 
which is quite different from the rest in shape, colour and con- 
tents (fig. 4). 


2Basionym: Schizoloma cordatum Gaud. Ann. Sci. Nat. Paris 3: 507. 
1824. 


Two PROBLEM-SPECIES 138 


The sorus in Syngramma pinnata is variable, a fact shown by 
Copeland in his plate 1. In what is regarded as the typical form 
of the species, the sori run along most of the veins, which form 
a network of oblique areoles (this venation is exactly as in 
Taenitis). In many specimens however the sori do not spread so 
much, and are more or less confined to the veins on a band about 
midway between the midrib of a leaflet and the margin, the 
sporangia more crowded than in the typical form and forming 
small patches. The sori in this variety occupy exactly the same 
position as the sori in Taenitis blechnoides, and if the gaps be- 
tween them were filled they would be indistinguishable from the 
sori of 7. blechnoides. 

In shape of frond, and in venation, S. pinnata is exactly like 
T. blechnoides; sterile plants would be indistinguishable. 
supposed difference in paraphyses is shown to be non-existent ; in 
paraphyses S. pinnata resembles T. blechnoides, not the other 
species of Syngramma. On the other hand, there is no other 
Syngramma which has the frond-form and venation of Taenitis. 
To me, the conclusion is that Syngramma pinnata is properly a 
species of Taenitis, and I have transferred it to Taenitis in a 
recent paper. 

Prof. I. Manton has already recorded the chromosome number 
n = 116 (4 & 29) for Syngramma quinata (Hook.) Carr., as 
found in Malaya; this is probably not the true S. quinata of the 
Pacific, but a distinct Malaysian species.* She has recently found 
the number n = 58 (2 X 29) ina plant of Syngramma alismi- 
folia (Presl) J. Smith brought to Kew from Sarawak; but an- 
other recent observation on Taenitis blechnoides gives a quite 
different chromosome number, n = 44. (Prof. Manton has in- 
formed me verbally of these two records, and I am grateful to 
her for permission to mention them here.) The chromosomes 
thus provide additional evidence that Taenitis and Syngramma 
are not closely related genera (though I believe that they are 
more closely related to each other than either is to Lindsaea). 


3 Kew Bull. 1958: 453. : 
4 Cf, Holttum, Ferns of Malaya, pp. 580, 627. 1954. 


114 AMERICAN FERN JOURNAL 


Dryopteris borreri in Bavaria 
J. PoELT 

The species Dryopteris borreri Newm., sometimes incorrectly 
called D. paleacea (Swartz) Hand.-Mazz., has attracted much at- 
tention in recent years since Doepp and Manton have shown 
that it is a distinct species. They have shown that it exhibits 
constant apogamy, in spite of funtioning antheridia. There are 
many connections with D. filix-mas through hybrids, which are 
also apomictic. Concerning these cytogenetic conditions we may 
refer to the publications of the above-mentioned authors. Wolf, 
v. Tavel, Rothmaler, Nordhagen, Reichling, Lawalrée, and 
others have dealt with its geographic distribution in Europe; 
they show that these plants are on the whole restricted to the 
oceanic regions of western and southern Europe. 

In Bavaria, it was mainly Paul who drew attention to these 
beautiful ferns. Since the Bavarian region lies on the north- 
eastern limit of the European area for this species, and since 
moreover the winters in this region are not mild but rather cold, 
it may be appropriate to deal briefly with this small but interest- 
ing part of the whole area. 

As indicated on the accompanying map, Dryopteris borreri 
has been found on the lower ranges of the northern rim of the 
Alps, in north-exposed valleys of the Northern Limestone Alps, 
and in some places of the adjacent region to the north. It 
is always found in glens of the northern slopes of mountains 
or in the hilly region mostly in deep holes, that is, in places 
where there is a continually high humidity during the vegetative 
season. The protection against frost, which is necessary for an 
oceanic species, is provided by the long-lasting cover of snow, 
which in glens on northern slopes and in holes, continues far 
into the spring, rather than by a mild climate, as it is in western 
Europe. There is a close parallel with Ilex aquifolium, whose 
survival on the northern range of the Alps likewise depends 
mainly on the snow cover in winter. 

On the other hand, these ferns are not bound to any special 


DRYOPTERIS BORRERI 115 


type of soil, since everywhere enough organic matter is formed 
to produce sufficient top-soil and the minerals contain enough 
lime to prevent the soil from turning acid. 

As is true elsewhere, it is not quite easy to bring the different 
forms of Dryopteris borreri occurring in Bavaria into a system. 
This is the result of the special cytogenetic conditions prevailing. 
As Doepp has shown, the transitional forms, hybrids with D. 
filix-mas and derived forms, are generally in the majority on 


& 
P 4  ) e 
eae ae ay € ( 
rg Se > 1 aes 
-@i 


Fig. 1. ASTERISKS, LOCALITIES OF DRYOPTERIS BORRERI; Dots, LOCALITIES 
or D. X TAVELIT AND UNCERTAIN ForMS; BROKEN LINE, NORTHERN BORDER 
or THE ALPS; BROKEN AND Dorrep LINE, SOUTHERN BorDER OF BAVARIA 
the borders of the area in which the species is found. This is 
Nea true in southern Bavaria. It almost looked as if genu- 
D. borreri could not be found any longer in this region. Yet 
a pat specimens found there have been proved to belong to this 
type both morphologically and in regard to the development of 
their spores. All these specimens are small plants with narrow, 
longish, pointed scales, with parallelogram-shaped pinnules, 
which are dentate at the apex but nearly smooth on the sides, 
and with brownish, strongly vaulted indusia. All the other spe- 
cimens show definitely, though in different degrees, the influence 
of Dryopteris filiz-mas, and must therefore have to be designated 
by the collective name Dryopteris < tavelii Rothmaler, which is 


116 AMERICAN F'ERN JOURNAL 


the name applied to the cross D. borreri X filix-mas. 

In some cases, it is quite impossible to arrive at an unambigu- 
ous decision. The reason is to be found in the facts stressed by 
Doepp that crossbreeds of D. borreri with D. filiz-mas can hardly 
be distinguished from D. borreri by their appearance. A cytolog- 
ical examination of all specimens found in different regions is 
not possible. It is therefore the task of plant geography to fix 
the geographic distribution of all those forms that show the in- 
fluence of D. borreri. This has in outline been done for Bavaria 
in the map in the present paper. We have to add that this spe- 
cies is not found in the other mountain ranges of Bavaria, like 
the Boehmerwald,! where a number of other oceanic species are 
found. The Bavarian area fits well into the general picture of 
the distribution of the species in the northern Alps, which ex- 
tends from Switzerland to eastern Austria. 

The few specimens of genuine Dryopteris borreri among an 
abundance of D. tavelii appear to prove that the present stock of 
specimens of the species in our region is a relic of an earlier 
climatically more favourable period, from which only the 
hybrids with D. filiz-mas have managed to survive on a large 
scale down to the present time. 


BIBLIOGRAPHIC REFERENCES 

Doepp, W. 1939. Cytologische oe genetische Untersuchungen innerhalb 
der Gattung Dryopteris. Planta 29: 481-533. 

—. 1941. Ueber Dryopteris meen Christensen (Dr. borreri Newm.). 
Ber. Deut. Bot. Ges. 59: 423-426. 

—. 1950. Zur Problematik von Dryopteris paleacea (Sw.) ©. Chr. und 
ihres Formen- und Verwandtschaftskreises. Ber. Deut. Bot. Ges. 62: 
1 


Lawalrée, A. 1950. Ptéridophytes, in Flore Gen. Belgique vol. 1, Bruxel- 


es. 

Manton, I. 1950. Problems of Cytology and Evolution in the Pterido- 
phyta. Cambridge, England. 

Nordhagen, R. 1947. Dryopteris a (Sw.) ©. Chr. og dens ut- 
bredelse i Norge. Blyttia 5: 89-11 


1 As has been shown by W. Freiberg, to whose kind help the author owes 
a great deal. 


PTERIDOPHYTES FROM AUSTRALASIA T1Y 


Paul, H. 1940. Botanische Kurzbeitriige. Ber. Bayer. Bot. Ges. 24: 50-58. 

Reichling, L. 1953. Dryopteris paleacea (Sw.) Handel- Mazzetti et Dry- 

opteris < tavelii Rothm. au Grand-Duché de Luxembourg et en Bel- 
0 39- 


Tavel, F. von. 1937. Denes borreri Newm. und ihr Formenkreis. 
erh. Schweiz. Naturf. Ges. 118: Jahresvers. 153, 1 
Wolf, H. 1936. Ein neuer Farn der Pfalz, seine vorkoumhon und seine sys- 
tematische Stellung. Pollichia, n. ser. 5: 80-92. 


Notes on Pteridophytes from Australasia and 
New Caledonia, I 


Mary D. TINDALE 
Due to the impending publication of two new floras which 
will deal with pteridophytes occurring in southeastern Australia, 
it is necessary to publish the following two new combinations 
and a new species which will be either included in or mentioned 
in these books. 
BLECHNUM MINUS SUBSP. SCA 
Bra be minus (R. Br.) Ettingsh. in Penksche: Ak. Wien 
: 63. 1864, ats scabrum (Domin) Tindale, comb. et 


vt ‘i nov. 
Bastonym: Blechnum eae (L.) Schlecht. var. scabrum 
Domin, in Bibl. Bot. 85: 116. 1913. 

Honorypr: Australia, Sieber Syn. Fil. exsie. No. 107. Domin 
does not state in which herbarium this specimen is located but 
I have examined isotypes bearing this label in the Natural His- 
tory Museum, Paris, and in the Riksmuseum, Stockholm. Both 
specimens agree well with Domin’s description 

A short description of B. minus ssp. panies is provided be- 
low: 

Rhizome consisting of a short, erect caudex up to 5 cm. high 
and a horizontal, stoloniferous portion ; caudex densely sealy, the 
seales long-acuminate, very thick, entire, very glossy, linear 
(more than 6:1) to narrowly lanceolate (6:1), chestnut or dark 


118 AMERICAN FERN JOURNAL 


red-brown, brown or dark brown with light chestnut-coloured 
borders, up to 2.5 em. long and up to 3 mm. broad, with a fibril- 
lose, twisted apex. Stipes usually red-brown or black, strongly 
tuberculate. Main rhachis red-brown or black, rarely fawn to- 
wards the apex. Fronds up to 120 em. in length including the 
stipes. Sterile pinnae gradually much smaller towards the base, 
thinly to thickly coriaceous, up to 25 em. long and 3 em. broad, 
closely spaced, often imbricate, prominently auriculate, the 
auricles up to 5 em. long and up to 1.5 em. broad, the apex of 
the pinnae acute or obtuse, the margin minutely serrulate. 
Fertile pinnae lomarioid. Spores bilateral, elliptical or sub- 
globose, with a slightly reticulate perispore and a narrow, erosely 
dissected wing. 

This variety is raised to subspecifie rank on ecological and 
geographical grounds. It ranges from Fraser Island, south- 
eastern Queensland, to the South Coast region of New South 
Wales, where subsp. scabrum occurs in swampy, low-lying land 
not far from the sea. However, B. minus subsp. minus is usually 
found on the tablelands of New South Wales and Victoria along 
river banks and creeks in narrow mountain gorges, at the edge of 
waterfalls, along the banks of streams in eucalypt forests and 
in swampy creeks in cleared rain-forest areas. In Victoria and 
Tasmania the type subspecies is found in more coastal regions, 
e.g. there is a very old record (NSW. P4372) collected in 1853 
on the banks of the River Yarra, Victoria, near Prince’s Bridge, 
now in the heart of the present city of Melbourne. The type of 
subsp. minus was collected by Robert Brown in 1804 at Port 
Dalrymple at the mouth of the River Tamar, Tasmania (BM). 

Some cig ecb "sl oben s of B. minus subsp. scabrum 
are - follows: QUEENSLAND: Fraser Island, Wide Bay ee 

E. Hubbard no. 4576. 17 Oct., 1930 (NSW. P8046). 


Oatley, J. H. Camfiel d, Feb., 1897 (NSW. P4356); Kogarah 
Bay, A. A. Hamilton, June, 1901 (NSW. oe Kurnell, 
Botany Bay, J. L. Boorman, May, 1906 (NSW. P1245 

Many of the swampy sites in the George’s River meee of 


PTERIDOPHYTES FROM AUSTRALASIA 119 


Sydney, N.S.W., where B. minus subsp. scabrum was frequently 
collected at the turn of the century, have been reclaimed and 
are covered with houses, so that this fern has become very un- 
common 

The evowhe key will serve to distinguish the above-men- 
tioned subspecies :— 


y 


Sterile and fertile pinnae not auriculate at the base or the lower pairs 
very slightly auriculate; scales of the rhizome es mostly concolorous, 
rarely with a paler border; stipes stramineous, sometimes mottled with 
rown, the base dark brown, smooth or slightly tre te. 

B. minus subsp. minus 
Sterile and fertile pinnae very prominently Rie tose at the base; 
scales of the rhizome glossy v Line a darker central band; stipes dark 
red-brown or black, very tubereulate —..-.-... B. minus subsp. scabrum 


‘¢ 


ADIANTUM CUNNINGHAMIL 


At my request the late Mr. A. H. G. Alston took photographs 
of the type of Adiantum affine Willd. in the Berlin Herbarium 
several years ago. These photographs left no doubt that this 
fern is the cosmopolitan Adiantum Capillus-veneris L., not the 
fern known as A. affine in New Zealand and Australia. The 
pinnules of the holotype of A. affine are cuneate-flabellate (see 
plate 12) instead of rhomboidal as in the fern hitherto known by 
this name. The latter species must now be called A. Cunning- 
hamii Hook. which was described! from a number of specimens 
collected in the North Island of New Zealand. After a compari- 
son of material from Australia and New Zealand, I have con- 
cluded that the Australian material is not sufficiently different 
to be regarded as a separate species. With one exception (i.e. 
NSW. P2636, from Rockingham Bay, Queensland) the seales of 
the rhizome in the Australian specimens that I have examined 
are eolden-brown instead of dark red-brown as in the New Zea- 
land material; also the scales on the rhizomes of the latter tend 
to be slightly larger 

There are two Wirieties in A. Cunninghamii, one with glabrous 


1 Spec. Fil. 2: 52. t. 86A. 1858. 


AMERICAN FERN JOURNAL VotumME 50, Puate 12 


ayy 


ADIANTUM AFFINE, TYPE. BERLIN HERBARIUM. PuHoTOGRAPH By A. H. G. 
ALST 


PTERIDOPHYTES FROM AUSTRALASIA 121 


rhachises and another in which the pele surfaces of the main 
and secondary rhachises are clothed w ark brown, crisped 
hairs. I have seen no intermediates ae the two types of 
rhachises in the Australian material, although specimens with 
he main rhachis glabrous and the upper surface of the second- 
ary rhachises hirsute do occur amongst the New Zealand speci- 
mens according to Cheeseman,” but these may be hybrids with A. 
fulvuwm. Raoul. 

Hooker’s original description of this species was drawn up 
from New Zealand specimens all of which had glabrous rhachises, 
so that this would now become the type variety, 7.e. A. Cunning- 
hamit Hook. var. Cunninghamii. The material with hirsute 
rhachises from eastern Australia may be placed in Adiantum 
Cunninghamii Hook. var. intermedium (Benth.) Tindale, comb. 
nov., which is based on A. affine Willd. var. intermedium Benth.,* 
the type specimen of which was collected at Rockingham Bay, 
Queensland, Australia, by Dallachy, and is preserved in the 
Herbarium, Royal Botanic Gardens, Kew, England. Dr. A. C. 
Jermy very kindly examined this specimen and checked that 
there are dark brown, crisped hairs on the upper surface of the 
rhachises. 

In the Australian material examined, specimens with glabrous 
rhachises are more common in northeastern Queensland but 
extend as far south as Bulgong Heights, North Coast, N.S.W. 
(e.g. NSW. P2624). In the southern part of its range in 
Australia the var. intermedium predominates especially on the 


Constable, 15 Sani ae The Dome, Dorrigo, M. Tindale, 


the Waters, Blue Mountains, Constable, 7 Feb., 1949 (NSW. 
P5655), and similar specimens are found in southeastern Queens- 
ant and on the Atherton Tableland, North Queensland. 


Man. N. Zeal. Fl. Ue a 
on Austr. 7: 725. 


122 AMERICAN FERN JOURNAL 


On the whole the specimens collected in the rainforests of 
New South Wales are larger, more robust and less glaucous than 
those from northeastern Queensland, although exceptions do 
occur such as NSW. P6721 collected at Linden, Blue Mountains, 
N.S.W., which is smaller and less branched, agreeing well with 
material from Queensland. NSW. P452 from the Cairns district, 
northeastern Queenland, is a very robust plant with hirsute 
rhachises. 

Dicrymia McKEE!I 

During several visits to New Caledonia between 1954 and 
1958, Dr. H. S. McKee made a number of interesting collections 
of pteridophytes several of which have proved to be undescribed 
species, one of them being described below :— 

Dicrym1a McKeei Tindale, sp. nov. Filix epiphytica. Rhizoma 
late repens 1.5-2 mm. crassu elaucum glabrescens, paleis 


nigris. Stipites remoti basi ad rhizoma articulati, erecti 
straminei vel pallide fusci, 1.7-6.2 em. longi, 0.5-1.2 mm. lati, 
praeter phyllopodium elabrati. Frondes_ ‘simplices lineari- 
e 


b 
cuneata, apice acuminata vel subacuta. poe teri TB43 
a 5 em. i e 


erumque u 
liberis in areolis inclusis. Costa utraque facie prominens, pallide 
fusca, supr ta. Sori 4-11 exindusiati perenne 


cellulo ; ferris pi: elongatis, angustissimis saepe castaneis, sur- 
sum 2 ¢ s latis, basi 1 cellulis latis. Sporae bilaterales reni- 
formes syalines minime profunde ranailate: tuberculat 

Epiphytie fern. Rbieoms widely hese 1.5-2 mm. in . diam., 
glaucous, glabrescent, paleaceous, the scales fugacious, appressed, 
clathrate, peltate- -based, suborbicular, contracted into a long- 


PTERIDOPHYTES FROM AUSTRALASIA 123 


acuminate apex, fimbriate, and brownish-black. Stipes remote, 
articulated at the base to the rhizome, erect 


subcoriaceous, narrowly cartilaginous-margined. Fertile lamina 
10-21 em. long, 1.2-1.6 em. broad, entire, subsinuate towards the 
apex, shortly cuneate towards the | base, the apex ae - 
subacute. Sterile lamina 7.5-13 ¢ long, 1-1.5 em. broad, 
apex obtuse. Veins hetitdlate. Sbuads, anastomosing, nes a 
series of about 4 areolae, the latter usually 4- or 5- angled, rarely 
with a very few, free, included veinlets in the areolae. Costa 
prominent on both surfaces, fawn, not grooved above. Sori 4-11 
on each frond, exindusiate, superficial, orbicular to oval, oeccupy- 
ing the upper part of the fro nd, in a single row close to the 
margin, pustulate on the upper surface, 3-7 mm. long, 2-5 mm 
road. Paraphyses absent. Sporangia glabrous the annulus 
with 12-15 indurated cells, 6 thin-walled cells and a 2-celled 
stomium, th dicel elongated, very narrow, often castaneous 
2-celled above, 1 cell broad at the base. Spores bilateral, reni- 
form, pedoaioni shallowly reticulate-tuberculate. 

Ho to l’Her ge Nouméa district, New 
Caledonia, 400 m. alt., climbing on a tree trunk, in rain-forest, 

he gag ae 2126, 20 Feb., 1955 (NSW. P8036). The holo- 

e is in the National Herbarium, Royal Botanic "Bardens 

ne. erat Ae ; isotypes ei US). 

OTHER SPECIMENS EXAMIN 

NEw be aap te ntA: Mount ‘Koghi, above l’Hermitage, 500-800 

alt. a tree trunk, on a forested slope, H. 8S. McKee 3941, 
4 Feb., "1956 (NSW. P8037) ; Koghi, 400 m. ‘alt., eens et arbres 
vivantes, foréts, Franc 470, 2 Feb., 1909 (NSW. "P80 

Dictymia McKeei belongs to a small genus of three or pee 
four species occurring in Australia, New Caledonia, and Fiji. 
It is closely related to D. Mettenii (Copel.) Copel., of New Cale- 
donia, and D. Brownii (Wikstr.) Copel., of eastern Australia, 
but is distinguished readily by the linear-elliptical laminae which 
terminate rather abruptly in the stipes, whereas in the other two 
species the linear-loriform laminae taper very gradually into the 
stipes. In the two above-mentioned species from New Caledonia, 
the scales of the rhizome are very fugacious and the sori are al- 


124 AMERICAN FERN JOURNAL 


most marginal, whereas in D. Brownii the rhizome scales are per- 
sistent and the sori are half-way between the midrib and margin. 
In D. McKeei there are a few, free included veinlets in the 
areolae which breaks down a supposed generic characteristic 
of Dictymia, since they do not occur in any of the hitherto de- 
scribed species. I have seen no material of this genus from Fiji. 
CKNOWLEDGMENTS 

In conclusion I should like to express my gratitude and appre- 
ciation for the facilities afforded by the directors and keepers of 
the following institutions: The Herbarium, Kew; the British 
Museum of Natural History, South Kensington; the Museum 
d’Histoire Naturelle, Paris; the Riksmuseum, Stockholm; the 
National Herbaria at Sydney and Melbourne, and the Botanic 
Museum and Herbarium, Brisbane. My thanks are also due to 
Dr. H. S. MeKee, Dr. A. C. Jermy, Mr. L. A. 8. Johnson, and the 
late Mr. A. H. G. Alston for their assistance in various ways. 

Royat Boranic GarpEns, SypNEY, AUSTRALIA. 


The Nomenclature of a Japanese Ophioglossum 
Hans P. Fucus 

In 1883, Carl Prantl (1883: 353) described a new species of 
Adder’s-tongue as Ophioglossum japonicum, basing his deserip- 
tion on material preserved in the herbarium of the Botanisches 
Museum, Berlin-Dahlem, that had been collected at Ujeno, 
Japan, by P. Hilgendorf! and near Tokyo by Wilhelm Doenitz, 
who collected in Japan during the period 1873-1879. Prantl 
overlooked or disregarded the existence of the earlier homonym- 
ous species Ophioglossum japonicum Thunb. (Thunberg 1784: 
329). Although the latter species is now referred to the genus 
Lygodium, as L. japonicum (Thunb.) Swartz, it still renders 
the later homonym OQ. japonicum Prantl illegitimate. 

Miyabé and Kudo (1916: 121) proposed the new name Ophio- 
glossum nipponicum, which is clearly a renaming of O. japoni- 

1 According to OS date, it seems that P. Hilgendorf collected 


in Japan as early as - The dates ‘‘1874-1876’’ given by Lanjouw and 
Stafleu (1957: 275) should therefore be corrected. 


JAPANESE OPHIOGLOSSUM 125 


cum Prantl (non Thunb.) from the synonymy cited and from 
the fact that it is indicated as a ‘‘nomen novum.’’ It seems like- 
ly that the specimens cited by Miyabé and Kudo from Hokkaido 
and Honshu belong taxonomically—at least in part—to 0. 
vulgatum L. This is, however, irrelevant, for the type of O. nip- 
ponicum must be the same as the type of O. japonicum Prantl. 
Therefore, as a species the name O. nipponicum Miyabé & Kudo 
is correct.” 

Overlooking the prior publication of O. nipponicum Miyabé & 
Kudé, William R. Maxon realized that O. japonicum Prantl was 
illegitimate and proposed the new name O. angustatwm (Maxon 
1923: 169). Two years later, Nakai (1925: 193) independently 
renamed OQ. japonicum Prantl as O. nipponicum Nakai, using the 

ame specific are nipponicum by a coincidence. One year 
cy Nakai (1926: 375) became aware of the apparent conflict 
and renamed is nipponicum as 0. Savatieri. All these later 
names are illegitimate, since they were superfluous when pub- 
lished, all being based ultimately on the same two syntypes cited 
by Prantl. 

There is however a name based on another type that has to be 
considered. In 1915, O. A. and B. A. Fedtschenko described an 
Ophioglossum ee L. var. bucharicum, based on a specimen 
collected by S. I. Korshinsky near Kabadiansk, in Bukhara, 
which is Lae in the herbarium of the Botanical Garden, 
Leningrad. Later, the same authors (1923: 8) raised their 
variety to specific rank, as O. bucharicum. Even though it is 
from quite a different locality, there is little doubt that this 
entity is conspecific with O. nipponicum. The illustration of 
Komarov (1934: pl. 3, fig. 9) matches perfectly material of O. 
nipponicum from Japan. Christensen (1934: 133) and Clausen 
(1938: 129) consider the names synonymous. 


2Specimens collected by Savatier (no. 1613) from the foot of Fudji 
Yama and reported by Franchet and Savatier (1876: 252) as 0. vulgatum 
belong to this species according to Nakai (1925: cee: 
also reported (erroneously) as O. nudicaule L. f. 
from specimens collected on the ‘‘Dunes de Sandai”’ by Pére Urbain Jean 
Faurie (no. 4294). 


126 AMERICAN FERN JOURNAL 


Recently, the Japanese student of Ophioglossum M. Nishida 
has considered the plant in question as only varietally distinct 
from O. thermale Komarov, a species name dating from 1914 and 
therefore having two years’ priority over 0. nipponicum, and 
therefore has proposed the new combination O. thermale var. 
nipponicum. It seems likely that he is correct that 0. nipponi- 
cum is only a morphologically slightly different form of 0. 
thermale, which ranges from Kamtchatka through Japan south to 

ormosa and India and west through China into the Pamir- 
Alai region in central Asia. However, as a variety the epithet 
bucharicum has priority and therefore the following new com- 
bination is neede 


OPHIOGLOSSUM THERMALE awa s Repert. Sp. Nov. oe 13 5 85. 1914, 
ucharicum (O. A. & B. A. Fedtschenko) H. P. Fuchs, comb, nov 
Ophioglossum japonicum Prantl, Ber. Deut. Bot. Ges. 1: 353. 1883; Jahrb. 
Bot. Gart. Berlin 3: 327, ve 8, fig. 29. 1884, non Thunb., 1784. 
oe vulgatum L. var. bucharicum O, A. & B . A. Fedtsechenko in 
Fedtschenko, Rastitel Sauk: imei 20. 1915. 
Ophiogossnm Se m Miyabé & Kudé, Trans. Sapporo Nat. Hist. Soe. 
, 1916 Be coll. partim 
Gonicsounee spa mO. A. & B. A. Vodicchenko, Not. Syst. Herb. 
Hort. Bot. cia sp ns Sak 15, 192 
eon angustatum Matey, Proe. Biol. Soc. Washington 36: 169. 


Dakine ‘Miseuie cum Nakai, Bot. Mag. Tokyo 39: 193. 1925. 
Ophioglossum Savatieri Nakai, op. as 40: 374. 1926. 

Ophioglossum thermale var. nipponi m Nishida ex Tagawa, Journ. Jap. 
Bot. 33: 203. July, 1958; ah ides op. cit. 34: 45, 124, 125. 1959. 
hase cae seipabion sensu auctt., e.g. ei anch. & Sav. Enum. Pl. Jap. 

252. 9, non L 
Ushisploane nudicaule sensu auctt., e.g. Christ, Bull. Herb. Boiss. 4 
675. Oct., 1896 et Makino, Bot. Mag. Tokyo 12: 376. 1897, non L. t 
BIBLIOGRAPHIC REFERENCES 
Curist, HERMANN. 1896. Filices Faurieanae. Bull. Herb. Boiss. 4: 664— 
675. Octo 
Sheba. ‘Cin. 1934. Index Filicum, Suppl. ITI, pp. [1]-219. October. 


Cuavusen, Ropert T. 1938. A Monograph of the Ophioglossaceae. Mem 
Torr. Bot. Club 19, pt. 2: [1]-177. July 


SCEPTRIDIUM IN JAPAN 127 


reat O. A. and B. A. 1923. De Generis Ophioglossum specie nova. 
ot. Syst. Herb. Hort. Bot. Petrop. 4: 8. Jan. 15. 

FRANCHET, eit and Lupovic avartke. 1879. Enum. Plant. Jap. [1]- 
9.3 


Komarov, V. L. 1934. Flora URSS. 1: [I]-XVI; [1]-302. 
Lansouw, J. and Frans Stariev. 1957. Index Herbariorum, Part. IT, no. 


bo 
4} 
oD 
i} 
oO 
+ 
~~) 
a 
=] 
= 
feo} 
=) 
e 
~ 
ee 
bo 


e; 95. December 
Maxon, WILLIAM R. 1923. Occasional Ni He on 1 Ola World Ferns, I. Proe. 
Bi oc. Washington 36: [169]-178. May 
Mivané, K. & Y. Kupo. Materials for a Flora of Hokkaido. Trans. Sapporo 
at. Hist. Soc. 6: 119-127. 1916. 
NaKAl, TAKENOSHIN. 1925. Notes on Japanese Ferns, II. Bot. Mag. Tokyo 
39: [176]-203. J 
—. 1926. Notes on rane Ferns, IV. Bot. Mag. Tokyo 30: [871]-400. 
July. 
PRANTL, ae 1883. Systematische Uebersicht der Ophioglosseen. Ber. 
Deut. Bot. Ges. 1: 348-353. 
isa. Beitrige zur Systematik des eke Jahrb. Bot. Gart. 
Berlin 3: [297]-350, pl. 7, 8. Preface dated Dece 
THUNBERG, C. P. 1784. Flora Japonica, pp. Yin tae 


A Synopsis of Sceptridium in Japan 
Maxoto NIsHIDA 

Lyon (1905) emphasized the presence of a suspensor in the 
embryo of Botrychium obliquum Muhl., which had been referred 
to the section Phyllotrichium, and took this up as the chief diag- 
nostic mark for the new genus Sceptridium, to which he trans- 
ferred all the species previously placed in the section Phyllo- 
trichium. However, almost all pteridologists since (Eames, 1936, 
for example) have not recognized the character of the suspensor 
as a valid diagnostic feature, for the gametophyte had been ex- 
amined only in the single species B. obliquum. Eames regarded 
it as a specific feature that is changeable, as it is in the order 
Marattiales within a genus or sometimes even within the same 
species, as in Angiopteris evecta Hoftm. I found a suspensor 
also in the embryo of B. japonicum and supported Lyon’s pro- 
posal for using the suspensor as a generic character (Nishida 


3 Concerning the dates ei Serge see Ohwi, Act. Phytotax. Geobot. 
2: 307, 308. November, 19: 


128 AMERICAN FERN JOURNAL 


1954, p. 55). Moreover, T have reached the conclusion that the 
genus Sceptridium would be acceptable by the appreciation of 
the significance of the vascular anatomy of the phyllomophore, 
the so-called ‘‘common-stalk’’ (Nishida, 1952, 

Tt is natural, I think, from the Agno of aur anatomy 
and sporangial structure, to divide the order Ophioglossales 
into two suborders, the Ophioglossineae and Botrychiineae, and 
three families—Ophioglossaceae, Helminthostachyaceae, an 
Botrychiaceae (Nishida, 1952, p. 59). The family Botrychiaceae 
contains the three genera Sceptridium, Botrychium, and Japano- 
botrychium (Osmundopteris). 


oy 


F BoTRYCHIACEAE 
Phyllomophore (‘‘common-stalk’’) oii: sterile blade with a long petiole, 
lon an or equal to the phyllomophore, producing a fertile blade 
from its basal part. Embryo endoscopic, with suspensor. 
ptridium Lyon 

Phyllomophore at least longer than the petiole of the sterile tases sterile 

blade sessile or with a short petiole producing the fertile blade. 
Small plant 5-20 (30) em. high, with pinnately or bipinnately divided, 
coarsely herbaceous, brownish or dirty green, glabrous sterile blades; 

u 


n 
took 
} 

o 
5 
° 
° 
iI 
tic} 
(os 
iJ 
(oz 
5 
oO 
Q 
° 
® 
=] 
= 
[=") 
oO 
4 
o 
&, 
io} 
S 
® 
f=") 
fo) 
i=} 
Q 
i=} 
tic} 
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i=} 
So 
[=] 
bale 
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=} 
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s 
of the basal parts of the phyllomophore; vascular divisions in the 
phyllomophore typically dichotomous; embryo exoscopic, without 
suupensor, 2 oe Botrychium Swartz 
Larger plants 30-60 (80) em. high, with 3- or 4-pinnately divided blades; 
sterile blades sessile, thin- beter eke or bright green, with 
seattered white hairs on the veins and veinlets on the dorsal (abaxial ) 
surface; buds densely ned nas white 05 developed in crevices 
opening at the bases of the phyllomophores; vascular branch-traces 
arising esi marginally ;! embryo exoscopic, without suspensor. 
anobotrychium Masamune? 
Scerrripium Lyon 
Seeptridium Lyon, Bot Gaz. 40: 457. 1905; Nishida, Journ. Jap. 
Bot. 27: 257. 1952. 
pide sect. Phyllotrichium Prantl., Bericht. Deut. Bot. 
Ges. 349. 1883; Jahrb. Bot. Gart. Berkin 3: 336. 1884; 
Seains Gen. Fil. 12. 1947 
1 Chrysler, 1945; Nishida, 1957. 
2 Tagawa, 1958. 


ScEPTRIDIUM IN JAPAN 129 
Botrychium subg. Sceptridium Clausen, Mem. Torrey Club 
92 2 


4.1 

Phyllo mibplhores short, 1-5 em. long (sometimes, in 8. dauci- 
folium, equal to the petiole of the sterile blade) ; medium-sized 
plants, (10) 15-40 em. high, with rather thick-herbaceous, dark 
green, 2—3-pinnately divided sterile blades, more or less hairy or 
glabrescent, densely covered with deciduous, long, white hairs in 
juvenile stages; vascular traces to sterile and fertile blades iso- 
marginal? or rarely and slightly extra-marginal (in S. multi- 
fidum) ; embryo endoscopic, with suspensor. 

Type: Sceptridium obliquum (Muhl.) Lyon, Bot. Gaz. 
458. 1905. Botrychium obliquum Muhl. ex Willd. in L. Sp. Plant. 
5: 63. 1810. Botrychium dissectwm Sprengel var. obliquum Clute, 
Fern Bull. 11; 89. 1903, sec. Clausen, Mem. Torrey Bot. Club 19: 
82. 1938 

Key T0 THE JAPANESE SPECIES OF SCEPTRIDIUM 
Spores with echinate exine; terminal segments of principal pinnae and 
pinnules elongate; ultimate indie segments of different sizes and 
shapes (Sect. toa 

Fertile blades not conspi eras overtopping the sterile and about equal- 

hyllo 


ling the latter in length; phyllomophore more or less elongate, about 
i ade S. daucifolium 
t, less than 


S. japonicum 
sterile e and pinnules not 
mate sterile segments all the same 


mish hairs on the dorsal 
; sterile blades with the 
short stalks of the deers: and pinnules not strongly tern 


ate. 
the principal sterile pinnae elongate; ultimate 
ey 


Terminal segments 
minus 


segments bob dentate or serrate. 
Terminal segments obtuse or acutish, not ade: “piniate segments 

entire or oe Het emmy roughly serrate 
Plants small, (20) em. high, not seni hairy; ultimate seg- 
ments eTuhaiy or rhomboid (or rarely ovoid), entire or crenate. 
S.m idee var. multifidium 


_ 
3 Nishida, 1957. 


130 AMERICAN FERN JOURNAL 


Plants large, (15) 20-40 em. high, sparingly hairy; ultimate seg- 
ments acutish, oblong or ovoid, crenulate or serrate. 
S. multifidum var. robustwm 
Sterile blades glabrous, rarely with a few, short (0.5-1 m ng), white 
hairs on the costae of the pinnae and pinnules; ene vane with 
the ear stalks of the pinnae and pinnules typically ternate 
Terminal segments of the principal pinnae and pinnules more or less 
elongate; ultimate segments acutish, with minutely serrate or cut 
margins; veinlets visible..__.......... S. ternatum var. nipponicum 
Terminal segments obtuse or acutish, not elongate; ultimate segments 
ovoid or oblong, with entire or evenly crenulate margins; veinlets 
not visible 8. 


ernatum var. ternatum 

A SYNOPSIS OF THE JAPANESE SPECIES 

I. Sect. Sceprripium. Botrychiwm eer Elongata Clausen, Mem. Torrey 
vie 19: 48. 1938 (as‘‘ Elongatae’ 


, Bot, Gaz. 40: 457. 1905. 
a daucifolium B japonicum nk Jatisbi Bot. Cart, Berlin 
3: 340. 1884, 


Botrychiuwm japonicum Underwood, ster Torrey Club 25: 538. 1898; 
Nakai, Bot. i Tokyo 40: 382. 
Nom. iri : O-hanawarabi. 


¢ Gentrat ae southern Japan, from Tanegashima Island, Kago- 
shima ne ea to Yamagata Prefecture; eastern China and Korea 
Hapitat: Shady bamboo thickets or forests 


eRe e261; ge 
agawa, Journ. Jap. Bot. 22: iat poe "Wakide Journ. Jap. B 
Si: 1956. 
Botrychium formosanum Tagawa, Act. Phytotax. Geobot. 9: 87. 1940. 
oM. JAP. Pelee 
Rance: Yakushim 


sland, Kagoshima Prefecture, and Aogashima 
Island, Izu Islands. Aogashima is the northern limit for this species. 
TA’ 


sts. 
da, comb. nov shan eine sect. Mul- 
orrey Club 19: 26. 1938 vi a jae’’). 


DIUM MULTIFIDUM (Gmelin) Nishida in Tagawa, eek ap. 
: 200. 1958. 


Osmunda multifida Gmelin Nov. Comm. Acad. Petr. 12: 517. 1768. 

Botryhi “malin m Rupr. Beitr. Pfi. ss. Reich. 717: 40. 1859; 
so Ged a Jap. Pteridophyta 19. 1957. 

a es yee anawarabi. 


ScEPTRIDIUM IN JAPAN 181 


Rance: Central and northern Honshu; Fagus and coniferous zones of 
the EBs hemisphere. 
Hasirat: Grassy open land and meadows, sometimes in deciduous 
Fagus-forests 
Var. ropustumM (Rupr.) Nishida in Tagawa, Journ. Jap. Bot. 33: 201. 


1958. 
Botrehiom rutifolium var. robustum Rupr. ex Milde, Nov. Act. Nat. 
es. Leop. 2 
Peet OH robustum Thilsrnood, Bull. Torrey Club 30: 51. 1903. 
Sceptridium robustuwm Lyon, Bot. : 458. 1905. 
Botrychium matricariae sensu iyibe et Kudo, Trans. Sapporo Nat. 
24.1 


Hist. Soe. 6: 
pcona hier gede sens tes aba: robustum ae Bull. Torrey Club 
64: 272. nis Torrey Club 19 . 1938. 


arabi. 
Rance: Common in Hokkaido (Yezo) and rare in northern Honshu; 
North Nace Alaska, ewidiete and Siber 
Hasitat: Shady woodlands in the cen zone, 
4. Sceprripium minus (Hara) Nishida in Tagawa, Journ. Jap. Bot. 33: 
201. 1958. 


Botrychium Japonioum var. minus Hara, Journ. Jap. Bot. 9: 127. 1933. 

Nom. Jap.: Usui-hanawarabi. 

Rance: Usui-toge Pass, near Karuizawa, Gumma eis and Mount 
Hakone, hegre aes in endemie in central Jap 

ABITAT: Rare in Fagus-z 

shat as the eae leraist segments of the pinnae and 
pinnules of this species are elongate, it would seem at first glance 
to belong to the section Sceptridium. However, the character of 
the spore-coat proves that it should be classed in section Multi- 
fida 


5. Sceprripium TeRNATUM (Thunb.) Lyon, Bot. Gaz. 40: 458. 1905. 
ar, TERNATUM 


Osmunda pats Thunb. Fl. Jap. 329, t. 32. 

ican ternatum Swartz in » ourn. Bot. aes 18002: 111. 1801; 
akai, Bot. Mag. kyo 40: « 1926. 

No p.: Fuyuno-hanawarabi 


RANG ntral and southern Jap: ranging from Sendai, Miyagi 
Prefecture, 40 Tanegashima, eeliigel. eae Korea, Formosa, China, 
and the coe ant Mountain 

fre vee grassy open lands, and woodlands, at elevations of 
less ee fe me 


132 AMERICAN FERN JOURNAL 


Var. NIpponicum (Makino) Nishida, Journ. Jap. Bot. 34: 137. 1959 
Botrychium nipponicum Makino, Journ. Jap. Bot. 1: 5. 1916; Nakai, 
Bot. M Tokyo 40 383. 2 
fe) a-hana 


2A 
AN! ie ae Give: poe Gifu, Nara, and Okayama Prefectures; 
es in eee Japan. 
Hasitat: In woodlands in lowlands. 


Dr. Makino indicated that Botrychium nipponicum was very 
closely allied to B. japonicum, differing in having a latericious- 
colored frond, conspicuous veins, and shorter and closer serra- 
tion. Inasmuch as the chief terminal segments of the sterile 
pinnae are somewhat elongate, he considered that this species 
must be allied to B. japonicum. However, the former has the 
segment margins minutely serrate instead of roughly dentate 
as in B. japonicum. I take the character of the spore-coat as 
a diagnostic feature distinguishing this species from B. japoni- 
cum. Going on this spore-character, B. nipponicum is closely 
allied to B. ternatum and I consider it only varietally separable. 
Sceptridium ternatum itself is so closely similar to S. multifiduwm 
that the two are very difficult to distinguish. Although the for- 
mer may be an ecological variant of the latter, I separate them 
tentatively by the diagnostic features mentioned in the key. 
In S. multifidum, the texture of the sterile blades is thick and 
coarsely herbaceous, rather than thin and more or less mem- 
branaceous as in S. ternatum. In S. multifidum there is a mem- 
branaceous sheath, the remnant of the withered phyllomophore 
of the preceding year, persistent at the base of the phyllomophore 
of the current year; such a sheath is lacking in S. ternatum. 

BIBLIOGRAPHIC REFERENCES 
Eames, A. J. 1936. Morphology of Vascular Plants, Lower Groups. New 
York. 
Curyster, A. M. 1945. Bull. fobs Club 72: 491-505. 
Lyon, H. L. 1905. Bot. Gaz. 40: 
Nisuipa, M. 1952. Journ. ee ha 27: 165-- 171, 271-278. 
954. 


—. cit. 29: 239-244, (In Japanese with English summary). 
—. 1955. Pistoinermhices 5: 449-456, 


ASPIDIUM DISTANS 133 
—. 1957. Journ. Coll. Arts & Sei. Chiba Univ. 2: 179-211. 


. o. 
Tagawa, M. 1958. Journ. Jap. Bot. 3. 
Curpa Untversity, CHIBA, JAPAN. 


The Identification of Aspidium distans Viv. 
Ropoutro E. G. PicHi-SERMOLLI 


Aspidium distans was described by Viviani! from specimens 
collected ‘‘in Corsicae sylvis Allo Spedale’’ by an old student of 
his, Dr. Stefano Serafini. Viviani gave a short description of 
his species and, following the conceptions of his time, properly 
referred it to the genus Aspidium. Nevertheless, his species was 
misinterpreted by succeeding authors since the first years of its 
publication. 

In 1827, Sprengel? regarded Aspidium distans as a synonym 
of Woodsia ilvensis (.) R. Brown, in spite of the fact that 
Viviani gave the dimensions of the fronds as two feet long or 
more. Bertoloni? studied a specimen sent to him from Viviani 
and treated A. distans as a good species; he described the fern 
more carefully and pointed out Sprengel’s mistake too. Never- 
theless, Aspidium distans continued to be referred to Woodsia 
ilvensis by Moore,* Milde,® Luerssen,® and other authors. 
Viviani’s species was considered again by Cesati, Passerini, and 
Gibelli,? who regarded it as an independent species and pro- 
posed for it the new combination Nephrodium distans.. How- 
ever, in a footnote, they observed that it remained to be seen if 
the species corresponded to Aspidium paleaceum Don. 


1 Appendix oa hee Somer Prodromum 8. 1825. 
yst. Veg. 41: 125. 


. 90. 1858. 
5 Fl. Ew urop. Atl. 165. 1867. 
6 Farnpfl. i a Krypt. Fl. Deutsehl. ed. 2, 3: 508. 1887. 
? Comp, Fi. "ie 1, 18, 1868. 
binomial was overlooked by Christensen in the gta a peat 
oan foe listed only in the first ““Supplemen ntum’? - 19138): Tt is 
later homonym of N. distans Hook., from Madagas' 


AMERICAN FERN JOURNAL VotuME 50, PLATE 13 


Ee 


LECTOTYPE OF ASPIDIUM DISTANS IN HERB, BERTOLONI, BOLOGNA 


ASPIDIUM DISTANS 150 


A more careful study was carried out by Trevisan, who ex- 
amined Viviani’s type, kept at that time in = Genoa herbarium. 
In the part of his very interesting paper ‘‘Sylloge Sporophy- 
tarum Italiae,® dealing with this fern, Trevisan gave some in- 
formation on the type material and on the identification of 
Viviani’s species. He pointed out that Viviani’s specimens in 
the Genoa herbarium, although very young, were fertile and that 
the sori were clearly smaller than those of Polystichum filix-mas. 
He regarded Viviani’s fern as nearer to Polystichum filiz-mas 
var. borreri (Newm.) Trevis. than to any other, differing in 
having the stipe very densely covered with scales, the blades 


erenate, and the piliform scales of the under-surface and the 
edges of the segments longer and denser. He coneluded that this 
fern belongs in the genus Polystichum (in his sense, i.e. Dryop- 
teris sensu Christensen) and is an independent species for which 
he proposed the new combination Polystichum distans.!° How- 


than those of Viviani, Trevisan clearly expressed some doubt 
that P. distans coud be kept distinet from P. filix-mas. 
Probably influenced by the uncertainty shown by Trevisan in 
the above-mentioned note added in proof, Fiori'! regarded 
Viviani’s fern as a variety of the male fern, and proposed the 
new combination Nephrodium filix-mas var. distans (Viv.) Fiori. 
Some time later, the same author’? treated this plant as a syno- 
nym of Polystichum filix-mas (.) Roth, and still more recently 
he considered Aspidium distans Viv. as a doubtful species.13 A 
different identification of the species was made by Briquet,'* who 


9 Atti Soc. Ital. Sei sata wee 213-258. 1875. 
10 Polystich um distans (Viv.) Trevis. was not listed in bahipiongce iy 
Pose uae Filicum”’ (1908) ont only in the first ‘‘Supplementum’’ (p. 
13)..0 ater homonym of P. distans Fourn, (1872) 
a In Fiori & ic gers Fl. Anal. on ao 907. 


Fiori . Ita 
18 ba Fi. Ital. Seat. D, Pharid. Sok oat, 
14 Prodr. Fl. Corse 1: 11. 1910. 


136 AMERICAN FERN JOURNAL 


regarded Viviani’s fern as a synonym of Dryopteris rigida 
(Hoffm.) Underw. var. australis (Ten.) Briq. In more recent 
times, Wolf!® regarded A. distans as doubtfully belonging to 
Dryopteris borreri Newm. Finally, Rothmaler'* referred Viviani’s 
fern to Dryopteris paleacea ‘‘(Sw.) Hand-Mazz.’’ 

In summation, Aspidiwm distans Viv. has been treated as fol- 
lows in terms of modern nomenclature: (1) As an independent 
species of Dryopteris allied to D. borreri; (2) as a synonym of 
Woodsia ilvensis; (3) as an independent variety of Dryopteris 
filix-mas; (4) as a synonym of Dryopteris filix-mas; (5) as a 
synonym of Dryopteris villarti (Bellardi) Woynar var. australis 
(Ten.) Fiori; (6) as a synonym of Dryopteris paleacea ‘‘ (Sw.) 
Hand-Mazz.’’; and (7) as a doubtful species. 

Trevisan!’ informs us that Viviani received four specimens 
from Corsica, evidently collected by Dr. Serafini from the same 
rhizome. Two of these, those studied by Trevisan, were kept 1 in 


completely destroyed during the war, these specimens were 
missing in the Genoa herbarium.'* Another specimen was pre- 
sented to Bertoloni by Viviani and is still present in the her- 
barium of the Botanical Institute of the University of Bologna 
(pl. 13). It bears the following label: ‘‘ Aspidiwm distant Viv.— 
Bert. Fl. Ital. Crypt. p. 53, no. 4—Dedit ipse Viviani Genuae 
1827 augusto mense.’’ The fourth specimen is preserved in 
Cesati’s herbarium in the Botanical Institute of the University 
of Rome. It is labelled as follows: ‘‘Aspidium distans Viv. 
(Corsica)—(ex Herb. Viviani)—Polystichum distans Trev. in 
Atti Soc. Se. Nat. Ital. vol. XVII, p. 227—Trevisan assicura di 


prossimo al Polystichum Filiz-mas var. Borreri. “GN pkpodtion 


15 Pollichia, ser. 2, 5: 97, 106. 

so erasers is 168, 1943. ef. ah Reichling, Bull. Soc. Bot. Belg. 86: 
17 Atti Soe. Ital. Sci. Nat. 173: 227, 228. 1875. 
18 Cf, Tlario, Arch. Bot. 13: 203. 193 t, 


ASPIDIUM DISTANS iY ¢ 


Viviani mihi cum jam extet N. distans Hook, sp. IV. p. 76. Bak. 
syn. 288. No. 160 ex Madagasear.’’® 

T have examined both the species of the Bologna and the Rome 
herbaria. They agree with each other closely and consist of 
young fronds destitute of rhizomes. Contrary to the plants 
examined by Trevisan in the Genoa herbarium, both are sterile. 
In spite of the lack of sori, a careful examination of these speci- 
mens showed me clearly that they belong to that fern which is 
named Dryopteris borreri. Consequently, Aspidium distans Viv. 
and Dryopteris borreri are taxonomic synonyms. Since Viviani’s 
epithet has priority over Newman’s binomial, a species ought 
to change its name, and ought to be named D. ans, but unfor- 
tunately for the principle of priority but senate for the 
stabilization of nomenclature, the combination Dryopteris dis- 
tans based on Viviani’s epithet would be illegitimate owing to 
the existence of the earlier homonym D. distans (Hook.) Kuntze. 
Consequently, the well-known name D. borreri will still stand 
as the legitimate name of this fern. The synonymy to be added 
to aa of Dryopteris borreri Newm. may be summarized as fol- 


bee BORRERI Newm. Hist. Brit. Ferns ed. 3, 189. 1854. 
Aspidium distans Viv. App. Fl. Cors. Prodr. 8. 182 
Nephrodium Pathe ites Guest Pass. & Gib. Comp. Fl. Ital. 18. 1868, non 


Hook, 
neat p Witans Trevis. Atti Soe. Ital. Sci. Nat. 173: 227. 1875. non 
ourn, F 
Nephrodium filiz-mas var. distans Fiori, in Fiori & Paol. Fl. Anal. Ttal. 


Dryopteris borreri has been considered as an independent - 
species by many authors, while it has been merged with D 
paleacea by others, e.g. Rothmaler (op. cit.). I do not intend to 
go into the taxonomic value of D. borreri here, since T am not 
acquainted enough with its extra-European allies, but in any 
case, as already shown by Alston, the names Dryopteris 

ently this note was written by Cesati after the publication of the 
part eating Bags the ferns in the above-mentioned Bhp mpendio della 
ali ? by Cesati, Passerini, and — belli. So far ee know, the 
name Nephrodium viviant was never publis hed. 


138 AMERICAN FERN JOURNAL 


paleacea (D. Don) Hand.-Mazz., Dryopteris paleacea (Swartz) 
. Chr., and D. paleacea “© (Swartz) Hand-Mazz.,’’? with ‘‘hy- 
bridized’’ authors as proposed by Rothmaler, must be rejected 
as illegitimate.2° If the European D. borrert Newm., the Ameri- 
ean D. parallelogramma (Kunze) Alston [= D. paleacea 
iyi ©. Chr.], and the Asiatic D. wallichiana (Spreng.) 
1. [= D. paleacea (D. Don) Hand.-Mazz.] are treated as a 
de species, the correct name for it is D. wallichiana (Spreng. ) 
Hyl. If only two species are recognized, D. borreri would be bet- 
ter treated in some infraspecific category under D. parallelo- 
gramma, since it is nearer to the American species than the 
Asiatic one. However, as far as I am able to see from examina- 
tion of some specimens, the Asiatic, the American, and the 
European appear to represent three distinct species. 
Istrruto Boranico ‘‘Hanpury,’’ Universita, Genova, ITALY. 


Hybrids in the Genus Asplenium Found in Northwestern 
and Central Europe 
EYER 

Hybrids in the genus Asplenium have been known for many 
years. Presumed parentages were at first purely speculative, but 
recent genetical and cytological work, including breeding and 
chromosome counts, have enabled us to determine the parentage 
with accuracy in many instances. The known hybrids are listed 
below, with some charts eee the probable ancestry. 


SPE 
1. ASPLENIUM ADIANTUM-NIGRUM ie Qn = 12, 2n = 144. 


2. A. onopteris L., 

3. A. FORSTERI Sadler. 2 at 72, 

4. A. rissum Kit., 2n = 72. 

5. A. opovatum Viv. em. Becherer, 2n = 144. 
6. A. FORESIACUM sina Christ, 2n = 7 
7. A. FONTANUM L. 2n = (A. hallert). 

8. A. LEPIDUM Presl, 2n = . 


- 207 This JournaL 47: 91, 92. 1957. 


—_ 


See eee 


ASPLENIUM HYBRIDS 139 


RUTA-MURARIA Li, 2n = 144. 
SEPTENTRIONALE (li.) Hoffm., 2n = 144. 
SEELOSII egy on S212, 

ARINUM in (2. 
tisonee ani L. em. Huds., 2n = 72, 2n = 144. 
ADULTERINUM Milde, 2n = 144, 
VIRIDE Huds., 2n = 72. 


. PHYLLITIS SCOLOPENDRIUM (L.) Newm., 2n = 72. 


CETERACH OFFICINARUM Lam. & DC., 2n = 144. 


HYBRIDS 


. ASPLENIUM X GERMANICUM Weis, Pl. Crypt. Fl. Goetting. 


299. 1770, 2n = 108. Asplenium septentrionale X_ tri- 
chomanes (n = 36). Figure: Eberle 1959b, pp. 72, 76. 
X HE i 


Abh,: 95. 1859, 2n — 144. A. septentrionale x tricho- 


manes (n = 72). Figures: op. cit. t. IV; Eberle 1959b, 
pp. 73, 14, 75; 77). 


3. A. X BAUMGARTNERI Doerfler, Oesterr.\ Bot. Zeitschr. 45: 


ou 


169, 1895, 2n = 144. A. trichomanes (n = 72).. X sep- 
tentrionale. Figure: Op. cit. 

. < MURBECKI! Doerfler, Genter. Bot. Zeitschr. 45: 223. 
1895, Qn — 144. A ruta-muraria * septentrionale. Fig- 
ure: Lunds Univ. Arsskrift 27, t. IZ. 1892. 


A. X suEvicum Bertsch ex Meyer, Ber. Deut. Bot. Ges, 72: 


40, 1959, 2n = 144. A. septentrionale X ruta-muraria. 
Figures: Veroeffentl. Wuertt. Landesst. f. Naturschutz _ 
(Stuttgart), Heft 79: 83; Eberle 1959b, pp. 78, 79 


6. A. X cLERMONTAE Syme, Engl. Bot., ed. 3, 12: 132. 1886. A. 


ruta-muraria X< trichomanes. Figures: Verh. Zool.-bot. 
Ges. Wien 55: 13. 1905; Alston 1940, p. 141, fig. 22. 


7. A. X Lusaticum D. E. Meyer, Ber. Deut. Bot. Ges. 71: 16. 


1958, 2n = 108. A. trichomanes (n = 72) X trichomanes 
(nm = 36), 


8. A. X BAvaricum D. E. Meyer, Ber. Deut. Bot. Ges. 71: 16. 


1958, 2n = 108. A. trichomanes (n = 72) X viride. 


140 AmeEricAN Fern JOURNAL 


A. X poscHARSKYANUM (Hofm.) Doerfl. Herb. Norm. no. 
3670, Schedae 234. 1898, 2n = 108. A. adulterinum X 
viride. Figure: Ber. Deut. Bot. Ges. 70: 59. 1957. 

TABLE I 


tre XX 
X hybrids 22 ma 
OQ not discovered 74 seel O 
till 1959 
u 


Po ote COCO OO Ox Oo g 
% 
Go 6%G 6 0 0 6 <0 © 9.0/0 OX & 


G ox 6 6.0 <6 0 0-0 9° 


0 
0 
x 
xX 
0 
0 
x 
ie} 
0 
x 
O 
x 


© ©. 0 10°76. 6756 9.9 
6 0 6.o.6°O O 0 9-9 6 


adi 0° 0. 0 0X 
ono! S fiss??° fore Mt ep ruta <6, Seel ng tri ai vir Phy Cet 
Hybrids between species of the Aspleniaceae 
in Northwestern Europe 
10. A. X TRICHOMANIFORME Woynar, Mitt. Naturwiss. Ver. 
Steiermark 49: 153. 1913. A. adulterinum X trichomanes. 
A. RUTA-MURARIA X virIDE D. E. Meyer, Ber. Deut. Bot. 
Ges.7.1: 49. 1958, 2% 108. 
12. A. < GAuTIERI Christ in Burnat, Matér. Hist. Alpes Marit. 
15. 1900, 2n = 54? A. fontanum X viride. Figure: Ber. 
Deut. Bot. Ges. 70: 59. 1957. 
13. A. < coRBARIENSE Rouy, Fl. France 14: 543. 1913. A. fon- 
tanum X trichomanes. 


ay 
~ 


ASPLENIUM HysBrips 141 


14. A. < REFRACTUM (Moore) Lowe, Ferns Brit. & Exot. 5: 103. 


Brenls x gi nig H BADENSE D. 


15, 


1858. A. obovatum * Tetelcsaguse Figures: 


A. < paGEesu Litard. Bull. Géogr. 


Lowe, op. cit. 


t. 835A; Alston 1940, p. 141, fig. 2A. 


Mey ER. a ASPLENIUM 


TA-MURARIA X CETERACH peters 
Bot. (LeMans) 20: 204. 
1910. A. foresiacum X< trichomanes. Figure: Brit. Fern 
taz. 6: 307. 1935. 
A. X costrr Litard. Bull. Géogr. 
1911. A. foresiacum X septentrionale. 
A. X sAvorKAr Kuemm. Mag. Bot. Lapok. 21: 1. 1923. A. 
lepidum < ruta-muraria. 
A. < woynartAnum Aschers. & Graebn. Syn., ed 2.4: 126. 
1913. A. forsteri X viride. 
A. X WACHAVIENSE Aschers. & Graebn. Syn., ed 2, 2: 125. 
1913. A. forsteri < trichomanes. Figure: Verh. Zool.-bot. 
yes. Wien 6: ¢. II, f. 1, 2. 1856. 


Bot. (Lemans) 22: 150. 


142 AMERICAN FERN JOURNAL 


TABLE I 
Species and hybrids in Central Europe 


Synthesized| and cytological investigated till 1959 


Ceterach officinarum 
4x,144 


x Asplenoceterach badense 
4x,144 


fontanum 
2x72 
¥ adulterinum 
4x,144 
(x ‘gdutiers) 
2,54 


N 
N 
‘ 


x poscharskyanum 
3x,108 
a “ 
x bavaricum viride 
(4) 


72 ~~ ruta-muraria x viride 
mats 3x,108 


trichomanes [so x heufleri 
4x, 144 4x, 144 


x baumgartneri 
4x,144 


x lusaticum 
3x, 108 


4x,164 4x,164 
ao $9 oe een 9 
trichomanes & g septentri G 
2%, 72 4 


ntrionale & o@ ruta-muraria 
x, 164 4x,144 


Asplenium 
x=36 


ASPLENIUM HYBRIDS 143 


20. A.  MURARIAEFORME Waisbecker, Oesterr. Bot. Zeitschr. 
49: 63. 1899. A. forstert * ruta-muraria. 

21. A. X prrarpi Litard. Bull. Soc. Bot. Deux- Sevres 1909- 
1910: 109. A. adiantum-nigrum X ruta-mura 

22. A. X soucHE! Litard. Bull. Soc. Bot. Deux- Sarre 1909- 
1910: 100, ¢. 1, 2. A. adiantum-nigrum X septentrionale. 

23. A. X poLosum Milde, Verh. Zool. Bot. Ges. Wien 14 (I): 
165. 1864. A. adiantum-nigrum X trichomanes. Figure: 
op. cit. t. XVIII 


Fig.2 


septentrionale 
mauane x heufleri 
4x,144 


x germanicum ‘ cn 
3x,108 x baumgartneri : 
Asplenium bx, 16h ete 7 lee 
24. > AspLENOPHYLLITIS sAcKSoNII Alston, Proc. Linn. Soe. 
London, Sess. 152 (2): 142. 1940, Asplenium adiantum- 
nigrum >< Phyllitis scolopendrium. Figure: op. cit. 
D. 


fig. 2D. 

25.  ASPLENOPHYLLITIS CONFLUENS (Moore) Alston, Proce. 
Linn. Soc. London, Sess. 152 (2): 139. 1940. Asplenium 
trichomanes X Phyllitis scolopendrium. 

26. )< ASPLENOPHYLLITIS MICRODON (Moore) Alston, Proc. Linn. 


144 AMERICAN FERN JOURNAL 


tum * Phyllitis iecsipemoutante Figure: op. cit. Fig. 2B 

ondon, Sess. 152 (2): 140. 1940. Aspleniwm obova- 

ato xX ub EAGG 'TERACH BADENSE D. EK. Meyer, Ber. Deut. Bot. 

Ges. 70: 61. 1857, 2n = 144. Asplenium Mies Nitibaess ~~ 
Ceterach officinarum. Figure: op. cit. p. 63. 


Fig. 3 
x Aspleno \ceterach 
badense \ 4x,144 
ehhh ruta-muraria 
: eckii 4x,144 3 
Asplenium bx, 166 . Ceterach ffofficinarum 


4x,166 


BIBLIOGRAPHIC REFERENCES 

Auston, A. H. G., 1940. Notes on the Supposed Hybrids in the Genus 
Asplenium Found in Britain. Proc. Linn. Soe. London, Sess. 152, 
pt. 2: 132—144, 

“a eeerea: P. & P. GRAEBNER, 1913. Synopsis der mitteleuropiischen Flora, 
ed. 2, vol. 1. Leipzig. 

Curist, H., 1900. ne rare der Schweiz. Beitr. Kryptogamenflora 
ae. vol. 1, 

EBERLE, G., 1957. hie ne Sane und Heuflers Streifenfarn (As- 
seta abi germanicum und A. heufleri). Jahrb. Nassau. Ver. Naturk. 
ee 


—, 1959a. Gtreliemeanne der Alpen und die Aufklirung der Entstehung 
. oe ae Mischlinge. Jahrb. Ver. Schutze Alpenpflanz. 
25—25. 


_, ue Parks | im Herzen Europas. Frankfurt a.M. 


Ferns KNown As RuMOHRA 145 


—, 1959¢. Altes und Neues vom Schriftfarm (Ceterach officinarum). Natur 
u. Volk 89: 229—236. 
Hota 1936. Histoire d’une Fougére Hybride de la France, Asplenium 
(Cossonianum) Murbeckii. Bull. Soc. Natural. Archéol. Ain (Bour Zz) 
50: 210—2 


Mever, D. E. 1952. Untersuchungen ueber Bastardierung in der Gattung 
Aspleniwm. Bibl. Bot., Heft 123. Stuttgar 
—, 1957. Zur gc wes der Asplenien Midbcloneensé: Ber. Deut. Bot. Ges. 


—, 1958. Ibid., nS cit. 71: 11—20. 
1959. nk cit. 72: 37—48. 
ivnende: Ss. a0. Tvenne Asplenier, deras affiniteter och genesis. Act. 
Univ. Lu na 
Hi accciahad Garren uNpD Museum, Beruin-DaHLEM, GER- 
MANY. 


Observations ad neat Ferns VI. The Ferns 
y Known as Rumohra 
nd V. Morron 

In ‘‘Standardized Plant Names’’ (1942), Dr. Maxon and I 
assigned (or I should probably say invented) the common name 
“Rast Indian Holly-fern’’ to a plant that has usually been 
known as Polystichum aristatum (Forster) Presl. It is perhaps 
not uncommon in cultivation in greenhouses and in the open in 
southern California. A related species, which can conveniently 
be known as Standish’s Holly-fern, was Polystichum Standishii 
(Moore) C. Christensen; it is less common, both in nature and 
in cultivation; it is however hardier, and I have a specimen at 
hand that is said by the collector, Mr. J. William Kingma, to 
have come from plants grown in the open in East Grand Rapids, 
Michigan. 

These plants are rather large and coarse. They differ from 
Polystichum in some respects: The indusium is attached lateral- 
ly, rather than peltately (i.e. centrally) as in Polystichum; the 
rhizome is creeping and the leaves are borne singly along it at 
intervals (rather close intervals, I might add), rather than being 
erect, with the leaves borne spirally in a crown as in Poly- 
stichum; the fronds are more or less broad and triangular, and 


146 AMERICAN FERN JOURNAL 


tripinnate to quadripinnate, with the basal pinnae enlarged on 
the basal side (i.e. basiscopically developed), whereas the fronds 
of Polystichum are mostly elongate (oblong to linear in outline), 
only once- or twice-pinnate, and the basal pinnae are not basi- 
scopically developed. More importantly, the structure of the 
blade is anadromous throughout, that is, the first basal pinnule 
to arise is that facing the apex of the blade, and the first second- 
ary pinnule to arise is that facing the apex of the pinnae, and so 
forth down to the ultimate veins. Polystichum (and true Dryop- 
teris generally) is at least partly catadromous, some of the first 
pinnules either facing the base or being opposite the anterior 
pinnules. Taken altogether, these differences are significant. 

R. C. Ching, in a paper entitled ‘‘A Revision of the Compound 
Leaved Polysticha and other Related Species in Continental Asia 
Including Japan and Formosa’’! decided that these two species 
and their allies could be separated generically from Polystichum, 
and he chose for them the generic name Rumohra Raddi.? The 
type of this genus is Rumohra aspidioides Raddi, a Brazilian 
species which is considered, I imagine properly, to be a synonym 
of Polystichum adiantiforme (Forster) J. Smith, a common 
species occurring in many widely-separated parts of the world. 
In cultivation and in dealers’ catalogues, P. adiantiforme is 
often listed as ‘‘leather fern’’ under the synonymous names 
Polystichum capense and P. coriaceum, or sometimes even under 
the old and entirely incorrect name Aspidium capense. Cope- 
land, in his Genera Filicum, followed Ching in uniting aristatum 
and its allies with adiantiforme, and even emphasized the ‘‘natu- 
ralness’’ of the resulting genus, but nevertheless such a con- 
clusion is highly debatable. 

Rumohra adiantiformis is primarily an epiphyte, with a long, 
stout, densely scaly, dorsiventral rhizome that climbs on tree- 
trunks, or occasionally on rocks. It has a strictly peltate in- 
dusium as in true Polystichum. It is very likely not phyloge- 
netically close to aristatum and its allies. Dr. R. BE. Holttum, in 

1 Sinensia 5: 23-9 

2 Raddi, Opuse. Bei, Hee S: 290. pl. 12, fig. 1. 1819. 


Ferns Known As RUMOHRA 147 


his fine paper on the classification of leptosporangiate ferns,® 
came to the conclusion that true Rumohra is generically differ- 
ent from aristatum. He would ally it with Davallia, which seems 
a rather remote alliance. He states that the structure of the 
rhachis is of the davallioid type, that is that the two ridges of 
the rhachis are continuous with the leaf-margins. He may be 
right, for the structure does approach that of Davalla, but it is 
not quite so obvious; the ridges seem to arise from an intermedi- 
ate area, neither from the margin as in Davallia nor from the 
margins of the costal groove as in Polystichum. 

Rumohra should therefore be considered as a monotypic genus 
of uncertain alliance. Its geographic distribution shows that it 
is a very old type, for it occurs in widely separated parts of the 
world, as in New Zealand, South Africa, and Chile and Argen- 
tina; it has also spread far to the north. Its synonymy is as 
follows: 


RUMOHRA ADIANTIFORMIS (Forster) Ching, Sinensia 5: 70. 1934. 


Polystichum capense J. Smith, Bot. Mag. Comp. 35. 1846. 

Polystichum adiantiforme J. Smith, Hist. Fil. 220. 1875. 

Inasmuch as Holttum rejected Rumohra as an available name 
for aristatum and its allies, he was obliged to find another, and 
he chose Polystichopsis, calling the genus Polystichopsis (C. 
Chr.) Holttum, with a reference to Christensen in Verdoorn, 


ence is given to Rumohra of Ching, in part, and going back to 
Ching’s paper we find the citation of Dryopteris subg. Poly- 


3 Journ. Linn. Soc. Bot. 63: 137, 152. 1946. 


148 AMERICAN FERN JOURNAL 


stichopsis C. Chr., with a reference to Christensen’s Monograph 
of the Genus Dryopteris, Part II. In tracing this citation back, 
we find that Christensen’s subgenus was based on Lastrea, sect. 
Polystichopsis J. Smith.® Therefore, the proper authority for 
the genus is Polystichopsis (J. Smith) C. Chr 

John Smith’s section Polystichopsis was designed to include 
all the species of Lastrea (in his sense equalling Dryopteris 
sens. lat.) with decompound deltoid fronds, and thus included 
quite a number of species that may not be closely allied, such as 
L. pubescens, L. hispida, L. decomposita, and L. funesta. Chris- 
tensen restricted the application of the name in his monograph 
to L. pubescens and its allies, and in Index Filicum, Supple- 
mentum Tertium,® he definitely cited Dryopteris pubescens (1.) 
Kuntze as the type of the section Polystichopsis (J. Smith). The 
type of the genus Polystichopsis is therefore the same species, 
D. pubescens, for Christensen’s choice of lectotype can not be 
changed or even challenged, for this is one of the original species 
and it agrees with the original brief characterization and there- 
fore qualifies as a lectotype on all counts. 

The determination of the lectotype is of some significance, 
because D. pubescens and its near allies D. chaerophylloides and 
D. lurida diverge in some respects from the group of aristatum. 
Christensen was content to leave D. pubescens and D. aristata in 
the same section, and Ching (and also Copeland) regarded them 
as unquestionably both Rumohra. Still, there are differences. 

Polystichum and Dryopteris are characterized by the absence 
of hairs in the channels of the upper leaf surface (and also else- 
where, except as capitate glands may be called hairs and reduced 
seales may be hair-like, as in D. Filiz-mas) ; the fronds are usu- 
ally evergreen, and of a coriaceous texture, shining 
upper surface and with mucronate or aristate teeth; the stipe 
bases are densely scaly. In all these characters retains agrees 
with Polystichum. On the other hand, Dryopteris pubescens has 

4In Dansk. Mae bey: Skr. Nat. Afd. VIII, 6: 101. 1920. 


5 Hist. os ms 
6p, 7 


Frrns KNown As RUMOHRA 149 


abundant, long, silky, septate hairs on the blades; the texture is 
soft-herbaceous and the teeth are scarcely aristate; the stipe 
bases are abundantly hairy, as well as sometimes scaly. It seems 
that consistency necessitates the generic separation of pubescens 
and aristatum; the latter is unquestionably a near ally of Poly- 
stichum and Dryopteris; the former must have its origin else- 
where. 

The group of aristatum is lacking an available generic name. 
The following is proposed. 
Byrsopteris Morton, gen. noy. 

Rhizoma crassum hypogaeum repens dense paleaceum, paleis 
magnis fibrosis elongato-lanceolatis, attenuatis, non ciliatis nec 
pilosis nee fimbriatis, subintegris, dentibus remotis ex eellulis 


et fimbriatis, pinnis primariis saepe paucis, duobus inferioribus 
maximis deltoideis basiscopice dilatatis, pinnula basali inferiore 
es 


fur e 
attingentes, apice saepe in hydathodis terminantes; sor1 globosi 
in venis terminales vel dorsales indusiati, indusiis crassis 
clauso a 


coriaceis subrotundis lateraliter in sin xis glabris 
integri pe persistentibus, parietibus 1 inerassatis 
glandulosis; sporangia numerosa longe pedicellata, eellulis in- 
duratis annuli el 14; paraphyses nullae; sporae bilaterales 


150 AMERICAN FERN JOURNAL 


Rhizome thick, hypogaeus, creeping, densely scaly, the scales 
large, fibrous, elongate-lanceolate, attenuate, not ciliate, pilose, 
nor fimbriate, subentire, the teeth remote, composed of two 
One agg en the internal cells of the scales elongate, with thick, 
br nslu ucent, lateral walls and thin, yaline, brownish 
arienise ails: leaves relatively few, distant or f adiacent on the 
creeping rootstock but not fasciculate, long-stipitate, the stipes 
straw-colored, thick, glabrous, suleate, strongly scaly at base, the 
vascular bundles three or more; leaf-blades decompound, deltoid 


r pentagonal, tripinnate to 5-pi ; achises straw- 
colored, glabrous, scaly, the scales entire or denticulate, some- 
times dila the and the briat $ ees innae 


margins dentate or mostly subspinulose; rhachises, rhachillae, 
and costae deeply channelled on the upper (adaxial) side, the 


indusiate, the indusium thickly coriaceous, subrotund, 

laterally attached at a closed sinus and thus appearing centrally 
peltate, entire, the cell-walls thickened and sometimes apparently 
glandular ; sporangia numerous, long-stalked, the annulus 13- or 
14-celled; paraphyses none; spores monolete, large, 43-61 p 
long,* obviously crested, the crests rch in some species. 

Typus: Polypodium aristatum For: 

The two ee that are known in pe may be distin- 
guished as follo 
Seales of the shield and rhachillae dark, broadened and 

fimbriate at the base, abruptly attenuate to an elongate, fili- 

form apex, one cell thick; sori terminal on the veins; blade 


7 Not measured in all species. 


AMERICAN FERN JOURNAL VoLuME 50, Piate 14 


2 


BYRSOPTERIS ARISTATA (TYPE OF POLYPODIUM ARISTATUM Forst. British 


Museum. PuorograPH BY C. V. Morton.) 


152 AMERICAN FERN JOURNAL 


coriaceous, very thick in texture, with spinulose-toothed seg- 
it aristata 

Scales of the blade pale, not much broadened or fimbriate at base, 
gradually attenuate, but two-celled almost or quite to the 
apex; sori dorsal on the veins; blades papyraceous, the seg- 
ments mucronate but hardly apiniiiose ee ee B. Standishii 


YByrsopteris aristata (Forster) Morton, comb. nov. ees a 
Polypodium aristatum Forster, Fl. Insul. Austr. 1786. 
Aspidium aristatum Swartz in Journ. Bot es rader ie. 4 1801. 

e 
ryopteris aristata ee Rev. Gen. Plant. 2: 812. "1891. ' 
Rumohra aristata Ching, Sinensia 5: 50. 1934, 
Polystichopsis aristata ee ttum, Ferns Malaya 486. 1954. 
Byrsopreris Standishii eg ie Morton, comb. 
Lastrea Standishii Moore, Gard. Chron. 1863: 292. 1863. 
Dryopteris Standishii C. Chr. s d. Fil. 587. 1906. 
Rumohra Standishii Ching, Sinensia 5: 64. 1934. 
The American species of Byrsopteris are: 

ByYRsopreris denticulata (Swartz) Morton, comb. 

Polypodium denticulatum Swartz, Prodr. Fl. Ind. Oce. 134. 1788. Type 
from Jamaica, Swartz. 

Dryopteris denticulata Kuntze, Rev. Gen. Plant. 2: 812. 1891. 

Rumohra denticulata Copel. Gen. Fil. 114. 1947. 


Byrsopreris formosa (Fée) Morton, comb. 


Aspidium formosum Fée, Gen. Fil. 296. 1852. Lectotype: Cuba, Linden 
2115, sletk ey Uleistosuen, Dansk. Vid. Selsk. Skr. Natur. Afd. VITI, 
7.119. 1920. 


Dryopteris formosa Maxon, Contr. U. 8S. Nat. Herb. 13: 17. 1909 (ad 
nom., excl. descr.) 
ByYRSOPTERIS leucostegioides (C. Chr.) Morton, comb. nov. 
acdignte apne see C. Chr. Dansk. Vid. Selsk. Skr. Nat. Afd. VIII, 
118. fig. 28. 1920. Type: Colombia, oe 234. 
helio eh attet (Hook.) Morton, co nov. 
Nephrodium dentiontatwm var. rigidissimum Hook. Sp. Fil. 4: 148. 1862. 
: Jam lson 


ype , Wi 
Pryopteis ge pts GC. Oe: Dansk. Vid. Selsk. Skr. Nat. Afd. VIII, 
118, fig. 27. 1920 


Ferns Known As RuMOHRA 153 


Some of the Old World species that I know are the following. 

A number of others which may or may not be distinct are recog- 
nized by Ching. Several species referred by Ching to Rumohra 

R. nipponica, R. sino-miqueliana, R. uadripinnata, and R. 
Miqueliana) are variously hirsute. Examination of the rhachis 
structure in those species available to me for study (the two 
last named) shows that these species are not con-generic with 
Rumohra or Byrsopteris, as might be expected. Their real af- 
finity (perhaps with Ctenitis ?) remains to be determined. 
Byrsopreris amabilis a Morton, comb. nov. 

Aspidium amabile Blume, Enum. PI. . 165. 

rhe tia esti J. Smith, Ferns peed & fag pe 1866. 

Dryopteris amabilis Kuntze, Rev. Gen. Plant. 2: 812. 1891. 

Rumohra ees Ching, Ginanata 5: 41. 1934 
BYRSOPTERIS assamica (Kuhn) Morton, comb. nov. 

Aspidium assamicum Kuhn, Linnaea 36: 108. 1869. 

Dryopteris assamica Rosenst. Med. Rijks ae Leiden 31: 6, 1917. 

Polystichum assamicum Ching ex C. Chr. Ind. Fil. Sup I: ots 1934. 

Rumohra assamica Ching, Sinensia 5: 47. 1934. 
Byrsopreris carvifolia (Kunze) Morton, comb. n 


Aspidium carvifolium Kunze, Bot. Zeit. 1843: 283. 1843 (as curvifolium 


pro err. 
Polystichum carvifolium C. Chr. Ind. Fil. 580. 1906. 
Dryopteris carvifolia C. Chr. Act. Hort. Goth. 1: 64. 1924. 
Rumohra carvifolia Ching, Sinensia 5: 60, 1934 
Byrsopreris coniifolia (Moore) Morton, comb. no 
ag: coniifolium Wall. ex. Kunze, Linnaea 24: 293. 1851, non Presl, 
1822. Type: Nepal, Wallich 341 (Isotype US). 
Lastrea coniifolia Moore, Ind. Fil. LXXXVITII. 1857. A new name, by 
t de Nomenel, (Art. 72, Nota) 
Bumohra Wallichii Ching, Sinensia §: 56. August 1934. Based on 


A ‘ 

rbporde goa himalayense Ching ex C. Chr. Ind. Fil. Sup. III: 163. Oct. 
1934. Based on Aspidium coniifolium Wall. 

As shown above, Article 72 of the present International Code 

of Nomenclature allows the use of the well-known epithet 

coniifolia. This Article is an extremely important one in stabiliz- 


154 AMERICAN FERN JOURNAL 


ing nomenclature, since it often results in the retention of well- 
known epithets, with merely a change in the parenthetical au- 
thorities, a minor matter so far as general usage is concerned. 
Byrsopreris Hasseltii (Blume) Morton, comb. 

Polypodiuwm Hasseltii Blume, Fl. Jav. Fil. 195. pl. 92. 1829. 

Dryopteris eae C. Chr. Ind. Fil. 269. 1905. 

Rumohra Hasseltii Ching, Sinensia 5: 61. 1934, 

piaeaimetne Shanes tii Holttum, Ferns Malaya 487. 1954. 
Byrsopreris Henryi (Christ) Morton, comb. nov. 

Polystichum Henryi Christ, Notul. Syst. 7: 36. 1909. Type: China, Henry 


Dryopteris Henryi C. Chr. Contr. U.S. ee Herb. 26: 282. 1931. 
Rumohra Henryi Ching, indie &: 57. 1934. 
Byrsopreris Maximowiczii (Baker) Morton, comb. nov. 
Nephrodium Mazimowiczii Baker, in Hook. & Bak. Syn. Fil. ed. 2. 499. 
1874. Type: Japan, Maximowicz 98. 
Dryopteris Maximowiczii Kuntze, Rev. Gen ae aah : 913. 1891. 
Rumohra Maximowiczii Ching, seus aan a 
- Byrsopreris mutica (Franch. & Sav.) Morton, comb. no 


Aspidium muticum Franch. & Sav. Enum. Pl. Jap. 2: 240, 635, 1879. 
di 


ype: 

Dryopteris mutica C. Chr. In a. Fil. ag 1905. 
Rumohra mutica Ching, Sinensia 5: 65. 

Byrsopreris speciosa (D. Don) Morton, comb. nov. 
Aspidium speciosum D. Don, Prodr. Fl. Nepal. 5. 1825. 
Polystichum speciosum J. Smith, Journ. Bot. 4: 195. 1841. 
Dryopteris speciosa C. Chr. Act. Hort. Goth. 7: 63. 1924. 
Rumohra speciosa Ching, Sinensia 5: 53. 1934 


Recently, Miss Mary D. Tindale® has revived the genus 
Lastreopsis Ching, which had been reduced to Ctenitis by Cope- 
land. She appears to be justified, for the type of Lastreopsis. 
L. recedens, and a number of allied species, differ from Ctenitis 
in the rhachis structure. In these species the two prominent 
ridges of the rhachis ar rhachillae are continuous with the 
leaf-margin (davallioid type), whereas in Ctenitis the rhachises 


8 Vict. Nat. 73: 180-185. 1957. 


Recent Fern LITERATURE 155 


are either not ridged at all or the ridges run to the center of the 
pinnules and not to the margins. Unfortunately, Miss Tindale 
includes in her enlarged genus Lastreopsis the species pubescens, 
chaerophylloides, ne lurida, the first of which is the type of 
Polystichopsis; if she is right, then the proper name is Poly- 
stichopsis, which oe ite over Lastreopsis. The matter needs 
to be investigated further, but it is my present opinion that both 
genera may stand. Polystichopsis has a rhachis structure of the 
dryopteroid type rather than the davallioid like Lastreopsis. 

Polystichopsis appears to be a small, exclusively American 
genus. e species, none of which are in BN he are: 
PoLYsticHopsiIs pubescens (L.) Morton, comb. 

Polypodium pubescens 1. Syst. Nat. ed. 10, 2: 13 97. 1759. 

Dryopteris pubescens Kuntze, Rev. Gen. Plant. 2: 813. 1891. 

Rumohra pubescens Ching, Sinensia 5: 35, 1934. 

Lastreopsis pubescens Tindale, Vict. Nat. 73: 185. 1957. 
Poysticnopsis chaerophylloides (Poiret) Morton, comb. 

Polypodium chaerophylloides Poiret in Lamarck, Hag sua 5: 542, 

1804 


Dryopteris chaerophylloides C. Chr. Dansk. Vid. Selsk. Skr. Nat, Afd. 
: 105. 192 


T, 6: 105. 1920. 
Rumohra chaerophylloides Ching, Sinensia 5: 35. 1934 
Lastreopsis chaerophylloides Tindale, Vict. Nat. 73: 185, 1957. 
PotysticHopsis lurida (Underw. & Maxon) Morton, com), nov. 
ryopteris lurida Underw. & Maxon in Slosson, Bull. cane Club 40: 


Rumohra lurida Ching, Sinensia 5: 35. 

Lastreopsis lurida Tindale, Vict. Nat. Be rie 1957. 
PoLYSTICHOPSIS ochropteroides (Baker) otihaiae comb, nov. 

Nephrodium ochropteroides Baker, Ann. Bot. 2820: 1801. 

Dryopteris ochropteroides C. Chr. Ind. Fil. bas 1905 

This is the only species of Polystichopsis occurring outside the 
West Indies; it oceurs in Jamaica, Panama, Colombia, and 
Surinam, Geary whites extremely rare and mostly collected only 
once in each disjunct area. Habitally, it is not very close to P. 
pubescens and the other species, all of which have the same facies. 

SMITHSONIAN INstrTUTION, WasHtNeToN 25, D. C. 


156 AMERICAN FERN JOURNAL 


Recent Fern Literature 


Ferns or ALBERTA—With the publication of the book by Dr. 
E. H. Moss on the flora of Alberta,! one of the large gaps in our 
knowledge of the Canadian flora has been filled. In the Pterido- 
phyta 55 species in 20 genera of seven families are considered. 
The author states that he has been conservative in his approach 
to taxonomy and a table claims treatment of only six subspecific 
entities. Actually 12 subspecific entities are considered. 

The most disappointing aspect of this Flora is the lack of in- 
formation regarding the distribution of species. There are no 
maps showing the occurrence of species (there are in fact no 
illustrations of any kind in the book) and phytogeographie in- 
formation is of the sketchiest nature. For Pellaea atropurpurea 
(L.) Link var. Bushii Mack. we are told that it grows ‘‘on 
calcareous rocks’’—nothing more. Actually, it has been collected 
only near the Bow Falls at Banff.2 Matteuccia Struthiopteris 
(L.) Todaro is found in ‘‘damp woods,’’ whereas Polypodium 
vulgare L. is found ‘‘on rocks, logs and banks.’’ The author 
states in the Preface that Alberta affords features of exceptional 
botanical interest with steppes, forests, mountains, prairies, ete. 
It seems a pity that there is no correlation of the species with 
geography. This is surely a major deficiency in a regional work. 
Also there is little indication as to whether species are common 
or rare with few exceptions, such as Asplenium Trichomanes L., 
which is included although no Alberta plants have been seen by 
the author. 

It seems a pity that in a $10.00 book of this nature that there 
could not be included a good colored map of Alberta. This 
would be useful in locating the few localities mentioned such as 
‘Waterton, Lake Athabasca, Drumheller, etc. Such a shortcoming 
is not uncommon, however, as the reviewer knows only too well 

1 Flora of Alberta, by E. H. Moss. 1959. University of Toronto Press, 
Toronto, Canada. 546 pp. $10.00. 


2The plants from this lo cality are bial considered to be var. 
occidentalis rather than var. Bushii-—C. V. Mor 


RecENT FERN LITERATURE 15ST 


when gnashing his teeth over the lack of a map civing the names 
of the counties of Michigan in Billington’s ‘‘Ferns of Michi- 
gan.”’ 

This reviewer was interested that both Dryopteris dilatata 
(Hoffm.) A. Gray and D. spinulosa (Muell.) Watt are included 
as separate species in the five species of Dryopteris considered. 
No subspecific categories are delineated for this genus. The in- 
clusion of D. Filiz-mas found on ‘‘wooded slopes’’ gives no in- 
formation as to the range of this northern species. 

Cheilanthes Feei Moore has evidently been added to the page 
proof in the introduction to the Polypodiaceae, as a species that 
was originally overlooked. It is not included in the alphabetical 
listing of genera. This species is stated to occur at Banff. 

The keys and nomenclature are to a large extent based on 
C. V. Morton’s treatment of the Pteridophyta in The New 
Britton and Brown Illustrated Flora, and are quite adequate. 
The paucity of fern species in the same genus will undoubtedly 
make the task of identification easier for the amateur. Of the 
16 genera of ferns described, only Botrychium (6 Spp-); 
Dryopteris (5 spp.) and Woodsia (4 spp.) have more than two 
species included. 

This book will be of value as an intermediate manual for stu- 
dents of the flora of Alberta. For visitors to the province, or as 
a field guide, it has definite limitations.—DonaLp M. Brirron, 
ONTARIO AGRICULTURAL COLLEGE, GUELPH, ONTARIO. 


Ferns or Catirornta.—The recently published ‘‘A California 
Flora’?! treats all of the vascular plants of the state. The vascu- 
lar cryptogams or Pteridophytes are included under three Divi- 
sions: Lepidophyta (Lycopodium, Selaginella and Isoétes), 
Calamophyta (Equisetum) and Pterophyta (the ferns proper). 
A total of 28 genera and 86 species are recognized. These num- 
bers compare with the 25 genera and 81 species of California in 


1A California Flora. By Philip A. Munz in collaboration with David D. 
Keek. pp. 1-1681, figs. 1-134, 2 maps. University of California Press. 1959. 
$11.50. 


158 AMERICAN FERN JOURNAL 


Abrams? and the 26 genera and 79 species in Jepson.* The slight 
increase in genera and species is, I believe, a reflection of the fact 
that most of the kinds of ferns in California have been known 
for some time. Taxonomic revisions and range extensions have 
altered our knowledge of them in the last few decades but not 
to the significant extent that they have in many groups of the 
flowering plants. 

Keys and diagnostic descriptions are provided for the families, 
genera, and species. The major groups of ferns and fern allies 
are illustrated by line drawings. The distribution in California 
is given for each species as well as a general statement of its 
entire range. An especially useful feature is that each species is 
related to the vegetation of the state by reference to the plant 
communities in which it grows. The twenty-nine plant communi- 
ties that occur in California are described in the introduction. 

The treatment of the Pteridophyta seems eminently usable and 
will undoubtedly provide, for most students, the standard treat- 
ment of this segment of the flora that the book itself will for the 
whole. For this reason it is unfortunate that, in following Cope- 
land’s Genera Filicum, the Flora does not always accurately re- 
flect our present knowledge. There are no serious reasons, for ex- 
ample, for the segregation of Blechnaceae, Pteridaceae, Aspidia- 
ceae, and Aspleniaceae from the Polypodiaceae. Also there are 
generic realignments that are not warranted by what we know, 
and do not know, about the species concerned. Some of the spe- 
cies of Notholaena are placed in Cheilanthes and one, N. cali- 
fornica, in Aleuritopteris; Cheilanthes californica is segregated 
as Aspidotis; and Cheilanthes siliquosa is placed in Onychium. 
This is not to say that the classification I would prefer (Notho- 
laena and Cheilanthes) is wholly justified, but rather that de- 
tailed and serious study must provide the basis for improving it. 
As a matter of nomenclature, it is also unfortunate that the name 
Lastrea has been adopted rather than the correct name Thely- 


2 Tilustr. Fl. Pacific States, vol. 1, 1923. 
3 Manual FI. Plant. Calif. 1925. 


AMERICAN FERN SOCIETY 159 


pteris. The names for the two Californian speciest are Thely- 
pteris puberula (Baker) Morton (Lastrea augescens) and Thely- 
pteris nevadensis (Baker) Clute (Lastrea oregana). 

A few inaccuracies that I have noticed are mentioned here so 
that users of the Flora may correct them. In a work of such 
scope and actual length it is inevitable that, in spite of a success- 
ful effort to maintain high standards, a few errors will remain. 
The correct name for the variety of Pteridiwm aquilinum in 
California is var. pubescens Underw.; var. lanuginosum is a 
later homonym. Pellaea compacta and Pellaeca mucronata var. 
californica are names for the same taxon; the recent revision of 
Pellaca treats it under the latter name. The var. compositum 
mentioned under Botrychiwm multifidum is a variety, not of that 
species, but of B. simplex. Notholaena californica (Ariz. and Ss. 
Cal. to Baja Cal.) is treated as Aleuritopteris eretacea (Liebm.) 
Fourn. but Notholaena cretacea Liebm. is a taxonomic synonym 
of the very distinct Notholaena sulphurea (Mexico to Chile). 
Notholaena californica has no name under Aleuritopteris and I 
hope one will not be made. 

To complete the list of California ferns, two hybrids (or prob- 
able hybrids) should be mentioned: X Adiantum Tracyi Hall ex 
Wagener (Humboldt, Sonoma, and Marin counties) and 
Cheilanthes Carlotta-halliae Wagner & Gilbert (Marin, San 
Benito, and San Luis Obispo eountries).—Rouua TRYON, GRAY 
HERBARIUM. 


American Fern Society 


New MEMBERS 
Mr. Mark Adams, 3816 22nd Street, San Francisco 14, California 
Mr. Frank Akutowicz, 2007 Harvey Road, Wilmington 3, Delaware 
Mrs. Barbara J. Bayless, 450 Dahlia, Ontario, California 
Mrs. R. O. Brown, 451 Melrose Avenue, Toronto 12, Ontario, Canada 
t 


Mrs. Eugenia Carpenter, Route 3, Box 620, Merritt Island, Flori 
Miss Ruth Castles, Forest Hill Apartments, Nashville, Tennessee 


4 Morton, This JourNAL 48: 138, 139. 1958. 


160 AMERICAN FERN JOURNAL 


Dr. W. J. Cody, Division of Botany & Plant anole Science Service, 
Canada Department of Agriculture, Ottawa, Provine mtario, Canada 
0 ifornia 


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~. As Gri ay, 
an Ernest C. fo 729 Canal Street, New Orleans, Louisiana 
Mrs. Lewis M. Hull, RFD 1, Boonton, New Jerse. 
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nzie, 495 Deloraine Avenue, Toronto 12, On svt Canada 
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Hanover, Indiana 

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lege, Amritsar, India 

Mr. Charles B, Miller, 705 Twin Oaks, Chula Vista, suet 

Mr. 


Ji gden, 207 E. Upland Road, Ithae: ew 
M Rennagel, 11226 South Painter Avenue ” Wai. California 
Miss Jennie Riggs, 2205 Capers Avenue, Nashville, Tenn nessee 
Mrs. M. S. Schoenberger, 20316 Seottedalé Boulevard, Cleveland 22, Ohio 
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Miss Mary Wood, Box 201, Woodbury, Tennessee 


iment 


eo Ae ee a 


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Successor to Henry George Fiedler 


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FOR THE HERBARIUM” 


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gen erously shared the s of his ] ong and successful experience in the 
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Help for the amateur botanist, and hints for the professional ¢ collector, 
abound in this thirty-six page illustrated treatise in which Dr. Johnston 
describes time-tested techniques for pressing, preserving and mounting 


flowering plants, ferns, palms, algae, mosses, lichens and fungi. Specific 
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Vou. 50 APRIL-JUNE, 1960 No. 2 


American Fern Yournal 


A QUARTERLY DEVOTED TO FERNS 
Published by the 


AMERICAN FERN SOCIETY 
ts 


Cc. V. MORTON 
. C. BENEDICT IRA L. WIGGINS 
A. C, SMITH 


ie 


CONTENTS 


JUL 18 ou 


Hunting Ferns in the Barrancas of Chihuahua, —— 
vING KNoBLOcH 161 


Ferns in Cultivation, TL _Synvia LEATHERMAN 168 
Some Fern Books for Different Regions of the United States 
C. V. Morton 169 


The Genesis of the American Fern Journal: aah ory Data 
C. Benepict 179 


Isoétes melanopoda in Southern Illinois 
Rosert H. Moniensrock, JR. 181 


The Smooth Scouring Rush and Its Complexities 
Ricnarp L, Hauke 185 


The Gametophyte and Young Sporophyte of Athyrium esculentum 
B. K. Nayar 194 


Recent Fern Literatare _—_§_§$_$__$_$_____—_ 208 
Notes and News: Ophioglossaceae Wanted ———______—__—- 


American Fern Society: R of President; Report of Treasurer; 
Report of Auditing Committee Report of Secretary; Report of 
Judge of Hissienes Report of Curator and Librarian; 

of Spore Exchange; aa Field-trip; a Field- 
trip; Vermont Field-trip; Obituaries: Brem r+ Whidden rend. 
ra a les Y. ry Snore Cueetr Jac “abo ‘a ell, — 


Che American Fern Society 


Council for 1960 
OFFICERS FOR THE YEAR 

Crarr A, Brown, — of Botany, Louisiana State University, Baton 
oe Louisia: President 
RayMonpD, rae Botanical Garden, 4101 East ayy ioe Street, 
gag atest Canada ice-President 

DonaLp Huttieston, Longwood Gardens, Kennett Square, iace 
Secretary 
Watter 8. Puinuirs, Department of Botany, University of Arizona, Tueson, 

i Treasur 


C. V. Morton, Smithsonian Institution, Washington 25, D. C 


Editor-in-Chief 
OFFICIAL ORGAN 
American Fern Journal 
EDITORS 

CY, —— eee seereTiom, Washington ve D, 0. 
R. C. BENED 5 Hall Street, Brooklyn 5, N. Y. 
Tra L. Wuens bata Herbarium, Btantoed University, Stanford, Calif. 
A. C, Smirz___....._____ Smithsonian Institution, Washington D.C. 


ond-class niaiees ae at Baltimorg, Ma 
Matter for ——— sho nag be addressed to C. V. Morton, Smithsonian 


gton 
Subsarigtiie: $2.50, poet of agency handling fees; sent free to mem- 
bers of the American Fern Society (annual dues, $2.50; sustaining m 
bership, poor 00; life membership, $50.00). Extracted reprints, if iy ge om 
advance, will be furnished authors at cost. They should be ordered 


returned. 
Back volumes $2.00 each; single back numbers 50 cents each; Cumulative 
Index to vols. 1-25, 25 cents, Ten per cent discount on orders of six volumes 
or more. 
oe of address should be sent to the Seeretary: Dr. Donald Huttle- 
ton, Longwood Gardens, Kennett 8: e, Pe lvania. 
seteee should allow two months for chan; nges to ~~ Lag ote 


maneetiont for membership, su — orders for back numbers, and 
other business communications should b. — —— vig gy Treasurer: Dr. 
bere 8. Phillips, Department of ge versity of A: 
rizona. 


LIBRARIAN AND CURATOR OF THE HERBARIUM 

Dr. H. W. Wagner, University of Michigan, Ann Arbor, Mich. 

regular loan department is maintained in connection with the library 
and herbarium, Members may borrow books and specimens at any time, 

© borrower paying all postal or express charges. The pages of the oe Journal 

- are open to members who wish to —— exchanges; a membership list 

published at — to assist those interested in obtaining specimens 
toca different localities 


American Fern Journal 


Vou. 50 APRIL-JUNE, 1960 No. 2 


Hunting Ferns in the Barrancas of Chihuahua, Mexico! 
Irvine KNosiocn 

Those who travel down Highway 45 from Juarez to Chihua- 
hua City and on to Mexico City are struck by the apparently 
endless desert on both sides of the highway that extends to the 
southern borders of the state. Few apparently know that the 
western half of the state contains high mountains covered with 
oaks and pines and generously provided with deep canyons or 
barrancas. There are ferns to be found on the mountains stud- 

ing the desert but they do not compare in numbers or variety 
with those in the wetter and shadier parts of the state. 

The writer has collected ferns in Chihuahua off and on since 
1938. At the present count there are over 112 species and varie- 
ties of pteridophytes here. Many of these are confined to the 
western half of the state, such as Pellaea Seemann, Pellaea 
Skinneri and Notholaena Weatherbiana. There are, of course, 
many others in the same category. Since some of the readers of 
this journal may be interested in Mexican ferns, we shall char- 
acterize some collecting grounds in the western part of the state 
and mention some of the more unusual ferns which are to be 
found there 

The Mojarachi-Maguarichi area is reached by driving (in a 
pick-up, jeep, or passenger car with high wheels and a second low 
gear) from Chihuahua City west to Cuauhtemoc, where one 
leaves the hard road. Then one follows the railroad to La Junta, 
Mifiaca, and on to San Juanito. Here one leaves the better- 
travelled dirt road and drives on an old mining camp road 


1 Contribution No. 60-16 from As Department of Botany and Plant 
Pathology, Michigan State Univers 
Volume 50, No. 1, of the petri pp. 1-160, was issued April 25, 1960. 


162 AMERICAN FERN JOURNAL 


leading eventually to Maguarichi. Recent torrential rains in the 
state have damaged the road and bridges. This is really a bad 
road at best with the average speed being five to eight miles per 
hour. I do not recommend this trip except for the most experi- 
enced campers. One can get gas at San Juanito but there is not 
likely to be any from there on. Food should be taken along and 
meals prepared unless one is willing to eat in very poor restau- 
rants. Water should be sterilized with iodine tablets and one 
should take two Entervioforme tablets every day to combat 
dysentery. At last reports, there was a new lumber camp beside 
the road, about two hours distance this side of Maguarichi. This 
locality is known as Las Lajas. There is an old road branching 
off to the left going to the abandoned mining camp of Mojarachi. 
Inquiries at Las Lajas will help one decide whether to attempt 
the road or to go on to Maguarichi. At the latter place, there are 
a few people living, one being an Austrian-American lady named 
Mrs. Zehtner who speaks very good English, as well as German 
and Spanish. She makes trips out quite often so one cannot be 
sure that she is there. Trinidad is the name of a local guide and 
mules can be rented from him for local trips or for the two hour 
trip to Mojarachi, south of Maguarichi. The latter place is about 
4,900 feet above sea level and the latitude and longitude are 
approximately 27°52’ N. and 108°—05’ W. respectively. Mo- 
jarachi is slightly south of Maguarichi and is at an elevation of 
about 7,200 feet. The latitude is about 27°-51’ N. and the longi- 
tude about 108°-00' W. At last reports there was no one living 
in the place. Maguarichi is lower and really out of the pine belt. 
There are some scrub oaks and many species of flowering peren- 
nials. It is not rich in fern species. Notholaena aurea, Notholaena 
sinuata var. sinuata and Asplenium Palmeri can be found here 
quite readily. The dainty Bommeria hispida is also common. It 
should be mentioned that the ferns are at their best during the 
rainy season and this coincides, naturally, with the muddiest 
roads. July and August are good collecting months but poor 
travelling months. By the middle of September travel is best 
and the plants are still in good condition. 


CHIHUAHUA FERNS 163 


On the way to Mojarachi by mule, one may find, by searching, 
Adiantum Capillus-veneris and Elaphoglossum pilosum. These 
are found pendant on wet cliffs, the former usually in the sun 
and latter usually in the shade. Mojarachi is higher in elevation 
and while there are many species of oaks the pines are also there 
in variety and abundance. There is more duff on the ground and 
there is more shade. About 50 species and varieties of ferns can 
be found here in a radius of two miles, at the base of cliffs and 
boulders and rising from the duff on the forest floor. This is the 
only station in the state for Selaginella Arsenei (from Segorichi, 
within the two mile radius), and Bommeria Knoblochti was also 
first found at Segorichi, Other good finds in the Mojarachi vicin- 
ity are Cheilanthes angustifolia, C. farinosa, Pellaea allosuroides, 
Pellaea sagittata, vars. sagittata and cordata, Asplenium exiguum, 
Woodsia mollis, Dryopteris augescens var. puberula, Plagiogyria 
semicordata, Polypodium aureum, and P. Hartwegianum. 

second region worth seeing is the La Bufa-Rio Batopilas 
area. To reach this one returns to San Juanito and follows the 
railroad line south to the town of Creel. Here one can get gaso- 
line and stock up with food supplies. There is a drug store here 
where one can get cold coca-cola. It is the drug store near the 
gasoline supply. One now is faced with a breath-taking five hour 
trip over a splendid but one-way road leading to the mining 
town of La Bufa, situated on the Batopilas river. If one speaks 
Spanish, one should go to the ‘‘Terminal’’ at the edge of town. 
This is an office of the Potosi Mining Company and here Senor 
Mendoza will tell you if the road is reasonably free of the diesel 
trucks that haul the ore from the mine to Creel. There are few 
turnouts on this road and one must give the right of way to the 
trucks by turning off or backing up. A Jesuit priest in Creel, 
Father Martinez, or Trompas as he is generally known, speaks 
good English and can be helpful. When everything is arranged, 
away you go through the pines, down into the Barranca de Cobre 
canyon, up out of this canyon, across a plateau, down into the 
Batopilas Canyon, and up the other side to the mine office itself. 
Here, at last reports, Mr. Robert Emmett was in charge and 


164 AMERICAN FERN JOURNAL 


here reasonable accommodations can usually be had. My trip 
there in 1957 may now be described but yours may be better or 
worse. 

I left Chihuahua on September 11 at about 9:30 a.m. in a 
station wagon through the courtesy of Mr. Martin Nesbitt, the 
manager of the Potosi Mining Company in the city of Chihuahua. 
We arrived in La Junta at 12:30 and ate at the Centro Viajero 
Hotel. I checked my baggage through to Creel, bought a ticket 
and got on the train from Chihuahua at 5:30 p.m. The train 
left La Junta at 7:30 p.m. and did not arrive in Creel until 
4:30 a.m. The Hotel Chavez was open but no beds were available. 
One should telegraph ahead for accommodations if one really 
wants to stay in the Hotel. I dozed in a chair in the lobby until 
awakened by the truck driver sent from the mine to pick me up. 
We ate breakfast and waited around until 10:00 a.m. for the 
station to open in order to get my baggage, and then set out for 
the mine, arriving at 2:30 p.m. It was a most thrilling ride 
indeed. At the mine office I was introduced to Mr. Emmett and 
shown a two-bedroom cottage for my use and the lunch-room 
where I was to eat. The country is beautiful but very precipitous. 
Every day a house boy brought me two cold beers and two cold 
cokes as well as a pitcher of ice water. The second day, I oe 
iodine tablets to the water and then had safe, ice-cold wa 
You see, it took me a whole day to wake up to the eine 
Fresh figs from a tree nearby in a garden were also a treat 

Needless to say, I botanized from early morning until dark 
in all the side canyons near the mine. Sunday was a red-letter 
day for I was taken to the Basigochi Country Club, back down 
the road, at an elevation of 7,400 feet. Here we played 18 holes 
of golf with Tarahumare Indians as caddies, truly a unique ex- 
perience. Returning to the mine in the evening, I collected a fine 
Elaphoglossum and a rattlesnake. Having read a great deal 
about Dr. Edward Palmer’s travels in southwestern Chihuahua, 
I had a burning desire to visit one of his choice collecting 
grounds, namely Batopilas. The late Edward Goldman had also 
been there in the days when the silver mine was booming. Ac- 


CHIHUAHUA FERNS 165 


cordingly, I arranged for a guide and three mules and set out at 
7 a.m. on September 18th. Batopilas lies down the canyon. Our 
cargo burros were slow and we did not arrive until 6 p.m., a 
long, hot, thirsty ride. One of the big men in this quaint town 
is Gregorio Bigler, to whom I had a letter of introduction and 
who, upon my arrival, immediately plied me with two cold 
bottles of Pepsicola. He kindly allowed me to sleep over the 
store on an army cot (which only proved comfortable after I 
had placed my air mattress upon it). The climate is hot and 
humid and malaria is prevalent despite governmental efforts. 

We meet here plants belonging to tropical families. It is 
not at all the type of vegetation one would expect in a northern 
state like Chihuahua. Mingled with the figs, the kapok trees, and 
Crescentia alata trees are giant cacti towering 40-50 feet into the 
air. I was very anxious to see the Hacienda San Miguel, the head- 
quarters of the old mine. What a sorry mass of ruins! The adobe 
buildings were rapidly returning to the soil from which they 
had come. Since I had sent my guide back to La Bufa for faster 
burros (for I had determined that the return trip would not take 
11 hours), I hired an 80 years old native named Joaquin Vega 
and together we tramped around the ruins of the Hacienda and 
surrounding valleys for two days. It was not enough that I was 
thoroughly exhausted from the trip down here but the town band 
insisted on playing mournful airs practically all night long. 

My explorations at Batopilas were only partially successful. 
I did relocate Notholaena Weatherbiana which Palmer had found 
around 1885, but I failed to find Asplenium modestum. How- 
ever, there were huge clumps of Notholaena Lemmonii on the 
adobe walls, as well as large quantities of Cheilanthes Pringlet 
and Selaginella pallescens. Back in La Bufa, after a seven hour 
trip, we spent several weeks all told in obtaining such ferns as 
Cheilanthes microphylla, Anemia anthriscifolia, Bommeria 
Knoblochii, Cheilanthes lendigera, and other more common 
species. Since a flu epidemic started to rage through the mining 
community, I left there and settled back in Creel. Just at the 
edge of town, where the new railroad line begins to the Pacific 


166 AMERICAN FERN JOURNAL 


coast, in pockets of a white-colored road cut, we found the prize 
fern of all, Asplenium Adiantum-nigrum. This had been found 
by Pringle and Dr. MeVaugh before, but it is a very rare plant 
that we were very glad to obtain. 

Incidentally, if one wishes to take a guided tour of the trip just 
described e can eontact ‘‘Wampler Trail Trips,’’ Box 45, 
Berkeley, California. These trips go once a year to the place 
just described and once to the Basaseachi Falls in Chihuahua. 
I believe the cost is about $350 to $400 dollars from the border. 

Describing trips further to the southwest, we might mention 
a trip we took in 1954. Five of us arrived in Creel in July. A 
small truck was hired here and we sped west on a new railroad 
grade being constructed. The road leads near the edge of the 
Urique Barranea in one place and we stepped out to take pictures 
here. This locale is known as the ‘‘Divisadero’’ and the canyon 
drops off 4,800 feet straight down to the river. Shortly there- 
after we arrived in the head camp of the railroad engineers, a 
settlement known as Areponapuchi. Here we were kindly given 
accommodations. The next day we descended into this canyon 
by mule back but the trail was too slippery even for mules. 
Camping above the river, we collected many interesting plants, 
such as Elaphoglossum pilosum and Cyrtomium auriculatum. 
Notholaeana Lemmonii and Pellaea Seemannii were also taken. 
I was greatly pleased to find Anemia anthriscifolia growing in 
great abundance on the rocky slopes and in shade. 

In 1958, a party of four of us came along this same railroad 
grade in a pick-up truck and went on down the road to Cuiteco, 
one of Rudolph Endlich’s collecting grounds. Here we branched 
off onto a dirt road and traveled several hours to Cerocahui, 
situated on the rim above the town of Urique, our eventual desti- 
nation. Cerocahui has about a dozen small houses and a magnifi- 
cent catholic church. We bedded down on the floor of one of 
the houses and set out to explore the neighboring canyons. Ferns 
that are absent or uncommon in other parts of Chihuahua are 
common and luxuriant here. e might mention Aspleniwm 
exiguum, Cheilanthes farinosa, Notholaena limitanea var. 


CHinvuaAHuA FERNS 167 


limitanea, and Polypodium subpetiolatum, the latter being very 
common at Cuiteco. The trip by mule to Urique consumed a 
back-breaking ten hours. Here we found another run-down 
mining town, all but in ruins. However, the people were as 
friendly as they usually are in Mexico and they did for us what 
they could. The weather was hot and humid; it rained every 
day and the air was malaria-ridden. Sanitary conditions were at 
a low ebb, to put it conservatively. We were in that unusual type 
of vegetation known variously as short-thorn or tropical decidu- 
ous forest where Acacias and Cacti mingle with representatives 
of tropical families, where some plants are covered with thorns 
and others devoid of them. My herpetological companions col- 
leeted large numbers of subtropical species and obtained all 
sorts of interesting records. Here we found two rare fern-allies 
in great abundance, namely Selaginella novoleonensis and 
Selaginella Wrightti. Notholaena candida var. candida, rare 
elsewhere, was also in quantity. This, however, was only a 
starter. In shade and on rocks we obtained two species not 
hitherto found in the state—Selaginella Sartori and Pellaea 
Skinneri. This Cerocahui-Urique area is one of the richest in 
ferns in the entire state and would be well worth further investi- 
gation. In a year or two at the most, the rails will be laid along 
the right-of-way and one will be able to get in here only by train 
or by air, If the new railway will operate like the present one 
from Chihuahua, it would not be advisable to go in by rail unless 
one is willing to be very patient and long-suffering. Flights from 
Chihuahua by light plane to Cerocahui (ten hours from Urique 
by mule) or to Naranja (four hours by mule from Urique) can 
be arranged. 

The writer is completing with Dr. Correll a manuscript on the 
pteridophytes of the state. My experience has been similar to 
that of other explorers, namely, that more time is spent in 
travelling than in collecting. This is a very wasteful procedure 
but nothing can be done about it unless granting agencies are 
willing to be more generous with funds so that helicopters can 
be rented. In the long run this procedure, while apparently more 


168 AMERICAN FERN JOURNAL 


expensive, might prove to be the most economical method yet. 
For example, one could sample dozens of areas in a state during 
a summer, gh that would take six to twelve summers to investi- 
gate by mu 

et 60 per cent covered by desert and only one third the 
size of Texas, Chihuahua has more fern species than Texas, and 
the number grows with each trip. Someday we hope to describe 
for our readers the ferns of the drier parts of the state. 

Micuigan State Universiry, East Lanstne, MIcHIGAn. 


Ferns in Cultivation, II 
SYLVIA LEATHERMAN 


Keeping potted ferns in good condition outdoors during hot 
summers has always been a problem. One solution is to actually 
plant them out in the garden, pot and all; the soil should be 
firmed closely around the pots. When the garden is watered 
regularly, the potted plants will also be watered, and the soil 
around the pots keeps them from drying out; this will, in all 
er save your prized ferns. 

re are a few words of caution about using this method. 


element ; most ferns will prefer a light shade. Secondly, remem- 
ber that the roots will go down through the drainage hole in 
the pot and anchor themselves in the soil under the pot. When 
the pot is lifted, the plants then wilt and sometimes die. Using 
the lid off the top of a coffee can or a similar barrier placed in 
the bottom of the hole will prevent this for a short period of time. 
However, if the pots are to remain in the ground for weeks, they 
should be turned and twisted once a week, which will prevent the 
roots from becoming established in the soil. If the pots are 
sitting on a barrier, they will be easier to turn. 

It is especially well to plant out the pots during a vacation 
period. Usually a person is hired to take care of the plants while 
the owner is away. Potted plants often suffer, since the hired 


Frrn Books 169 


persons do not understand them and do not devote enough time 
to them. Planting the pots out will save them, since the garden 
is watered vaniialy and plants in the ground almost always 
come through in good condition 

2637 NortaH Ler AVENUE, En veo CALIFORNIA. 


Some Fern Books for Different Regions of the United States 
C. V. Morton 


Some, but not all, of the descriptive accounts of the ferns of 
various regions of the United States have been reviewed in the 
JourNAL. A number of readers have suggested a general listing 
of the best works, and such a summary is perhaps timely. 

Dr. S. F. Blake’s ‘‘State and Local Fern Floras of the United 
States’?! is most useful, being almost complete and extremely 
accurate. The Supplement? brings this list up to 1950. The pres- 
ent list is not in any sense a second supplement, for I have made 
no attempt at completeness, and purely local lists are not noted 
at all 


GeneraAL Works: M. L. Fernald’s account of the ferns in 
Gray’s Manual of Botany, Eighth edition,® is the most complete 
treatment, representing a lifetime of intense study by an out- 
standing botanist. The area covered is from Newfoundland west 
across Quebee and Ontario along the 49th parallel to the north- 
western corner of Minnesota, southward along the western 
border of Minnesota and Iowa, and along the 96th meridian 
through Nebraska and Kansas to Missouri and then eastward 
along the southern borders of Missouri, Kentucky, and Virginia ; 
thus all the northeastern and midwestern states are included, 
but the prairie regions of the Dakotas, Nebraska and Kansas are 
excluded. Fernald’s work is topnotch and there is hardly a real 


1 This rat 31: 81-90, 131-148. 1941 

2 This Journal 40: 148-165. 1950. These lists of Dr. Blake’s were re 
Lp by the American Fern Society, and are still available for sale, at 

5 cents for the two. Orders should go to the Treasurer. : 

Pp, 1-1632. 1950. American Book Company, 55 Fifth Avenue, N.Y, 
$12. 50. Reviewed, this JournaL 40: 229. 1950. 


170 AMERICAN FERN JOURNAL 


error in it, except the confusing and erroneous adoption of the 
name Cheilanthes vestita for the plant usually and properly 
called (. lanosa and the adoption of C. lanosa for the plant cor- 
rectly called C. tomentosa Link. The name Pteretis for the 
ostrich-fern may have been correet when Fernald’s work was 
published (although this is debatable) but it is definitely not 
correct now, since at the 8th International Botanical Congress in 
Paris in 1954 the name Pteretis was rejected and the name 
Matteuccia conserved. 

The second large-scale work on the ferns of the eastern states 
is that of Morton in ‘‘The New Britton and Brown Illustrated 
Flora of the Northeastern United States and Adjacent Canada.’” 
Although this was actually printed two years later than Gray’s 
Manual, the manuscript had been prepared several years pre- 
viously and was in press before the publication of the Manual. 
The area covered is essentially the same, and the treatment is 
quite similar in many ways, although there are some differences 
in nomenclature. Although, according to the general policy of 
this work, minor varieties and forms are not mentioned, as they 
are in Gray’s Manual, the present work has the great advantage 
of fine drawings of all the species. Another complete work for 
the east is ‘A Field Guide to the Ferns,’ by Boughton Cobb ; 
the illustrations in this work can be especially recommended for 
the natural way in which the plants are depicted. 

An old standby, still available and to be recommended 
Herbert Durand’s ‘‘Field Book of Common Ferns.’’ Otkck 
general works on eastern ferns are Dr. Edgar T. Wherry’s ever- 
popular ‘‘Guide to Eastern Ferns,’” still as useful as ever to the 
beginner, Farida A. Wiley‘s ‘‘Ferns of Northeastern United 


4In n thr ee volumes. Published by the New York Botanical Garden, Bronx 
Park, New York 58, N. he ae cond, slightly revised printing, 1959. $30.00. 
Not reviewed in th 

5 Published ‘ How oiten Miftin Company, 2 Park St., Boston, Mass. 1956. 
3.95. Reviewed, this sores a6: 161: 1 

_ 6 Revised ed., pp. 1-219. . Putnam’s Sons, 210 Madison Ave., New 
ork 16, N. Y. $3.50 


d. 2, pp. 1-251. 1942. University of Pennsylvania Press, Philadelphia, 
Pa, $2. 00. Reviewed, this JourNaL 33: 76, 1943. 


FrrN Books 171 


States,’’® rather similar but not quite as good, and Willard N. 
Clute’s ‘‘Our Ferns, Their Haunts, Habits, and Folklore,’”? 
which is to be recommended highly for the amount of informa- 
tion not obtainable elsewhere. 

The following older, much-loved books are now out of print 
but can sometimes be picked up from dealers in second- hand 
books: ‘‘Ferns; A Manual for the Northeastern States,’’ by C. E. 
Waters (1903); ‘‘How Ferns Grow,’’ by Margaret Slosson 
(1906) ; ‘‘Ferns and How to Grow Them,”’ by Grace A. Woolson 
(1905, rev. ed. 1914); ‘‘The Fern Lover’s Companion,’’ by 
George H. Tilton (1922) ; and ‘‘ American Ferns; How to Know, 
Grow and Use Them,’’ by Edith A. Roberts (1935). 

The states of North and South Dakota, Nebraska, Kansas, Min- 
nesota, and Iowa, and the southern parts of Manitoba and Sas- 
katchewan are treated in ‘‘Flora of the Prairies and Plains of 
Central North America,’’!® by P. A. Rydberg; this is not a 
region in which ferns are abundant, except in the easternmost 
part. The entire Rocky Mountain area is covered only in Ryd- 
berg’s ‘‘Flora of the Rocky Mountains and Adjacent Plains,’’! 
in which the ferns were treated by Margaret Slosson very 
aE For the Far West there is only the treatment by Wil- 
ha _ Maxon in Abrams, ‘‘Illustrated Flora of the Pacific 
aca 12 a highly professional treatment, in fact a model in 
every way. A less elaborate work covering a part of the same 
area is ‘‘Ferns of the Northwest,’ by T. C. Frye. 


8 Revised pation pp. 1-1 108. ee yas the author, The 
American Mus f Natural History, New 2 ¥, Kev awed: this 
JOURNA ery - 


9 Ed. 2, pp. 1-388. 1 938. Frederick ‘ ec Company, New York, N. oe 
Out of print. Sits this JOURNAL 57. 

10 Pp. 1-969. Published by the New York "Botanical Garden, Bronx Park, 
New York 58, N. Y., 1932. Out of print. Not reviewe 

11 Pp. 1-1110. Pu plished by the author, 1917. Reiss sued in faesimile edi- 
tion, 1954, Hafner Publishing Co., 31 E. 10th Street, New York 3, N. Y. 
$15.00. Not ee ae : 

oe Vo ok; aoe es Published by Stanford University Press, Stan- 
~ California $17. 

Ate tropolitan Press, Portland, Oregon. Current 
availabe eee Binfords and Mort, 124 N. W. Ninth Avenue, Portland, ad 
. Reviewed, this JouRNAL 25: M65. 1935 


172 AMERICAN FERN JOURNAL 


The ferns of the southern states were treated by William R. 
Maxon, in J. K. Small’s ‘‘Flora of the Southeastern United 
States,’’!4 now somewhat out of date but still useful; the area 
covered includes all the region south of the northern boundaries 
of North Carolina, ne and Arkansas, and west to eastern 
Oklahoma and Texas, i.e. the northern boundary coincides with 
the southern io of Gray’s Manual and Britton and Brown. 
A more recent, but in some ways eccentric treatment, is J. 
Small’s ‘‘Ferns of the Southeastern States,’’!° which covers a 
similar area except that it extends west only to the Mississippi 
River. The southwest is more or less covered (unsatisfactorily) 
in ‘‘A Flora of Arizona and New Mexico,’’!® by I. Tidestrom 
and T. Kittell. 

ALABAMA: No recent work. 

AuaskKa: ‘‘Flora of Alaska and Yukon,’’!? by E. Hultén, with 
keys but no descriptions; ‘‘Flora of Alaska and Adjacent Parts 
of Canada,’’!8 by J. P. Anderson, with keys and descriptions, not 
altogether trustworthy, and with poor illustrations. 

RIZONA: ‘‘ Arizona Flora,’’!® by T. H. Kearney and R. H. 
Peebles, the treatment of the ferns by C. V. Morton, with keys 
but no descriptions or illustrations. 

ARKANSAS: No recent _ 

Cauirornia: Maxon, in Abrams, mentioned above; ‘‘Manual 
of the Flowering fen of rien Sakai by W.. L. Jocecn, the 


14 Ed. 2, pp. 1- se 1913. Published by the author, now deceased. Out 
of oe Not review 
. 1-517. 1938, "Published by the author. Out of print. Reviewed, 
1939. 


16 Pp, 1-897. 1941. Catholic University of America oe Washington, 
D.C. Reviewed, this Journan 32: 119. 1942. Out 2 Ue 
17 Lunds Univ. Arsskr. Ei, 437: -1-108, 1941... Per haps still oe 
through Dr. E. Hultén, Saag es Riksmuseet, Stockholm, Sweden. 
1942. 


L 
18 Towa State Coll. ea Ba 18: 137-175. a Rey Ligeia bay JouR- 
NAL 34: 64. 1944. Reprinted, Iowa State University, Ames 1959. 
19 Pp, 1-1032. 1951. University of California Pane Berkeley, "California. 
Tours print. A new edition vd be printed soon. Not reviewed in the 
A 


0 Pp. 1 238, fig. 1-1028. ese 3rd printi U. ty of California 
Pitie Tlorkeey, California, $6.5 x ees CEG AE DON 


Fern Books 13 


fern part out of date in concept and nomenclature and not on a 
par with the rest of the work. For southern California, ‘‘A 
Manual of Southern California Botany,’’?! by P. A. Munz, can 
be highly recommended for its accurate and careful treatment. 
A new treatment, nee Munz & Keck, was reviewed in the last 
number of the JOURNAL. 

CoLorapo : <iColorade Ferns,’’22 by H. D. Harrington and 
L. W. Durrell, with keys, descriptions, and line drawings, mostly 
good. A similar but abbreviated treatment, without the draw- 
ings, is found in Prof. Harrington’s ‘‘Manual of the Plants of 
Colorado.’’?8 

Connecticut: No recent treatment. 

ELAWARE: ‘Flora of Delaware and the Eastern Shore,’’** by 
Robert R. Tatnall, an annotated list; see also Maryland, below. 
Still available from Greenwood Book Shop, 110 West 9th St., 
Wilmington, Del. $3.50. 

District or ConumBia: see Maryland, below. 

Forma: ‘‘Ferns of Florida,’’> by J. K. Small. See also the 
reference above to Small’s larger book. 

rorerA: ‘‘Ferns of Georgia,’’> by R. MeVaugh and P. H. 
Pyron, an altogether admirable treatment, that will be useful 
throughout the sout 

Ipano: ‘‘Contributions toward a Flora of Idaho,’’? by Ray J. 


. 1-642, 1935. Claremont Colleges, Claremont, California. Out of 

print. aon reviewed in the JOURNAL. 
ae 1950. Colorado Agricultural and Mechanical College, Fort 
Collins, Uolorade, $1.00. Reviewed, this JouRNAL 41: 93. 1951. Out of 


Shp 1-666. 1954 (offset). Sage Books, Denver, Colo. Now gag Ph 
from The Spied ent Be 79 So. York St., Denver 10, Colorado. $8.0 
Bes eviews in the Jou 

Pp. 1-313. 1948. Published by the Society of Natural History of 
Dele Wilmington coh : Free Library, Wilmington 28, Delaware. 
Reviewed, this JOURNAL 38 

25 Pp. 1-236. Illus. Soe cblished by the author. Out of print. Not re- 


OURNA 
20'Pp. 1-195. Tilus. 1 1951. sae es - Georgia Press, Athens, Ga., $5. 
27 Leaflet 27, Pter ido phytes, pp. 1-37. 1949. Published by the ous 
Dept. of Botany, University oft Tia ho, Moseow, Ida. Not reviewed in the 
JouRNAL. Probably obtainable from William ©. Brown Company, Dubuque, 


174 AMERICAN F'ERN JOURNAL 


Davis, the pteridophyte treatment by Seville Flowers, a mimeo- 
graphed publication, useful but without illustrations. 

Tuurwors: ‘An Enumeration of Hlinois Pteridophyta,’’°§ by 
G. N. Jones, with keys and localities but no descriptions or illus- 
trations; ‘‘Vaseular Plants of Illinois,’”® by G. N. Jones and 

Fuller, similar but with maps showing the distribution. 

Tnprana: ‘‘Flora of Indiana,’’®° by Charles C. Deam, the most 
famous of the local floras of the eastern states, and deservedly 
so, representing as it does a lifetime of the most careful field and 
herbarium study by Dr. Deam. 

Towa: ‘‘Native Ferns of Iowa,’’#! by I. E. Melhus, with de- 
scriptions and maps but not the most trustworthy data. 

Kansas: No recent complete publication. 

Kentucky: No recent publication. 

Lovurstana: ‘‘Ferns and Fern Allies of Louisiana,’”? by Clair 
A. Brown gud Donovan S. Correll, an admirable work, with fine 
photographs 

Maine: ‘‘The Ferns of Maine,’’? by Edith B. Ogden, by far 
the best work for New England, with good descriptions and 
drawings. 

Maryuanp: ‘‘The Ferns and Fern-allies of Maryland and 
Delaware Including the District of Columbia,’’*4 by Clyde F. 


28 Am Midland Naturalist 38: 76-126. 1947. The es eracda Press, 
Notre Dame, Ind. Not reviewed in the JourNAL. Out of 

°9 Tilinois State Museum Sci. Ser. 6: 1-593. Maps 1— 1376, Cur 
rently available from Univ. of Illinois Press, Urbana, Ill. $10. 00. coe flawed, 


5. 1956. 
3 1-1236. 1940. gel Pai of pies hey Division of Forestry, 
Indianapolis, Ind. Out of print. Not reviewed in t OURNAL, 
1 Towa State College Redoet Cireular 225. on. Reviewed, this JouR- 
35 


32 Pp. sg 1942. Still available from Louisiana State University Press, 
00. 


33 Maine rah rel 41, 3 [University of Maine Studies no. 62], 
p. 1-128. 1948. Sti £ obtainable from University of Maine Library, Orono, 
Maine, * ee Review this JouRNAL 39: § 

34 Pp. 1-7 2, maps 1-58. 1953. Published by the author, 10105 
Harford ‘Roa’ Cee Md. $3.00. Reviewed, this JourNAL 44: 90. 


FrrN Books 175 


Reed, interesting and unusual in some ways, especially by the 
inclusion of photographs of the spores. 

Massacuusetts: No recent work. 

Micnigan: ‘‘Ferns of Michigan,’’®* by Cecil Billington; al- 
though not altogether adequate, still one of the better books that 
ought to be in fern students libraries. 

Minnesora: ‘‘Ferns and Fern Allies of Minnesota,’’*° by 
Rolla M. Tryon, Jr., a fine, professionally executed book, notable 
also for the fine drawings and unusual silhouettes of fern fronds. 

Mississtppt: No recent work. 

Missourr: ‘‘Ferns and Fern Allies of Missouri,’’8’ by M. E. 
Pinkerton, a descriptive account (containing a number of er- 
rors) ; ‘Ferns and Fern Allies of Missouri, 38 hy BH. J. Palmer 
and J. Steyermark, a cheek-list only, with commentaries. 

Montana: No recent work. 

Nepraska: No recent work. 

Nevapa: ‘‘Flora of Utah and Nevada,’”® by I. Tidestrom, the 
ferns contributed by William R. Maxon; a brief but authentic 
treatment, without descriptions or illustrations but with keys. 

New Hampsutre: ‘Ferns and Fern Allies of New Hamp- 
shire,’#° by Edith Scamman, a careful treatment, with good 

rawings. 

New Jersey: ‘‘The Ferns of New Jersey,’ by M. A. 


35 Cranbrook Institute of Science Bulletin 32, 1-240, pl. 1-16, fig. 
1-79. 1952. Available from Cranbrook a ce ‘ot tebe: enunnie 
Hills, Michigan. "$5.00. Reviewed, this JOURNA 

36 Pp, 1-166, fig. 1-207, maps 1-85. 1954. Univ Sreity of Minnesota Press, 
pre pole Minn. Paper $2.75, Cloth $4.00. 

7 Anna eee hier Gard. 20: 45-78, map, pl. 1 1933. Reviewed, 
nite focake 24: 1934. Available from Minouit” Basile Garden, 
2315 Sag Grove Rowe St. Loui uis, Mo., 

s JOURNAL 22: 105-122. 1933. Reviewed, this JouRNAL 24: 18. 


- 
39 Contr. U. S. Nat. Herb. 25: ade pl. Ane are Out of p 
40 New Hampshire Academy of Science Bull. 2 4-06. Pt. ba; Not? 
pate sah from Dr. R. L. Blickle, Secretary Fo died ett a pert 
f a by wipe N. A. $1. 95. Rey tones this JouRNAL 40: 8. 195 50. 
aj p. 1-201, 1-110, mage 1? 76. 19. Rutgers University Basel New 
Beaueriek, Nod “out of ‘print. bs this JOURNAL 38: 95. 1948 


176 AMERICAN FERN JOURNAL 


Chrysler and J. L. Edwards; especially notable for the fine 
pees ig of habit me of individual fronds. 

aes : “‘The Ferns and Fern Allies of New Mexico,’’#? 

As oa oui HK. F. Castetter, and O. M. Clark, with keys, 
pal ao tan and well-drawn illustrations. 

N. kK: No general treatment, except ‘‘ Annotated List of 
the Ferns and Flowering Plants of New York State,’’4? by 
Homer D. House, a list only, without keys, descriptions, or il- 
lustrations ; a more local but better treatment is ‘‘Ferns of the 
Vicinity of New York,’’#! by J. K. Small, in which ‘‘vicinity’’ is 
broadly interpreted as being ‘‘within a hundred miles of Man- 
hattan Island’’; this is a complete manual, with fine descriptions 
and drawings. 

NortH Carouina: ‘‘Ferns of North Carolina,’’#? by H. 
Blomquist ; a useful book, although not without some defects, 


Nortu Dakota: ‘‘Handbook of North Dakota Plants,’ ee I 
O. A. Stevens, a general manual in which the ferns are treated 
very briefly. 

Onto: ‘‘Ferns of Ohio,’’8 by H. H. Vannorsdall, an admir- 
able and useful book, with fine photographs. 

OxLanHoma: ‘‘Ferns of Oklahoma,’’!? by H. I. Featherly and 


42 Univ. of New Mexico Publ. Biol. 6: 1-139. 1954. Available from Univ. 
of ‘ae tte mbes Albuquerque, New Mexico $1.00. Reviewed, this 
JOURNAL 46: 34. 

43 New York pine abae Bulletin 254, pp. 1-759. 1924. Not reviewed 
in pe one Out o 

44 Pp. 1-285. Illustr. 1935, abies by the author. Out of print. Re- 
viewed, this JourNAL 26: 

45 Pp. 1-131, - g. 1-79. es Dak e University Press, eas North 
Carolina. Out of p int. Reviewed, this JourNAL 25: 59. 

46 “© A County Check Li a of North Carolina coe and Bex = Aiea 7? by 

oT and Donovan §. Correll. Journ. Elisha Mitchell Sci. Soc. 


. 1-324, fig. 1-319. 1950. Available from North Dakota eae 
for Re a, onal Siu, State College Station, Fargo, North Dakota, $4.50. Ni 
reviewed in the 

ote 1-298, “fa. sia 2165 1956. Obtainable from rage na Store, 
Wilmington, Ohio 0, $3.09. Reviewed, this JouRNAL 47: 957. 

; Oklahoma Agr. & Mech. Coll. Ex xper. Station Cire, 20 a 
1-24. Tllustr. 1939. Reviewed, this Journan 30: 102. 1940. Out of print, 


Fern Booxs 177 


Clara Still Russell, is brief and inadequate. 

OrEcon: ‘‘A Manual of the Higher Plants of Oregon,’’®° by 
M. E. Peck, an adequate ‘‘manual-type’’ account ; I understand 
that a new, revised edition is imminent. 

PENNsyLVANIA: There is no descriptive manual. Dr. E. T, 
Wherry’s ‘‘The Ferns and Lycosphens of Pennsylvania, ’’°! is an 
annotated check-list; Elsie Deane Canan’s ‘‘A key to the Ferns 
of Pennsylvania,’’®? is merely a key, although a rather full one, 
with rather primitive line-drawing illustrations. 

Ruope Isuanp: No recent treatment. 

SourH Carotina: No recent treatment. 

Sourn Daxora: No recent treatment. An annotated list in 
“‘Plora of South Dakota,’’®? by W. H. 

TENNESSEE: The finest of all state floras and a book that 
should be in every fern student’s library is ‘Ferns of Ten- 
nessee,’’=+ by Jesse M. Shaver; it is a model work, not easily 
imitated, since it represents many years of careful devoted work. 

Texas: ‘‘Ferns and Fern Allies of Texas,’”°> by Donovan S. 
Correll, is one of the newest and best manuals, professionally 
executed. 

Uran: ‘‘Ferns of Utah,’’®* by Seville Flowers, with keys, de- 
scriptions, and accurate, tasteful drawings; to be recommended. 

Vermont: No recent specialized flora. An annotated list®? is 


50 Pp, 1-866. 1941. Published by the author, Willamette University, 
Salem, Oregon. Out of print. Not reviewed in the JOURNAD, 

51 Bartonia 21: 11-37. 2. Reviewed, this JournaL 32: 117. 1942. 

52 Pp, 1-110, fig. 1-59. 1946. Published by The Science Press Printing 
Company, Lancaster, Pa., $1.50. Reviewed, this JOURNAL 36: 124. 1946. 
Available from Miss Canan, 1023 Menoher Blvd., Johnstown, Pa., $1.50. 

53 Pp, 1-16. Ilustr. 1932. Out of print. ae 

54 Pp. 1-502. fig. 1-243. 1954. Available from Bureau of Publications, 

. $6.00. Reviewed, 

this JouRNAL 45: 21. 1955. at 

55 Pp, 1-188. pl. 1-38, fig. 1-3. 1956. [A reprinting of ‘‘Flora of Texas, 
vol. 1, part 1. 1955]. Texas Research Foundation, Renner, Texas. $5.50. 
Reviewed, this JouRNAL 47: 79. 1957. 


56 Bull. Univ. Utah 35: pp. 1-87, fig. 1-164. 1944. Reviewed, this 
JourNAL 35: 61, 1945. Still obtainable from the University of Utah Press, 
Salt Lake City, Utah. $1.00. : - 

57 3rd rev. ed., 1-353 37. Still obtainable from Dr. H. - ae 


Vv. pp. monk’) , 
mann, Department of Botany, University of V ermont, Burlington 


178 AMERICAN FERN JOURNAL 


presented in ‘‘The Flora of Vermont,’’ edited by E. J. Dole. 

Vireinia: ‘‘The Ferns and ‘Fern Allies’ of Virginia,’’®® by 
A. B. Massey, a usable pamphlet. 

WasHIneton: No recent state treatment. See above the more 
general works of Maxon (in Abrams) and Frye. 

West Vireinta: ‘‘The Pteridophytes of West Virginia,’ by 
M. G. Brooks and A. 8. Margolin; a more recent treatment, in 
abbreviated style, is in ‘‘Flora of West Virginia,’’®° by P. D 
Strausbaugh and Earl L. Core 

Wisconsin: ‘‘The Ferns gad Fern Allies of Wisconsin,’’® by 
R. M. Tryon, Jr., D. W. Dunlop, N. C. Fassett, and M. E. Diemer, 
a fine work with beautiful photographs, somewhat diminished in 
effectiveness in the second edition (by offset). 

yomIna: ‘‘The Ferns and Fern Allies of Wyoming,’’®? by 
C. L. Porter, a mimeographed publication, showing thought and 
care; to be recommended. 


8 2nd ed., Virginia Polytechnic Institute Agricultural Extension Fgptlom: 
Bulletin 256, pp. 1-78, fig. 1-21. 1958. Available gratis from Prof. 
i Reviewe J 
1958 


ie, eksburg . Vv. 

Bi. f 1e first edition reviewed this JouRNAL 34: Q : 
; e irginia University Bulletin, Series 39, no. 2, pp. 1-60, i106. 
8. Reviewed, this JouRNAL 1940. A few copies edietic for 
lens gratis by E. L. Core, Waiveraity of West Virginia, Morgantown, 
Win Part 1. West Virginia Univ. Bulletin, Ser. 52, 12-2. 1952. Not re- 
viewed in the JouRNAL. Available from Tablas Boskatocs atic ite 
W. Va., $1. op 

61 Kd. 2, pp. 1-158, ig 1-213, maps 1 1953. University of Wisconsin 
Press, Madison! ci 50. Reviewed, ‘his jJovwwan, 433 177. 1953. [First 
bcos: reviewed this Doceeat Sis 24 
2Pp. 1-18: ‘The saad Mountain Horba yarium Leaflet 27. 1957. Not re- 
viewed n the JourRNAL. To be obtained — on lata: tocrrot. 

orter, UneniG at Wyoming, Laramie, Wyoming. 


SMITHSONIAN InstTITUTION, Wasuineton, D. C. 


GENESIS OF FERN JOURNAL 179 


The Genesis of the American Fern Journal: 
Supplementary Data 


Raupeu C. BENEDICT 


The article in the preceding number of the Fern Journal, 
“<The Genesis of the American Fern Journal,’’ was written at 
Pilot Knob, New York, without benefit of correspondence or 
Fern Society files. Reference to these files has not disclosed any 
factual errors in the earlier account, but it has added consider- 
able data from official files which seem worth recording at this 
time. In particular, a circular, mimeographed letter sent to all 
members during 1909 by Mr. Evelyn J. Winslow, President of 
the Society for that year, makes very explicit the problem that 
faced the officers of the Fern Society with respect to a possible 
new, Society-owned publication. The letter, which follows, car- 
ries its own message. It may be noted here that Society members 
responded to this letter with a vote of sixty to ten in favor of a 
new journal. 

“The Fern Bulletin has heretofore been furnished to mem 
bers of the Fern Society for 60¢ per member. For the year 1910, 
the publisher [Willard N. Clute] offers it at 64¢ with a discount 
of 4¢ if the whole subscription is paid in advance by Jan. Ist. 
The Executive Committee is in doubt whether to accept these 
terms, and takes this way of getting the sentiment of the mem- 

ers. 

‘‘The proposition to issue a Society journal has been for some 
time under consideration and the contingencies involved investi- 
gated. The cost of printing and mailing a 30 page quarterly 
would be about $130.00. We should pay about $100.00 for the 
Bulletin [The Fern Bulletin] and $20.000 for the Annual Re- 
port, which could be included in the Society journal. This leaves 
a very small margin to be made up by advertising and outside 
subscriptions. Of course it would be the business of the publish- 
ing committee to keep the size of the paper within the means of 
the Society ; but in case of a deficit, several members have already 
volunteered to make it good. 


180 AMERICAN FERN JOURNAL 


““We have several members who are entirely competent to 
do the editorial work, and one [Philip Dowell], who is associate 
editor of one of the leading botanical publications of the coun- 
try [Bull. Torrey Botanical Club], has signified a willingness to 
assist in this way, if called upon to do so. 

““By owning our publication we should gain immunity from 
the risk of being suddenly left without an official organ through 
no act of our own; freedom from the necessity, often embarrass- 
ing and sometimes humiliating, of periodically making terms 
with a publisher who has us at his mercy; the assistance of sev- 
eral of our ablest fern students, who, for reasons personal or 
otherwise, have long refused to contribute to the Bulletin; a 
stronger sense of common interest and responsibility among the 
members, increased growth, and an improved stature among the 
botanical organizations of the world. 

“It is important that every member should vote, and do not 
forget to sign your name to the card.’’ 

The President 


Officers’ reports published during the next two years (1910 
and 1911) provided considerable additional data for an under- 
standing of the events that led to the institution of the Journal. 
As to its reception, while there is no information available about 
possible withdrawals from the Society, the first year of the 
Journal’s adoption recorded a gain of some thirty in the total 
membership, to pass the 200 mark for the first time. 

ow, fifty years later, perhaps a few reminiscent comments 
may be in order. From the first year, the editorial policy was to 
try to maintain a fair balance in the Journal between technical 
articles and those of special interest to members just starting 
fern study. William R. Maxon, a tower of strength to the Jour- 
nal through the years, transferred his taxonomic ‘‘Notes on 
American Ferns’? series to the Fern Journal. About the same 
time, he arranged for the series entitled ‘‘Notes of naturalist 
afloat’’ by Safford. During those early years, friendly amateurs 
used to write in occasionally asking for more articles of a popu- 


ISOETES MELANOPODA 181 


lar nature. Anyone interested may decide whether a fair balance 
has been kept by reviewing the 198 quarterly issues and the 
7,400 pages which constitute the Fern J ournal up to 1960. 
There can be no question that in the many pages of the back 
numbers of the Fern Journal beginning as well as advanced 
fern students will find a wealth of significant articles. 

Now, to start the fiftieth volume we have the outstanding 
Golden Anniversary Number which is a tribute to the efforts of 
the editor in furthering the Fern Journal and the Fern Society. 
Although this first 1960 issue is predominantly technical, not a 
few of its articles will be of interest to members whose fern study 
is only a few years old. It is my hope, however, that this fiftieth 
volume may also see a plentitude of articles which will provide 
immediately interesting reading for the newest tyro among our 
members. 

Pitor Knos, New York. 


Isoetes melanopoda in Southern Illinois 
Rosert H. MoHLensRocK, JR. 


Just as the original collection of Lsoétes melanopoda Gay and 
Dur. by Elihu Hall in 1853 was accidental, so was the recent 
discovery of this fern ally in southern Illinois. 

Engelmann in 1886 described the original discovery by Hall 
at Athens, Menard County, Illinois, ‘‘. . . Mr. Hall was acci- 
dentally led to the discovery of this plant on his farm in 1853 
by finding its trunks and spores in turning up the soil for brick- 
making; ...’’ 

On June 13, 1955, the author made one of his frequent stops 
along Illinois Route 3 in western Jackson County to study the 
flora that occurs in the roadside ditches along that highway. On 
this particular date, the site selected was approximately three 
miles south of the junction of Illinois Routes 3 and 144, and 

1§ee, Benedict, R. C. The American Fern Journal through Thirty 


Years. This JouRNAL 31: 41-48. 1951, and also Wherry, E. T. Cumulative 
Index Volumes 1 to 25. [Available for sale from Treasurer, 25¢.] 


182 AMERICAN FERN JOURNAL 


about one mile south of Worthen Bayou. As I walked down the 
slope from the highway into the ditch, it was apparent that a 
fresh mowing had occurred along the length of the twenty-foot 
wide ditch. Grasses and sedges had been made destitute of their 
crowns. Many of the bunch-plants which had precarious anchor- 
age because of the soggy character of the ground in which they 
grew fell easy prey to the mower and were knocked over, some- 
times being detached completely from the soil. Thus was the case 


Fie. 1. Inunpatep Dircn wirn ISOETES MELANOPODA, PHOTOGRAPH BY 
N W. Voter 


with the Isoétes melanopoda. Here staring at me with their 
blackish ‘‘eyes’’ were the megasporangia at the base of the 
megasporophylls or ‘‘leaves.’’ Only ten reports have been re- 
corded for this species in Illinois, most of them not since the turn 
of the century. And this was the farthest south in Illinois that 
the Black-based Quillwort has been found. On further exami- 
nation, hundreds of specimens were observed. 


AMERICAN FERN JOURNAL VoLtuME 50, PLatE 15 


MELANOPODA FROM SOUTHERN ILLINOIS; 3/5 NATURAL SIZE. 


PHOTOGRAPH BY JOHN W. VOIGT 


184 AMERICAN FERN JOURNAL 


The ditch is from one to six feet lower than a highway and 
averages nearly twenty feet wide (Figure 1). much as one 
foot of water may stand in it for a short oe ok year, al- 
though the water depth is generally only a few inches. During 
the driest seasons of the year, there may be no standing water, 
although the ground always remains spongy underfoot. 

A luxurious growth of vegetation occurs along the entire 
length of the ditch. Several rare species for Illinois are found 
here. Grasses and sedges abound, although the grasses seldem 
are allowed to grow large enough to flower. Spike rushes 
(Eleocharis Smallvi and Eleocharis tenuis) and rushes (Juncus 
bufoniwus and Juncus effusus) are common. Species of Carex in- 
clude Carex caroliniana (the only station for it in Illinois) and 
Carex granularis. The beaked rush (Rhynchospora corniculata) 
has its most northern station in Illinois along this ditch. 

tyledons are not uncommon. Clammy hedge hyssop 
(Gratiola virginiana), water primrose (Jussiaea diffusa), and 
water starwort (Callitriche heterophylla) occur in the more 
deeply inundated areas, while buttonweed (Diodia teres), Am- 
manma coccinea, and Phyllanthus caroliniensis are found in less 
moist situations, 

Most of the specimens of Isoétes melanopoda were robust 
(Plate 15). As many as eighty leaves were counted on some of 
the larger plants; these reached a length of 25 em. The corms 
measured 2.5 em. across in a few specimens, indicating a ie 
age for these individuals. Growing with the very common dark- 
based plants were scattered individuals of forma pallida Fernald, 
with pale bases. In these specimens, the megasporangia were 
cream to pinkish in color, although in some, traces of black could 
be observed. Those referable to forma pallida were fully as 
robust as the darker ones. 

Subsequent visits to the ditches along Highway 3 have led to 
the discovery of this species on both sides of the highway for a 
distance of two miles. 


REFERENC 
ENGELMANN, G. The genus techn in neki Ameriea. Trans. St. Louis 
Aead. Sci. 4: 358-390. 1886, 


EQUISETUM LAEVIGATUM 185 


The Smooth Scouring Rush and Its Complexities 
Ricuarp L. Hauke 


The smooth scouring rush, Equisetum laevigatum, is the only 
species of Equisetum endemic to North America and thus should 


cerning this species that most botanists, amateur and pro- 
fessional, are uncertain about whether they actually know it. As 
part of a recently completed monographie study of the genus 
Equisetum subgenus Hippochaete,' I devoted special attention 
to this species, and wish to discuss the results of that study 

In 1840, Nicholas Riehl collected a smooth-stemmed scouring 
rush along the banks of the Mississippi river below St. Louis, 
which Alexander Braun named E. laevigatum in 1844. A fruit- 
ing specimen of Riehl’s collection is to be found in the New York 
Botanical Garden herbarium and another is in Vienna. George 
Engelmann, having seen Braun’s manuscript apparently, went 
out along the Mississippi banks to collect this new species. The 
specimens he collected in August, 1843, have since been mis- 
takenly considered the type. They are without cones and show 
the autumnal condition of the species, with colored sheaths 
reminiscent of those in E. hyemale. However, all have the in- 
ternal structure as well as the smooth stems characteristic of 
E. laevigatum 

In 1902, A. re Eaton described a new Equisetum intermediate 
between E. hyemale and E. laevigatum, which he named £. 
hyemale var. intermedium. According to Eaton it had the exter- 
nal appearance of the former species and the internal structure 
of the latter species. He stated that Milde, certainly, and A. 
Braun, probably, based their descriptions of EF. laevigatum part- 
ly on this plant. Eaton later (1903) described another new 
Equisetum of this alliance, E. funstonii. 


1 Doctoral thesis, Department of Botany, University of Michigan, and 
available on microfilm from University Microfilms, Inc., Ann Arbor, Michi- 
gan, at moderate cost. 


. 


186 AMERICAN FERN JOURNAL 


Willard N. Clute collected a plant at Joliet, Illinois, that he 
named E. ferrissti in 1904. He wrote to Eaton that it seemed 
much like the descriptions of E. hyemale var. intermedium but 
did not agree with the co-types of that variety, differing mainly 
in the very long internodes. Clute later (1928) stated that the 
spores of E. ferrissii appeared abortive. An isotype in the Gray 
Herbarium of Clute’s species bears a close resemblance to E. 
hyemale var. intermedium and E. ferrissii is thus a synonym for 
Eaton’s variety. 
The most widely recognized American authority on Equisetum, 
John H. Schaffner, named a new species in 1912. He considered 
. hyemale var. intermedium to be the same as H. laevigatum roy 
segregated the vgn with smooth, deciduous stems as LH. - 
anum. He not study the internal structure, to which uk 


floras and manuals, and his interpretation has become the widely 
accepted one. Farwell (1917) protested that Braun’s original 
description of E. laevigatum was clearly of the same species 
Schaffner was describing as EF. kansanum, except for a misunder- 
standing of the annual nature, but Farwell’s protest went 
unheard. 

n a recently completed study (1958), Emily L. Hartman 
worked with plants identified as Z. lacvigatum, E. hyemale var. 
intermedium, E. funstonii, and E. kansanwm, and reached the 
conclusion that the western American H. funstonii and the east- 
ern 17. kKansanum were identical, that the two together form a 
subspecies, subsp. funstonii, of EB. laevigatum, and that E. hye- 
male var. intermedium is a synonym of EL. laevigatwm. She sepa- 
rated the subsp. funstonii from subsp. laevigatum on the col- 
lenchyma ratio and cone apex. The first has carinal collenchyma 
(she called it sclerenchyma) equal to or less than vallecular 
collenchyma and the cone apices are blunt to acute. The latter 
has carinal collenchyma exceeding the vallecular collenchyma, 
and apiculate cones. 


EQUISETUM LAEVIGATUM 187 


Dr. Hartman stated (p. 140) : ‘‘The distribution of the three 
intergrading taxa throughout the same geographical range, and, 
more significantly, their occurrence in identical habitats alone 
invalidates the recognition of the entities in question as distinct 
species.’’? However, it is just as difficult to conceive of sympatrie 
subspecies. Partial isolation of a portion of the range of the 
species in question would be expected to precede the development 
of any discontinuous variation sufficient to be recognized as of 
subspecific importance. Stebbins (1950, p. 50) said: ‘‘The sub- 
species, on the other hand, is usually conceived of as a group of 
populations with a common origin and a more or less integral 
geographic distribution, which has acquired its distinctive mor- 
phological characteristics partly through the influence of similar 
environmental factors, but also to a large extent through partial 
isolation from other subspecies.’’ 

Perhaps if Dr. Hartman had studied all of the species of 
Hippochaete in North America and sought as many characters 
as possible, such as spore size and appearance, her conclusions 
might have been different. She might have suspected that the 
two groups of plants intergraded not necessarily because they 
are subspecies but possibly for some other reason. Even were 
these two really subspecies, it is impossible to understand how 
Dr. Hartman attached the name laevigatum to the group she did. 
The specimens she erroneously considered the type for this name 
all have the vallecular collenchyma reaching the vallecular canal, 
which is the key character for her subsp. funstonii. In fact one 
of the specimens of Engelmann’s August 1848 collection was 
annotated subsp. funstonii by Dr. Hartman. Conversely, she 
annotated one of the specimens of the type collection of subsp. 
funstonii as subsp. lacvigatum. Apparently she considered any 
specimen with the carinal collenchyma exceeding the vallecular 
by the slightest amount as subsp. laevigatum and any specimen 
with the vallecular collenchyma equal to the carinal as subs 
funstonii, Since within a single clone of E. lacvigatum the 
depauperate stems and the upper portions of normal stems may 
have slightly greater carinal collenchyma, whereas the lower 


188 AMERICAN FERN JOURNAL 


portions of the same stems may have slightly greater vallecular 
collenchyma—this is especially true of many specimens in the 
Southwest—this single variable character is insufficient for de- 
limiting two subspecies. 

Thus it appears that a misconception of the nature of specia- 
tion and a lack of sufficient information led Dr. Hartman to an 
erroneous conclusion. The consideration of E. funstonn and E. 
kansanum as synonymous is probably correct. The placing of 
this taxon as a subspecies of another taxon consisting of EL. hye- 
male var. intermedium and E. laevigatum as synonyms is ques- 
tionable. 

The discovery of aborted spores and irregular meiosis in some 
specimens identified as HE. hyemale var. intermedium and E. 
laevigatum first led me (1958) to the suspicion that hybridization 
between E. hyemale var. affine and E. laevigatum was a factor in 
the confusion attendant upon these species. I examined more 
than two thousand herbarium specimens of these ae collected 
in the United States. Specimens with aborted spores and those 
vegetative specimens resembling them were segregated into a 
group. A comparison of this group of suspected hybrids with 
the parents, involving statistical analysis of measurements from 
more than one thousand of the specimens examined, revealed that 
the suspected hybrids were intermediate between the parental 

types, as shown in Table 1 


TABLE 1 
E. hyemale E. 
Character var. affine Putative Hybrid laevigatwm 
Stem diameter ! 7.80 5.80 4.90 
Stem height 2 7.84 7.46 5.80 
dge nu 30.14 22.04 21.97 
Sheath length 1 9.21 10.98 10.47 
Sheath width 1 8.34 6.74 5.90 
th ratio l:w 1.06 1.68 1.80 
omatal length 3 84.9 87.1 91.4 
Stomatal width 3 TES a 67.3 
Stomatal ratio 1:w 1.09 1.24 36 
Cone apex — apiculate PT iproighies apiculate blunt 
Stem duration evergreen s only ep deciduous 


g 
Collenchyma ratio (1:8)1:7-1:4(1:3) srt ay re 3-1:2(2:3) (2:3)1-3:2 


1JIn millimeters. 2 In decimeters. 3 In microns. 


EQUISETUM LAEVIGATUM 189 


All values in Table 1 are means except the last. This is a 


cur 
under the middle of the grooves (vallecular collenchyma) an 
ridges (carinal collenchyma). These strands extend radially 
toward the center of the stem. The collenchyma ratio is the ratio 
of the radial measurement (measured in an internode somewhat 
below the middle of the stem) of the vallecular collenchyma to 
the carinal collenchyma. Thus, Z. hyemale has extensive devel- 
opment of the carinal collenchyma and slight development of the 
vallecular collenchyma, whereas in EL. laevigatum they are both 


Statistical analysis of herbarium specimens, as shown in Table 
1, reveals the intermediacy in many characters of the specimens 
suspected of hybrid origin on the basis of their aborted spores. 
That the suspected parents, E. hyemale var. affine and E. laevi- 
gatum, could cross has been shown by culture of isolated gameto- 
phytes on an inorganic agar medium and controlled crossing of 
antheridial gametophytes of one species with archegonial gameto- 


demonstrated that the sperm from either species could fertilize 
eges from the other with the production of viable sporophytes. 
One fact which at first seemed inconsistent with the interpre- 
tation of the existence of a hvbrid between E. hyemale var. affine 
and EF. laevigatum was the presence of the supposed hybrids in 
areas where one or both parents were absent. Such is the case in 
New York, Pennsylvania, New Jersey, Connecticut, West Vir- 
ginia, and Virginia. All specimens from these states identified 
as E. laevigatum appear to be of hybrid origin although the 
species FE. laevigatum ranges only as far east as Ontario, Michi- 
gan, and Ohio. The answer is apparently distant dispersal by 
vegetative means, particularly by water transport of fragments 


190 AMERICAN FERN JOURNAL 


of living plants. In Michigan, the sandy shores of many lakes 
are overgrown with various Hippochaete, providing an abundant 
supply of living stem fragments. Victorin (1927, p. 94) de- 
scribes the vegetative dispersal of Z. x litorale and E. hyemale 
var. jesupii (E. X trachyodon), two other hybrids, by ice action 
along the shores of the St. Lawrence river. Since most of the 
localities of the hybrid east of Michigan are in contact with the 
Great Lakes waterway it is quite conceivable that the plants 
were carried there by water. Many localities from central New 
York seem to be exceptions until it is noticed that they occur 
along a barge canal connecting Lake Erie to the Hudson River, 
and are thus also in contact with the Great Lakes waterway. 

That vegetative propagation can readily occur was shown by 
an experiment in which over a hundred segments of green stems 
of the hybrid containing at least one node each were placed 
around the edge of an old gravel pit and more than twenty small 
plants were produced from them. 

The occurrence of widespread hybridization between E. hye- 
male var. affine and E. laevigatum is therefore established. It 
has been shown that there are many specimens morphologically 
intermediate between those two species, and that these intermedi- 
ate specimens have aborted spores. The ability of sperm from 
one species to fertilize eggs of the other has been demonstrated, 
as has also the ease of vegetative dispersal of the hybrid. Thus 
can be explained the presence of the sterile hybrid in cases 
where one or both parents are absent. 

This knowledge of the existence of a hybrid between EF. hye- 
male var. affine and E. laevigatum permits a clarification of the 
nomenclature of these widespread and long confused taxa. The 
correct name for the hybrid is ZH. * ferrissii Clute, the first 
legitimate binomial applicable to the hybrid. The name LE. hye- 
male var. intermedium, given by Eaton, cannot be used because 
an interspecific hybrid can hardly be considered a variety of one 
parent. The synonomy of EZ. >< ferrissii and its parents is as 
follows. 


EQUISETUM LAEVIGATUM 191 
EQuUISETUM HYEMALE VAR. AFFINE (Engelm.) A. A. Eat. Fern Bull. 11: 
Wid 


903. 
EF. prealtum Rat. Fl. Ludovie. 13. 1817. 
E. laevigatum y elatum Engelm. Amer. Journ. Sci. 46: 87. 1844. 
Br. ibid. 88. 


E. robustum B minus Engelm. ibid. 
‘ Engelm. ibid. 
E. hiemale var. californicum Milde, Verh. Zool.-Bot. Ges. Wien 12: 1264. 
E. hiemale var. iaponicum Milde, Ann. Mus. Bot. Lugd.-Batavi 1: 68. 
1864. 
E, robustum var. drummondii Milde, Equiset. 539. 1867. 
E. hiemale f. pl i Prager ea A. A. Eat. in Gilbert, List N. 
id 


m. Pterid. 26 
. hiemale var. evade As A. Bat. roe Bull. 11: 108, 1903. 


Sees 
= 
SS 
z 
= 
m] 
= 
= 
° 
= 
=: s 

ON 
=? 
{sab 
a 
o 
Vw 
>. 
aa 
2o8 
© 
> 
he 
“3 
R 
hm 
H 
sa 


ee var. robustum (A. Br.) A. A. cm pie 
E. hiemale var. prealtum (Raf.) Clute, Fern Bull. 16: 18. 1908 
Maporhacte ena var. californica (Milde) Farw. Mem. ew York 
6: 


HH: oa (Raf.) Park ibid. 

H. prealta var. affinis (Engelm. . pols ibid. 

it, Fated var. pseudohyemalis Farw. Amer. n Journ; 3: 76) 1917. 

E. affine Rydb. Fl. Rocky Mts. and Adj. ra is 1917. 

E. hyemale var, affine f. pumilum Viet. Contr. Lab. Bot. Univ. Montreal 
S: 89, 1927. 

E. hyemale var. ramosum Honda, Bot. Mag. Tokyo 47: 435. 1983. 

E, iomared Iljin, Flora URSS 1: 110. 1934. 

H. prealta subvar. coe ae Rarw: Amer. Fern Journ. 27: 17. 
1937, 


H. prealta var. pseudohyemalis subvar. polystachya (Prager) Farw. ibid. 
. californicum (Milde) G. N. Jones, Univ. Wash. Pub. Biol. 7: 23, 
T&.1986, 


byt 


. kansanum f. elatum (Engelm.) Broun, Index N. Am. Ferns. 89. 1§ 
hyemale var. californicum f. iar m (A, A. Eat.) Broun, ibid. 
prealtum f. drummondii (Milde) Broun, ibid. 93. 

realtum var. affine (Engelm id. 
. affine f. neopo Bl eivem (Farw.) Broun, ibid. 
prealtum var. affine £. polystachyum (Prager) Broun, ibid. 94. 
prealtum var. affine f. pumilum (A. A, Eat.) Broun, ibid. 
prealtum var. affine f. ramosum (A, A. Eat.) Broun, ibid, 


Ss ae 
2 
= 
S 
4 
9 
2 


192 AMERICAN FERN JOURNAL 


E. hyemale var. elatum (Engelin.) Morton, Leafl. West. Bot. 16: 156. 
1951 
E. hyemale var. pseudohyemale (Farw.) Morton, in Gleason, New Ill. 
.and Adj. Can. 1: 16. 1952 
HQUISETUM LAEVIGATUM A. Br. Amer. Journ. Sci. 46: 87. 1844. 
E. laevigatum B scabrellum a ae Amer. Journ. Sci. 46: 87. 1844. 
funstoni A. A. Eat. Fern Bull. 11: 10. 190 
funstoni f. caespitosum A. A. Eat. ibid. 11. 
funstoni f. nudum A. A. . ibid. 
. funstoni f. ramosum A. A. ee ibid. 12. 
funstoni f. polystachyum A. A. Eat. ibid 
laevigatum f. ramosum A. A. Eat. ibid. 49, 1903. 
. laevigatum f. caespitosum A. A. Eat. ibid. 43. 
. laevigatum f. variegatoides A. A. Eat. ibi 
oe f; ee A. A ane ibid. 44. 
fin. Ohio Nat. 13: 9 
aes: iui (A. Br.) Farw. ee Nee York Bot. Gard. 6: 
469. 1916. 


peeks Hw 


H. prealta var. scabrella (Engelm.) gree ibid. 

H. laevigata var. eatonii Farw. aes 

H. laevigata var. funstoni (A. Aga sy iid. 471. 

H. laevigata var. polystachya (A. A. Eat.) Farw. ibid. 

E. funstonii var. caespitosum Jeps. Man. Fl. Pl. Calif. 40. 1923. 

E. funstonii var. nudum Jeps. ibid. 

E. funstonii var. ramosum ees ibid. 41. 

E. fontinale Copel. Madrofio 3: 367. 

Hf. laevigata var. ramosa (A. A. Eat.) aes: Amer. Fern Journ. 27: 17. 
1937. 


H. laevigata var. caespitica Farw. ibid. 
. laevigata var. caespitosa (A. A. Eat.) Farw. ibid. 

H. laevigata var. variegatoides (A. A. Eat.) Farw. ibid. 

E. kansanum f,. caespiticum (Farw.) Broun, Index N. Am. Ferns 89. 
1938. 

E. kansanum f. paige as (A. A. Eat.) Broun, ibid. 

E, kansanum f. eatonii (Farw.) Broun, ibi 


E. kansanum f. patie ak (Gea: Fat.) Broun, ibid. 
E. kansanum f. ramosum (A. A. Eat. ) atten ibid. 

E. kansanum f. variegatoides fe x at.) rae ibid. 
E. laevigatum f. scabrellum Meta pron . a0. 


E. laevigatum subsp. funstonii (A. A. Eat.) "chal Trans. Kansas 
Acad. Sci. 61: 144, 1958 


EQUISETUM LAEVIGATUM 193 


EQuIsETUM X F¥RRISSII Clute, Fern Bull. 12: 22. 1904. (pro. sp.). 

E. hyemale var. affine X laevigatum 

. hiemale intermedium A. A. Eat. Fern Bull. 10: 120. 1902. 
Eat. ibid. 122. 

: AS 10 

oe iatile L. var. siccwm Lunell, Bull. Lied Heeb: 25-75. 1908; 
Hippochaete siatey var. intermedia (A. A. Eat.) Farw. Mem. New York 

Bot. Gard. 468. 1916. 


(ees 
> 
~ 
d 
3 
= 
a 
= 
= 
Dd 
3 
3 

s 
d 
2 
= 
lac) 
PS 
= 
Se 
nm 
Saal 
S 
S 
> 
e 
= 
= 
> 


H. prealta var.  kaaech (A. A, Eat.) vig ibid. 
vee laevigata var. eatonii sees ibid. 470. (pro. te). 
E. intermedium Rydb. Fl. Rocky Mts. ret Adj. fe 1053. 1917. 
E. lacigatum # cbneieitia Haberer, Bull. New York State Mus. 243— 
: Siw cinterdécilinire f. proliferum Haberer, 7 
oy ene var. affine f. intermedium Vict. nalts Sah Bot. Univ. Mon- 
Ls 80. 
E. ae ds iain (A. Br.) Bush, Am. Midl. Nat. 12: 111. 1930. 
E. laevigatum auct. non n A. Br.: Schaffner, Ohio Nat. 13: 19-22. 1912; 
and most subsequent author 
E, seereey subsp. pecnates of Hartman, Trans. Kansas Acad. Sci. 
5-148. 1958 


LITERATURE CIT 
Braun, A. 1844. A monograph of the Hod American species of the 
us Equisetum. Amer. Journ. Sci. 46(1): 81-91. {Dated October- 


CLuTE, W. N. 1904. A new species of Equisetum. Fern Bull. 12: 20-23. 


f inois. 
Eaton, A. A. 1902. A new Equisetum. Fern Bull. 10: 120-122. 1902. 
—, 1903. The genus Equisetum in North America, XII. Fern Bou. tit" 7- 


‘1p. 

Hartman, E. L. 1958. The taxonomy = ret - the Equisetum 
laevigatum complex. Trans. Kansas Acad. Sei. 61: 148. 

Havkg, R. L. 1958. Is iia laevigatum a genet wees Fern Journ. 
48: 68-72. 

Scuarrner, J. H. 1912. An undescribed Equisetum from Kansas. Ohio 
Nat. 13: 19-22. 

Sreppins, G. L. 1950. Variation and evolution in plants. 
versity Press, New Yor 

Marir-VictTorin, F. 1927. Les Equisétinées du Quebec. Contr. Lab. Bot. 
Univ. Montreal No. 9 
DEPARTMENT OF BoraNy, UNIVERSITY OF RuopeE ISLAND, KING- 

ston, RHopE ISLAND. 


Columbia Uni- 


194 AMERICAN FERN JOURNAL 


The Gametophyte and Young atest of 
Athyrium esculentu 


B. K. Nayar 


Athyrium esculentum (Retz.) Copel. is one of the species of 
ferns that has suffered the most in nomenclature, having been by 
different authors attributed to at least eight different genera 
(Henuonitis, Diplazium, Asplenium, Anisogonium, Microstegia, 
Callipteris, Digrammaria, and Gymnogramme). It extends from 
Polynesia to India, growing as a straggling weed in marshy, or 
just moist, areas which are not necessarily shaded. A. esculentum 
lacks the elegance of most other species of Athyriwm and con- 
sequently is not favoured as an ornamental fern, although it is 
comparatively easy to cultivate and is one of the most important 
of ferns as human food. The tender leaves of the plant are used 
as a vegetable in preparing tasty salads, pickles, ete. 

Comparatively little is known regarding the gametophyte of 
Athyrium esculentum or for that matter any species of Athyrium. 
In view of this, spores were collected from plants growing at the 
National Botanic Gardens (Lucknow) and sown in September, 
1955, on sand beds irrigated from below and maintained in a 
glass house. The technique followed is as described earlier 
(Kachroo & Nayar, 1953; Nayar, 1954). 

The spores of A. esculentum (Figs. 2, 3) are bilateral,! ani- 
sopolar, with a single linear short proximal laesura, of medium 
size, monolete, concavo-convex in equatorial view, with one of the 
equatorial ends narrower than the other and with a brown 
seabrate exine (having small irregular elevated patches). The 
exine pattern (Fig. 1) is discernible only in acetolysed and 
bleached preparations. The average size of the spores is P 29.16 p, 
KE; 44.00» and Ey 30.24. The size variations are: P 25.00 to 
32.50 w, E, 39.50 to 50.50 » and Es 25.00 to 36.00 p. 

In culture the spores germinate within a week. The first 


acetolysis method (Erdtman, 1952) was used in the study of spores 
oe iow termpanites used in spore description is after Harris (1955). 


ATHYRIUM ESCULENTUM 195 


rhizoid protrudes as a papillose structure, the exine rupturing 
at the laesura, and is soon cut off from the body of the spore by a 
basal wall. The rhizoid in early stages may have a few included 
chloroplasts. The germinating filament originates laterally to 
the rhizoid towards the narrower end of the spores (Migs. 4, 5 ). 
As it grows the exine splits longitudinally into two and remains 
attached to the basal cell till very late in development. The 
germinating filament becomes 3 to 4 cells long before the forma- 
tion of the prothallial plate begins (Fig. 6). The cells are 
broader than long and densely chlorophyllous. Rhizoids originate 
laterally. Some of the cells of the filament may form antheridia 
at this stage. 

An obeonical apical meristematic cell is established in the usual 
way and a spatulate prothallus is formed within a month after 
germination (Fig. 7). Soon the gametophyte develops a cordate 
apex with the meristematic cell lodged at the bottom of the 
apical notch (Fig. 8). Antheridia are formed continuously from 
the filamentous stage onwards and are both marginal and super- 

cial. 

The prothallus remains naked till it becomes distinctly cordate, 
when marginal unicellular, elub-shaped hairs are formed (Figs. 
9, 10). Bach hair originates as a mammilliform protuberance, 
which is cut off by a basal wall from the parent cell. The pro- 
tuberance elongates and the apex becomes highly vacuolate. 

oon, a greenish-yellow extracellular cap is secreted (Fig. 11), 
which in older hairs may be shed. 

As the gametophyte becomes distinctly cordate the apical 
meristematic cell becomes replaced by an apical meristem of 
conical cells (Fig. 12). Formation of a midrib is initiated by two 
months’ old gametophytes and archegonia are produced con- 
tinuously thereafter. Superficial hairs resembling the marginal 
ones but sometimes two cells long (Figs. 13, 14) are also devel- 
oped sparsely over the midrib and wings. 

The mature gametophyte (Fig. 15) is cordate, broader than 
long (ca. 10 mm in diameter) with a deep apical notch usually 


AMERICAN FERN JOURNAL VOLUME 50, PLaTE 16 


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ATHYRIUM ESCULENTUM 197 


overlapped by the lateral lobes and with a prominent midrib 
bearing sex organs on the ventral surface. The wing cells are 
uniformly thin-walled (Fig. 16) and densely chlorophyllous. 
The sex organs are of the usual type in higher ferns. Antheridia 
are globular and generally sessile (Figs. 17, 18). Occasionally 
the opercular cell is divided into two, three (Pig. 18a), or four 
cells. In liberating the sperms the opercular cell is entirely 
th 


Wagner, 1952b). The simplest cotyledonary leaf is cuneate 
(Figs. 20, 21) with a short petiole and a single vein forking 
equally twice. Their bases generally form an angle of less than 
90° and the veinlets near the middle of the lamina run parallel 
to each other. The apex is usually truncate or shallowly notched. 

Generally the second leaf (in some cases the first leaf itself) 
marks the next stage in development. It has a broader lamina 
with a distinct notch at the apex and a wider angle at the base. 
The veins fork three times (Fig. 22) with the branches towards 
the middle of the lamina more pronounced and sometimes forking 
once again in such a way as to give an appearance of pinnate 
branching (Fig. 19-ii). In such cases usually one side of the 
leaf is larger than the other (the right hand half in Figs. 19-0 
and 22). 

The third stage is usually met with in the fourth or fifth leaf 


Figs. 1-18. Spore- and gametophyte-morphology of A. esculentum. Fig 
1. L. O. pattern of the spore exine; Fig. 2. Equatorial view of spore; Fig. 3 
Proximal polar view of spore showing laesura; Figs. 4, 5. Origin of the 
germinating filament; Figs. 6-10. Early stages in the development of the 

3 Ft 1 


Ss 
H 


MSs Nea 8 i 
mature prothallus showing meristem and adjoining tissue ; 8. 
Superficial hairs on mature prothallus; Fig. 15. Mature prothallus (diagram- 
matic) ; Fig. 16. One of the wing cells showing thickenings at corners; Fi 
17, 18. Stages in development of antheridium (dotted line represents surface 
pattern) ; Fig. 18a. Surface view of mature antheridium showing a divided 
cap cell. 


3 
Pa 


VoLUME 50, PLarre 17 


AMERICAN FERN JOURNAL 


ATHYRIUM ESCULENTUM 199 


or in weak individuals even later. The lamina broadens and the 
apex becomes elongated giving an oval shape to the leaf. The 
main vein entering the leaf base instead of dichotomising pro- 
ceeds towards the tip as a midrib and gives off lateral veins 
alternately. The lower lateral vein on each side dichotomises 
once or twice, the branches towards the middle being longer than 
the others (Figs. 19-1, 23). The transition to the midribbed 
stage is rather sudden and no intermediate stages have been ob- 
served. The leaf margin is wavy, the depressions corresponding 
with the spaces between vein tips. 

In the fifth or sixth leaf the lamina broadens considerably and 
becomes trilobed (Fig. 24), the midrib and its upper branches 
oceupying the middle lobe and the basal pair of lateral veins 
occupying the lateral lobes. The lateral veins develop in the 
same manner as the midrib. Just below the sinus on either side 
are formed the first areoles, by the basal adaxial tertiary veinlet 
of the lateral lobes joining with the basal secondary veins of the 
middle lobe or a branch of it. After joining, the fused vein pro- 
ceeds towards the base of the sinus. 

In later formed leaves the middle lobe becomes more pro- 
nounced and the basal secondaries of it begin forming the next 
pair of lobes with an areole at the base of each sinus (Fig. 25). 
The sinuses separating the first pair of lobes become deeper and 
almost reach the midrib making the leaf pinnatisect. Conse- 
quently the areole at the base of the sinus is not formed. 

Further expansion of the lamina is by a pronounced increase 
in length of the leaf and formation of successive lateral lobes on 
Figs. 19-39. Morphology of the juvenile leaves of A. esculentum. Fig. 
19. Gametophyte with attached sporeling (growing apex of the sporeling 
not shown; i, ii, éii-the first, second and third leaves); Figs. 20-26. Lea 
succession in the young sporophyte; Fig. 27, Portion of adult lamina show- 
ing venation pattern (mr- midrib, Iv- lateral vein); ig. 28. Portion of 
margin of first leaf showing hairs (t— elub-sha ed hair, ah— acicular hair) ; 
Fig. 29. Superficial hair on first leaf; Fig. 30. Mu ticellular hair on the 
fourth leaf; Figs. 31, 32. Superficial hairs on the sixth leaf; Figs. 33, 

4, 35. Hairs on the petiole of the seventh leaf; Fig. 36. Palea on the 
petiole of the same; Figs, 37, 38, 39, Hairs on the lamina of adult leaf. 


Fig. 28. 


q 


200 AMERICAN FERN JOURNAL 


either side of the midrib (Fig. 26). Though areoles are formed 
on both sides of the midrib connecting successive secondary 
veins, they do not form later as the lobes become separated. In- 
stead, areoles of the same pattern are formed on either side of 
the secondary veins in the lobes. The margins of the lobes remain 
wavy. The leaves pass on from the pinnatifid to the pinnate 
condition by gradual deepening and broadening of the base of 
the sinuses, until the lamina is reduced to inconspicuous wings 
on the midrib (now the rachis) and finally to a deep green line 
lodged in an inconspicuous groove on the sides . The leaf-lobes 
(now the pinnae) develop a narrow stalk-like base and the vena- 
tion becomes more complicated. 

Increase in the size of the leaf is more marked in the longi- 
tudinal plane until an oblong deeply pinnatifid lamina is ob- 
tained. The midrib becomes grooved on the upper surface. The 
lateral lobes elongate, become oblong with almost parallel sides 
and a tapering apex. The secondary lateral veins of the lobes 
produce alternating tertiary branches which run obliquely to the 
secondary veins. The first formed tertiaries from nearby sec- 
ondaries fuse to form a single vein which runs parallel to the 
secondaries for a short distance and ends blindly. As the leaf- 
lobe expands, more tertiaries are produced by each secondary 
vein and the lower ones fuse in pairs. The fusion vein from each 
basal pair of tertiaries while proceeding towards the margin fuses 
with successive tertiaries on either side, thus forming two regular 
rows of obliquely placed areoles between the nearby secondaries 
(Fig. 27). Tertiaries formed towards the tips of secondaries are 
free. The fusion veins above the last pair of areoles either end 
blindly below the marginal sinus of the lamina or in some cases 
fork just below the sinus, the branches running parallel to the 
sides of the sinus for some distance. 

Once-pinnate leaves characterize the young plants of A. escu- 
lentum for quite a long time, and adult plants may revert to this 
leaf form under adverse conditions of growth. The bipinnate 
leaves of the adult plants are formed by the pinnae of the 
juvenile leaves undergoing the same pattern of development as 


ATHYRIUM ESCULENTUM 201 


the main leaf itself. 

The cotyledonary leaf bears unicellular, thin-walled, acicular 
hairs (Fig. 28, ah) all over the lamina and petiole. Mixed with 
them, unicellular club-shaped hairs (Fig. 28, h; Fig. 29) re- 
sembling those on the gametophyte, but without the caps, occur 
sparsely. The fourth leaf bears also club-shaped, uniseriate, 
multicellular hairs (Fig. 30) toward the base of the blade and 
on the petiole. These hairs are much bigger than the unicellular 
hairs and become pale brown when fully developed. The multi- 
cellular hairs become more numerous in succeeding leaves. The 
cell at the apex of the hairs becomes more prominent being 
globular, much bigger than other cells, and sharply marked off 
from the main body (Figs. 31, 32, 83). 

The seventh or the eighth leaf is the first one to bear paleae. 
The multicellular, club-shaped hairs on these bear lateral uni- 
cellular glandular branches resembling gametophytic hairs (Figs. 
34, 35). Later, the cells near the basal region of the hair expand 
and divide longitudinally, initiating the formation of a flattened 
base. The apical region in all cases remains narrow, uniseriate, 
and elongate, terminating in a globular or ovoid cell with very 
dense contents. Repeated longitudinal divisions of the cells of 
the basal half result in an oval palea with an attenuated hair- 
like tip and bearing superficial and marginal club-shaped hairs 
(Fig. 36). The adult leaf bears uniseriate, multicellular, club- 
shaped hairs with a prominent globular terminal cell having 
dense dark contents (Figs. 37-39). 


COMPARISON 


Little is known regarding the gametophyte and much less 
about the young sporophyte of Athyriwm and related genera, 
so much so that it is well nigh impossible to make many com- 
parisons. Stokey (1951), Wagner (1952a) and others have 
shown the importance of characters of the gametophyte and the 
young sporophyte in assessing the phylogeny of the different 
genera of ferns. The author (Nayar, 1956) has shown the 
probable correlation between the gametophytie and sporophytie 


202 AMERICAN FERN JOURNAL 


hairs in some of the polypodiaceous ferns and has suggested 
that it may be true for other ferns also. The study of A. 
esculentum lends further support to this statement. The acicular 
hairs on the young sporophyte seem from present observations 
to be entirely new structures, the paleae and hairs being com- 
parable to gametophytie trichomes in their ontogeny and funda- 
mental morphology. 

Thickening of the walls at the corners of the cells of the mature 
gametophytes was reported in Athyrium filix-femina (L.) Roth 
by Stokey (1951). Athyrium esculentum differs from this in 
having uniformly thickened walls. Unicellular, club-shaped hairs 
of the A. esculentum type oceur in A. angustifoliwm (Michx.) 
Milde but are reported to be absent in A. filix-femina and A. 
alpestre Rylands (Stokey, 1951). Unicellular, club-shaped hairs 
with an apical cap, mixed with two- to three-celled, elongate, 
acicular hairs (comparable to the acicular hairs on young juve- 
nile leaves of A. esculentum) occur profusely on the prothallus 


in these genera also are comparable to those of A. esculentum. 

The first juvenile leaves in the related genera of ferns (Dryop- 
teris, Polystichum, ete.) are usually dichotomous in plan and 
four- to eight-lobed (Wagner, 1952a). The early juvenile leaves 
of A. esculentum, though dichotomous in plan, have a cuneate, 
almost entire lamina, resembling in some respects the first leaf 
of some species of Asplenium (Slosson, 1906). The early leaves 
of those Aspleniaceae that are known have a tendency toward a 
simple, single vein pattern in the simplest frond condition and 
generally a more or less obcordate shape and a dichotomously 
divided vein in the first several fronds (Wagner, 1952b). All of 
them possess the ability to produce dichotomous vein patterns up 
to at least 4 to 6 vein termini. Among the Aspidiaceae, Tectaria 
alone, as far as known, is comparable to A. esculentum in the form 
of the juvenile leaves. 

ACKNOWLEDGMENTS 

Thanks are due to Prof. K. N. Kaul, Director, National 
Botanic Gardens, Lucknow (India), for the keen interest he has 
in this work. 


ReEcENT FERN LITERATURE 203 


LITERATURE CITED 


ErprMan, G. 1952. Pollen Morphology and Plant Taxonomy. Waltham, 
Ma 


Be: 

Harris, W. F. 1955. A Manual of the Spores of New Zealand Pteridophyta. 
New Zealand Dept. Sci. Ind. Res. Bull. 116. 

KAcHRo0o0, 56. iainaniiptistes of Tectaria variolosa and T. fuscipes. 
Sci. ee Cult. » 22(2):. 103-105. 

—. 1957. Ga aE ie of phepiiales pees (Wall.) Ching and C, 
moulmeinensis (Bedd.) Copel. Sci. & C 

Kacuroo, P., and Nayar, B. K. 1953. Meee in Pteridaceae-I. Gameto- 


phytes of Some Species of Adiantum L. Phyt set 3: 240-248. 
Nayar, B. K. 1954. Studies in Polsaelegastt: Morphology of Pseudo- 
.) C. Chr. Ibid. 4: 379-390 


drynaria coronans (Wall 
—. 1956. Correlation between Sie gia and Sporophytie Trichomes in 
t 57. 


3 York. 
Stoxry, A. G. 1951. The Contribution by the Gametophyte to the Classifica- 
tion of the Homosporous Ferns. Phytomorphology, 1: 39-58. 
Waener, W. H., Jr. 1952a. The Genus Diellia. Univ. Calif. Publ. Bot. 


12 

—. 1952b. ues of Foliar Dichotomy in Living Ferns. Amer. Journ. Bot. 

39(8):. 578-599. 

Recent Fern Literature 

CenTRAL European Ferns.'—In a collection of natural his- 
tory books published by the Senckenberg Natural History So- 
ciety, Georg Eberle has just written a book of high quality on 
the ferns of central Europe that is likely to attract new friends 
to these plants. It contains a summary on ferns in general— 
their organography and life cycle, a well-documented exposition 
on hybridity in central European ferns, taking account of the 
work of Manton and D. E. Meyer, an account of apogamy and 
apospory, and finally a treatment of the various species and their 
hybrids in the territory studied. The illustrations consist of two 
drawings and 92 magnificent photographs taken by the author 
between 1926 and 1958, showing the plants in their natural 
habitats —A. Lawatrée. 


1 Eberle, Georg. Farne im Herzen Europas. pp. I-VIII, 1-116, ill. Ob- 
tainable from Verlag Dr. Waldemar Kramer, Frankfurt am Main, Ger- 
Many, 1959. Priee DM 8.50 


204 AMERICAN FERN JOURNAL 


Notes and News 


MEMBERS OF THE OPHIOGLOSSACEAE WANTED FOR TRANSPLANT 
EXpERIMENTS.—Samples of from six to twelve whole living plants 
of Ophioglossaceae with roots and a clod of soil attached are 
desired, these to be sent in a plastic bag to the undersigned. 
These specimens will be carefully transplanted into various 
BPDEE Tats habitats to test whether or not the ‘‘distinguishing 
characters’’ are genetically fixed or are merely environmental 
modifications. The following are especially desired : Botrychium 
multifidum (Gmel.) Rupr. ssp. multifidum (=ssp. typicum 
Clausen) ; B. multifidum ssp. californicum (Underw.) Clausen ; 
B. australe R. Brown; B. dissectum f. pda (Gilb. & 
Haberer) Weath.; B. schaffneri Underw B. japonicum 
(Prantl) Underw.; B. simplex var. De canis (Lasch) Milde ; 
B. simplex var. sombroaune (A. A. Eaton) Clausen; B. matri- 
cariifolium ssp. hesperiwm Maxon and Clausen; B. lanceolatuwm 
(Gmel.) Angstr. ssp. lanceolatum (=ssp. typicum Clausen) ; B. 
virginianum ssp. europaeum (Angstr.) Clausen; and Ophioglos- 
sum vulgatum var. pycnostichum Fernald.—W. H. WAGNER, JR., 
Department of Botany, University of Michigan, Ann Arbor, 
Michigan. 


American Fern Society 
Report of the President for 1959 

Moderate growth in the membership of the American Fern 
Society continued through 1959, with applications for member- 
ship slightly exceeding our losses through death of members, 
resignations, and lapsing of dues. The membership stood at 764 
on January 1, 1959, and 797 by the end of December. This in- 
crease is less than that of recent years, but it indicates that 
many of our members are still actively working in behalf of the 
Society and calling it to the notice of their friends. This type of 
growth is healthy and indicative of the esteem many hold for 
the ferns of this and other countries. 

Growth in the number of members in good standing is gratify- 


AMERICAN FERN SOCIETY 205 


ing, but in itself is not sufficient to insure long term benefits to 
the Society as a whole and to its individual members. Promo- 
tion of knowledge about ferns and expansion of awareness of 
their beauty and charm among green thumbers, gardeners, and 
householders does bring such benefits. That the members of the 
American Fern Society have fostered such studies and interest 
is shown by the continued flow of papers to the Editor of the 
American Fern Journal. These papers, a good sample of 
which have appeared in the Fern Journal’s last four numbers, 
have been varied in scope, subject matter, style, and emphasis. 
Some of our members have lamented the small number that deal 
with the practical problems they meet in growing ferns in their 
own gardens, greenhouses, or homes. The large proportion of 
space devoted to articles dealing with the systematic position 
of certain ferns, with necessary changes in scientific names, and 
other technical subjects, is owing to the fact that most of the 
people who submit articles to the Editor are professional 
botanists. A member chiefly interested in growing, or just en- 
joying, ferns rarely wants to devote the necessary time and 
effort to preparing a paper for publication. To make the situa- 
tion more difficult, the professional botanist all too often can 
not spare the energy required to grow many of his favorite ferns, 
so he seldom is in a position to write a gardening type of article. 

During the year strong regional sub-organizations have fune- 
tioned smoothly and effectively on both the Atlantic and the 
Pacific coasts. The ‘‘New England Section,’’ under the capable 
leadership of such staunch supporters as Dr. R. C. Benedict, Dr. 
Benjamin Allison, Mr. Boughton Cobb, Miss Clara Hires, and 
others living in and near this area, has sponsored informal meet- 
ings to discuss ferns and to share information, arranged visits to 
gardens, negotiated with the Director of the New York Botanical 
Garden looking toward a fuller utilization of ferns in that justly 
famous garden. Owing to the distances between the New Eng- 
land-New York area and the official Society field-trip on the 
north shore of Lake Superior, the New England group spon- 
sored an eastern fern foray during July. All of these activities 


206 AMERICAN FERN JOURNAL 


have added to the prestige of the American Fern Society, and I 
sincerely believe the Society as a whole appreciates such excel- 
lent work. 

Diagonally across the continent, members of the American 
Fern Society living in and near Los Angeles several years ago 
organized the Los Angeles Fern Society, open to anyone inter- 
ested in ferns. But the founders stipulated that officers of that 
organization also must hold membership in the American Fern 
Society. These fern enthusiasts in the southwest have held regu- 
lar monthly meetings for lectures on various aspects of fern 
taxonomy, ecology, structure, and culture, with vigorous discus- 
sion often following the presentation of papers. Their meetings 
have provided opportunities for the participating members to 
‘‘swap’’ plants and to compare notes on the distribution of varl- 
ous native ferns, sources of supplies, and names of reliable 
dealers. Mrs. Sylvia Leatherman, who has been a member of 
the American Fern Society for a number of years, served as 
President of the Los Angeles Fern Society during 1959, suc- 
ceeding Dr. W. C. Drummond, who was the first president of 
their organization. 

In order to allow members planning to attend the Ninth Inter- 
national Botanical Congress in Montreal to participate in a fern 
foray just prior to the International meetings, Dr. Olga Lakela, 
Emeritus Professor of Botany at the University of Minnesota, 
led a two-day foray from Duluth, Minnesota, along the north 
shore of Lake Superior on August 7th and 8th. Eighteen mem- 
bers and friends enjoyed two fine days under Dr. Lakela’s guid- 
ance. Several of those present went on to Montreal and took part 
in a fern excursion to the Rougemont area south of Montreal on 
August 21. This foray, although not officially sponsored by our 
Society, gave some of us a rare opportunity to become acquainted 
with pteridologists from England, Scandinavia, Italy, Belgium, 
France, India, China, and Africa. At least three of those on the 
Rougemont foray became members of the American Fern Society 
as a direct result of these contacts! 

Negotiations with the American Institute of Biological Sci- 


AMERICAN F'ERN SOCIETY 207 


ences, begun in 1958, continued into 1959 and resulted in our 
full affiliation with that organization. Henceforth, the American 
Fern Society will be able to hold its annual summer meeting in 
conjunction with those of the A.I.B.S. as an affiliate rather than 
a guest, and have the major portion of the local arrangements 
and provisions for the printing of the program handled by the 
staff of the A.I.B.S. The A.I.B.S. Bulletin, which will list the 
American Fern Society among the Affiliated Societies, reaches 
approximately 80,000 biologists in the United States and Canada 
five times each year. 

Rising costs for publishing the American Fern Journal have 
been a source of much concern to the Editor and to the Council. 
Several times during the past five years the printers have served 
notice that the rates for printing the Fern Journal would be 
advanced. Each time the advance has been accepted, although 
without enthusiasm. Finally, in 1959, another such notice re- 
sulted in wide inquiry among various publishing houses by Mr. 
Morton, with the result that arrangements have been made to 
have the Fern Journal printed, beginning with the January- 
March number of Volume 50, by the Monumental Printing Com- 
pany, of Baltimore, Maryland. The Council appreciates the 
many favors that the Business Press, Inc., extended to the So- 
ciety during the 35 years it printed the Fern Journal. We hope 
that the association with the Monumental Printing Company 
may be as long and as free of difficulties. 

It is a pleasure to commend the Council members and other 
representatives of the Fern Society for the loyal support each 
has accorded the President and the Society throughout 1959. 
When prompt action was required to carry through a project of 
importance to the Fern Society, each member of the Council has 
voted in accordance with his conviction on what was in the best 
interests of the Society. Our Representative to the Council of 
the American Association for the Advancement of Science, Dr. 
A. C. Smith, has attended each meeting, without cost to the Fern 
Society, and promptly furnished a resumé of the recommenda- 
tions made and actions taken by that body. Dr. Rolla Tryon, 


208 AMERICAN FERN JOURNAL 


representing us on the American Horticultural Council, called 
on the secretary of that organization to ascertain how our inter- 
ests fitted in with the aims of the Horticultural Council. It ap- 
pears that there are several spheres of interest held in common 
by our Society and the Horticulturalists. It is conceivable that 
their support in promoting protection and conservation of 
threatened fern localities might be of inestimable value. 

Mrs. Boydston has prepared a report covering the Spore Ex- 
change. I commend her for her faithful and efficient handling of 
that segment of the Society’s activities ! 

Dr. Herbert Wagner continues to serve as Librarian of the 
Fern Society’s collection of books and reprints on ferns, and to 
take care of the herbarium of dried and pressed fern specimens. 
(Permit me to call attention to the notice printed inside the front 
cover of the Fern JOURNAL, giving the terms on which loans and 
exchanges are conducted.) 

As in past years, our Editor-in-Chief, C. V. Morton, works 
harder and longer each year on the affairs of the American Fern 
Society than any other member of the Council. It is he who reads 
all papers and articles submitted for publication in the FERN 
Journal, carries on correspondence with authors and the print- 
ers, makes the necessary editorial corrections and provides direc- 
tions to the printers for the make-up of each issue of the Ameri- 
can Fern Journal. Nor is his task finished when the galleys 
have all been read and the final printing completed. He still has 
to dispatch dozens of extra copies and back numbers to indi- 
viduals and libraries requesting such material, a task that in- 
volves withdrawing the wanted items from the stock of reserve 
numbers stored where the temperatures soar in the summer time 
and approach or reach the freezing point in winter. So again, I 
am happy to extend to him the sincere thanks of the entire mem- 
bership of the Fern Society. 

Two of the Associate Editors, Dr. R. C. Benedict and Dr. A. C. 
Smith, have rere aided the Editor-in-Chief with care- 
ful reading of galley proofs and in checking manuscripts on 
which he wished their sari advice. Their help is appre- 
ciated fully. 


AMERICAN FERN SOCIETY 209 


Finally, to every member of the Fern Society, please let me 
express gratification that he or she has continued to support the 
Society morally and financially during the period of the indi- 
vidual’s association with our organization. Some of you have 
been members for over half a century, others for only a few 
months. The important point is that you have had, and continue 
to have, a genuine interest in ferns—in how to grow them, how 
to protect and conserve the wild ones still to be found in places 
of beauty and charm, and how to learn more about them through 
continued observation and study of this intriguing group of 
plants. It is my firm belief that each of you will provide Dr. 
Clair Brown, your new President, with the same steadfast sup- 
port you have accorded me. May the fortunes of the American 
Fern Society and of its individual members, collectively and in- 
dividually, continue to prosper. 

Respectfully submitted, 
. Wiaarns, President 
Report of the Treasurer for 1959 

The end of 1959 found our financial condition approximately 
the same as the preceding year. The balance on hand as of the 
first of the year was slightly less but still sufficient, so that we 
had no bank charges except for Unesco Coupons. Sale of back 
numbers continued to supply a substantial sum. The increase of 
postage rates, not anticipated when the 1959 budget was made 
out, caused some over-spending. No funds were withdrawn from 
the reserve accounts and the Society stands in good financial 
condition 

Subscription rates, both domestic and foreign, are scarcely 
sufficient to cover the cost of the Journal. Most of these sub- 
seribers require special attention and several letters for each 
transaction. I suggest that consideration be given to raising the 
subscription price to equal the membership fee. During the year 
we receive many checks for subscriptions from individuals which 
require special handling. The cost of back volumes could also 
be raised to the current annual value. These two raises would 
eliminate some of the current collection problems. 


210 AMERICAN FERN JOURNAL 


Receipts 
Amount Total 


Cash on-hand, January. 1, 1959. 2s et $1,401.12 
85 


2,808.99 
$4,210.11 
Disbursements 
A.F.J. Vol. 48, No. 4 612.78 
A.F.J. Vol. 49, No. 1 785.49 
A.F.J. Vol. 49, No. 2 2 366.11 
ABS. Vol. 49, No.3. 397.34 
Reprints 2. 357.31 
Envelopes and Mailing 106.70 
Spore Exchange 20.00 
American Horticultural Council 1959 dues — 20.00 
Dues to AIBS 100.00 
verpayment to Book Agent 2.35 
Bank charge on Unesco Coupons 51 
efund, Roy Bloemer—delay delivery back Nos. 9.45 
Expenses 

Secretary ii.3400 0 5.55 
BUSOP a 199.37 
Editor, 1958 and 1959 31.65 

3,014.61 

Cash in Southern Arizona Bank, January 1, 1960 —_...---.---- $1,195.50 


AMERICAN FERN SOCIETY 211 


STATEMENT DECEMBER 31, 1959 
Assets 


Cash in Southern: Arizona; Bank... 22 ee 
Cash in Green Point Savings Bank (Bissell Herbarium Fund) — 


Cash in Green Point Savings Bank (Life Membership Fund) - 801.11 
Cash in Green Point Savings Bank (Reserve Fund) — 1,910.04 
Cash in Green Point Savings Bank (Una Weatherby Fund) —. 3,125.75 
Inventory, American Fern Journal ——.--—------------------__--- 3,150.08 
American Fern Society Library —-—--------------__-__----- 396.00 
$11,276.21 
Liabilities 

dvance Dues Collected _......---------- Dee 382.75 

Accounts Payable (Vol. 49, No. 4) —--------—-—-----—--- 487.43 
870.18 

Fund Balances 

Bissell Herbarium Fund 697.73 
Life Membership Fund 801.11 
Reserve 1,910.04 
3,125.75 
3,871.40 


Genera 
$11,276.21 

Respectfully submitted, 
Water S. Paris, Treasurer 


Report of the Auditing Committee 


We hereby certify that we have seen the books and accounts of Dr. Walter 
American Fern Society, Inc., and have obtained 
t 


S. Phillips, Treasurer of the 
d as se 


onfirmation of the correctness of the Society’s balances on han 


fuk in detail in the accompanying report of the Treasur 
Av anes eiomnedied 


CHarLEs T. Mason, JR. 
Ricoarp H. HEVLY 


Report of the Secretary for 1959 
Your secretary has, he hopes, gotten through his first year at 
the post without seriously antagonizing anyone by his ineptness. 
At any rate, he has enjoyed the associations made and hopes that 
he is carrying out his duties satisfactorily. 


212 AMERICAN FrRN JOURNAL 


The Society membership has continued to increase to 797, as 
compared with 764 at the beginning of 1959. California con- 
tinues in the lead in number of members with 162. New York 
comes second with 90. Two possessions and 24 foreign countries 


It is with evant regret that I report the loss of several long- 
time members by death—Mr. Chauncey Jackson Newell (1902), 
Prof. Bremer W. Pond (1910), Mr. Peter Osterlund (1920), Mrs. 
Charles Tanger (1930), Mr. F. N. Irving (1940), and Mrs. F. G. 
Dunham (1941). 

Due to the International Botanical Congress held in Montreal 
in August, the Society did not hold a meeting last year. It is 
hoped that this year’s meeting in Stillwater, Oklahoma, will have 
a good representation of the membership and that a number of 
interesting and informative papers on ferns will be presented. 

Respectfully submitted, 
Donatp G. Hurrieston, Secretary 


Report of the Judge of Elections 


The results of the balloting for officers of the American Fern 
Society are as follows: 
For President 


Claire AD Brown Ls 331 

Ira Ty Wieeins 4 

Warren Ho Wagner = = Zz 
For Vice-President 

Marcel Raymond 335 

Rolla M. Tryon 1 

Frieda L. Wertman 1 

Hugh C. Cutler 1 
For Secretary 

Donald G. Filion mie ee ie ee Bay 
For Treasurer 

Walter S: Phillips 2 338 


I therefore declare the following candidates elected to the 
several offices: President, Clair <A. Brown; Vice-President, 


AMERICAN FERN SOCIETY zis 


Marcel Raymond; Secretary, Donald G. Huttleston; Treasurer, 
Walter S. Phillips. 
Respectfully submitted, 
Hueu C. Cuturr, Judge of Elections 


Report of the Curator and Librarian for 1959 
Compared to the year 1958, in matters dealing with the Her- 
barium and the Library of the Society, the business has been 
very slow. The bulk of consultation of the collections, and such 
few loans as were made, involved persons in the local area of 
Michigan. Mr. David B. Lellinger, a graduate student in the 
Department of Botany of the University of Michigan, has pro- 
vided much valuable assistance in routine work with the litera- 
ture and specimens. I should like, once again, to call the atten- 
tion of readers who have not yet utilized our collections to the 
description of the procedure given briefly in the bottom para- 
graph of the inner cover of the JOURNAL. 
Respectfully submitted, 
ARREN H. WaGNER, JR. 
Curator and Librarian 


Report of the Spore Exchange for 1959 


Contributions to the Spore Exchange during 1959 included 
five samples from Mr. L. J. Brass, who collected them on the 
Archbold Expedition to New Guinea, 22 from the University of 
California Botanical Garden, through Dr. Herbert Baker, who 
promises more, 38 from Mr. Jury, our member in New Zealand, 
15 from Mr. Aarestup (Denmark), 5 from Mr. Fisher (London), 
7 named and a number unnamed from Mrs. Faithful (New Zea- 
land), through Mrs. Eva Alexander, Birmingham, Alabama, 7 
from Japan contributed by Mrs. Barbara Joe Hoshizaki, 22 from 
Japan from Mr. Sekido (bearing names in Japanese but not 
English), and 164 from Fern Society members living in the 
United States. Among all these there were very few duplicates. 
Highty-one were fresh supplies of species already listed; the 
others were new to the list. 


214 AMERICAN FERN JOURNAL 


These 1959 contributions to the Exchange have been sent in 
by 43 kind people, at the cost, no doubt, of considerable time and 
trouble. To each one goes the gratitude of the officers and mem- 
bers of the Fern Society. The exchange, of course, could not 
function without these faithful, interested people 

Though there have been well over a hundred reees and letters 
written about the exchange, only 22 members actually requested 
spores in 1959, and a total of 267 packets was mailed to them 
Since the procedure for growing ferns from spores is easy (at 
least as this amateur does it anyway), it seems strange that more 
of our members do not take advantage of the Spore Exchange. 
Once in a while the results are disappointing, but far oftener 
are gratifying as well as enjoyable. 

A new list of spores now available is in preparations and will 
be sent to all those who have requested or contributed spores or 
otherwise shown an interest in the Exchange. It will, of course, 
be sent also to any other member requesting it. The new list will 
contain only those species sent in during 1958 and 1959. From 
the old list, 41 have had to be omitted because they were sent in 
earlier or because the supply is exhausted. Of these, the ones 
that we are most anxious to receive and list again are: Adiantum 
pedatum var. aleuticum, A. peruvianum, A. Wagneri, Asplenium 
fontanum, A. marinum, A. monanthes, A. resiliens, A. Tricho- 
manes (crested), Cheilanthes alabamensis, Dryopteris viri- 
descens, Lygodium japonicum, Polystichum acrostichoides (bi- 
pinnate form), and P. tripteron. There have been requests for 
these. If you can supply these, please do so, or make a note to 
make collections when the 1960 spores ripen. Others often re- 
quested and nearly always in short supply are the Appalachian 
spleenworts, especially Bradleyi, montanum, pinnatifidum, and 
Ruta-muraria. 

It would be fine if conservatories and botanic gardens general- 
ly could be interested in growing ferns, as are Longwood Gar- 
dens and the University of California Botanical Garden. Con- 
ceivably, an exchange, perhaps through the Fern Society, be- 
tween such places with large areas under glass could be most 


AMERICAN FERN SOCIETY 215 


interesting and worthwhile. We need finer and more comprehen- 
sive collections about the country for those of us who love to see 
and study the various warm-climate species but have no chance 
to grow them and no opportunity to travel and see them in their 
native haunts. 

With sincere thanks to all for the past and future interest, we 
enter 1960 with high hopes for a bigger and better exchange pro- 
gram. 

Respectfully submitted, 
KATHRYN E. Boypston 
Route 3, Niles, Michigan 


Report on the 1959 Minnesota Field-trip 


The 1959 field-trip of the American Fern Society took place 
on the north shore of Lake Superior, Minnesota, on August 8 
and 9. From Duluth to Grand Portage, a distance of some 160 
miles, scheduled stops were made. 

The attending members, with reservations for the night of 
August 7 in the Residence Halls of the Duluth University, were 
Dr. and Mrs. Ira Wiggins, Dr. and Mrs. Ralph H. Benedict, 
Mrs. George Gardner and a companion, Mrs. Frieda Wertman, 
Dr. Herbert Clarke, Dr. Robert Lommasson with four children, 
and Dr. J. W. Moore. Local personnel assisting with arrange- 
ments were Miss Mary I. Elwell, Miss Helen Heino, Dr. Paul 
Monson, and Dr. J. B. Carlson. 

Saturday, the following morning, the enthusiastic group left 
the campus in six cars, heading for the North Shore drive. At 
Knife River, on Lake Superior, a stop was made on the property 
of Miss Olive Prine and Miss Jessie Wells to observe the rela- 
tively undisturbed crevice flora of the shore outerops, and 
especially Botrychium multifidum and B. simplex on the shady 
upper terrace. 

In the scenic gorge of Gooseberry River State Park, below the 
falls, on high cliffs of diabase, a search was made for crevice 
ferns, species of Woodsia, Dryopteris, and Cryptogramma Stel- 


216 AMERICAN FERN JOURNAL 


leri. Cystopteris bulbifera filled moist hollows of overhanging 
ledges near the river level. 

The Palisade Head, one of the highest bluffs of a commanding 
view of the lake, is accessible by car. Field lunches with hot 

e were enjoyed there. Facing the cooling breezes under 
lowering clouds, those luscious tomatoes of the Benedicts, delecta- 
ble in their colorful ripeness, seemed a bit exotic. The forested 
slopes and the sliding talus of porphyritic felsite, were explored 
for species of lycopods. Opportunities for collecting in general 
were not overlooked. Of special interest were the less common 
species, e.g. Lycopodium Selago and var. patens. 

The remaining afternoon was absorbed in exploring Temper- 
ance River Gorge for woodland ferns. Among others, a fine 
colony of Dryopteris spinulosa var. americana was encountered. 
Due to impending rain and the late afternoon hours, it seemed 
advisable to proceed, without further stops, to Naniboujou Lodge 
where reservations for overnight lodging and meals had been 
arranged. 

Fair skies and sunshine were in store for the Sunday morning 
trip to Grand Portage. One of the mountainous bluffs of slate, 
overlooking the historic portage trail and the lake, with shady, 
moist, ledges, and broad talus slopes, afforded opportunity for 
observing Woodsia scopulina, W. Cathcartiana, and the rarest 
of all, spotted by Dr. Benedict, W. glabella. There ended the 
foray with a list of 28 species of ferns, and eight species of fern- 
allies 


The writer is indebted to Dr. J. W. Moore for assisting in the 
identification of species in the field, and to each and all for 
gracious cooperation and enthusiasm in sharing the fern trails 
of Lake Superior coast—Ona@a LAKELA. 


Report on the Rougemont Field-trip 


On Sunday, August 23, 1959, those who had registered for 
Field Trip 4 (Pteridology) of the Ninth International Botanical 
Congress climbed into a bus before 8 a.m. and waited expectantly 
to drive to Rougemont, Quebec, southeast of Montreal, Canada. 


AMERICAN FERN SOCIETY 217 


The trip was oversubscribed and two enthusiasts were forced to 
follow the bus in a private car. Some of the participants from 
abroad included Drs. ‘iit Manton, England; W. Zimmermann, 
Germany; Pierre Mar France; R. E. G. Pichi-Sermolli, 
Italy; and EK. A. Se ae Saath Africa. Those from the Ameri- 
can Fern Society were Drs. Ira L. Wiggins, Alice and Rolla 
Tryon, Clair Brown, T. M. C. Taylor, Donald Huttleston, Wil- 
liam Cody; Mrs. Lenette Aveinass, Mrs. Fern Crane, Miss Eva 
Sobol, and Miss Muriel Hegwood, among others. 

The trip was ably conducted by Lionel Cing-Mars who kindly 
supplied everyone with seventeen mimeographed sheets listing 
the vascular plants that he had collected at Rougemont from 
1948 to 1959, two pages of descriptions of the various habitats 
and important species that we would encounter, and a one-page 
list of some interesting plants that have not been collected at 
Rougemont as yet, but that might logically occur there. 

The first stop was a pine-wood in sand. Here we collected 
Dryopteris marginalis, D. intermedia, Athyrium Filiz-femina, 
Botrychium virginianum and in a moist location, Dryopteris 
Thelypteris, D. noveboracensis, and Osmunda Claytoniana. 
Lycopodium complanatum var. flabelliforme, L. lucidulum, and 
L. obscurum were collected in these woods also, before we took a 
short walk down a hill to a dark, wet deciduous woods where we 
found Dryopteris spinulosa, D. cristata, D, Phegopteris, D. dis- 
juncta, Osmunda cinnamomea, and Athyrium thelypterioides. 

At the second stop we looked at a large clump of Matteuccia 
beside the road and then proceeded through a pleasant maple 
sugar bush up the north side of the mountain. There were some 
fine clumps of Polystichum acrostichoides, Adiantum pedatum, 
and a fine stand of Dryopteris heaagonoptera. It seemed a 
strange twist that this last fern should be growing on the north 
side of the mountain in perhaps its most northerly location in 
Canada, whereas D. Phegopteris was found on the southern 
slope. 

After an enjoyable picnic lunch beside a duck pond near a 
local Quebec inn, we returned to Rougemont and one party of 


218 AMERICAN FERN JOURNAL 


active oldsters climbed a steep limestone cliff while the other 
party of young laggards went up a less precipitous trail through 
rich deciduous woods alongside a small stream. Those who took 
the former course brought back Woodsia ilvensis, Cystopteris 
fragilis, and Asplenium Trichomanes. The others saw some fine 
specimens of Dryopteris Goldiana, and strong clumps of 
Athyrium pycnocarpon. Polypodium virginianum was collected 
along rocky ledges above the stream. One fern that eluded the 
collectors here was Cryptogramma Stelleri; presumably because 
of the drought of August it could not be found. Dennstaedtia 
punctilobula and Onoclea sensibilis were seen along a roadside 
near an orchard. In all 33 species were seen and collected and 
another twelve have been collected by Mr. Cinq-Mars in the last 
ten years. Unfortunately, we were unable to add any new 
species to his list of pteridophytes. 

One further stop was made when our leader graciously offered 
us some apple juice and cider at his field-station in the center of 
the apple growing area. After this pleasant interlude we re- 
turned to Montreal with memories of a beautiful day and a fine 
field-trip—D. M. Brirron. 


Report of the 1959 Vermont Field-trip 


Under the leadership of Dr. Benjamin R. Allison, a New 
England field-trip of the American Fern Society was planned 
this summer to take place in Vermont, July 17 to 20. Walling- 
ford Inn (Wallingford, Vermont) and St. Johnsbury House (St. 
Johnsbury, Vermont) were headquarters for the field-trip. 

With Mr. Henry Potter of Clarendon as guide, 20 members 
and friends gathered at the Wallingford Inn for the Friday and 
Saturday trips which included an exploration of Proctorsville 
Gulf where 24 species of ferns were seen, 23 of which had been 
seen by the group two years ago and in addition, as a climax to 
the trip, several specimens of Botrychium matricariifolium in 
fruit. 

After lunch, being assured by our guide that we were safe 
from the onslaught of a railroad train, we explored about half a 


AMERICAN FERN SOCIETY 219 


mile down the tracks in a cut blasted out of solid gneiss by the 
Rutland Railroad in 1848. During the interval of 100 years 
many ferns had taken over on the banks and the rocks in the cut. 
The most exciting find was a good stand of Cryptogramma 
Stelleri clinging to the face of the solid rock in the cut. In addi- 
tion, we saw quantities of Cystopteris fragilis, Phegopteris con- 
nectilis and some Gymnocarpium Dryopteris. 

Friday evening Ralph C. Benedict joined us and during the 
evening gave an informal talk on fern nomenclature and num- 
bers of fern species. Florida claims the largest number with 150 
species ; Texas comes next with 120; and Vermont and Michigan 
tie for third place with about 80 species each. Ralph’s guess is 
that there may be 400 species in the United States. (These fig- 
ures include the fern-allies.) 

On Saturday morning we climbed to the ice-beds in White 
Rock Park to see quantities of Dryopteris dilatata growing in 


the open caves on a very hot July day. Also in this park we 
added several species of Lycopodium to our list as well as 
Dryopteris spinulosa and the hybrid Dryopteris fructuosa. 

n the afternoon on a rugged hillside in Dorset we added 
seven other species of ferns to our list—Pellaca atropurpurea, 
Asplenium Ruta-muraria (very erisp because of the dry weath- 
er), A. Trichomanes, A. platyneuron, Camptosorus rhizophyllus, 
Woodsia obtusa, and a Botrychium (possibly obliquum). 

With a visit to Mr. Potter’s meadow to see Ophioglossum 
vulgatum in fruit, the Wallingford part of the Field Trip ended, 
and on Sunday 12 members of the group went to St. Johnsbury, 
so as to be ready to explore the Lake Willoughby area on 
Monday. 

We were looking forward eagerly to the exploration of the 
Lake Willoughby area, having heard that this was the richest 
fern area in New England, but we suffered some disappointment 
in the number of species that we found. In explanation of this 
Mr. Potter reminded us that the area had been over botanized 


220 AMERICAN FERN JOURNAL 


some years ago. However, the base of the cliff on Mt. Pisgah, on 
the east side of Lake Willoughby, yielded two rare species— 
Woodsia glabella and Pellaea glabella. 

Two score or more years ago our guide, Mr. Henry Potter, 
was taken across Lake Willoughby by E. J. Winslow (former 
president and one of the founders of the amt JOURNAL) to see 
Asplenium viride and Dryopteris fragrans on the West Cliff. 
We hoped to see these on the East Cliff but were not able to find 
them, and it was not possible for the group to explore the West 


iff, 

All those present felt that the trip was a great success due to 
the planning of Dr. Allison and to the expert guiding of Mr. 
Potter, who knew where the ferns grew. 

In all we identified 41 species of ferns, six species of Lycopo- 
diwm, and three species of Equisctum. The following is a list of 
the ferns seen, the names taken from a check-list prepared by 
Ralph C. Biaudicd. 


Dennstaedtia punctilobula Thelypteris noveboracensis 
Pteridium aquilinum egopteris connectilis 
Adiantum pedatum Gymnocarpium Dryopteris 
Cryptogramma Stelleri Dryopteris margin 
Pellaea atropurpurea D. cristata 
P. glabella D. intermedia 
Polypodium vulgare D. spinulosa 
Masses Struthiopteris D, dilatata 
Onoclea sensibilis D. Goldiana 
Asplenium Ruta-muraria . fructuosa 
A, A ag Polystichum acrostichoides 
A. platyneur ze, unit 
Ca cnpieabnds iniesbiiias Osmunda regalis 
Athyrium Filia-femina O. cinnamomea 
+ pycnocarpon O. Claytoniana 
A, thelypterioides Ophioglossum vulgatum 
Woodsia glabella Botrychium matricariifolium 
. obt virginianum 
Cystopteris bulbifera B. multifidum (?%) 
C. fragilis B, obliquum (?) 


Thelypteris palustris 


AMERICAN FERN SOCIETY 221 


The following members and friends were present at one or all 
of the sessions: Dr. and Mrs. Benjamin R. Allison, Mrs. Louise 
Adams, Dr. Ralph C. Benedict, Miss Alice Bristow, Miss Helen 
Bristow, Mrs. L. L. Delafield, Mr. and Mrs. Burton Dezendorf, 
Mrs. Dolt, Mr. and Mrs. Murray Evans, Mrs. H. KE. Kineaid, Mr. 
Henry Potter, Miss Elsa Potter, Miss Anna E. Seudder, Dr. and 
Mrs. A. V. Smith, Miss Eva Sobol, and Miss Margaret Timp- 
son olgaied E. ScuppE 


Obituaries 


BREMER WHIDDEN Ponp, June 23, 1884—-September 2, 1959.— 
Bremer Whidden Pond was born in Boston, the son of Charles 
Choate Pond and Annie Louise (Whidden) Pond. He received 
his high school education at Winchester, Massachusetts, and at- 
tended Dartmouth College, receiving his bachelor’s degree in 
1906. After spending a year studying in Germany, he began 
graduate work at Harvard, receiving in 1911 the degree Master 
in Landscape Architecture. For the next three years he served 
as secretary to Frederick Law Olmsted, thus rounding out the 
preparation for his life’s work. 

n 1914, Bremer Pond began his teaching career at Harvard. 
Except for a brief period during World War I when he served as 
Captain, then as Major, in the Construction Division of the 
Q. M. C. in Washington, he taught landscape architecture at 
Harvard until his retirement in 1950. He served as Chairman of 
the Department from 1928 on. His contribution to the profession 
Was a very valuable one. 

In addition to his teaching he maintained a private office which 
was very active until about 1940. There are many significant 
examples of his work. For fourteen years he served his pro- 
fessional society, the American Society of Landscape Architects, 
well as Seeretary and Trustee. In addition, he performed in- 
valuable service in many committees and helped solve many 
perplexing problems to insure a healthy growth of his profession. 

Professor Pond was a member of the American Fern Society 
for almost fifty years, having joined in 1910, at about the time 


222 AMERICAN FERN JOURNAL 


when the AmERICAN FERN JOURNAL was started. Aithough he 
was never a pou student of ferns, he maintained a lifetime 
interest in t 

Although Ener Pond gave untiringly of himself to students 
and friends, his interest in nature was equal to his interest in 

ankind. In his earlier years there were many trips to Europe 
and in his later years much time was spent in the woods—tramp- 
ing through the wild and unspoiled country at ‘‘Lost Horizon,”’ 
his summer home in northern New Hampshire. Fishing was a 
major recreation and the artistry of his fly tieing has been the 
envy of many. His interests in life were wide and he was a most 
stimulating, friendly, and understanding companion who will 
be missed greatly by many of us for a long, long time.—W ALTER 
L. CHAMBERS. 


Mrs. Cuarues Y. TANGER.—Members of the American Fern 
Society will regret to learn of the death on September 28, 1959, 
of Mrs. Charles Y. Tanger, of Lancaster, Pa. She joined the 
Society in 1930, and since that time welcomed many of its mem- 
bers to study the rarer species of that region, and to enjoy her 
small but fascinating back-yard fern garden—Epcar_ T. 
‘WHERRY, 


CHaUNCcEY Jackson NEWELL.—We regret to note the passing 
of one of our oldest and most faithful members, Mr. Newell, a 
member for 57 years. An account of Mr. Newell and his interest 
in ferns was published in the JourNAL some time ago under the 
heading ‘‘Undistinguished’’ Fern Lover.’’1—C. V. Morton. 


FRANK N. Irvina.—Washington has lost one of its most avid 
fern lovers with the passing of Mr. Frank N. Ir ing. He was 
primarily a collector, and took great pride in the appearance of 
his specimens, justifiably so, for they are very likely the hand- 
somest specimens ever prepared. Only material in absolutely 
top condition was collected, and each leaflet was carefully and 


1 This JouRNAL 46: 169, 170. 1956. 


AMERICAN FERN SOCIETY 223 


raat pressed. His herbarium has been presented to the 

_ S. National Herbarium, through the kind offices of Mrs. 
i. Skougaard. It is being especially prepared for preser- 
vation as a memorial to Mr. Irving—C. V. Morton. 


New MEMBERS 


Mrs. Harry B. Allen, 334 Golden Gate Avenue, Belvedere, California 

Mr. William B. Armitage, 1159 Beach Street, Vancouver 5, British Colum- 
bia, Canada 

Mr. oe Breier, 150-06 Goethals Avenue, Jamaica te, New York 

Mr. Robert G. Bruce, P. O. Box, Jamestown, Virgin 

Mrs. Neil Chrisman, 63 West Ross St., Wilkes-Ba as Ma 
J 


“Ch a jana 
Mr. Pierre C. Fischer, Route 7, Box 147A, Tucson, Arizona 
Mr. Neal Edwards Vietabee, Jr., P. O. Box 1215, Williamsburg, Va. 
Mrs. Joseph Germanio, Box 87, Tuckahoe, New Jersey 
Mrs. A. H. Hosier, R.R. 2, Waverly, Tennesse 
Mr. A. Clive Jermy, British Museum coe History), 
London §.W. 7, England 
Mrs. ae L. Jones, pee Avenue, New Goss Manor, Dallas, Penna. 


Cromwell Road, 


Mr. J. Geoffrey Kramer, Egy 

Mr. lis oa Narita-shi, Chiba-ken, a 

Mrs. C. Ervin Ross, Merry Gardens, oe et Mai 

Mr. Normand J. Thompson, 514 Nort bh Shey TAB Ne Calif. 


bo 
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CHANGES OF ADDRESS 


Dr. Ralph D. Ade, 1535 10th Avenue, N.E., Rochester, Minneso 
rs. Geoffroy Atkinson, 415 South Pleasant Street, ereaited sfiua husalia 
Mr. Howard J. Boyd, 6624 8. E. 38th St., Po rtland, 
Dr. Daniel F. Burton, 512 Hickory St., Mankato, Minn. 
Mrs. canny Burton, 512 Hickory St., Mankato, Minn 
Mr. Orville W. Crowder, Box 7149, Baltimore 18, itaey yland 
« Mr. “hae K. Eastwood, % W. S. Phillips, Botany Department, University 
of Arizona, Tucson, Arizor 
Miss .. D. Feigley, 550 Chectntit Street, Winnetka, Illinois 
Mrs. Spencer ©. Flo, 772 Douatey Club Road, Green ome Mass 
Mr. Neal - Gilbert, 141 Rounds, Buffalo 15, } New 


224 AMERICAN FERN JOURNAL 


Mrs. Winifred Harley, Merry Point, R. R. 1, Lancaster, Va. 

Dr. Richard L. Hauke, ee ie oe of Botany, University of Rhode Island, 
Kingston, Rhode Isl 

Miss Muriel P. Hegw vr, 402 Randolph St., Radford, Va. 

Mr. A Ist. Street, New York 21, New York 


y 
5 
e 
be 
w 
oO 
a é 


Mrs. 

Mr. Albert E. Lownes, 16 Barberry Hill, ee 6, as Island 

Mrs. Arthur Luhr, Box 371, Murphys, Calif 

Mr. Mulford Martin, 622 West 137th Bice i llliuaitt 24, New York 31, 
New York 

Mr. Ray McAdams, 2001C Northwood aoe. North Campus, Univer- 
sity of Michigan, Ann Arbor, Michiga 


Mr. Robert McLean, P.O. B x 7745, Sede 1, Penr 
Mr. John T. Mickel, papain of Botany, University ‘a Michigan, Ann 
Arbor, Michigan 


Mr. Robert Millspaugh, eas Rock Road, Box 272, Roslyn, New York 
Prof. Dwight M. Moore, 506 Vandeventer Ave., Fayetteville, Ark 
Miss Jewel Moore, Rite 5, Box 357, Hot Springs ee 


Mrs. Mamie M. Peterson, 24461 Mission Bouleva rd, Hayward, California 


Mrs. I. M. Renton, Route 1, Box 111, Snoqualmie, Washingto 
Mr. Ray A. pols: 18328 East Foothill Bae PLO, ne 276, Azusa, 
Califor: 


Mr. Mastin ‘dassdalies P. O. Box 805, Los Gatos, California 

Mr. Milton E. Scherer, 830 North Tech Road, Sault Ste. fog vida © 
Mr. poate Simon, Bluemont Nurseries, Box 219, Monkton, Mar 

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Dr. pie a Steyermark, Apartado 2156, Caracas, Venezuel 
Dr. John H. omas, Dudley Herbarium, Stanford University, Stanford, 
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Dr. Clara G. Weishaupt, Botany Department, Ohio State University, 1735 
Neil age Columbus 10, Ohio 

Mrs. Frieda 5240 France Avenue, ear pia 10, Minnesota 

Dr. Eula Wied. 3033 Fondren Drive, Dallas 5, Texa 


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CAMBOSCO SCIENTIFIC COMPANY 
37 ET e BRIGHTON STATION 
BOSTON, MASS. U.S.A 


— 
Vou. 50 JULY-SEPTEMBER, 1960 No. 3 


American Fern Journal 


A QUARTERLY DEVOTED TO FERNS 


Published by th 
AMERICAN FERN SOCIETY 
EpITors 
Cc. V. MORTON 
R. C. BENEDICT IRA L. WIGGINS 
A. 0. SMITH 
CONTENTS 
How I Became Interested in Ferns Epear T. WHERRY 225 
Ferns in Oubangi, Congo._________-----@. J. DE JONCHEERE 228 « 
Ferns Cultivated in California: Sadleria___.____.__-BarBara Jor 232 


Ferns in Cultivation, III]. Growing Fern Balls 
Syiv1a LEATHERMAN 235 


A New Native Hardy Plant Area at the New York a #78 Garden 
©. BENEDICT 236 


‘ “To New c coeraie Stations for ———— complanatum 
Mehelliforme 27 _Tuomas A. Hurto 240 - 


Vein Patterns in Microsorium seandens and Its Allies 
Mary T. TINDALE 241 + 


A New Station for Trichomanes Petersii in Georgia 
Henry Booxout 246 ° 


Recent Fern Literature 47 
~~ — — er Fee meer Pere oS. 253 
an Bee 


American Fern Society_—_§—§$§$_____— 


The American Fern Society 


Council for 1960 
OFFICERS FOR THE YEAR 

Cua A. Brown, — of Botany, Louisiana State —— Baton 
Rouge, Louisia: esident 
eet May YMOND, Monten Botanical Garden, 4101 East Sherbronke Street, 
eatredd, Cana sident 

DonaLpD omd ang Longwood Gardens, Kennett Square, Phe 
Secretary 
—— 8. Pumuies, Department of Botany, University of —— Tucson, 
reasurer 

ON. ig bsnl Smithsonian Institution, Washington 25, D. C. 


Editor-in-Chief 
OFFICIAL ORGAN 
American Fern Journal 
EDITORS 

OY; ee ee by ahr Washington 25, D. C. 
R. C. BENEDICT 5 Hall Street, Brooklyn 5, N. Y. 
Ira L. WieeIns... _ Herbarium, Ste cob Seen, Stanford, Calif. 
A. C, Smira__ Smithsonian Institution, Washington 25, D. C. 


An pre exagp: ) postr d ae to the general study of ferns, published 
quarterly by the American Fern Society, 3110 Elm Ave., Baltimore 11, Md. 
Secsent-aises ¥ ostage ees y at Baltimore, Md. 

Matter tet publication should be addressed to C. V. Morton, Smithsonian 
Institution, Wa: 

Sabaniotion $2.50, exclusive of agency handling fees; sent free to mem- 
bers of the American Fern Society (annual aoe, $2.50; sustaining mem- 
bership, $5.00; ne m Bed ily 950, 00). Extracted pea om if ekaek in 
advanee, will be furnished qners at cost. They should be ordered when 


urned, 
Back volumes $2.00 each; single eng numbers 50 cents each; Cumulative 


Index ~ vols. 1-25, 25 cents, Ten per cent discount on orders of six volumes 
or mo: 

Chanaee of address should be sent to the Secretary: Dr. Donald Huttle- 
ston, Longwood Gardens, Kenne are, ogni eae Members and sub- 


seribers should allow two months for chan nges to take effect. 

Applications for membership, subscriptions, orders for face — and 
other business communications should be addressed to the Treasurer: Dr. 
bbl 8S. Phillips, Department of Botany, University rs ‘<yaaiai, Tucson, 


LIBRARIAN AND CURATOR OF THE HERBARIUM 
Dr. H. W. Wagner, University of Michigan, Ann Arbor, Mich. 


- : loan ed 

and herbarium. Members may — books and specimens at any time, 

the borrower paying all postal or ee The pages of the Journal 

on are open imhek who wiek to to arrange es; a membership list 
specimens 


is pu ublished peg interv, Mees to e 
p Regge yng assist those interested in obtaining 


American Fern Journal 


Vou. 50 JULY-SEPTEMBER, 1960 No. 3 


How I Became Interested in Ferns 
Epa@ar T. WHERRY 


For some years my professional activities were in the fields of 
chemistry and geology, and my acquaintance with ferns was zero. 
Then in 1913 I was appointed Assistant Curator of Mineralogy 
in the U. S. National Museum. Having built a house on a large 
lot the next year, I decided to develop a wild flower garden there, 
and I made the acquaintance of amateur botanists who guided 
me to some of the notable plant localities around Washington. 
In a ravine along the Potomac I was shown a colony of Walking 
Fern growing on a gneissoid rock, although according to ‘‘the 
books’’ it was supposed to grow on limestone. Having in my 
laboratory work just become familiar with the then new con- 
ception of hydrogen-ion concentration, I wondered if this phe- 
nomenon might have a bearing on the habitats of such plants. So 
I started looking for Walking Fern on different rock forma- 
tions, and found it on many kinds in the states within easy 
reach—Maryland, Pennsylvania, Virginia and West Virginia. 
Tests of the soil at the roots showed that on the average its re- 
action was indeed essentially neutral (as limestone is), and a 
paper on this subject was published in the Journal of the Wash- 
ington Academy of Sciences in 19 

Then it seemed of interest to study other ferns in like manner, 
and in order to learn the various kinds I bought a book called 
‘“Who’s Who Among the Ferns,’’ by W. I. Beecroft, of Boston. 
Whether this author ever did any scientific work on ferns I do 
not know, but at any rate his book—unlike too many more recent 
ones—was both simple and accurate, and made the recognition of 
Species an easy matter. In doubtful cases I took specimens to a 


nein aiglace es 


Volume 50, No. 2, of the JouRNAL, pp. 161-224, was issued June 28, 1960. 


AMERICAN FERN JOURNAL VOLUME 50, PLATE 18 


Epear T. WHERRY, AGE 70. Courtesy or ACADEMY OF NATURAL SCIENCES, 


PHILADELPHIA 


How I Became INTERESTED IN FERNS 297 


hew acquaintance, Dr. William R. Maxon, then Associate Cura- 
tor of the U. S. National Herbarium, who generously set me 
straight. He invited me to join the gale Fern Society and 
to contribute articles to its journal, which I began to do in 
1920. Travelling farther and farther afield, I gradually got to 
know in life most of the pteridophytes of the United States. 

My writings on these plants having made me well known, the 
members of the American Fern Society elected me president of 
the Society for 1934. The finances of the Society were in bad 
shape, the Treasurer having used its reserve funds in business 
ventures which failed, and a committee was appointed to 
straighten things out. Its members—R. A. Ware, C. A. Weath- 
erby, and myself—worked out plans for handling the Society’s 
funds which have proved successful; we also had the Society 
incorporated. After being in office for five years I withdrew my 
name from renomination. 

A cumulative index of the 20 volumes of the earlier Fern 
Bulletin had been published and proved useful, but there had 
been none of the American Fern Journal. As a result of the 
severe business depression, it was possible to obtain workers 
whose wages were paid by the government, and I accordingly 
employed a young lady to compile a 25-year index. The bill for 
printing this called for most of the Society’s funds, and I fear 
gave the Society’s then President, Dr. R. T. Clausen, and 
Treasurer, Dr. H. K. Svenson, considerable worry. However, the 
Society was soon in favorable financial condition again. 

An inaccurate popular fern book having appeared in 1936, 
I decided to prepare a more realistic one based on my own ob- 
servations in the field, herbarium and library. To illustrate this 
I was fortunate to obtain the services of Miss Olive Stoner and 
Mr. Cyrus Feldman, each of whom drew about 50 species. The 
book was published in 1937 under the title ‘‘Guide to Eastern 
Ferns.’’ The first edition of 2000 copies was sold out in four 
years, and a second was then issued, containing some improved 
drawings, text changes, and a diagnostic key. Besides the 2000 
printed copies of this, I had a like number copied by a photo- 


228 AMERICAN FERN JOURNAL 


graphic process and published by the University of Pennsylvania 
Press. The bulk of these issues have now been sold, but a few 
remaining copies have been deposited at the Academy of Natural 
Sciences of Philadelphia, which sells them at $2.00, plus 10c¢ 
sales tax. It has been my pleasure to donate annually the small 
““profit’’ received from this sale to the American Fern Society. 

BoranicaL LABORATORIES, UNIVERSITY OF PENNSYLVANIA, 
PHILADELPHIA, PENNA. 


Ferns in Oubangi, Congo 
G. J. DE JONCHEERE 


It was my good fortune to visit recently a part of the Belgian 
Congo that is hardly ever seen by tourists, especially those with 
an interest in ferns. I am speaking of the Gemena district, in the 
Province of Oubangi, in the northern part of Belgian Congo, 
east of the market town of Libenge, which is situated along the 
Oubangi River. This large stream is a tributary of the Congo 
and forms the border between Belgian Congo and French Equa- 
torial Africa. 

Thanks to the good offices of my host, Mr. Hans Kooiman, who 
is an agricultural advisor for several plantations there, I had 
the opportunity of making some extensive excursions in the 
neighbourhood of the plantations Bogbulu and Bala Bala. I 
made a collection of about 150 numbers of ferns, comprising 
about 40 species. The determination of the ferns was made easy 
by Madame Tardieu-Blot’s beautiful publication ‘‘Les Pterido- 
phytes de 1’Afrique intertropicale francaise,’’ in which the 
numerous plates are an example of good botanical illustration. I 
should like to thank Madame Tardieu also for her personal as- 
sistance in a few difficult identifications. Messrs. Adams and 
Alston’s ‘‘List of the Gold Coast Pteridophyta’’ was useful also. 

As far as the landscape around Gemena is concerned a certain 
monotony can not be denied. Mile after mile one travels through 
a slowly undulating peneplain, cut through by brooks and rivers 
and covered by grass fields and jungles. The deforested areas no 


FERNS IN OUBANGI, ConGo 229 


doubt owe their origin to the intensive and repeated burnings by 
the primitive Negro tribes that populate the country. The transi- 
tion from grass field to forest is immediate and with hardly any 
fringe area. Although the rainfall is certainly adequate, it 
strikes the observer that the rain-forest is not so lush and humid 
as the primeval forest in Java and Malaya, well-known to the 
writer. A certain scarcity of epiphytes and especially the infre- 
quency of Hymenophyllaceae confirms this impression. 
Although native agriculture is still on a very low level, and 
even near the settlements fruit trees are not often encountered, 
the oil-palm is frequently found in anthropogenic situations, al- 
ways yielding a rich harvest of epiphytes, even in exposed places. 
It is interesting to notice that the ferns found on this palm really 
belong to the natural vegetation of sun-loving epiphytes that are 
found on the top branches of the giant jungle trees. During my 
stay, a small clearing was made on Bogbulu and on the branches 
of the fallen trees practically the same species were found as 
were previously collected on the Elacis palm near the ground. 
list of the species collected in the area, subdivided according 
to habitat, is given below: 
SHADE-EPIPHYTE! 
ee on the bases of jungle trees in the cere of the primeval forest) 
CHOMANES vee AERUGINEUM van den Bosch. This is the 


m 
LOMARIOPSIS HEDERACE ston cl faving vertically on mostly small 
trees, and also trailing on a ground in wet, shady places; common, 
fertile fronds mostly absen 
OMARIOPSIS GUINEENSIS ies Alston. Growing like the preceding 
cane: remarkable for the diversity of its fronds: very young plants with 
simple lobed tate older plants with imparipinnate fronds; also common. 
jpeles M Mannianum Hook. Growing on thick trunks of old trees 
in ep ay 
ASPLENIUM BARTERI Hook. Found in the wettest localities, in deep shade; 
the ‘‘queue de rat’’ is a striking feature of the leaves, adorned with a bud. 
SuN-EPIPHYTES 
(found on the top branches of large jungle trees and on single trees in the 
open, mostly Elaeis 


230 AMERICAN FERN JOURNAL 


Lycopopium Sraupti (Nessel) Adams & Alston. Found only once, on 
ge ae in a shady situation; certainly not typically a sun-epiphyte. 
TERIS vpsusaa mee (Gmelin) Posth. A well-known tropical fern 
of ae ea 
OLEANDRA DISTENTA Kunze. Apparently deciduous and just getting new 
leaves in Septem 
LLIA CHAEROPHYLLOIDES (Poiret) Steud. One of the commonest 
epiphytes on Elaeis, and — also on trees in Leopoldville and elsewhere. 
‘VITTARIA GUINEENSIS Des 
NIUM AFRICANUM Saal Although often compared with the Bird’s- 
nest fon splenium nidus, this species does not form real nests, and is 
Comm 


Ww. 
PLATYCERIUM STEMARIA (Beauv.) Desy. These common ‘‘staghorn’’ ferns 
are no doubt the most impressive in the sun-loving vegetation in the high 
jungle trees. Especially, the huge entire fronds of a angolense, more than 
0 em. in circumference, dangling from the branches and surrounded by the 
erect brown nest-leaves, make an unforgettable mons 
rwell. t common 


LATA ‘ 
YNARIA LAURENTII (Christ Hieron. Another interesting fern with 
po and foliage lea 

Microsorium Punctatum (L.) Cop 

HYMATODES SCOLOPENDRIA (Bu a Che. Well-known tropical epiphyte. 

MICROGRAMMA LYCOPODIOIDES (L.) Copel. Extremely variable: Small and 
strongly dimorphic on high jungle-trees, much more lush in the shade in 
anthropogenic situations, sometimes covering whole stems of Elacis. 

SHADE TERRESTRIAL FERNS 
(in rain-forests and shady situations) 

PTERIS ATROVIRENS Willd. Rhachises and costae spiny. 

Loncuitis Currort (Hook.) Mett. Huge fern, more than 2 meters high, 
in wet and shady situations. The young plants are completely different 
from full-grown specimens and have been described as a different species. 

CTENITIS EFULENSIS (Baker) Tardieu 

CTENITIS LANIGERA (Kuhn) Tardieu 

Desai PROTENSA (Afzel.) Copel. The two last-named species of Ctenitis 

e to be found on shady banks and well-drained situations in the forest; 
pe to C. efulensis, they have the typical Ctenitis form and can not be 
istaken. 


Creniris JENSENIAE (C. Chr.) Tardieu. Found in several locations and 
apparently common in the primeval forest. The specimens found have been 
examined by Madame Tardieu, since they do not conform to the e in 
being less divided, not proliferous, and in having a slight anastomosis of 


FERNS IN OUBANGI, CONGO 231 


the veins; aa may represent a new species, but need further study. 
LASTREOPSIS BARTERIANA (Hook.) Tardieu. The nomenclature of Madame 
Tardieu is followed here, although this ters is not a Lastreopsis in my 
opinion. 
BOLBITIS GEMMIFERA Sranns ) C. Chr. The interesting fertile leaves look 
a bit those of Lomari 


ATHYRIUM iEWeeonn (Heol ) Tardie 
a YRIUM SAMMATII (Kuhn) Tardieu. ae last two species belong to the 
Diplazium grow 


‘p- 

ASPLENIUM SUBAEQUILATERALE Hieron. I compared my specimens (which 
have a bud below the upper pinnae) with the type of Hieronymus in Dahlem 
which is not proliferous, but otherwise identical. Alston calls the prolifer- 
ous plants Aspleniwm gemmascens Alston. This fern is near Aspleniwm 

i ial, 


ASPLENIUM CEEI Pichi-Sermolli. Found in deep shade in the jungle, cover- 
ing quite an area with deep green leaves; milan aie common. The identi- 
fication has been checked by Madame Tardie 

TERRESTRIAL FERNS 
in open situations, secondary forest, and roadsides) 


re. 
DICRANOPTERIS LINEARIS (Burm.) indeiw. The five preceding species are 
ubiquitous tropical species of weedy nature. 
WAMP STRIAL FERNS 
in open swamps and riversides) 
Lycopopium cerNuUM L. 
ELAGINELLA MYyOSURUS (Swartz) Alston 
CycLosorus striatus (Schum.) Copel. 
CycLosorus goaeiLopus (Schkuhr) Link 
NEPHROLEPIS BISERRATA (Swartz) Schott. The two last named ferns are 
very common and sometimes cover entire swamps; N. biserrata is also a 


epiphyte. 
RIS SIMILIS Kuhn. This is an interesting fern, with short, thick 


ry mAs as adopted by Madame Tardieu, which is a synonym of 
P. atrovirens Willd. My specimens were kindly compared by Dr. D. E. 
Meyer Aste the type of P. similis Kuhn in the herbarium of the Botanisches 
Musewn, Berlin-Dahlem. 

BURCHARDSTRASSE 19, HamBure 1, GERMANY 


232 ' AMERICAN FERN JOURNAL 


Ferns Cultivated in California: Sadleria 
BARBARA JOE 


Among the ferns arriving from the Hawaiian Islands is a 
group of tree ferns belonging to the genus Sadleria. Known to the 
Hawaiian natives as Amaumau and Amau, they have been known 
among some nursery dealers merely as dwarf tree ferns. Smaller 
than the true tree ferns, these ‘‘dwarfs’’ may still grow to a 
height of 12 feet beneath the shade of taller tree ferns in their 
native homes. The Hawaiians are said to have prepared a red 
dye from the trunks of Amaumau, and used the young leaves 
and pith of Amau for foo 

Sadlerias are not entirely of recent introduction to this coun- 
try, for large mature specimens have been known in Santa 
Barbara for some time. However, the majority of the cultivated 
specimens have arrived with the introduction of Cibotiwm trunks. 
When confronted with the usual selection of bare tree fern 
trunks, one can separate the Sadlerias by the protective covering 
of long narrow chaffy scales, which are distinctly different from 
the limp hairs of the Cibotium. Trunks bearing foliage are readily 
separated from each other. The fronds of Sadleria are oblong, 
and the pinnae are only pinnatifid, whereas the Cibotium 
fronds are broadly triangular, with pinnae two times further 
divided. 

Ideal in tubs or other containers, Sadlerias are better than 
Cibotiums when a smaller plant with a more open crown is 
desired. The trunks are planted about 14 to 14 their length in 
well-drained and aerated soil. They may be set in a temporary 
rooting medium until new roots appear and then transferred to 
the permanent planting. The planting media must be kept moist ; 
good drainage will help prevent rotting. Successfully rooted 
trunk specimens have the advantage of being large, mature 
plants in a relatively short time. Spore grown plants of Sadleria 
are also available in the trade. Though their trunks are not so 
developed, the foliage is usually more symmetrical than that of 
the trunk-grown specimens. 


AMERICAN FERN JOURNAL VoLUME 50, PLate 19 


Top: SADLERIA HILLEBRANDII, SEGMENTS; BortoM: SADLERIA 
CYATHEOIDES, HABIT. 


234 AMERICAN FERN JOURNAL 


The cultural requirements of Sadleria are very much like 
those of Cibotiuwm. The plants are more tolerant of heat and 
dryness than many other tree ferns. Along the coastal areas they 
will endure almost full sun. The evergreen foliage is, however, 
more lush and green when light to medium shade is provided 
along with ample water. Their rate of growth is moderate. The 
extent to which these plants will endure freezing temperatures 
has not been determined. Short periods of temperatures slightly 
lower than 32°F have been borne without injury to the plants. 

The genus Sadleria contains seven species which are native 
to the Pacific Islands. All are tree-like in habit with bipinnatifid 
or bipinnate fronds. The veins are netted to form meshes along 
the midvein; elsewhere they are free. The sori are linear and 
continuous, Bonie borne on both sides of the midvein of the 
segment. The flap-like indusium is shaped like the sorus; it is 
attached to a vein of the areole (mesh) and opens toward the 
midvein of the segment. This genus is very closely related to 
Blechnum, from which it differs mainly in being tree-like. 

At present only two species have been recorded in cultivation 
from Hawaiian shipments, but the introduction of the other 
species endemic to the Hawaiian Islands is to be expected. These 
two species may be separated by the following key: 

A. Veins not conspicuous, immersed in the frond; leaf stalk 
naked except at the base _S. eyatheoides 
AA. Veins raised and prominent on the underside of the frond; 
leaf stalk covered with scales __S. Hillebrandii 

Sadleria cyatheoides Kaulf. Amaumau. Trunk 3-5 feet ; leaf- 
stalk essentially naked except at the base, where densely covered 
with long-linear scales; midrib naked or with a few scattered 
scales; surface of frond essentially naked; veins immersed and 
inconspicuous. Hawaiian Islands, Sumatra. Semi-hardy; of easy 
culture; to 12 feet high. 

Sadleria Hillebrandii Rob. Amau. Trunk 2-3 feet; leaf-stalk 
and midrib covered with scales; surface of frond bearing long 
hairs; veins raised and conspicuous; texture less rigid than that 
S. cyatheoides. Hawaiian Islands. Semi-hardy; presently not 


or 


GROWING FERN BALLS 23 


common in the trade; to 10 feet high. 
Sadleria is easily separated from all other cultivated ferns 
by the small-tree-like habit and the presence of long-linear sori. 
DEPARTMENT OF BoraNy, UNIVERSITY OF CALIFORNIA, Los 
ANGELES, CALIFORNIA. 


Ferns in Cultivation, III. Growing Fern Balls 
SytviA LEATHERMAN 

Living ‘‘fern balls,’’ large baskets completely covered with 
beautiful billowing fern fronds, are spectacular, and always 
attract a great deal of attention. Fern growers may be interested 
in knowing how I obtain these specimen plants. 

The best ferns for fern balls in ae California are the 
species of Davallia, commonly known as ‘‘rabbit’s-foot fern’’ 
and ‘‘squirrel’s-foot fern.’’ The plants imported from Japan 
are the species Davallia Mariesii; the other common species in 
cultivation is Davallia trichomanoides, which is often sold under 
the incorrect name D. canariensis, which belongs properly to 
another species, also in cultivation, but not so common. Most 
people like to obtain small plants and grow these on to maturity. 
As the plants grow, they are shifted to larger pots. When they 
nicely fill a six inch pot, they are ready to transplant to baskets. 
The ‘‘foot’’ (rootstock) should be placed at an angle across the 
top of the pots and green sphagnum moss tucked under it. 
The ‘‘foot’’ is anchored to the moss by using hairpin-shaped 
wires; this encourages them to send out side ‘‘feet,’’ which are 
also anchored down by wires. When the six inch pot is full of 
‘‘feet,’? the plant is ready to be shifted to the basket. In the 
basket, the ‘‘feet’’ are trained across the top of the sphagnum on 
top of the basket and eventually down over the sides. It is amaz- 
ing how fast the plants will grow when given this moss cushion. 

The common Boston Fern (Nephrolepis) will respond also to 
this type of treatment. Many people cut away the runners (the 
long, wiry strands), as they think they are unslightly and un- 
necessary, and then they wonder why their ferns do not fill out. 
If these runners are wound around inside the top of the pot and 


236 AMERICAN FERN JOURNAL 


anchored to the soil by using hairpin-shaped wires, they will 


should be ned with green sphagnum moss. As the runners grow 
again, they can be wound around the soil on the top of the 
basket, and after an abundance of growth has been obtained 
on the top of the basket, the runners can be pinned down on 
the moss on the outside of the basket. After a period of time, 
the basket will be completely covered, and the result will be a 
spectacular living fern ball 

2637 NortH Lee AVENUE, SoutH Eu Monte, CALIFORNIA. 


A New Native Hardy Plant Area at the New York 
Botanical Garden 


R. C. BreNeEpict 


An informal committee of the American Fern Society is happy 
to announce that plans are well advanced for the cooperation of 
members of the American Fern Society in the development of a 
hardy native plant area at the New York Botanical Garden. The 
background on which this cooperation is based may be outlined 
as follows: 

Several members from the New York area who had been ac- 
tively interested in the past in sponsoring field trips and local 
meetings came together to explore the possibilities of further 
activity in the New York area. Under the Chairmanship of Dr. 
B. R. Allison, Chairman of the New England Field Trip Com- 
mittee of the Fern Society, with Miss Clara S. Hires and Messrs. 
Boughton Cobb and Ralph C. Benedict, the Committee got in 
touch with Dr. William Steere, who began his service as Director 
of the Botanical Garden in 1958. Two questions were asked: 
Could the Fern Society hold oceasional meetings at the New York 
Botanical Garden? and Could the Fern Society be of assistance 
in building up the collections of living fern plants at the New 
York Garden? 


NEw York BoranicaL GARDEN 237 


To both questions a cordial affirmative was received. 
garding meetings, Dr. Steere said that with due notice ti 
avoidance of conflicts, the Fern Society would be welcome to 
meet in the Museum Building. With respect to the living fern 
collections, Dr. Steere reported that the Garden had already in 
active work a special committee engaged in developing a native 
hardy fiora area under the Chairmanship of Mrs. Perey Douglas, 
and the cooperation of the Fern Society in the assemblage of 
appropriate fern species would be immediately welcome. 

The upshot of preliminary conferences with Dr. Steere was 
the calling of a meeting on May 23, 1959, at the Botanical Garden 


Society Committee had had two chances to visit the site of the 
hardy plant development, once with Dr. Steere and a second 
time with Mr. John Crawford, landscape architect in charge for 
the Botanical Garden Committee. Dr. Allison had asked Dr. 
Benedict to present a plan for Fern Society cooperation in the 
Selection and assemblage of appropriate ferns. Such a list and 
a tentative plan of action were prepared and sent in advance to 
Dr. Steere and Mr. Crawford. 

The meeting, under the auspices of a sunny but comfortable 
day, was held in three successive places: At the old Snuff Mill 
Restaurant, on a patio overlooking the Bronx River gorge, with 
a wooded bank, partly fern-clad opposite; in a classroom of the 
Museum Building; and in an inspection of the actual site of the 
new native plant area. 

The Museum portion constituted the business meeting of the 
afternoon. Twenty-two were in attendance, including Dr. Steere, 

ve members of the Botanical Garden special committee, and 
Mr. Crawford, landscape architect for the project. With Dr. 
Allison presiding, Dr. Steere outlined the Garden’s interest in 
having a “‘living museum’’ of native plants and said that the 
Garden would provide staff for the actual planting of con- 
tributed specimens. Mrs. Douglas and Mrs. Donaldson of the 
Garden Committee spoke for that committee. Emphasis was 


: 


238 AMERICAN FERN JOURNAL 


placed on the avoidance of depleting native sources of rare 
species. 

Dr. Benedict prefaced his remarks by noting that the Fern 
Society had a special connection with the New York Botanical 
Garden because it was there that the first two experimental 
privately financed issues of the American Fern Journal had been 
issued. These two were incorporated as part of Vol. 1 when the 
Fern Journal was officially adopted by the Fern Society in 1911. 

As part of a plan by which the Fern Society might contribute 
the fern plants for the new native plant area, he suggested that 
the planting list of species should consist of some thirty to forty 
species, the successful installation of which could be reasonably 
certain. This would exclude the rarer species and others known 
to be difficult of transplanting and establishment. Some time in 
the future the rarer species might be added to the planting > 
oo if specially raised spore culture plants could 
made ava 

As a means of implementing the assembling of the proposed 
list, he suggested that a special committee be designated from 
the Fern Society to act in cooperation with Mr. Crawford and 
the Botanical Garden Committee. As appropriate areas in the 
hardy plant area become determined and well prepared, it would 
be the function of this Fern Society committee to arrange for 
the collection of plants contributed by Fern Society members. 
This would be a continuing program 

ree irrigation system is about to a installed, but rock ferns 
would require special sites. At this time the best site for a 
Dryopteris species group requires the clearing of the big Jap- 
anese Polygonum. 

Dr. Robert K. Lampton, 25 North Terrace, Maplewood, New 
Jersey, agreed to accept the appointment as chairman of the 
Fern Society committee the duty of which will be to carry for- 
ward in liaison with Mr. Crawford the actual process by which 
the desired fern plants will be obtained and transmitted to the 
Botanical Garden. Members of the Fern Society who are in- 
terested in helping are advised to get in touch with Dr. Lampton, 


New York Boranicat GARDEN 239 


and ask what they may contribute and tell him what they may 
have to offer. There will probably be definite days scheduled 
when such contributions can most readily be taken care of. 

The final stage of the May 23rd meeting was held in the actual 
hardy plant area. At this time two members showed their readi- 
ness to start Fern Society contributions by bringing in actual 
plants. Miss Hires brought eight species; Dr. Allison brought in 
a large ‘colony’ of the narrow-leaved chain fern and others. 
These plants were heeled in until specific sites are ready. 

The question of possible contributions of tropical plants for 
greenhouse collections was not dealt with specifically at the 
meeting, but the promise of a substantial source of such plants 
has recently come in a letter from Mrs. Sylvia Leatherman, 
President of the Los Angeles Fern Society. At a meeting of that 
Society recently, readiness was expressed to offer any species 
from their collective list of ferns which the New York Botanical 
Garden might wish to obtain. Meanwhile, Mr. L. P. Politi, Horti- 
culturist of the New York Garden, has a very considerable col- 
lection of species under cultivation from spores obtained from 
various European and American Botanical Gardens. 

he planning committee of the Fern Society, of which Dr. 
Allison is Chairman, hopes that a second meeting may be held 
to observe results of and to provide additional contributions of 
fern plants. Mr. Politi of the Botanical Garden promises to have 
individual fern plants from his spore cultures for those in at- 
tendance. Due notices will be sent. 

In addition to the Architect-Engineer, Mr. John Crawford, 
the following members of the American Fern Society were in 
attendance: Dr. and Mrs. Benjamin R. Allison, Dr. Ralph C. 
Benedict, Mr. and Mrs. Burton Dezendorf, Dr. and Mrs. F. 
Gordon Foster, Mr. Robert Gaede, Miss Clara Hires, Dr. Robert 
K. Lampton, Dr. Elva Lawton, Mrs. Charlotte Learned, Miss 
Eva Sobol, Dr. William C. Steere, and Mr. and Mrs. George 
Tierney, and the following from the Wild Flower Group: Mrs. 
John W. Donaldson, Mrs. Perey Douglas, Chairman, Mrs. Marcus 
Fair, Mrs. Charles Hoffman, and Mrs. Martha Innes. 


240 AMERICAN FERN JOURNAL 


Two New Georgia Stations for Lycopodium complanatum var. 
flabelliforme 


Tuomas A. Hutto 


Although several species of Lycopodium occur in abundance 
in certain areas of the eastern United States, these plants have 
been reported infrequently in Georgia. This is particularly true 
for Lycopodium complanatum var. flabelliforme, which, accord- 
ing to Dr. Wilbur H. Duncan, Curator of the University of 
Georgia Herbarium, has been reported from only ten of Geor- 
gia’s 159 counties. 

In July, 1958, while on a botanizing expedition with Mr. Tom 
Jackson, of Rinehart College, I came across a small stand of 
L. complanatum var. flabelliforme in a wooded area at the base- 
of Georgia’s famous monadnock, Stone Mountain, in Dekalb 
County. According to Dr. Duncan this is the first report of this 
species from Dekalb County, so this raises the total to 11 counties. 

More recently, on January 24 of this year, my father, Mr. 
Jim Hutto, accompanied me on a search for the grapefern Bot- 
rychium dissectum var. obliquum in a wooded area in Hall 
County about six miles south of Gainesville, Georgia. I had 
previously located an extensive growth of B. dissectum in this 
locality in 1958 and I was interested in reexamining it. Several 
feet from the bank of a small stream running through the area 
I came across a patch of L. complanatum var. flabelliforme. A 
search of the area revealed that this small patch was all that 
was present. Dr. Duncan informs me that this is a new station, 
although he has previously reported this Lycopodium from Hall 
County from a station about six miles further south. 

It is my opinion that this and other species are probably more 
abundant in Georgia than we realize, particularly in the north- 
ern part of the state. With increased botanizing on the part of 
Georgia fern enthusiasts I believe the picture of the distribution 
will become more complete. 

RosweE.u, Grorata. 


VEIN PATTERNS IN MicrosoRIUM 241 


Vein Patterns in Microsorium scandens and Its Allies 
Mary D. TINDALE 


The following new combination is proposed 
Microsortum scandens (Forst. f.) Tindale, comb. nov. 

Polypodium scandens Forst. f., Prodr. 81. 1786; Benth. FI. 

Austral. 7: 770. 1878; Moore & Betche, Handb. Fl. New So. 
Wales 515. 1893. Lectotype: Without locality, labelled 275 
and 437, Polypodium scandens (BM); ‘‘Society Islands’’ 
has been added to the label later. 

Phymatodes scandens (Forst. f.) Presl, Tent. Pterid. 196. 

1836; Pichi-Sermolli, Webbia 8: 222. 1951. 

Pees scandens (Forst. f.) Fée, Gen. Fil. 271. 1852. 
Illustration: Domin, Bibl. Bot. 85: 179, fig. 40. 1913, as Poly- 

podium pustulatum. 

In Se ncabae Australia there are only two species of Micro- 
sorium, namely scandens, which ranges from southeastern 
Guscristand to Dela and M. diversifolium (Willd.) Copel., 
occurring in southeastern Queensland, New South Wales, Vic- 
toria, and Tasmania. Both species are quite common also in 
New Zealand. 

Prof. R. BE. Pichi-Sermolli has discussed in detail’ the reasons 
why we should revert to the use of the epithet scandens for this 
Australasian fern, After an examination of the type specimens 
of Polypodium scandens Forst. f. and P. pustulatum Forst. £. 
in the British Museum of Natural History, London, I agree with 
Pichi-Sermolli that the common, sweet-scented species of poly- 
pody in the rain-forests of eastern Australia and New Zealand 
is identical with Polypodium scandens Forst. f. Unfortunately, 
Domin? adopted the name P. pustulatum Forst. f. for Micro- 
sorium scandens and used Polypodium scandens Labill. for the 
closely allied species Microsorium diversifolium. Following him, 
Copeland published the new combination Mi icrosorium pustula- 
tum (Forst. f.) Copel. 


1 Webbia 8: 212-222. 1951. 
2 Bibl. Bot. 85: 178. 1913. 


VEIN PATTERNS IN MICROSORIUM 243 


The following key will serve to separate the two species found 
in southeastern Australia. They are closely allied to M. novae- 
zealandiae (Baker) Copel. of New Zealand, and M. sibomense 
(Rosenst.) Copel. of New Guinea. The latter species is char- 
acterized by a thin lamina with one or two rows of scarcely 
impressed sori. 

Seales of the rhizome squarrose, persistent, dark brown or purplish brown, 
narrowly lanceolate to lanceolate; rhizome (when living) tough and wiry; 


lamina dark green, membranous to herbaceous, with a distinct musk scent 
when fresh or freshly dried; segments of the pinnatifid fronds 1.2-10 ¢ 
lon M. adie 


eee of the rhizome appressed, deciduous, black, purplish-brown or dark 
to light grey, with a brown or light brown border, narrowly lanceolate or 
vate; rhizome (when living) fleshy and often very glaucous; lamina 
ae green, herbaceous to coriaceous, unscented; segments of the pinna- 
tifid fronds 1.8-15 em. long. M., diversifolium. 
As Copeland? points out, the type species of the genera Micro- 
sorium and Phymatodes are very unlike each other, but it is 
impossible to draw a dividing line when all the species are con- 
sidered. I concur with this view. I believe that the group of 
species allied to MW. commutatum Scuet Copel. form a con- 
necting link between these two gener 
M. scandens has a very cceehen 4 vein-pattern in the lamina 
whereas in M. diversifolium the venation is essentially of the 
type characteristic of Phymatodes, i.e. the sori of the only (or 
in some species the lower) row are each borne at the junction 
of several minor veinlets and a strong acroscopic vein (arising 
from the base of or near the base of a main lateral vein). In 
M. scandens the sori are borne at or close to the junction of the 
main lateral vein or its acroscopic branch and the horizontal vein 
approximately parallel to the costa. 


3 Gen. Fil. 195. 1947. 
Upper Lerr. PoLypopDIUM FORMOSANU 
VEINS AND Sori; HAMATE VEINS Aco sitar IN AREOLE; X 6. ER RIGHT. 
MicrosortumM SCANDENS; FERTILE LOBE; HAMATE VEINS IN po ES; x 4. 
Lower Lert. MIcrosorIuM SYLVATICUM; FERTILE Lope; SoRAL ARRANGE- 
MENT AS IN MicROSORIUM IN ONE AREOLE, AS IN PHYMATODES IN OTHER 
5 MICROSORIUM SCOLOPENDRIA; COMPLEX 


M; FERTILE LOBE WITH DETAILS OF 
Up 


PHYMATODES-TYPE VENATION; X 3 


244 AMERICAN FERN JOURNAL 


In some members of the M. commutatum group the sori are 
borne in box-like areolae which occur in alternating rows. The 
sori in the costal row are arranged as in Phymatodes but the 
upper row of sori are mostly irregularly placed. These points 
may be easily seen in the type of Microsoriwm multijugatum 
(Copel.) Copel., which was collected by Copland King, No. 228, 
at Goodenough Bay, New Guinea, in 1903 (NSW. P8078). 

Specimens such as NGF. 6788, Microsorium sp. aff. multijuga- 
tum, collected at the Baiyer ee New Guinea, by Womersley 
and Floyd in November, 1954, appear to be midway between 
Phymatodes and Microsorium. The thickened vein on which the 
sorus is borne in the costal areolae arises in some cases from 
the costa in the middle of the areola but in others from the base 
of or close to the base of the lateral veins. 

In M. sylvaticwm (Brack.) Copel. the venation is somewhat 
similar except that the parallel lateral veins are more promi- 
nent and the cross-veins parallel to the costa are less pronounced 
than in M. multijugatum. The position of the sori is sometimes 
as in Phymatodes. A specimen of M. sylvaticum collected at 
Upolu, Samoa, by E. Betche, No. 57, in February, 1880, is char- 
acterized by a thin lamina with the sori not immersed in pits. 
Many of the species such as M. scolopendria (Burm.) Copel. and 
M. alternifolium (Willd.) Copel. that have been traditionally 
placed in Phymatodes have deeply impressed sori. Other species 
such as M. sibomense and M. parksii (Copel.) Copel. with the 
typical Phymatodes-type of venation have scarcely immersed 
sori. 


The dividing line between Polypodium sens. strict. and Micro- 
sorium* may also be determined by their venation. In the latter 
genus the species usually placed in Phymatodes appear to be 


4 As recently pointed out by W. A. Sledge, Bull. Brit. Mus. (Nat. Hi st.) 
2, No. 5: 142. 1960, the generic name was ori ginally ators by Link in 
Hort. R. Bot. Berol. 2: 110: 183 33, both in the text and coe as a ies 
sorum, but later Link corrected this to Microsorium (Fil. Spee. Hort. 
rol. Cult, Recens. 116, 135. 1841), as pointed out to me by c. Vv. 
Morton 


VEIN PATTERNS IN MicrosoRIUM 245 


closer in affinity to Polypodium than those traditionally placed 
in Microsorium. The vein-pattern in Polypodium formosanum 
Bak., of Japan, is very similar to that of Microsorium scandens 
except that hamate and half-hamate veins are not given off from 
the basiscopie side of the cross-veins approximately parallel to 
the costa. In fact these cross-veins only occur in some of the 
areolae near the main rhachis, otherwise the sori are terminal 
on free veins in the areolae. These free veins arise at or near 
the base of the lateral veins on the acroscopic side. This type of 
venation is characteristic of the group of P. loriceum Li. which 
includes the following species :—P. plesiosorum Kunze (Mexico), 
P. chartacewm Bak. (Ecuador), P. falearia Kunze (Costa Rica), 
P. limbatum Brade (Brazil), P. lachnopus Wall. ex Hook. (Him- 
alayas), P. nipponicum Mett. (Japan, China and Tibet), P. pwne- 
tulatum Hook. (Ecuador) and P. amoenum Wall. ex Mett. 
(China and Tibet). I rarely observed cross-veins from the sori 
in any species of this group except P. formosanum. 

Carl Christensen® discussed and illustrated many aspects of 
the vein-patterns in the groups of Polypodium, ranging from 
P. vulgare L. and its allies which have free-veins to P. manmet- 
ense Christ. and P. microrhizoma Clarke, where areolae are 
formed to enclose the sori in some eases but the rest of the veins 
are free. He stated that this was just a step to the vein-pattern 
in the group of P. loricewm. 


ACKNOWLEDGMENTS 

I should like to express my appreciation for the facilities 
afforded by the directors and keepers of the following institu- 
tions: The Herbarium, Kew; the British Museum of Natural 
History, London, and the National Herbarium, Sydney. The 
writer also wishes to thank Dr. R. E. Holttum for a very helpful 
discussion on the subject of the above paper. 

NatTionAL Herparium, Royau Botanic GARDENS, SYDNEY, 
AUSTRALIA, 

5 Dansk. Bot. Arkiv. 5, No. 22: 1-10, 1928. 


246 AMERICAN FERN JOURNAL 


A New Station for Trichomanes Petersii in Georgia 
Henry Booxour 

In 1901, A. B. Seymour! collected Trichomanes Petersii, for 
the first time in Georgia, in a deep ravine at Tallulah Falls, 
Rabun County, of the Blue Ridge Mountain Province. Wilbur 
Duncan? made the second collection in Walker County of the 
Cumberland Plateau in 1952. On July 22, 1958, Frank Snyder 
and I discovered Trichomanes Petersii growing on rock cliffs in 
Butternut Cove of the Cedar Creek ravine, Stephens County, 
about ten miles south-southeast of the original site (now lost) at 
Tallulah Falls. The collection is Bookout & Snyder 82, 14 mile 
off Yearwood Drive up the Cedar Creek ravine, elevation 825 
feet. Specimens are in the University of Georgia Herbarium, 
the Harvard University Herbarium, and the author’s private 
collection. 

I had invited Mr. Snyder, an amateur botanist from Toccoa, 
out to Camp Mikell to see Camptosorus rhizophyllus (Bookout 
81), which I had discovered three days before on July 19. The 
walking fern itself is not common in Georgia, and this new site 
was a new record for Stephens County and a considerable ex- 
tension of its range southward for the eastern part of the state. 
Mr. Snyder and I were examining the nearby cliffs when I pulled 
off something that looked like a small liverwort or a flat-leaved 
moss, but noting the leafy rhizomes I said, ‘‘This looks like a 
filmy fern.’’ When the fruiting bodies were discovered, the 
identity was assured. The fern is common below Cedar Fork in 
the Cedar Creek ravine at isolated stations on both slopes. It 
grows on siliceous cliffs and loose rocks where they are shaded 
and kept continually moist by seepage. There are several places 
where the filmy fern might have been found growing in the 
“spray of a waterfall,’”’ but not a single instance was noted. It 
seemed to prefer, instead, the protection afforded by overhanging 
rocks. 
on B. Seymour, 1903. Trichomanes Petersii Found Anew. Torreya 3: 
-< Mises Dunean, 1955. New Records for Georgia Ferns. This JOURNAL 


Recent Fern LITERATURE 247 


On August 21, I revisited the site of Camptosorus rhizophyllus 
where I discovered something that originally had gone unnoticed 
—that on three different rocks the filmy fern and the walking 
fern were growing in close association. Since the filmy fern is 
always found on siliceous rocks* and the walking fern more com- 
monly on caleareous rocks,‘ their association seemed at first sight 
to be remarkable. Dr. A. S. Furcron, the state geologist, identi- 
fied one of the rocks as hornblende gneiss, a siliceous rock con- 
taining a calcium silicate. Perhaps this calcium silicate, Dr. 
Fureron suggested, releases sufficient calicum ions to make the 
walking fern at home. However, the other two rocks were 
muscovite quartzite with a granitoid appearance. In any case, 
the dry soil separated from the intertangled rhizomes of the 
filmy fern and the walking fern when tested with a Sudbury 
Soil Test Kit, model D, showed a rather strongly acidie pH of 
4.5. The filmy fern, apparently, is typically located. The walk- 
ing fern, however, here demonstrates its greater impartiality to 
environment in forming a self-perpetuating colony under rela- 
tively atypical conditions of an acidic soil. 

ATLANTA, GEORGIA 


3. T. Wherry, 1955. The Substratum of Trichomanes Petersii. This 
JOURNAL 45: —95. ; sa 
Rogers MeVaugh and Joseph Pyron, 1951. Ferns of Georgia, p. 72. 


Recent Fern Literature 


PouLEN AND Sporn MorpHowocy.'—The first part of this vol- 
ume is devoted to illustrations and legends for 57 genera of 
gymnosperms, 113 genera of pteridophytes, and 69 genera of 
bryophytes; the second part to a discussion of the ultra-micro- 
scopic structure of pollen and spore walls; and the third part to 
methods of preparation of ultra-thin sections. 

“oy Sr Pterido- 


Erdtman, 
151 pp. lus. 


1 Pollen and Spore Morphology/Plant Taxonomy. Gym SI 
phyta, Bryophyta. (An introduction to Palynology, vol. 2.) 
Ed. Almquist & Wiksell, Stockholm; Ronald, New York, 1957. 
$8. 


248 AMERICAN FERN JOURNAL 


Readers of the American Fern Journal who are interested in 
fern spores will find illustrations of spores representing world- 
wide distribution of fern genera, but not a complete coverage of 
all the fern genera of the world. The arrangement of the illus- 
trations is alphabetical by genera with a few exceptions when 
several genera of a family are illustrated in one figure. These 
figures usually include a view of the spore at a magnification of 
250%, a palynogram with the distal face on the left and the 
optical section on the right at a magnification of 1000, plus 
sketches to show the LO pattern at high and low focus. The LO 


slightly darker field; then, as the focus is moved downwards, the 
pattern reverses. Thus, through the differential absorption of 
light, details of the sclerine stratification are predicated. 

Unfortunately the text to these illustrations is to be published 
in a separate volume that has not yet appeared. It is to be hoped 
that the descriptions will tell how other species of a genus differ 
from the one illustrated. One of the few instances in which more 
than one entity of a genus is illustrated is Cystopteris fragilis 
which is depicted by the typical, spiny spore and its forma 
Dickicana by a perine surface of irregular folds and crests like 
many species of Dryopteris. The taxonomists in the past have 
neglected characters of the spores which may be useful in classi- 
fication and identification of ferns, 

These questions may be asked: How reliable is the identifica- 
tion, in the absence of the citation of voucher specimens? How 
accurately do these palynograms represent the spores? The 
author had the privilege of studying in Professor Erdtman’s 
laboratory. Most of the spores were secured from herbarium 
specimens. Although a record was kept of the data on the her- 
barium label and the herbaria which furnished samples, Pro- 
fessor Erdtman had to depend upon others for the reliability of 
the identification of specimens. Citation of voucher specimens is 
expected in the forth-coming volume of text. Slides of the spore 


Recent FERN LITERATURE 249 


material were prepared by the acetolysis process. Briefly, the 
dry spore sample was boiled for 1 minute in a mixture of 9 parts 
acetic anhydride plus 1 part of concentrated sulphuric acid, then 
centrifuged, washed, and mounted in glycerin jelly. This process 
removed the protoplasmic contents of the spore (also the perine 
at times), thus making it easier to study details of the wall struc- 
ture. Professor Erdtman studied the slide, made a rough sketch 
to indicate the details he wanted the artist to illustrate, and 
mailed the slide to the artist who then made the illustration. The 
writer has studied some of these slides and has actually located 
the grain the artist drew. These stylized drawings are a faithful 
representation of the grains studied. The method of prepara- 


lactophenol, or other mounting media will and do look different 
from those prepared by acetolysis. There is a danger associated 
with removing spores from a herbarium sheet which should not 
be over looked—that is, the spores may be contaminated with 
those from other specimens. This is serious if only a few spores 
are taken from the specimen. Contamination is probably respon- 
sible for the opinion that a given species may produce both 
trilete and monolete spores. 

This and its companion volumes will be necessary reference 
works for the palynologist—CLamr A. Brown. 


New Garpen Encyciopepia.—Under the editorship of T. H. 
Everett, of the New York Botanical Garden, the ‘‘New Illus- 
trated Encyclopedia of Gardening, Unabridged’’ was published 
in April, 1960. It is in six volumes, bound in buckram and boxed. 
The work contains about 3,000 pages, well over 3,000 black and 
white pictures, and more than 600 colored illustrations. It is 
primarily a gardening rather than a botanical work. The pub- 
lishers retail price of the set is $49.50, but it is being offered to 
members of the Fern Society at the special price of $33.00, plus 
$1.50 mailing charge. Dr. Ira L. Wiggins (Natural History 
Museum, Stanford University, Stanford, California) has agreed 
to handle orders. 


250 AMERICAN FERN JOURNAL 


Some REcENT Papers ON CHINESE FERNS by R. C. Ching.— 
The well-known pteridologist R. C. Ching was inactive botanical- 
ly for many years during and after the last war, but he is now 
at work again at the Academia Sinica, Peking, Peoples Republic 
of China. A recent letter from him will be quoted in the Notes 
and News section. Some of Ching’s recent publications have been 
kindly shown to me by Dr. Egbert H. Walker and Dr. Clyde F. 
Reed. These are noted below. They are all in Chinese, with 
English summaries, and Latin descriptions of the novelties. 

“‘Cyrtomidiectyum Ching, A Yet Little Known Chinese Fern 
Genus.’’! The genus Cyrtomidictyum was proposed by Ching in 
1940, but due to war conditions, it was not known to Copeland 
when he was preparing his Index Filicum. It was based on 
Aspidium lepidocaulon Hook., which was placed in Polystichum 
in the Index Filicum. It differs from Polystichum in being al- 
ways simply pinnate, in exindusiate sori, borne dorsally on the 
veins, in the pinnae being entire and not aristate-serrate and not 
tipped with a spine, and in having the sterile blades prolonged 
and rooting at apex. All these characters occur occasionally in 
other species of Polystichum but are combined only in Cyrto- 
midictyum. e€ genus was originally monotypic, but three 
species are now added: C. basipinnatum (Baker) Ching, op. cit. 
262 (based on Aspidium basipinnatum Baker (from Kwang- 
tung), C. conjunctum Ching, sp. nov., op. cit. (type: Hsiung 
Yao-Ko 06466, from Hwang-Kan Shan, Kiangsi), and (. Faberi 
(Baker) Ching, op. cit. 265 (based on Nephrodium Faberi Baker, 
from Chekiang; oceurs also in Taiwan). Also described is C. 
lepidocaulon (Hook.) Ching var. inciswm Ching, op. cit. 265 
(type: Taquet 2456, from Quelpaert, Korea). 

‘On the Genus Adiantum J. of China with Notes on Some 
Related Species from Neighbouring Regions.’’? Ching argues 
that Adiantum belongs to a distinct family, Adiantaceae, which 
ought not to be joined with the Pteridaceae following Copeland 


1 Acta Phytotaxonomica Siniea 6: 255-266. pl. LI-LIV, August, 1957. 
* Acta Phytotax. Sin. 6: 301-354. November, 1957, : 


ReceNtT FERN LITERATURE 251 


or with the gymnogrammoid and vittarioid ferns as proposed by 
Holttum. He recognizes two genera, Adiantum and Hewardia; 
H. olivacea (Baker) Ching, H. phyllitidis (J. Smith) Ching, and 
H. lucida (Cav.) Ching are proposed as new combinations but 
not validly published by the Code, since the place of publication 
of the basionym is not given. A key in English is given to the 
34 species of Adiantum recognized from China. If space is avail- 
able in the Fern JourNAL, perhaps we shall reprint this key, 
which should be more widely accessible. 

“The Fern Genus Plagiogyria on the Mainland of Asia.’” 
This paper is fortunately completely published both in Chinese 
and English. It discusses Plagiogyria in detail as to its history, 
its systematic position (as the representative of a monotypic 
family according to Ching), its origin and geographic distribu- 
tion, its adaptive peculiarities, and its taxonomic division. The 
genus is divided into two sections termed Carinatae and 
Euplagiogyriae, the latter with the subsections Huphlebiae, Ad- 
natae, and Pycnophyllaec. Unfortunately, although these are 
carefully provided with Latin diagnoses they are not validly 
published by Art. 35 of the Code, since the nomenclatural types 
are not indicated. Altogether, 33 species are recognized from 
China, including a large number of new ones. 

“A Revision of the Fern Genus Archangiopteris Christ & 
Giesenhagen.’’* Archangiopteris is said to be distinguished from 
Angiopteris by its smaller, simpler, slenderer habit, by having 
elongate, suberect dorsiventral rhizomes provided with thin- 
fleshy ovate-oblong bivalved persistent stipules, a stipe with a 
fleshy nodose swelling below the middle (or 4 or 5 nodes in A. 
hokouensis Ching), by the presence, especially on the lower part 
of the stipe, of elongate, clathrate, coarsely dentate, peltately 
affixed scales, by the simply-pinnate blades (bipinnate in A. 
bipinnata Ching), with few pairs of large, broadly lanceolate 
pinnae with inflated petiolules, by the longer sori, consisting of 


3 Acta Phytotax. Sin. 7: 105-154, pl. XXVIII-XL. May, 1958. 
4 Acta Phytotax. Sin. 7: 201-224, pl. XLIX-LU. August, 1958. 


woe AMERICAN FERN JOURNAL 


many sporangia (40-240), which are not inframarginal but 
medial, by the numerous, articulate, freely-branched paraphyses, 
and by the echinate spores. Along with Macroglossum, these two 
genera constitute the family Angiopteridaceae, which Ching be- 
lieves ought to be kept distinct from the Marattiaceae. Ten 
species of Archangiopteris are now known, confined to a small 
area in Yunnan, Kwangsi, Annam, and Tonkin, except for A. 
tonkinensis (Hayata) Ching, which has been found on Hainan, 
. Somai Hayata, of Taiwan. 

‘Materials for the Pteridophytie Flora of Hainan.’’> The 
text of this series of notes on the ferns of the island of Hainan is 
not translated into English, but the new species are given Latin 
diagnoses. Some of the species, e.g. Dicranopteris ampla, are not 
validly published because they are based on more than a single 
collection and the nomenclatural type is not indicated. The new 
species, both valid and invalid, are®: Selaginella scabrifolia 
Ching & C. H. Wang, Angiopterss acutidentata, A. caudipinna, 
a haces A, Howti Ching & Wang, A. neglecta Ching & 
Wang, A. oblanceolata Ching & Wang, A. renioti Ching & Wang, 
A. subintegra, A. venulosa, Osmunda an gustifolia, Dicranopteris 
ampla Ching & Chiu, Hicriopteris simulans, Mecodium haina- 
nense, Hymenophyltum spinosum, Vandenboschia assimilis Ching 
& Chiu, V. hainanensis Chin ng & Chiu, Gonocormus australis, 
Orepidomanas dilatatum Ching & Weae: C. hainanense, C. 
Smithiae, Microlepia scyphoformis Ching & Wang, Hypolepis 
gigantea, Lindsaya neocultrata Ching & Wang, Oleandra haina- 
nensis, Pteris crassiuscula Ching & Wang, P. hainanensis, Dipla- 
zium serratifolium, D. submettenianum, D. viridescens, Cyathea 
hainanensis, Dryopteris acutidens, D. caudifrons, D. livida [as 
“‘lividis’’], Tectaria media, Lepisorus affinis, L. longifolius, 
Phymatodes lancea Ching & Wang, Colysis intermedia Ching 
_ C. triphylla, Vittaria latifolia, and V. lauana—C. V. 


5 Acta Phytotax. Sin, 8: 125— 171, 
pl. XVI-XXIV. August, 1959. 
6 All by Ching unless otherwise specifie d. : 


NotrEs AND NEws 253 


Notes and News 

The Los ANGELES FERN Soctety continued to meet regularly 
throughout the last year. The programs were devoted to dis- 
cussions of ferns and their cultural requirements. Culture from 
fern spores was capably discussed and demonstrated by Frank 
Sobas and Frank Pauker, the Program Chairman and the Mem- 
bership Chairman, respectively; this created more interest in 
the fern spore bank. 

Dr. W. C. Drummond gave a series of lessons on the technical 
words used in describing ferns. He has prepared a mimeo- 
graphed list each month for distribution to the members. 

The Southern California Horticulture Society invited the Los 
Angeles Fern Society to present a program. The speaker, Sylvia 
Leatherman, discussed requirements as to light, soils, watering, 
and so forth, and gave ideas on the use of ferns in landscaping ; 
her talk was illustrated by specimens. 

During the year members have brought specimen plants to the 
meetings for exhibition. One of the most spectacular displays 
was Mr. Glen Seofield’s huge specimen Platyceriums. 

During the summer, meetings are held at members’ gardens 
and also in the Fern Dell in Griffith Park, with pot luck dinners 
and a member participation program. These garden meetings 
are always a treat from the superb collections of living ferns 
on display. 

The present officers are: Sylvia B. Leatherman, President; 
Dr. W. C. Drummond, Vice President; Mabel Anderson, Secre- 
tary, and Hertha Solmitz, Treasurer. Marie Zachau is the Li- 
brarian, and lends books to members.—Sytvia B. LEATHERMAN. 

Excerpr rrom LETTER By R. C. Caine To Ciype F. REEep :-— 
‘*”.. Mr. Morton is right in saying! that Asplenium conmixum 
Ching is an err. script for A. commiztum. I am not the author 
of the pteridophytes of north-eastern China. It was written by 
a group of young men of the Forestry and Soil Institute in 
Mukden. In the course of preparation for the book, they came 
to consult the fern specimens in our herbarium. I did not see 


1This JourNaL 49: 127. 1959. 


254 AMERICAN FERN JOURNAL 


their manuscript before going to the press. pata aes 
bium C. Chr., based upon Harry Smith no. 6130, m Shensi 
was published for the first time in Dansk Bot. Hee "Vol 3, no. 
3, pp. 56, 1936, a fact which I failed to know until a few weeks 
after I had conveyed to Mukden my short Latin diagnosis; we 
have a cotype in our herbarium. So that the correct citation for 
this species should be: C. Chr. in Dansk Bot. Arkiv and not FI. 
Herb. Bor-Orient. China. As era keene by Christensen, the type 
is now in Herb. Uppsala, Sweden. 


Lerrer rrom R. C. Curne.—Dr. R. C. Ching published a 
number of important papers on Chinese ferns and on fern classi- 
fication during the thirties but was inactive botanically during 
and after the war. A recent letter from him may be of interest 
to others: 


Dear Mr. Morton: November 5, 1959 

We have known each other by name for many years. I remem- 
ber I once received your papers on ferns, long ago. As you know, 
I am an old acquaintance of Dr. William Maxon of your Institu- 
tion, and at one time we worked together in the herbarium at 
Kew. Since the outbreak of the war waged by the Japanese in 
1937, we were cut off. Fortunately, I barely had time enough to 
return to your institution all the herbarium species of Dryopteris 
of the Himalayan and Chinese regions that Dr. Maxon had sent 
to me on loan for my monograph of the genus. 

I did not publish much on ferns between the years 1942 and 
1953, for in that period I was teaching at the University of 
Yunnan, as a professor of forestry, which was my original pro- 
fession. Can you tell me where Dr. Maxon is now and also Dr. 
E. B. Copeland? I received his Genera Filicum in 1948 but I 
have heard from him no more since. 

I am now engaged in preparation of the Fern Flora of China, 
which is a part of the Flora of China, which when completed 
will be in eighty volumes, of which five volumes are now in press. 
Volume 2, the first part of the Fern Flora, was published on the 
first of October, 1959. I am sending you under separate post a 


AMERICAN FERN SOCIETY 250 


copy of it with my compliments.t There will be four more 
volumes on the ferns, containing in all about 2,300 species in 
China, to be published in the next few years. 

With best regards, I am, 

Sincerely yours, 

R. C. CHIne 

Division or TAXONOMY AND PHYTOGEOGRAPHY 

ACADEMIA SINICA, PEKING, CHINA 


American Fern Society 


New MEMBERS 
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1Now received, and to be reviewed later. C.V.M. 


256 AMERICAN FERN JOURNAL 


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CHANGES OF ADDRESS 


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Help for the amateur botanist, and hints for the Lgnearage collector, 
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Vou. 50 OctoBEr-DecemBer, 1960 No. 4 


American Fern Journal 


AME CAN FERN SOCIETY 
Res 


C. V. MORTON 
anrabe: BENEDICT IRA L. WIGGINS 
A. C, SMITH 


CONTENTS 
On the Lava Fields of Rangitoto..............:00:00 MARGUERITE CROOKES 257 - 


Feeding Hardy Ferns: Wise, Safe, and Risky Methods 
A. J. MacsELr 263 


Azolla caroliniana Willd. in Georgia.................. Winsur H. Duncan 266 - 


The Lycopodiaceae and Selaginellaceae of Iowa 
Tom S. CoOpeRRIDER 267 - 


Psilotum in Louisiana Wi.uiAM D. Reese 269 - 


A Se Characters of the 
. ne Fern WarD CRANE 270 - 


Taxonomic Notes on Ferns, I. C. V. Morton 275 - 


nae yy Observations on the Himalayan Species of Athyrium 
d Comments on the ee gel of the Genus 
N. Meura anv 8, 8S. Brr 276°: 


Notes and News: Fern Photograph 296 


American Fern Society 296 


Index to Volume 50 sad 


The American Fern Society 


Council for 1960 
OFFICERS FOR THE YEAR 

Ciam A. Brown, Department of Botany, Louisiana State University, Baton 
Rouge, Louisiana President 
— RayMOND, oe Botanical Garden, 4101 East Sherbrooke Street, 
Montreal, Cana Vice-President 

DonaLp patesay Longwood Gardens, Kennett Square, Penna. 
Secretary 
WALTER 8. Puituips, Department of Botany, University of Arizona, Tucson, 
Arizona Treasurer 
C. V. Morton, Smithsonian Institution, Washington 25, D. C. 
Editor-in-Chief 


OFFICIAL ORGAN 
American Fern Journal 


EDITORS 
a, OO ee. aeiitotion, Washington 25, D. C. 
R. C. BENEDICT 5 Hall Street, Broo 5, N. Y. 
Tra L. Nabeccinten eT —— Start nford University, Stanford, Calif. 
A. C, Suz ..Smithsonian Institution, Washington 25, D. C. 


lustrated quarterly devoted to the general study of ferns, published 
anime — a the ae n Fern Society, 3110 Elm Ave., Baltimore 11, Md. 
postage aid. at Baltimore, Md. 
Matter for publieation should be addressed to C. V. Morton, Smithsonian 
Institution, W: 
Sabasintion $2.50, ‘xelaive of agency handling fees; sent free to mem- 


bers of the erican Fern Society even! soa $2.50; sustaining mem- 
bership, $5.00; life member ships $50.00). racted reprints, if ordered in 
> bodes be prbbeareciht authors at es They should be ordered when 


r 
Baek ' vonanas $2.00 each; single back numbers 50 cents each; Cumulat 
Index den vols, 1-25, 25 cents. Ten per cent discount on orhans of six ae 
or mi 
Chan, nges of address should be sent to the Secretary: Dr. ae Huttle- 


ston, Longwood Gardens, Kenne ware, Pennsylvania. Members and sub- 
seribers should sion two et and for changes to take effect. 

= ~~ stor ip, a orders for back numbers, and 

other busin ms should be addressed to the Treasurer: Dr. 


Walter 8. “ration Danes of Botany, University of Arizona, Tucson, 


agit AND CURATOR OF THE HERBARIUM 
W. H. Wagner, University of Michigan, Ann Arbor, Mich. 

A pbaet loan department is maintained in — —_ the grind 
and herbarium. Members may borrow ag 
the borrower paying all postal or express coal The saree wang the 5 Be 
pagelinn tryed : a on ee exchan; nges; a membe rship list 

interv: th 

fe Stee eee assist those interested in obtaining specimens 


American Fern Fournal 


Vou. 50 OcropeR-DECEMBER, 1960 No. 4 


On the Lava Fields of Rangitoto* 
MARGUERITE CROOKES 


Surrounding the areas and ‘‘islands’’ of vegetation on Rangi- 
toto Island, New Zealand, one encounters masses of black lava, 
treacherous to scramble over and ugly to behold. But a promising 
spot indeed is this ‘‘cinder heap’’ for a fern hunter. One would 
expect a few rock ferns and some of the hardy and ubiquitous 


variety of its fern flora, and the greatest surprise of all is the 
great quantity of filmy ferns ( species of Hymenophyllaceae ) 
which one associates with the shady forests and deep valleys of 
the mainland. 

Exploring the islands one will find growing on them, right out 
on to the open lava, huge masses of Kidney Fern, Cardiomanes 
reniforme (Forster) Presl, with its beautiful clear green kidney- 
shaped fronds, fringed when fertile with tiny bright brown 
columns of sporangia. The kidney fern is also found completely 
covering the floor and rocky sides of small gullies where the 
lava has consolidated. Along with it is found in many places an 
abundance of Piri Piri, Mecodium sanguinolentum (Forst.) 
Presl, sometimes called the Scented Filmy fern because it scents 
the paper when drying. I well remember the astonishment of 
the late Sir Arthur Hill, Director of Kew, when he saw these 
exquisite delicate ‘‘filmies’’ flourishing on the open lava. ‘‘Good 

eavens,’’ he exclaimed, ‘‘Why at Kew we grow those things 
under three layers of glass!’’ Nor are the filmies confined to 


these two species. One sometimes meets with the handsome 


*Invited paper for Fiftieth Anniversary Volume. [ Ea. ] : 
Volume 50, No. 3, of the JOURNAL, pp. 225-256, was issued Oct. 18, 1960. 


AMERICAN FERN JOURNAL VOLUME 50, PLATE 21 


RANGIT 


0 ISLAND, AT ENTRANCE TO AUCKLAND Hargpour, ABoUT 3 MILES 
Wiper, 854 Freer Higu, Area 6400 Acres. PHorograpH py L. H. MILLENER 


Lava Fretps or RANGITOTO 259 


Mecodium dilatatum (A. Rich.) Copel. with its large bright 
green frond. This species is a tree climber, and so also is the 
occasionally met Hairy Filmy Fern, Mecodium scabrum (A. 
Rich.) Copel., with its rather bristly stipe and dark dull green 
frond, and less commonly, the finely cut Meringiwm multifidum 
(Forst.) Copel. with serrate edges. Occasionally, Mecodium 
demissum (Forst.) Copel., one of our larger filmy ferns, is also 
met. Smaller fry are not entirely absent though not common. 
Here and there we find the little dull green Hymenophyllum 
revolution Col. (long confused with the widespread Tunbridge 
fern, Hymenophyllum tunbridgense), the rather yellow-green 
fronds of Mecodium flabellatum (Labill.) Copel. and the small 
ereyish-green Mecodium rarum (R. Br.) Copel. 


But although the filmies are the most astonishing of its ferns, 
Rangitoto yields many others of great beauty and interest. 

ost attractive is the common Microsorium diversifolium 
(Willd.) Copel. with its shining bright green pinnatifid fronds 
often bearing large bright brown or orange sori. It climbs most 
energetically over rock and tree by means of its fleshy green 
rhizome, ‘‘spotted like a snake’’ owing to the presence of the 
black appressed scales. Of similar habit and catholic tastes is 
the little Pyrrosia serpens (Forst.) Ching, but its rhizome is 
more slender and its simple fleshy fronds have a whitish or buff 
coloured tomentum on the under side. 


That large, complex and fascinating genus Asplenium is 
represented by four species. The beautiful tufts of the glossy 
bright green, simply pinnate fronds of the Shining Spleenwort 
(Asplenium lucidum Forst.) are to be seen, large and luxuriant 
in the shade and smaller, tougher and more yellowish-green in 
exposed places. The tufts of Asplenium falcatum Lam., also 
with simply pinnate fronds, are found in somewhat similar 
situations. But perhaps most fascinating of all is the Hanging 
Spleenwort, Asplenium flaccidum Forst., a most puzzling com- 
pound species which still poses many problems for the sys- 
tematist. Much work has been done on it by the late David 
Knowlton, who was engaged in carrying out a detailed study 


AMERICAN FERN JOURNAL VoLUME 50, PLATE 22 


An ISLAND or VEGETATION, SHOWING SomE OF SURROUNDING LAVA FIELD 
AND CONSOLIDATED Forest anout BASE or THE CONE. PHOTOGRAPH BY 
ae. MILLENER 


Lava Fieups or RAaneiroro 261 


of the New Zealand Aspleniums when he so tragically lost his 
life while fern hunting on the Little Barrier Island, Hauraki 
Gulf. The Hanging Spleenwort on Rangitoto seems quite nor- 
mal, though as always varying greatly according to situation. 
In the forest it sends down long, pendent, thick fronds rather 
like green leather, sometimes reaching five feet, but when on the 
rocks it produces smaller stiffer tufts of a rather yellowish- 

een. The Hanging Spleenwort apparently hybridizes with 
several other species with considerable enthusiasm, but 
Raugitoto I have only ever encountered one plant that looked at 
ali. suspicious. 

he common bracken, Pteridium esculentum (Forst.) Diels, 

is encountered here and there, but as there is no room for it to 
run it presents a somewhat disgruntled appearance. Much m 
at home is its not distant relative sometimes called eutihtie 
Bracken or Shaking Bracken, Pteris tremula R. Br., which, in 
spite of its timorous name, is hardy, and sends up its tall rather 
light green tufts of fronds in all sort of odd attractive corners. 
The more beautiful Pteris macilenta A. Rich. has also been 
encountered but only rarely. One might also see occasionally, 
though only where its rhizomes have room to run, the somewhat 
sticky Scented Fern, Paesia scaberula Kuhn, whose sun-loving 
tastes and exuberant growth make it extremely unpopular with 
the New Zealand farmer. 

If one investigates suitable rocks where humus has accumu- 
lated in the crevices, two small Rock Ferns Cheilanthes Siebert 
Kunze and C. distans R. Br. will be found. They are only a few 
inches high and are not at all easy to distinguish when mature, 
but the unrolled fronds of C. distans are very hairy giving a 
‘‘eottonwool’’ effect to the young Nae Another Rock Fern 
rather larger and with a simply pinnate frond and rounded 


when young has beautiful autumn colouring. Also to be seen 
on Rangitoto is our sole New Zealand member of the Comb Fern 


262 AMERICAN FERN JOURNAL 


genus, Ctenopteris Blume. Our little New Zealand Comb Fern, 
Ctenopteris heterophylla (Labill.) Tindale, is a tough little 
ufted fern with deeply pinnatafid fronds. Two more little 


fronds and is moreover stoloniferous, and so if it finds a suitable 
tree trunk or rock face can cover considerable areas. It is not 
particularly common on Rangitoto as it likes a certain amount 
of shade. The exact systematic position of this monotypic genus 
has been a matter of earnest debate among pteridologists. It 
has on occasion even been included among ‘‘jointed ferns’’ to 
the dismay of Copeland who very firmly named it Anarthrop- 
teris (an, not, and arthron, a joint) so that such unseemly errors 
should not again be committed. 

A few odd species will crop up where the land has become 
more consolidated and more soil has collected. Two maidenhairs, 
Adiantum affine Willd. and A. hispidulum Swartz, have been 
recorded, and also a tree fern, Cyathea dealbata (Forst.) Swartz, 
and Shield Fern, Polystichum Richardii (Hook.) J. Smith, but 
these are not characteristic. 

I feel I cannot conclude without some word of the ‘‘hidden 
ferns’’ of Rangitoto, which are a never-ceasing source of wonder 
to the visitor. One pursues a track through forbidding masses 
of black brittle lava rocks some quite small, some large or 
forming slabs, part of the cooled surface of the once molten 
flow, an area seemingly of complete and barren devastation. 
But if one steps aside on to the treacherous lava—carefully for 
it has an unmannerly habit of giving way and badly scored 
ankles are not hard to acquire—then bends down, moves aside a 
clump of rock, and peers into the miniature cavern below one 
will be astonished at what is seen. First is noticed the delicate 
green fronds of the Necklace Fern, Asplenium flabellifolium 


$Me 


FrrpiInc Harpy FERNS 263 


Cay., which sends its little simply pinnate fronds from rock 
to rock; on finding suitable substrata these fronds will root at 
the end and continue on their fragile way. Tiny clumps of 
filmy ferns, probably Mecodium sanguinolentum (Forst.) Copel. 
may be encountered; the clump will never extend very far, but 
it is amazing to find it there at all. Liable to be found also are 
small plants of Shining Spleenwort, Hanging Spleenwort, and 
Asplenium falcatum Lam., and the Trembling Bracken may also 
put in an appearance. These ferns are not likely to grow very 
large and indeed some may not survive at all when they grow 
out into full sunlight. But they will gradually add their 
modicum of humus and in the meantime are a lasting source of 
delight to the wandering botanist. 

Other areas in New Zealand may be found rich in the num- 
ber of fern species but I know of no other spot that can rival 
Rangitoto with a fern wealth that is at one so beautiful and so 
entirely unexpected. 

Hout, Mountain Roap, HEenperson, AUCKLAND, 
New ZEALAND. 


Feeding Hardy Ferns: Wise, Safe, and Risky Methods' 
A. J. MACSELF 

The majority of our members are experienced cultivators of 
ferns who require no instruction from me, but so often, when 
friends have seen my ferns in their full dress, the question has— 
been asked, ‘‘What do you feed them on?”’ that I am daring to 
place my views on the subject before readers in order that they 
may pull them to pieces, ruthlessly, if they wish, so long as they 
will offer alternatives from which we may all learn something. 

Ferns, hardy or exotic, do not relish rank animal manure. 
That much I proved in the early days of my horticultural 
career, when I ruined a batch of several thousand market ferns 
by mixing cow and horse manure in the potting compost. I have 
since witnessed the torture and piecemeal murder of a planta- 


tion of hardy ferns which a gardener of the know-all type in- 


1Reprinted from the British Fern Gazette, by permission. 


264 AMERICAN FERN JOURNAL 


sisted, despite all advice, upon planting in richly manured soil. 
Whilst my opinion is that they dislike rank manure such as we 
use in the preparation of rose beds and herbaceous borders, I 
would say that hardy ferns positively loathe and hate compound 
fertilizers if applied in the manner usually adopted in the feed- 
ing of flowering plants. 

I am well aware that it is possible, when a fern in a pot has 
filled the receptable with roots, to compel it to produce extreme- 


Why? Simply because it was well known at the nursery that 
they would soon be but lumber occupying s 

Thus far, it doubtless appears to the a that my idea of 
feeding ferns is to starve them, but no; I want rather to urge 
that in dealing with hardy ferns dieting is a matter for careful 
consideration, and is very different from cramming, choking or 
giving one’s plants dyspepsia. Ferns, like most plants, require 
potash, nitrogen, and phosphates, but the proportions of these 
which will make a well-balanced food for ferns is not on all 
fours with the perfect blends for a dahlia, chrysanthemum or 
an onion. Long before I learned of the existence of the British 
Pteridological Society I carried out a good many tests to ascer- 
tain the effect of various kinds of foods upon various kinds of 
ferns, and here are some of my findings. 

Scolopendriums will assimilate and benefit from more liberal 
feedings than other kinds of ferns. They will make wonderful 
fronds if given light pepperings of a mixture consisting of two 
parts superphosphate of lime, one part sulphate of potash and 
one part sulphate of ammonia. Athyriums treated with the same 
mixture grow rapidly out of character, and become gross and 
coarse. The fronds of Polystichums fed thus are prone to irregu- 
lar development, but by using a weak liquid made alternately 
with nitrate of soda and muriate of potash—not more than a 


Frepine Harpy FERNS 265 


teaspoonful of either salt to the gallon of water—Polystichums — 
may be made to produce great fronds, even in development and 
rich bright green. Personally, I would adopt this feeding only 
in the case of plants to be grown as specimens for a particular 
purpose, such as for exhibition, and I would not continue the 
diet for two successive years unless I was prepared to throw the 
lant away after the second year. Polypodiums will gain 

strength and substance from superphosphate, but sulphate of 
ammonia soon causes rotting of the rhizomes. 

echnums seems to turn sick at anything in the chemical 
line, and Asplenium Trichomanes say no to anything richer than 

rcoal, from which there is little obtainable in the way of 
slable food although it serves a useful purpose in keeping com- 
post open and sweet. Adiantums make big fronds if fed with 
nitrate of soda, but the texture is soft and flabby, and the centres — 
of the crowns are disposed to die after a season’s production of 
lush growth. 

If I were bent upon growing a dozen assorted ferns in pots to 
show in great form, my first step would be to mix some bone 
meal and crushed oyster or cockle shell with the potting soil. 
If the potting were done in August or September, plain water 
only would be given until the following April. Thenceforward 
one watering would be with very dilute nitrate of soda and the 
next equally weak muriate of potash. 

In the open ground, where I want plants to make good steady 
growth and to retain health and vigour year after year, my plan 
is to dig in bone meal when preparing the site. Thereafter all. 
the feeding done is to spinkle old soot over the ground not more 
than once a year, and each winter to spread the ashes from a 
garden fire which has consumed a good deal of woody material. 
I do not object to a coating half an inch thick, for it never seems 
to do any harm, and supplies potash in the forms ferns seem to 
relish, 

Once in about three years I have thrown a few crystals of 
sulphate of iron around my Lastreas and Polypodiums, and the 
fronds stand longer and take on a darker tone of green. A dust- 


266 - AMERICAN FERN JOURNAL 


ing of finely powdered lime once in a while is acceptable to the 
Lastreas (except dilatata), the Polypodiums, and the Scolo- 
pendriums, and that about sums up feeding ferns so far as I am 
concerned. I know some peuple use Blank’s celebrated fertilizers 
according to the instructions on the tins, and I know they can 
boast about the length and breadth of fronds produced, but they 
also have frequent reason to complain that this plant or that 
was grand last year, but something has gone wrong with it and 
it looks like going home. 

Occasionally I have had a Polystichum or a Scolopendrium 
which has gone hard and stubborn in the crown, refusing to 
send up new fronds. A drenching with water in which sulphate 
of ammonia, 1% oz. to the gallon, has been dissolved for six hours 
or more has been given, and growth has soon broken away, but 
having achieved that much no more has been given; it is all 
right to give a gentle fillip, but all wrong to whip and goad to 
second exhaustion. 


Azolla caroliniana Willd. in Georgia 
Witsur H. Duncan 


Azolla caroliniana Willd. was reported by MeVaugh and 
Pyron' from Camden, Chatham, Echols, and Liberty Counties. 


barium add stations in Decatur (R. F. Thorne 17105) and Me- 
Intosh (Duncan 19969 from Sapelo Island) Counties. These and 
the previous collections seem to indicate that this species occurs 
only in the counties along the southern boundary and the At- 
lantie Ocean. 

It was, therefore, of considerable interest to find in the July- 
September 1958 issue of the American Fern Journal a statement 
that the original supply of Azolla used in physiological studies 
was collected in northern Georgia by R. A. Benedict.2 I wrote to 

1 MeVaugh, Rogers and J oseph Pyron. 1951. Ferns of Georgia. University 
of Georgia Press, Athens, Geor orgia. 195 


ickell, Louis G. 1958. Physiological Studies with Azolla under Aseptic 
Conditions. Amer. Fern Journ. 48: 


LYCOPODIACEAE AND SELAGINELLACEAE OF [OWA 267 


Dr. Nickell to obtain more precise information concerning the 
locality where the Azolla was obtained. He kindly sent dried 
specimens from the culture and stated (letter of 26 November 
1958) that the original material was obtained from a roadside 
ditch near the main highway just before crossing the state line 
leaving Georgia. From additional correspondence it was learned 
that Mr. Benedict, to the best of his recollection (letter of 27 
January 1959 from Dr. Nickell), obtained the Azolla about 20 
miles before leaving Georgia on route #301. Examination of a 
highway map determines that the station in question is probably 
in Screven County of the central Coastal Plain. 
or the present, therefore, it seems that this Azolla is not 
known from northernmost Georgia. Students should look for 
this species in northwestern Georgia, however, for it is reported 
by Shaver? from an adjacent county (Bradley) in Tennessee. 
DEPARTMENT oF Botany, UNIVERSITY OF GEORGIA, ATHENS, 
GEORGIA. 


3 Shaver, Jesse M. 1954. Ferns of Tennessee. Nashville, Tennessee. 502 pp. 


The Lycopodiaceae and Selaginellaceae of Iowa 
Tom S. CoOPERRIDER 

Until recently only four species of the Lycopodiaceae and one 
of the Selaginellaceae were known to be a part of the Iowa flora. 
None of these species is common in the state, but each has been 
collected at least once in the last decade. Lycopodium clavatum 
L. var. clavatum is known from three woodland stations in John- 
son County, in the east-central part of the state.1 L. complana- 
tum L. var. flabelliforme Fern. has been collected from nine 
woodland stations in eastern Iowa. L. Iucidulum Michx. has been 
collected from thirteen stations, most of them in wooded ravines 
and slopes in northeastern Iowa. L. obsewrwm L. var. dendroi- 
1 Statements oe ie distribution are based on specimens in the follow- 
ing Iowa herbaria: Davenport Public Museum, Grinnell College, Iowa State 
College, Iowa State Teachers College, and the State University of Towa. 
Thanks are extended to their curators, Dr. L. F. Guldner, Dr. N. H. Russell, 
rT RoW. Pohl Dro M. Lb: Grant, and Dr. R. F. Thorne, respectively, for 
permission to study these specimens. 


268 


AMERICAN FERN JOURNAL 


dewm (Michx.) D. C. Eaton, is known but from a single station 
in White Pine Hollow State Forest, Dubuque County, in the 
northeastern part of the state. There, a vigorous colony grows 
among the branches of a dense stand of Canada yew on a cool, 
north-facing, limestone-talus slope. 


: LYON OSCEOLA 


DICKIN 
SON 


SIOUX OBRIEN 


CLAY 


PALO 
ALTO 


KOSSUTH] WINNE- | WORTH | MITCH 


BAGO ELL SHIEK MaKe 


FLOYD | CHICK 
anon ASAW 


FAYETTE |CLAY 


PLYMOUTH | CHERO- 
KEE 


BUENA 
VISTA 


POCA- 
HONTAS 


HUM- WRIGHT |FRANK-| BUTLER |BREMER 
BOLOT LIN 


EBSTER 


BLACK | BUCH- AWARE 
ANAN 


wee 


SAC 


CAL 
HOUN 


| 


TON | 


| 
TAMA NTON 


LINN JONES [JACK 


= 


CRAWFORD 


CARROLL | GREENE 


BOONE 
CLINTON 


bole 
vis 


CEDAR 


SCOTT 


JASPER Powe- | iowa 
SHIEK | 


TINE 


x 
SA 


e 


MONT- 
|GOMERY 


ADAMS 


UNION |CLARKE | LUCAS |MONROE ge apes 
| 


HENRY 
| ‘MO! 


55 


cases 


} 
VAN. 


DAVIS 
BUREN |LEE 


WAYNE os 


RING-~ SCATUR 
GOLD 


Map oF ees 


ATEN 


oe InpicatTe Stations FoR LycopopIUM SELAGO VAR. 
PODA 


, Cross STaTION FoR SELAGINELLA AP 


Collections of Selaginella rupestris (L.) Spring have been 
made from twelve Iowa stations. Most of these are located in 
the northeastern part of the state. The plants grow on open 
fields of sand, in the crevices of sandstone exposures, and on 
talus from sandstone and quartzite exposures. 

One more species of each of these families may now be re- 
ported. The writer recently re-examined the specimens identi- 
fied as Lycopodium lucidulum in the herbaria of Iowa State 
College and the State University of Iowa. Two specimens from 
the former herbarium and four from the latter were found to 


_ be L. selago L. var. patens (Beauv.) Desv. Data on the labels 


indicate that these were collected from five different stations in 


PsmorumM IN LOUISIANA 269 


the extreme northeastern corner of the state, usually on or near 
sandstone bluffs or ledges. The most recent of these collections 
was made in 1931. 

The first known Iowa collection of Selaginella apoda (L.) 
Spring was made by R. F. Thorne and R. L. Hulbary, of the 
State University of Iowa, in September, 1958. An abundant 
stand of the species was discovered in a seepage bog at the foot 
of a sandy bluff along the Cedar River in Muscatine County. 

The distribution of these two additions to the known Iowa 
flora is shown on the accompanying map. 

DEPARTMENT orf Brotogy, Kent Strate University, Kent, 

10 


Psilotum in Louisiana 
WiuuiAm D. REESE 

Recently, after hearing Mr. John Lynch, of Lafayette, de- 
seribe the area in southeastern Louisiana in which he gathers 
the fibrous bases of Osmunda regalis for use in his orchid houses, 
it occurred to me that the habitat he described must be very 
much like the ones in which I had seen Psilotum nudum in Flor- 
ida. Subsequently, I took advantage of an offer to accompany 
Mr. Lynch and his sons on an Osmunda-gathering expedition to 
the area in question and did indeed discover Psilotum in the 
cypress swamps, at two stations approximately eight miles apart. 
The plants, although mostly rather small, were fairly vigorous 
and healthy in appearance, but all were sterile. They were not 
of general occurrence in the swamps but were locally abundant 
on mounds of pure humus which, emergent about 6 to 8 inches 
above the surface of the water, were well interlaced with roots 
of other species of plants. Growing with the Psilotum were a 
few plants of fruiting Botrychium dissectum var. obliquum 
and the mosses Pa hes texanus, Leucobryum albidum, and 
Climacium kindber: 

The collections, Aah of which are deposited in the 
United States National Museum, Univ. of Southwestern Louisi- 
ana, Louisiana State University, Florida State University, and 


270 AMERICAN FERN JOURNAL 


Harvard University are the following: About 100 yards north 
of the Gibson-Schriever road, about 1.3 miles east of the junc- 
tion with U.S. Highway 90, Terrebonne Parish, on hummock of 
humus at base of gum tree in cypress swamp, about 8 inches 
above water, Jan. 23, 1960, Reese (with John, Semmes, and Dick 
Lynch) No. 2576. About 0.5 mile west of U.S. Highway 90 
bridge over Bayou Boeuf, about 0.25 miles north of the high- 
way, St. Mary Parish, on hummock of humus in cypress swamp, 
about 6 to 8 inches above water, Jan. 23, 1960, Reese (et al.) 
No. 2577 

Dr. Clair Brown, of Louisiana State University, has informed 
me of one other record of Psilotum from the wild in Louisiana. 
According to Dr. Brown the name of the finder is unknown, and 
no specimen was preserved. The plants were found in the vicin- 
ity of Schriever, in the same general area dealt with in this note. 


_ Psilotum is otherwise known from Louisiana from specimens 


found at a nursery near New Orleans, and from Lynch’s orchid 
houses in Lafayette, where it was recognized by Mr. Lynch after 
being found in the swamps. Doubtless it was brought in initially 
with the Osmunda bases and very likely is of rather wide occur- 
rence in the state in greenhouses. 

University or SouTHWESTERN LOUISIANA, LAFAYETTE, LOUISIANA. 


A Key to American Dryopteris whey Based 
on Characters of the Peri 
FERN WarD Ga 
In 1954, Dr. Edgar T. Wherry collected a number of Dryop- 
teris plants to be used specifically for cytological and palynologi- 
cal studies. Among the rhizomes sent to Dr. Stanley Walker,” 
University of Liverpool, there were some D. X Leedsii from the 
type locality in Maryland.’ It was surprising to learn from him 


1Presented at the Ninth International Botanical Congress, Montreal, 
August, 1959 
2Wa Iker, z Cytotaxonomie Peisiges of Some American Species of Dryop- 
teris. This JOURNAL 49: 104-1 
ates Thomas Jr. Heit Fiala Notes. This JourNAL 49: 117-122. 
1959. 


’ 


Key To AMERICAN DRYOPTERIS 241. 


that only half of these plants were diploid hybrids, the re- 
mainder being fertile tetraploids, but an examination of the 
spores confirmed this report. In another set of these ferns, col- 
lected in 1956, it was a simple matter to predict the results of 
chromosome counts when palynological evidence was considered. 
This tetraploid Dryopteris is the new species described below: 
Dryropteris Wherryi F. W. Crane, sp. nov 


fo les ca. 120-140 em. longa, eoubas: 30-40 em. longis, 
diam., viridi-stramineis, basi compris paleis brun- 
neis translucide late marginatis dense obtec , eis sursum pe 


oribus et minoribus; laminae late Crearlaeeclatae 90-100 ¢ 
longae, 25-30 em. ‘latae, apice attenuatae, basi 

angustatae, pinnato-pinnatifidae, rhache colore stipitis, paleis 
capillifo nian us ; pinnae 16-2 -jugae, inferiores ee 


ta Q- ongae, 3-5 em. latae, we lanceolatae, ne 


triangulares, costam versus ampliatae, alas + distincte for- 

mantes, cppesitas vel suboppositae, regulariter serratae, dentibu 
acutis vel subacutis, in spinulos sensim flere coarctatis, 
supra olivaceo-virides, subtus griseo-virides; costa albido- 
virescens, + late alata, ealee filiformibus translucent hie 
inde obtecta, supra canaliculata; sori usque ad 8-jugi, evidenter 
inframediales, indusio lato glabrato obtecti; sporae magnae, 
nigrescenti-brunnea ae, 28 X 46p, perisporio excluso; perisporium 

glabrum, saepius alis latis instructum, hine inde alis angustiori- 
Bas interspersis, + continuis et qua de causa seulpturam for- 
mantibus. A Stanley Walker Universitatis Liverpoolensis mihi 
relatum filicem tetrapolideam esse, chromosomatibus normaliter 
conjungentibus. Orta, ut videtur, neuploations chromosomat- 
ium hybridae diploideae Dryopteridis X Leedsu. Detecta a 
Edgar T Wherry, cui de di icata. 

14 


greenish-straw co olor, t he bas onset cov ered W ith brown scale’ 


with wide tran et marg wardly the ie rap an 
rather fewer; blade eeadly near lanceolate, 90-100 em. long, 
25-30 em. wide, iba at ap eareely narr eae towar 


t 
base, pinnate-pinnatifid, baie hair. Tike scales; pinnae 16-20 
pairs, the lower elongate- petiolate, the upper "nearly sessile, 
alternate, the lower scarcely reduced, 12-18 em. long, 3-5 


AMERICAN FERN JOURNAL VOLUME 50, PLATE 23 


§ 7S + oT " > 
SPORES or Dryoprerts Species, FiaurE NUMBERS CORRESPONDING TO 
f 1G8 


Species NuMBERS IN Key; Figs. 6, 9 PREVIOUSLY UNPUBLISHED 


Key to AMERICAN DRYOPTERIS 273 


wide, elongate-lanceolate, slightly spaced, pinnatifid, the tip long 
acute-toothed, the basal and terminal pinnatifid-serrate ; pinnules 
2-3 em. long, 5-10 mm. wide, linear-lanceolate or narrowly tri- 
angular, enlarged toward costa, wing + distinctly developed, 
opposite or subopposite, uniformly serrate, the teeth acute or 
subacute, the spinules distinctly folded back, olive-green above, 
beneath gray-green; costa whitish-green, + broadly winged, 
covered with translucent filiform scales, grooved above; sori up 
to 8 pairs, manifestly inframedial, the indusium broad, glabrous; 
spores large, dark brown, 28» X 46, excluding the perispore; 
perispore glabrous, usually furnished with wide wings inter- 
spersed with narrow wings, + continuous so as to appear sculp- 
tured. 


SS ae 
: 30n 
Fig. 6, DRYOpTERIS WHERRYI. Figs. 14, la. DRYOPTERIS X LEEDSII; TYPICAL 
Hysrip Spores, UsuALLY A Few LARGE AND MISSHAPEN, THE Masority 
SMALL AND ABORTED 

Houorypus in the United States National Herbarium, nos. 2, 
258,784 and 2,258,785, collected by Edgar T. Wherry, August 
15, 1956, 3 miles below Conowingo Dam, Harford County, Mary- 
land. (Herbarium F. W. Crane, no. 5615). Isotypus: Herbarium 
of the University of Pennsylvania. 


274 AMERICAN FERN JOURNAL 


This fern may be distinguished from D. Leedsw by the 
seales extending in abundance only to the mid-stipe, the lower 
segments of sterile pinnae little-spaced and the gap-width less 
than half the segment-width, the sori tending to lie nearer the 
midrib than the margin, and the spores being normal. Dr. Stan- 
ley Walker, University of Liverpool, reported to the author that 
the fern is tetraploid, the chromosomes pairing normally, and 
that apparently it haa arisen through doubling of the chromo- 
somes of the hybrid diploid, Dryopteris < Leedsii Wherry. It 
was discovered by Dr. Edgar T. Wherry, to whom it is dedi- 
cated. 

Dryopteris Wherryi was illustrated by Dr. S. Walker recently 
as Dryopteris ‘‘Leedsii’’ (tetraploid) ; it is the left hand figure 
of Plate 11 of his article,t which by a printer’s error is marked 
“Wie. 10 (left) : Dryopteris celsa’’ but which is actually fig. 8, 
D. ‘‘Leedsii’’ (tetraploid), the right hand frond being the true 
D. Leedsii (diploid). The correction is made in the Errata.® 

The author wishes to thank Dr. E. T. Wherry and Dr. 8S. 
Walker for their cooperation, and Dr. H. P. Fuchs for the 
Latin version of the description. 


: EY 
A. Spinules abse 
I. Spores or 22-2 os x 32-37. 
Wines RIG, WEG cs eae . Goldiana 
ings narrow, more or less continuous so as to produce a sculptured 
appearance 
Wings hi aoateebl us Z craic’? 


ings few, some wide ones interspersed 
Wings numerous, the appearance tuberculate 
Wings mostly wide, though with some smaller ones, continuous as in 
li. 


4. Filix-mas 
. fragrans 


marginalis 6. Wherryt 
B. Spinules pres 
I. Spores onal a x 36-41p. 


” 4Walker, 8. Cytotaxonomie Studies of Some American Species of Dryop- 
teris. This OURNAL 49: 104-112. 1959, 
5This Journal, 49: 160. 


Taxonomic NOTES ON FERNS 275 


oe few, simple. 
ings narrow, set with small sharp-tipped spinules —_- 7. intermedia 
Wings wide, with a few large blunt-tipped spinules 8. celsa 
Wings more numerous, wide, simple; smaller wings also present; 
Bpmuses Ditiit-tep ped es ee ee Tad onibance 


Wings fairly wide; spinules mostly large, decidedly blunt-tipped 
A ie eS 10. spinulosa 
Wings wider; spinules mostly small, sharp-tipped_...11. ‘‘dilatata’’ 
Size large, 31-334 x 48-52 


Wings numerous, small, reunited: spinules many, agi ac hee 


ea ee ee ee cristata 
Wings fewer, wide, characteristically angular; spines widely 
aced, plant: tipped, Clintoniana 


sp 
174 Summit Avenue, Summit, New Jersey. 


6Proper name for eastern North American plant uncertain. 


Taxonomic Notes on Ferns, I 
. V. Morton 
Arayrium Lilloi (Hicken) Morton, comb. 

Nephrodium Lilloi Hiecken, Anal. Soe. Cient, Argent. 63: 8. 
1907. Type: La Casita, Valle del Rio Canasor- 
cona, Province of Tucuman, Argentina, 1700 meters alti- 

tude, Jan. 26, 1903, M. Lillo 2982. 
Drypopteris Lilloi Hicken, Apuntes Hist. Nat. Buenos Aires 

1: 151. 1909 

This species has never been placed. In Christensen’s ‘‘A 
Monograph of the Genus Dryopteris’’! it is listed as dubious, 
said to be probably a valid species allied to Dryopteris connexa, 
following Hicken’s original comparison. Hicken himself men- 
tioned in his comments that the indusium appeared to be lateral, 
recalling that of Asplenium or Athyrium. 

Material from Tucuman in the U. S. National Herbarium 
which agrees entirely with the original description and with the 
illustration (both the drawing and the photograph of the type) 
shows that this species really is an Athyrium, and not a Nephro- 


1Part 2: 125. 1920. 


276 AMERICAN FERN JOURNAL 


dium (or Dryopteris). The indusium is elongate and sometimes 
bent, where it runs past a forking of the vein on which it is 
borne, as is general in Athyrium. It is extremely broad and 
thin, and completely covers the sporangia when young: The sori 
are then ‘‘allantodioid’’ (sausage-shaped). The species thus 
belongs to Sect. Kee and is allied to Athyrium umbro- 
sum (Aiton) Presl, adei 


@Q 
me 
fort 
& 
=) 
Qu 
b 
= 
~~ 
=> 
< 
>. 
= 
= 
8 
Ss 
wH 
~ 
Bs 
i] 
~~ 
ic 
= 
=e) 


Allantodia R. Brown, which was characterized largely on the 
shape of the indusia. The section forms a link between Athyrium 
and Diplazium, and its affinities need to be investigated thor- 
oughly. The peculiar distribution (Atlantic Islands, Australia, 
and southern South America) suggests an ancient origin; it is 
not exactly paralleled among ferns, although the distributions 
of the genera Pleuwrosorus and Culcita are somewhat similar. 


Cytological Observations a8 the Himalayan Species of 
Athyrium and cee on mad opto tionary 
Status of he Gen 
P. N. Mreura anp §. . Bir 

Athyrium Roth is one of the most widely distributed ferns 
in the Himalaya Mountains, where about 30 species occur,! out 
of a total of 180 species in the genus.2. This number is bound to 
increase as the taxonomy of these ferns is better understood. 
_ Conspicuous disagreement among various seid ae exists re- 
- garding the systematic position of Athy , as a survey of the 
various systems of classification sel e since the beginning 
of the present century reveals.? Recently, Copeland and Holt- 
tum have followed Milde* in uniting Diplaziwm Swartz with 
Athyrium, which has in fact added to the existing confusion, 
for Athyriwm so construed (and also including Deparia, Cornop- 
teris, and other segregates) is then a genus of about 600 species. 

a poe for the Fiftieth Anniversary Volume of the Journal. [Ed. ] 

1Clar ; Beddome, 1892; Hope, 1899-1904. 

20ning, 

okt aii 1906, 1938; Bower, gee 8 1940; Dickason, 1946; 


Holttum, 1949; Copeland, 1947; Alston, 
4Bot. Zeit. 1866: 373. 


HIMALAYAN ATHYRIUM : Qt 


In the present paper the genus is treated in the traditional sense 
as proposed by Roth® and followed by Diels, Christensen, Ching, 
Dickason, and others. 

Some species of Athyrium are difficult to separate morphologi- 


of cytology is significant here. Cytological information is avail- 
able regarding 22 clear-cut species, of which eleven are from 
the Himalayas,® five from Ceylon,’ three from Europe,* two 
from North America,® and one from South India.4° Chromosome 
counts show that all are based on x = 40. Thus on cytological 
grounds Athyrium differs consistently from Diplazium, which 
has 41 as a base number;" this fully justifies the retention of 
Diplazium as a genus distinct from Athyriwm. 

The present paper deals with cytological observations on 16 
previously unstudied species of Athyriwm, and with the evolu- 
tionary status of the genus 

MATERIALS AND METHODS 

Fourteen species have been studied from the Eastern Himala- 
yas, all from Sikkim State except A. macrocarpum (Blume) 
Beddome (from Darjeeling, 6,000 feet elevation’) ; the other 
two species are from the Western Himalayas. Material of 
Athyrium pectinatum (Wall.) Presl from both regions has been 
studied. The species show an altitudinal range from 3,000 to 
14,000 feet. Two more species, A. Schimperi and A. thelypteri- 
oides that have already been studied from material from Darjeel- 
ing and Mussoorie respectively (Mehra and Verma, 1957) have 
been reinvestigated from cytogeographical considerations. 

The material of four species came from the Western Himala- 
yas. Two of these, A. thelypterioides and A. dentigerum 


5 Mag. 21: 105. 1799 
Mehra ee Wage) 1957; 
TManton, 1 ; Manton aa ge 1954. 


*Britton, 195 3; Wiener 1955. 
au Mahabale i ‘al 

11Manton, es and oe 1954; Brownlie, 1958; Bir, 1959. 
Seedentica by Mr. S. C. 


AMERICAN FERN JOURNAL VOLUME 50, PLATE 24 


_ UATUM, DipL., n = 40, * 500. Fic. 2, Ipp., 
TETRAPL., 2 80 )0. Fic. 3, Inmp., Spores X 150 (a, Dipi.; b, 


Fig. 1, ATHYRIUM ATTEN 
me SO Ne. SI 


RAPL.). Fig. 7, DIAGRAM FROM Fig. 4. Fig. 8, D1aGRAM FROM Fig. 6. 
Fig. 9, Spore Moruer Cent, A. MEHRAR, 40 BIVALENTS, LATE DIAKINESIS, 
« 500. 1 id 80) 5 


Fig. 10, A. SPINULOSUM, n’ = 


. 
HIMALAYAN ATHYRIUM 279 


(Clarke) Mehra et Bir,!* were collected near Dhanolti, along 
the Mussoorie-Tehri Road, at an altitude of 7,000 feet. Athyrium 
pe, is a rather rare fern in this area and only a 
w plants were seen, growing in a clump; it is never so abun- 
aint here as in the Eastern Himalayas between 8,000 and 13,000 
feet altitude. Athyrium falcatum Beddome is extremely rare 
at Nainital (7,000 feet). Athyrium pectinatum (Wall.) Presl 
is a comparatively low altitude fern; it is abundant near Mossy 
Falls (5,000 feet) and Sainji (3,000 feet) on the Mussoorie- 
Chakrata Road, where it forms extensive beds because of its 
ereeping and branched rhizomes. Two individuals of another 
Athyrium were found near Magra (6,000 feet), on the way to 
Nag Tiba (Mussoorie) that are morphologically similar to A. 
pectinatum; they are suspected to be of hybrid origin, since 
they are cytologically abnormal, and are here designated A. X 
pectinatum 
The most beautiful of the Sikkim species is A. Tsavi Ching, 
which covers vast areas in the Lachen Valley at elevations be- 
tween 10,000 and 12,000 feet, especially near Simdong (11,000 
feet); the fronds and stipes are characteristically yellowish in 


colour. Another fern having almost the same distribution is - 


A. attenuatum (Clarke) Tagawa. Two closely allied species 
with creeping rhizomes are abundant around Thangu (13,000 
feet), namely A. subtriangulare (Hook.) Beddome, with a con- 
spicuously yellowish lamina, and A. spinulosum (Maxim. ) 
Milde, with chaffy and markedly pinkish stipes. Still higher 
up (14,000 feet), on the hill at the back of Thangu Dak Bunga- 
low, A. subtriangulare var. sikkimense Bir grows occasionally 
under rhododendron trees in rather exposed situations. One of 
the rarest species is A. Mehrae Bir, which has very fragile 


feet elevation, but a thorough search of the ‘area may possibly 


WS eehser M dentigerum (C. B. vont Fghenis et Bir, comb. nov 
Asplenium filix-foemina var. dentigera C. B. Clar ke, Trans. Linn, Soe. 
London, IT, Bot. 1: 491. 1880 


280 AMERICAN FERN JOURNAL 


reveal more individuals. The other species—A. rupicola (Hope) 
C. Chr., A. parasnathense (Clarke) Ching, A. himalaicum 
Ching, A. Biri Ching, A. rubricaule (Edgw.) Bir, A. poly- 
sporum (Clarke) Ching, and A. aff. flabellulatum (Clarke) 
Tardieu—grow at comparatively low et hotes around Lachen, 
8,000-9,000 feet altitude. In eastern Sikkim, A. rupicola at- 
tains much higher elevations, and some specimens were collected 
at Changu (about 13,800 feet). Athyrium Schimperi and A. 
pectinatum grow at still lower elevations than the other species 
from Sikkim reported here; they are common along the road 
between Chungthang and taoha at about 6,000 feet. Full de- 
scriptions have been published only for A. spinulosum, A. 
Schimperi, A. pectinatum, A. faleatum and A. macrocarpum,4 
A. atienuatum,” A. subtriangulare,’® and A. rupicola.™ The 
taxonomic observations and complete descriptions of the others, 
which are either new species or varieties or new combinations, 
will be published separately. 

The material was collected in July and August, 1958 and 
1959, and fixed in 1:3 acetic alcohol and modified Carnoy’s 
Fluid (1 part glacial acetic acid, 3 parts absolute alcohol, and 
4 parts chloroform). Chromosome counts have been made from 
‘the spore mother cells entirely by the squash technique; the 
counts have been confirmed from a large number of cells in each 
case. All the photomicrographs are from permanent aceto- 
carmine preparations. Voucher specimens are preserved in the 
Panjab University Herbarium. 

BSERVATIONS 

The course of meiosis in all cases is perfectly normal, except 

as mentioned below in A. thelypterioides and A. x pectinatum. 

the species are sexual, since 64 normal and apparently 
viable spores were counted within a sporangium in each case 
except as noted below in A. thelypterioides, and apogamy has 
not been observed in any of them. Both perisporiate and 

Me rang 1892. 

Tagawa, 1956, p. 177. 


Ee oe and Baker, 1874, p; 225. 
\7Hope, 1899, pp. 531, 532. 


HIMALAYAN ATHYRIUM 281 


non-perisporiate spores (cf. Figs. 3 and 17d) are present. The 
spores are broadly perisporiate in eight Lei ae A. 
subtriangulare (and var. sikkimense), A. spmnulos . Mehrae, 
A. polysporum, A. dentigerum, A. faleatum, A. er and 
A. macrocarpum. The spores of A. subtriangulare (n = 80 

are small; those of the rest are of the same general size. The 


any clear-cut perisporium. The spores of A. rubricaule (n = 40) 
are the largest in this group. 

The cytological results are summarized in Table I.'* It is 
clear that 15 of these species are diploids, the haploid chromo- 
some number being 40, and that three are tetraploids (n — 80). 
In the Athyrium species worked out here, polyploidy has been 
noted only up to the tetraploid level. 

A previous report of the chromosome number of A. thelyp 
terioides was by Mehra and Verma’? who reported both diploid 
(n = 40) and tetraploid (n = 80) races. The authors have 
since studied several populations from the Lachen Valley, 
8,000-12,000 feet, in northern Sikkim, also, and noted only the 
diploid race. Further scrutiny has shown that the tetraploid 
individuals differ from the true A. thelypterioides in the follow- 
ing characters and probably represent a different species: 

1) The individuals are smaller. 

(2) The rhizome scales are smaller, and linear in outline, 
as compared with the larger, linear-lanceolate, broad-based 
scales of A. thelypterioides. 

(3) The complete absence of uniseriate fibrillar hairs on the 
primary and secondary rachises, which is an important charac- 


18The names of the species marked * and ** are entirely on the authority 
of Prof. ea C. Chin, 
19Mehra and Verma, 1957. The name here used, Athyrium acrostichoides 
aeatiay “Diels riseey. t is Pek wert being a later anges m of Athyriwm 
acrostichoideum Bory ex Mérat (1836). The Gr ges for ‘‘aerostichoides 
é atinized f 


r able 
‘pteroideus,’’ cited as examples of orthographic variants in the Inter- 
national Code of Botanical Nain enaiatune (Paris Edition, Art. 75, 1956). 


Fig. 4, A. DENTIGERUM, SPORE ang oo CELL, 40 BIVALENTS, X 600. 5, os 
SUBTRIANGULARE VAR, SIKKIMENSE, n = 40. & 600. 6, A. FALCATUM, ” oe 0, 
x 600. 11, A. Tsau, os RE Mo jaar CELL, 40 BIVALENTS, x cae 12, ve 


HIMALAYAN ATHYRIUM 283 


ter of the species. 

(4) The stipes and rachises are straw-coloured and almost 
naked, a few scales being present at the base only, in strong 
contrast to the dull-brown stipes, covered with hairs and scales, 
of A. thelypterioides. 

(5) The lower pinnae are never reduced to mere auricles as 
in A. thelypterioides. 

(6) The sori are typically ‘‘athyrioid’’ and not ‘‘diplazioid,’’ 
as in A. thelypterioides. 

(7) The spores are non-perisporiate rather than broadly 
perisporiate as in A. thelypterioides. 

Therefore, in the authors’ opinion the tetraploid individuals 
represent a different species; they are probably very near A. 
MacDonellit Beddome 

e diploid A. Shalypterioides is morphologically variable. 
The individuals collected from Tonglu (8,500 feet) and near 
Thangu (12,000 feet) in the Eastern Himalayas differ from 
each other in the size of the pinnae and segments, the extent 
of the marginal crenations, the texture of the blade, and the 
amount of hairs and scales. 

In the Mussoorie area the species is extremely rare and no 
sexual race has been found. All the individuals of A. thelyp- 
terioides growing near Dhanolti, 7,000 feet, Mussoorie, are 
cytologically abnormal. There are no signs of chromosome 
pairing in any of the spore mother cells, and at late diakinesis 
80 univalents were clearly seen (Fig. 12). The chromosomes 
are longish and mitotie in appearance, and show median or 
submedian constrictions; soon, however, they become contract 
and ovoi e further course of meiosis is highly a hae: 
The aekpheek plate is seldom properly organized, and the 
univalents lie scattered in the cell. At A-I, the univalents 
divide longitudinally and the auc chromatids reach either 
of the poles or may be left undivided and remain as laggards 
which are ultimately organized into micronuclei (Fig. 13a). 
The two nuclei resulting from the first meiotic division may 
be irregularly shaped and sometimes highly disproportionate, 


AMERICAN FERN JOURNAL VoLuME 50, PLATE 26 


HIMALAYAN ATHYRIUM 285 


as a consequence of the unequal distribution of the chromatin 
material (Fig. 13b). The homoeotypic division is also irregular. 

those ae that divided in the first division being 
oes to divide for the second time are left as laggards; at 
the ‘‘tetrad’’ stage numerous micronuclei are present (Fig. 
3c). The cytokinesis may be complete or incomp ae e 
young spores are of variable shapes and sizes and possess micro- 
nuclei (Fig. 13d). Obviously as a result of these rosea 
the mature spores are unequal in size, some even shrivelled up 
and abortive (Fig. 13e). 

Although the Dhanolti (Mussoorie) specimens just ae 
that show abnormal meiosis differ markedly from those 
Sikkim (diploid, n = 40) in the more elongate, deeply Bee 
faleate segments, with a much broader sinus between them, the 
fall within the range of variation noticed in the species in 
various regions of the Himalayas. The differences ee ned 
are probably of little taxonomic importance, and the ‘‘sterile”’ 
specimens — Dhanolti can hardly be segregated as a variety 

orm. almost complete similarity between the sexual 
and the ssebersle?? individuals of A. thelypterioides leads us to 
nies that the cytological abnormalities in the Mussoorie 
populations are not due to hybridity but are of genic origin. 
The individuals of the fern designated as A. X pectinatum 

in Table I possess characteristically pinkish stipes when fresh. 
They differ from specimens of A. pectinatum from Sikkim, 
Nainital, and Mussoorie BEE in only two characters: 
Lamina only bipinnate, with lower pinnae more reduced (tri- 
pinnate in A. pectinatum) (Figs. 17a, b), 20 and spores tuber- 
may be pointed out that these cytes exactly resemble the figure 
given by Beddome (1 sei t. 154), for A. Filix-foemina from southern Tae, 


which later on ve ddome, 1892) was ronal as a small form of 4. 
foemina var. pect se 


13, A. THELYPTERIOIDES, MEIosis; a, ANAPHASE I; b, TeLopHase I, 
UnrquaL Nucuet; ec, ‘‘TreTRap,’’ Many Mrcroxt CLEI; d, YOUNG SPORES, 
VARIABLE SizE, WITH MICRONUCLEI; e, M RE SPORES a-d 535, e xX - 
20. 15, / ECTINATUM, z i A <. PECTINATUM, 2n = 80, 
LE. 15 (II) anp = (1), < 800. 16, Ipm., Metosis; a, MrerapHasE I; 
b, ANAPHASE T; BInucLEATE Spore, MANY Micronvuciet; d, TETRAD, 
Wicsosecna: e, YouNG SPORES, Micronucnel; ALL X 540 


AMERICAN FERN JOURNAL VoLUME 50, PLATE 27 


ae isk (bu 


Fia. 17. a, b, PINNULES oF A. X PECTINATUM AND A. PECTINATUM, X 3. 
¢. d. SPORES OF A. X PECTINATUM AND A. PECTINATUM RESPECTIVELY, < 220. 
18. A. ATTENUATUM, Lower EpIperMIs WITH STOMATA, a, DrpLorp, b, TET- 


? 


HIMALAYAN ATHYRIUM 287 


culate-thickened on the surface (smooth perisporium in A. 
pectinatum) (Figs. 17c, d). In this fern the course of meiosis 
is highly irregular. In the majority of the cells only a partial 
pairing of the chromosomes is noticed at diakinesis. One such 
cell is reproduced in Fig. 15, which shows 15, + 50;, giving 2n 
= 80. M-I and A-I are also abnormal and laggards are common 
(Figs. 16a, b). Very rarely, the second division fails and a 
binucleate spore results (Fig. 16c). The four nuclei at the tetrad 
stage are often unequal and numerous micronuclei are organized 
(Fig. 16d). The cytokinesis is irregular, resulting in spores of 


ee they resemble those of Diplazium japonicum (Thunb.) 
Beddome, although morphologicially there is no similarity 
whatsoever between D. japonicum and A. < pectinatum. There- 
fore at present nothing can be inferred regarding the other 
parent of this sae: 

Only one species, A. attenuatum, shows intraspecific poly- 
ploidy. Diploid and tetraploid individuals grow side by side, 
but although the smallest individuals of the diploid race are 
shorter than the smallest tetraploids, morphologically the two 
races are similar in leaf outline; there are hardly any quantita- 
tive differences that could be detected, either in the size of the 
pinnae or their segmentation. The spores do show differences 
in size and in the quantity of stored food material, the diploids 
being smaller and less filled with food material than the tetra- : 
ploids (Figs. 3a, b). There is a little difference in stomatal sizes, 
the stomata of the tetraploids being slightly bigger. The number 
of stomata e Bae’ area in the tetraploid is lesser than in the 
diploid (Fig. 18a, b). The epidermal cells of the tetraploid are 
somewhat tae 

Another species that draws one’s attention is A. subtriangu- 
lare. The typical variety (n = 80), with fronds 50-70 em. long 
and 20-40 em. wide, grows at lower elevations than var. sik- 
kimense, which is diploid (n = 40), with fronds only up to 


\ 


288 AMERICAN FERN JOURNAL 


em. long and 12 em. wide. Curiously enough the variety, 
although only diploid, has spores somewhat larger than the 
ee variety, which may very well speak for its separate 
ident 

Shonomanor of apogamy, aside from the classical ex- 
aoe A. Filix-foemina var. ‘Clarissima Jones,’ var. ‘Claris- 
i and var. ‘Uncoglomeratum’ Stansfield,?! has not 


disproves Mahabale’s observation that most species of Athyrium 
are apogamous. Both the examples of apogamous Athyriums 
worked out by Manton (1953) from Ceylon, A. maximum and 
A. pinnatum, are Diplaziums as that genus is now understood 
with the present day availability of chromosomal information. 
The names of these ferns are Diplazium maximum (Don 
Chr. and D. silvaticwm (Bory) Swartz respectively. 
DISCUSSION 

A regional analysis of all the species thus far worked out is 
presented in Table II.?? For the purpose of calculations, species 
with two cytological races have been counted on the higher side 
of ploidy. This Table shows that three sexual species have been 
found with different cytological races—A. attenuatum (2x and 
4x, from the Himalayas), 4. gymnogrammoides (4x and 6x, 
from Ceylon), and A. macrocarpum (2x from the Himalayas, 
4x and 6x in Ceylon). No major morphologi 
evident in A. wp ant but the 4x and 6x individuals of A. 
markedly different ont each other, as 
has bean iihieteated Oy Manton and Sledge ;?* nothing can be 
said about A. macrocarpwm until a ihoroagh comparison of 
Himalayan and Ceylonese specimens is made. 


21Farmer and Digby, 1907. 

22The information Peinded { in this table and the subsequent ones is based 

n Manton, 1950, 1953, 1954; Britton, 1953; Manton and Sledge, 1954; 
Washo ie 1955; Mehra and Verma a, 1957; Brownlie, 1958; Bir, 1959; and 
the Present investigation. 

and Sledge, 1954. It may be mentioned that the taxonomie 

status ae er gymnogrammoider, which neon to a group of species allied 
to A. solenopter onfused, and it ean not be said —— the 4x and 
6x individuals fall within the Tonasation of ‘the species or 1 


TABLE I. 


Name Locality and altitude n chromosome Fig. Ploidy 
number no 
A m attenuatum (Clarke) Tagawa Simdeng, N. Sikkim, 11,000’ 40 1 loid 
Athyrium eames (Clarke) Tagawa Simdong, N. pty 11,000’ 80 2 Tetraploid 
*A. Birii Ching (mss.) Near Lachen, N, Sikkim, eae 40 Diploid 
A. dentigerum (Clack) Mehra et Bir Dhanolti, Mussoorie, 7,000 40 4,7 Diploid 
A. faleatum Beddome Nainital, 6,000’ 40 6,8 iploid 
A. aff, flabellulatum (Clarke) Tard. Lachen, N, Sikkim, 8,500’ 40 Diploid 
*A. himalaiewm Ching (mss.) Lachen, N. Si aan , 8,500" 40 Diploid 
A. macrocar Dey latina Beddome Sioa 7,00 40 Diploid 
A. Mehr Near Thangu, N. ‘skkim, tide 000’ 40 9 Diploid 
eae © pastures ie aso: Pi Lachen, N. Sikkim 40 Diploid 
A: a um ll.) Sainji, Mussoorie, pe - 40 14 Diploid 
A. pectinatum owas Boer Chungthang, N. Sikkim, 6,000’ 40 Diploid 
Ax een (Wall.) Presl Magra, Mussoorie, 6,000’ 15(11) + 50(1) 15 Diploid 
hybri 
**A. polysporum (Clarke) Ching Lachen, N. Sikkim, 8,500’ 80 Tetraploid 
A. rubricaule (Edgw.) Bir Lachen, N. Sikkim, 8,500’ 40 Diploid 
A. rupicola get! Cc. Ch Near Lachen, N. Sikkim, 8,500’ 40 iploid 
A. Sehimperi Mou Near Chungthang, N. Sikkim, 6,000’ 40 loid 
A, subtriangulare took. Beddom Thangu, N. Sikkim, 13,000 80 Tetraploid 
A. subtriangulare var, sikkimense Bir Above Thangu, N aes: ae 000" 40 5 Diploid 
A. spinulosum (M: ilde aN: Sikkim, 1 80 10 Tetraploid 
*A. Tsaii Ching Guay Simdong, N. Sikkim, ihe gee 40 11 Diploid 
New oe a eae by 8. S. Bir during the present investigation, as 
named by Prof. R. C. 


New Fas (Ching, ined.). 


WOIYAHLY NVAVIVNIF, 


296 AMERICAN FERN JOURNAL 


TABLE II. REGIONAL TABULATION OF DIPLOID, TETRAPLOID, AND HEXAPLOID 
ATHYRIUMS 


: rth America 
Himalayas Ceylon _ Europe (outae ote aaa 


Diploid 
thelypterioides Filia-foemina Fitia-foemina var. 
os eae alpestre auxti 
Atkinson ava flexile thelypterioides 
Ande saat pycnocarpon 


attenuatum, p.p. 


rie sum 


= Clarkei) 
py es raed var, 


Sp. "(ealled L Leucostegia 
yaklaensis Beddome) 


Tetraploid 
attenuatum, p.p. anisopterum 
anisopterum erythrorachis 
Pn sabre mnogrammoides 
spinulos macrocarpum, p.p. 
Webi sandalacd 

Hexaploid 


gymnogrammoides 
‘ocarpum, p.p. 


HIMALAYAN ATHYRIUM 291 


Only four instances where the same species occurs in different 
climates have been investigated. The temperate species A. thelyp- 
terioides (Himalayas and North America) and A. Filix-foemina 
(Europe and North America) show the same chromosome num- 
ber, n = 40. Athyrium anisopterum has n = 80 in both the 
Himalayas and Ceylon, but A. macrocarpum shows a higher 
ploidy in Ceylon, being diploid (n = 40) in the Himalayas and. 
tetraploid or hexaploid in Ceylon (n = 80 or ea. 120). 

Table III, which gives a comparison of cytological data from 
various regions, shows that the three European and three North 
American species are all diploid, but that no Ceylon species are 
diploid, two being hexaploid, two tetraploid, and one a hybrid. 


TaBLeE III. ReGgioNaAL COMPARISON OF CyToLogicAL Data 
No. Dipl. Tetrapl. Highest Hybr. Percent 
pol: of 


spec. dipl. 
__ Region — counted 
Himalayas 291 24 5 4x — 82.74 
Ceylon 5 a 2 Two 6x 1 
Europe 3 3 ae — aa 100 
No. Amer. 3 3 Sea — — 100 | 


14. x pectinatwm, from Mussoorie, not included. 
In the Himalayas of the 29 species that have been studied, 24 
are diploid (82.7%) and 5 are tetraploids. The highest grade of 
euploidy encountered in the Himalayas is only up to the tetra- 
ploid level (as compared with hexaploid in Ceylon), and the 
preponderance of diploids is significant. The Himalayas are 
similar to Europe and North America (Ontario and Michigan) 
in the temperate climate, whereas Ceylon has a tropical climate. 
Therefore, the above analysis of the situation in Athyrium ap- 
pears to give support to Manton’s (1953) observation that, 
‘Evolution is proceeding faster in the tropics than in temperate 
latitudes. ’’ 

Whether Athyrium has been placed in the subfamily Aspleni- 
oideae of the Polypodiaceae,2* or in the tribe Athyrieae of the 
Aspleniaceae”®, or the subfamily Athyrioideae of the Dennstaed- 


*4Christensen, 1906, 1938. 
25Ching, 1940; Dickason, 1946, 


Taste IV. Comparison or CyTotogicaAL DATA FoR ATHYRIUM WITH ALLIED GENERA 


No. of No. of pee Dipl. Tetrapl. Highest Hybr. Apogamy 
species species polypl. 
Genus counted 
Athyrium1 180 38 40 27 8 Two 6x 1 — 
Diplaziwm? 380 27 41 11 8 Two 6x One 5x Four 3x 
One 5x 
Diplaziopsis*® 2 z 41 1 - —_ = — 
Callipteris* 4 A 41 a -— — —_ = 
Cornopteris® 13 z 41 — 1 —_ _ a= 
Cystopteris 17 10 42 2 4 Two 6x One 3x t 


One 8x 


Percent 


22.25 


1Pata ean Manton, 1950, 1953; Mahabale et al., 1953; ae 1953; Manton & Sledge, 1954; Wagner, 


1955; a & Verma, 1957; Bir, 1959, and present investiga 
a aay 1954; Manton & Sledge, 1954; Brownlie, rie: st 1959, and Bir ined. 


3Bir, 1958. 
iach: 1954 (under name Athyriwm accedens). 
5According to Copeland, 1947. 


ir, ined. 
7Manton, 1950; Britton, 1953; Wagner, 1955; Bir, ined. 


G66 © 


TVNUNOP NYG NVOINAWV 


HIMALAYAN ATHYRIUM 293 


tiaceae,?° or in the family Aspidiaceae,”’ or recently in a separate 
family Athyriaceae,2* it has usually been grouped with Cystop- 
teris Bernh., Diplaziopsis C. Chr. Diplazium Swartz, Cornopteris 
Nakai, and Deparia Hook. & Grey. Kept separate by Ching, 
Diplazium, Cornopteris, and Deparia, were merged with Athy- 
rium by Copeland, who on the other hand maintained Bory’s 
genus Callipteris, which is not done by other authors. Calliptertis 
was based on C. prolifera (Lam.) Bory, which is Diplazium pro- 
liferum (Lam.) Thouars [Athyriwm accedens (Blume) Milde, 
of Holttum’s treatment2®]. The cytological data on these genera, 
which for the purposes of this discussion are treated separately, 
is presented in Table IV. The number of species referred to each 
genus follows Ching’s estimate,®° except for Callipteris. 

The data in Table IV show that the basic chromosome num- 
bers of these genera are 40, 41, or 42. The larger numbers may 
have evolved through aneuploidy. Deparia, a monotypic genus 
endemic to Hawaii, has not been studied yet. Diplazium seems 
to be actively evolving compared to Athyrium, because of the 
comparatively lower percentage of diploids, the greater number 
of hybrids and the consequent establishment of apogamy. The 
few hybrids recorded, the higher percentage of diploids, and the 
almost total absence of apogamy reveals that from an evolution- 
ary viewpoint Athyrium is a static genus in this respect, and 
that evolution in it is principally by genic mutations. 


SUMMARY 

Chromosome counts of 16 species of Athyrium from the Hima- 
laya Mountains show that 13 species are diploid (n = 40), and 
3 tetraploid (n — 80). All are sexual, apogamy being absent. 
Only one species, A. attenuatum, shows intraspecific polyploidy 
(diploid and tetraploid races). The counts of 29 species of the 
genus known from the Himalayas show that 82.74% are diploids 
and only 17.26% tetraploids, a significant abundance of diploids 


26Holttum, 1949, 1954. 
27Copeland, 1947. 
28Alston, 1956. 
2°Holttum, 1954. 
80°Ching, 1940, 


5 


294 AMERICAN FERN JOURNAL 


compared with data from Ceylon. From comparisons with allied 
genera, especially Diplaziwm, it is abundantly clear that Athy- 
rium is a statie genus as far as evolution by hybridization and 
polyploidy is concerned. 


ACKNOWLEDGMENTS 
The authors express their grateful thanks to Prof. R. C. Ching, 
of the Academia Sinica, Peking, for the identifications of many 
of the species. They are also thankful to Mr. R. 8. Pathania for 
the photomicrographs and to Mr. B. Khanna for help with the 
drawings. 
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ALSTON, in H. G. 1956. The Sub-division of the Polypodiaceae. Taxon 
5: 23-1 


BEDDOME, a H. 1863. The Ferns of Southern India. Gantz Brothers, 
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—. 1892. andbook to the Ferns of British India, Ceylon, and the 
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—.. 1959. ptctssonsite Notes on Some Aspleniaceae from ee Valley. 
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Bower, F. O. 1928. The Ferns. Volume III. Cambridge University 
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Burros, ae ie 1953. Chromosome Studies on Ferns. Amer. Journ. Bot. 
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Cuine, R. C. 1940. On the Natural Classification of the Family Poly- 
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CHRISTENSEN, C. 1906. Index Filicum. Copenha 
1938. Filicinae, in Verdoorn, Manual of Peadeboy. pp. 522-556. 
The ue. 
CLARKE, a By 1880.4 ibe of the Ferns of Northern India. Trans. 
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CopeLANnD, E. B. 1947. on Filicum. Waltham, neon U. S. 
946. A Piviosanstie Study of the Ferns of Burma, 
and The Ferns of Burma. Ohio Journ. Sci. 46: 73-141. 
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a 
3 
hej 
ao 
i] 
B 
a 
b& 
3S 
= 
S) 
AE 


. 200. 
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Notes 4ND NEWS 295 
Hooker, W. J. and ery PG piesa bay 58 safeecaineal Filicum, ed. 2. London. 


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ie es et al. 1953. Life History of Athyrium Hohenackerianum Bedd. 
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Manton, I. 1950. Pribions: of hinted and Evolution in the Pterido- 
phyta. oe University Pre 

—. 1953. The Cytological Brotution of the Fern Flora of Ceylon. Sym- 
posia Soc, ak Biol. No. 7, Evolution. pp. 174-185. 
. 1954. Cytological NE n One Hundred Species of Malayan Ferns. 
Appendix to oe E. Holttum, se of Malaya, Vol. II. Ferns. Singapore. 

A 54. 


Meura, P. N. and Vea A ©. 4957. Tite abies . Some Athyrium 
Species from Northern India. Ann. Bot., , 21: 455-459. 
AGAWA, M. 1956. sian Notes on ponee Poco 7. Acta 
Phytotax. Geobot. 16: 174-178. 

Waenerr, W. H., Jr. Tee Cytotaxonomic Observations on North Ameri- 

ean Ferns. Rkcdor ra 57: 219-240. 

Botany DrePAaRTMENT, PaNngaB University, AMRITSAR, INDIA. 


Notes and News 


Fern PHorocrapHs.—An article entitled ‘‘A Technique for — 
Close-Up pipe Bae of Freshly-Collected Fertile Fern Speci- 
C 


American Fern Journal has published other examples of Mr. 
Neidorf’s work. The photographs are notable for the amount 
of detail (some of it inevitably lost in the reproductions) re- 
vealed in the shape, arrangement and structure of the sori, 
indusia, sporangia, glands, scales, ete., details which are charac- 
teristic of the various species. Many of the photographs are 
quite striking in appearance and would be suitable for framing 


296 AMERICAN FERN JOURNAL 


or would serve as useful adjuncts to herbarium specimens. 

In addition to the article Turtox News includes a notice that 
sets of Mr. Neidorf’s fern photographs are available for sale 
exclusively from Turtox Products. Initially, a set of 12 prints 
of familiar northeastern species is being offered. The prints are 


8 inches by 10 inches in size, mounted on mats suitable for wall 


A ee et eS a 


display or framing, priced at $36.00 per cet. It should be noted 
that while only twelve prints are being offered at this time Mr. 
Neidorf has photographed some 68 species of ferns and fern 
allies, as part of a project which is still in progress. Inquiries 
should be addressed to Mrs. Ruth L. Shepherd, editor of Turtox 
News, General Biological Supply House, Inc., 8200 South Hoyne 


es Avenue, Chicago 20, Illinois. 


1This JOURNAL, 37: 74. 1947; 43: 5, 59. 1953; 44: 17, 67, 112. 1954; 
45: 18. 1955; 46: 29. 1956. 


American Fern Society 
New MEMBERS 
Mr. Merle Brush, Chelsea, 
_ Mr. Don S. Caldwi fee a5 - mere and Harman Attorneys, 112 West Fifth 
Street, Odessa, 
Mr. Lynn H. onde ests Lee Avenue, Downers Grove, Illinois 
Mrs. Henry Enoch, 127 East 55th Street, New York 22, 
Mrs. Effie D. Famalett, 2 Cleveland Place, Glen Cove, i: 9, New York 


Mrs. A. E, Holman, San Juan Darlington, Santuree, Puerto 
n 58, 


Miss Maria Teresa Murillo P., Carrera 5a-A, no. 27- uates Colombia 
} 109 Armagh Drive, Baltimore 12, la: 
Mr Set eat Oliver, 74 Charter Road, Acton, Massachusetts 


s. James A. Stewart, 118 Bryant Avenue, Franklin, Ohio 


CHANGES OF ADDRESS 
Mr. Sarmukh Singh Bir, Punjab University Botany Department, Khalsa 
‘ollege, Amritsar, India 
r. M. G. Degra' oe Bulls Eye Lake Road, Valparaiso, India: 
Miss E. Irene Graves, 237 Summer Street, Bridgewater, Massachusetts 


levard, H. , Calif. 
_— Mrs. Elizabeth Valoutine P.O, Box 186, Furlong, Bucks Country: Penna. 


INDEX TO VOLUME 50 297 


Index to Volume 50 


Abnormal Nuclear Division in Fern Pro- 


119; 
dium, 121; deflectens, 49; S ipitatin: 
54, 55; fructuosum, 51; fulvum, te 
tance Ett seein deg ae Pes 
513 v0: 


48, 51; Praca, "18, "4 63, 64, 75, 
217, Peat 5 aleutic m, 42; peruv 
num, 51; petiolatum, 51; platyphyl 
lum yi ieee 54, 70, v. hirsutum, 
55, v. sulphureum, 55; pulverulentum, 
51; dap vorbis: 55; terminatum, 51; 


tetraphyllum, 51: tomentosum, 51; 
Tracyi, 18, 159; eeahery ma kG 

Aglaomorpha Meyen 6: 

pe ‘topteris, 158, 159: eretacea, 22, 159 
lanto 

Ageovin” iechypievun: 

se, phila elongata, 48; 

icrodonta, 48 

Agastereytere Dictyopteris, 262 

Anemia, 79; anthriscifolia, 165, 166; 
phyllitidis, 79, 80 


23 
excelsa, 80, 81; 


Angiopteris 7, 251; evecta, 8, 127 
Anisogoni ; it 4 
pte ig leptophylla, 55 
ers, 


‘Aotouny um, 112; Mannianum, 


Archangiopteris, 251, 252; Sinema: 
251; ee 251% Somai, 252; 
tonkinensis, 251 

Arthropt - macrocarpa, 83; orientalis, 
1 


ie 
e 146 47; pacha bin’ 153: ” 
coniifolium, 53; riaceum, 147; 
cristatum, ‘os, v. eeviana: 105; dis- 
ans, 133— ie = 152; i lepido- 
caulon, 350 ; nitu nudatum, 24; 
muticum, 15 ‘: alescaut ce 


133: 
ee ae ‘Sieboldii, 98; speciosu 
45 nulosum ulig Shr 39, 
08, i 06: aHnionueh: 06; yunna- 
98, 04 
Aspidotis, 158; californi re, 19 
Aspleniopsis decipiens, 

a creed 75, ba mh 494, 202. 275; 
LL 138, 166, x 
x Paget 


s, 140. 
40; africanum, 230; auriculatum, 48; 
Barteri, 229; Baumgartneri, 139, 
142, 143; X bavarieum, 139, 142; 
borneense. ; castaneum 0; Ceci 
31; X Clermontae, 139; commixtum, 
3 Costei, 141; corbariense, 


140; X dolosum, 143; ebenoides, 1 

exiguum, 70, 163, 166; er oteows, ace. 

263; Filix- femina v. dentiger: 279: 

fissum, 138; flabellifolium, 562° acci- 

dum, 259, 261 263; fontanum, 8 
x 


x jos atic 
G) 


20, f. Hortonae, 34; X Poscharskya- 

num, 140, 2; Pringlei, 70; radicans, 

9; X refractum, 141; ns, 63, 

70; rhizophyllum, 12; Ruta-muraria, 
te: 


xX suevicum, 120, 
1 


265, x Phyllitis ” Scolopendrium, 143, 
x Trichomanes, 139, v. vespertinum 

18, X viride, 139; X trichoma niforme, 
140: viride, 20, 139, 142, 220; ves- 
are inum, = x ow wachaviense, 141; 4 


4 
ge tala Pach baden 141, 144 
x ee ae ecniiiens: 143; Jack- 
143 icrodon, Z 


1 
282, 289, 290; Filix-femina, 43, 66, 
202, 217, 220, 285, 288, 290, 291, v. 
A — = 70, v. Michauxii, 290, v 


Aone 
Habel m, 280, 28 
foliolosum, 29) 


anogram- 


2 


298 


moides, 288, 290; himalaicum, 280, 
281, 289, 290; Lilloi, 275; macro- 
carpum, 277, 280, Ly 288-2915 
maxinum, 288; Mehrae, 278, 279, 281, 
289, 290; oxyphyllum, 290; pachyphle- 
bium, 254; parasnathense, 280, 281, 
289, 290; pectinatum, 277, 279-281, 
284-287, 289, 290; pinnatum, 288; 


brosum, 27 28 Cot 
Azolla caroliniana 266, 267; mexicana, 66 
Azolla caroliniana Willd. in se 266 
Ballard, Francis. The Correct Name for 
the Hybrid yopteris viatata: 


et, KR. C. The Genesis of the 
ne n Syma 2, Supple- 

mentary Dat. New Native 

eg “area at the New York 
236 


Bir, S. 8. See Mehra 
ee 119, 


yi ata, 4 g 

Lindigii, 49; quoyana, 82, 83 
Bommeria hispida, 162; Knoblochii, 163, 
Bookout, Henr A New Station for 

sl chamanee: Deiereli in psa 246 
Britton, D. M. Report of Rougemont 


oneidense, 43; sect. ongata, 130; 
formosanum, 130; japonicum, 127, 130, 
132, 204, v. minus, 131; lanceolatum, 
41, 204, v. angustisegmentum, 41; 


38-40, 130, 204, 215, 220 ssp. cali 
204, v. {compas n, 


robus 
ies sedate: 

okey 0; mong ok 32, 38—4 
sect. Phsiodrichiam, 128; robus- 
tum, 131; rutifo as hts Nese By 131; 
subg. Geoperiiia 82, 129; Se haff- 


AMERICAN FERN JOURNAL 


simplex, 42, 159, 215, 


mers, 77, 204 
204, v. te nebrosum 42, 


4 
° 
s: 
Ee 
gx 
z 
2 
oo 
Ei 


- rum, 

204 ; tenuifolium, ‘40; chong? 32, 

38-40, 131, 132; virginianum , 63, 

4, 217, 220, ssp. shat Aw ae 204 
Boydston, Kathryn Report of Spore 


Byrsopteris, 149, ; amabilis, 1533 
assamica, 153; aristata, 151, 1525 
carvifolia, 153; coniifolia, 153; denti- 

ata, 152; formosa, 152; Hasseltii, 
54; Henryi, 154; leucostegioides, 15: 
Maximowiczii, 154; m a 543 
rigidissima, 152; speciosa, 154; an- 
ii, 15 
Callipteris, 194, 292, 293; prolifera, 29 
01 


93 
s Dhiasplviine, 64, 219, 220, 
Cam mpyloneuron phyllitidis, 85 
ocean ee “Suengrs a 257 
ropteris cosa, 23 
Cet erach De ne py 84; officinarum, 82, 
84, 139, ce 14 


seariosa, 54;  siliquosa, 
entosa, 170; Seatita, 170 
Letters, 253-255; Recent 
252 
23 


gz 
Cornopteris, 6, 292, 293 
a ar Name for the Hybrid Dryop- 
ata X 


Crane, Pera 
Dryopteris Species Based on 
s of the 


ters of t! eitorctgdy 270 
Crookes, Marguerite. On the Lava Fields 
of Rangitoto, 257 
Crotty, William J. Abnormal Nuclear 
ivision in Fern Prothallia, 92 
Cryptogramma is) 41, v.  acrosti- 
choides, 41; Spigot! 215, 218-220 
Ctenitis, 153, ; eful a 230; Jen- 
seniae, 230 ne nigera, 230; protensa, 
Ctenopteris, 262; oy maori 82; hetero- 
phylla, 262; jubiformis, 86; sus- 
sa, 86 


Report of Judge of 
81; dealbata, 262 


6 
Jutler, ig Ni 


21 
Cyathea, 26; Brawl 


INDEX TO VOLUME 50 


Seyenaiagl crenata, 80; ee Fi 80, 83 
; goggilo- 


ulon y. incisum, 25 
German mvienlhtai” 166; 
0; Fortunei, 10 
Gystopteris, 26, 292 293; bulbifera, 66, 
220, X fragilis, 43; Dickieana, 
aes fragilis, 38, 54, 75, 218-220, 
. laurentiana, 43, y. protrusa, 
S 64, v. simulans, 63, v. tennesseen- 


falcatum, 


the Hima- 
and Com- 
yolutionary Status of 


4 
Gyuiogical Observations on 
layan Species of Athyrium 
ments ie th 
a Gen 2 
Dan EeMbribiané , 49 
Davaite: 147; canariensis, 235; chaero- 
phylloides, 230; denticulata, 81; Marie- 
sii > ‘tricho noides, 
G. J. Ferns in Oubangi, 
Denustacdaw arborescens, 49; cicutaria, 
48; nage tae 218, 220 
1 252; Curranii, 3s 
linearis, 13, 231; pectinata, 48; pu 
Dictymia Brownii, 123, 124; McKeei, 122, 
123, 124; Mettenii, 123 


Dielli aby - 

Digra , 194 

Diplasiopss, "292, 293 

orn 194, 276, ue 292-294; 
an; 9; japonicum, 277, 287; 


4 
main, 288; Dic titerni, 293; sil” 
Dipteris chinensis, 11 
Doodia media oe 
pagar de yedat, 7 


Dry _ Laurentii, 230; ” scandens, 241 

Diy a MOpa heraclea, 

Dryopteris, 7, 10, 26. 87-92, 95, 98, 
104, 135, 146, 149, 157, 202, 215, 


52; assamica, 153; 
104; atropatustris, 89; augescens Vv 
puberula, 163; Bodinieri, 99, 104; X 
Boottii, 88, 89, 106; ri, Has 
136-138, x Filix: 
153; celsa, 274, 275: chaerophylovies 
148, 155; cinnamom Th* 
ana, 275; connexa, "275; te a. 


1 
x-mas, 20, 114 —116, 13 6, 148. 
Os , 


Goldiana, 
in termedia, 


89, x feet anuiic 90, x 


299 


spinulosa, 905 oecgatyt 154; Henryi, 
154;  heteroneura, 100, 101, 104; 
poceennooaen. " dia, - 
88, 90, Ae 0, 275, v. fructuosa, 
44, X marginalis, 44, 90, X spinulosa, 
43, 90; khasiana, 103, 104; x edsii, 
88, 90, 270, 271, 273, 274; leuco- 
stegioides, 152; Lilloi, 275; ludovici- 
ana, 275; lurida, 148, 155; marginalis, 
64, 88, 220, 274, X spinulosa, 91; 
Maximowiezii, 154; mutica, 154; neo- 
podophylla, 103; noveboracensis, 217; 
ochropteroides, 155; p as LS 
136—138; parallelogramma, 75: 8 
patula, ; ossii, 70; egopteris, 
SiTs pittsfordensis, 44, 88, 91; 
podophylla, 102, 103; eune. Poly- 
stichopsis, 148; X Poyseri. 90; 
pseudosieboldii, 101-103; sere 
14 5; subg Deauontecie, 98-104; 
rigi vy. australis, 136; rigid sima, 
152; separabilis, 89; serrata, Sie 
boldii, 98-100, 102, 10 ero- 
neura, 100, v. pseudosieboldii, 102; 
X Slossonae, 88, 89; spinulosa, 43, 88, 
90, 105, 154, 157, 217, " \ 296; 
y. americana, , v. fructuosa, 91, v. 
glandulosa intermedia, 3 


03, 
Deyoters Borreri in Bayari 114 
Duncan, Wilbur H. = zolla  havouninee 
Willd. in ig cae 26 
ages Georg. Farne cn Herzen Europas 


( 
The Be ology of Peruvian Ferns, 46 
sa gre TT OE 48, 164; pilosum, tay 


The. Endemic cae te of the Cali- 
f 


107, v. arcticum, 107, Vv boreale, 107; 
bogotense, 47; ¢ lif ea 
Calderi, 107; X Ferrissii, 186, 190, 
1938; fluviatile v. siccum, 193; fon 
tinale, 192; Funstonii, 185, 186, 188, 
192, f. caespitosum, 19 v. caespito- 
sum, 192, f. nudum, 192, v. nudum, 
192, f. polystachyum, 192 f. ramosum, 
192, ramosum, 19 subg. Hipp 
chaete, 185; hyemale, 185, L89 
affine, 188-191, v. affine X laevigatum, 
193, v. californicu 191 Dru 
mondii, 191, v. elatum, 192, f herba 
ceum, Hie Hates cs 191, 
japonicu : inte:medium, s 
cuatermediums 40, 186, 186, 185, 190 
193, esu 190 kansanum, 43, 
f. polystachyum, 191, 193, v prealtum 
191, p rum, SAN seudo 
hyemale, 64, 192, f. pumilum, 191, v. 
robustum, 191, v. Suksdorfii, 193; in 
termedium, 193; k num, 186, 188, 
192, f. caespiticum 192, f. caespito- 
sum, 192, f. Eatonii, 192, f. elatum, 


300 
191, f. poiystachyum, 192, f. ramosum. 

- 192, £3 varisentaides, 192; ‘Komarovii, 
191; laevigatum, 66, 185-190, 192, 
£93, +f: caespitosu kOe, tum, 
191, ssp. Funstonii, 65, 186, 187, 192, 
f. polystachyum, , f. proliferum, 
193, f. ramosu f brellum, 


192 192, f. variega- 
toides, 192; tnd litorale, 190; pratense, 

191, -v. affine, 7 : 
Drummondii 191, v. laevigatum, 
townie 
1 


aes! 
tra don, 190; Zslochaetum, i 
Erdtman, G. oe nd Spore Morphol- 
ogy (Rev.), 7 
Evergreen ti actaeon and the Meanings 
i fs traspecifie Categories as Used in 
American tes ie Pa 32 
ipnawaraes mme p' 
wan, Siginsthons "Ele Hlustations of Ferns 
d 26 


Faust, Mildred E. Survival of Hart’s- 
e Fern in Central New York, 55 
Fern gee 
Ferns and Miss in Kansas, 62 
Ferns Cultivated in California: 
Sadleria, 232 
Ferns in Cultivation, II, 168; iif, 


Growing Fern Balls, 335, | 
Ferns in bangi, Congo, 
Feedin ardy Ferns: Wine. Safe, and 


Risky Methods, 263 
Fi-st Tuigteattons of Ferns from Peru 
and oe eS 
Fuchs, Han e Nomenclature of a 
Japanese “opihioglossum, 124 
and Youn, : Piege halonhed 
voi ‘eutentan m, 
f mate eee Jour- 
nal, 2; Susiplerasntes Data, 179 


Gein ee 48; bifida, 48; 
80 

Ginn gh Billardieri, 262 

enemies ium Dryopte Tis, 43, 219, 220, 


y. Robertianum, 43 
Gymnogramme, 194; oe 61; 
Ss ¥ a. 


5-62 
Hauke, Richar The Smooth Scour- 
ing Rush and Its Consplaxities; 185 
Sit ae gs marginatum, 48 
Hemioniti ae 
Hemipter 
Heterog: ania pinnatum, 84 
Hevly, Richa rd H. Report of Auditing 


re 
lucida, 251; olivacea, 
o 


9, 80 
Hippochaete, 187, 190; hyemalis y. — 
‘ i » 192, v.. caes- 
192,°.v; os 
. Funstonii, 192, 
vy. polystachya, 192, v. ramosa, 192, 
¥. variegatoides, 192; prealta, 191, 


trian- ° 


AMERICAN FERN JOURNAL 


affin 91, vy. intermedia, Bae 
one pon Are titess 191, subv. poly- 
pans AO SLOL, * Ye pseudohyemali pees 88 


aabr ella, 192, SS: Suksdorfii, 193 
Histiopteris incisa, 2 

Holttum, R. E. Poke Problem Peete 

Se! pact ee cordatum brn lag Syn- 


gramma pinnata J. Smith, 

How I Became Interestel in Ae 

How ell, ohn Thomas, The Endemic 
Slip an of the Cailfornia Floral 
Province, 

Humata he sie Hla, 81 

Hunting Ferns in the Barrancas of Chi- 
huahua, Mexico, 161 

Huttleston, Donald. Report of Secretary, 


Sate, Thomas A. New Georgia 
Stations for Ly Stam complanatum 


var. res ae cn rme, 240 

Hybrids in the Genus Asplenium Found 
in ln Peet oa and Central Europe, 
138 

Hymenoplytlum revolutum, 259; Ruizi- 

m, mice 15,259 

Bente vg i hoe 19; hostilis, 48; 

apalisioerain 


The Hentification io Aspidium distans 
13 


(Obituary), 222 
Butleri, 63; 
—184, f. allida, 184; 
ae v. Oe bra He Orcuttii, 18 
Isoétes melanopoda in Southern Llinois, 
181 


Isoloma, 109, 111 

Japanobotrychium 12 

Joe, Barbar Fe: ns Tuloeded in Cali- 
fornia: Sadler 

A Key to dmeceas Dryopteris Species 
Based on Characters of the Perispore, 
270 

Knobloch, Irying. Hu g Ferns in the 

Barrancas of Saeaban: Mexico, 161 

akela. a. Report of 1959 Minnesota 

Field-trip, 215 

Lastrea, 98, 148, 158, 26 66; auges 
cens, 159; coniifolia, oo decomposts. 
148: dilatata, 266, v. glandulosa, 91; 

t 48; hispida, 148; 

sect. Polystichopsis, 148; 

Standishii, 152; atigiaues, 

6 


Lastreopsis, 154, 155; jeu 2315 
chaerophylloides, 155; — 155; 
; recedens 
sin Galtivation, 
‘n Balls, 235 
euneata, 13; 
ensifolia, 110; viridis, 13 
Lomariopsis Dincenee 229; hederacea, 
sai 


Lone s Currori, 230 

parts oe nde sum, 10 

408 rsd ied Fe eas Society, $353 
Loxogram 86; nia, 86; Parksii, 82 
Loxsoma, a3 


INDEX TO VOLUME 50 


Loxsomopsis costaricensis, 
The a and pe SES of 


Eso Rabid tT, 151i, 3219, 0220 5 alope- 
curoides, 42, v. adpressum, 43; cer- 
uum, 231; clavatum, 51, 7; com- 
planatum, 51, v. fi abelliforme, 217; 
240, 267; inundatum Jussiaei, 
51; Iucidul “a 21 ‘67, 268; ob 
securum, 2 — dendroideum, 267; 
pendulinum, Selago, 216, v. pa 
tens, 216, 268 "Btanal 230 

Lygodium japonicum, 4; palmatum, 
79; volubile, 48 

Macroglossum, a 

Macself, A. J. Hardy Ferns: 
Wise, aon ne Methods, 263 


Safe, 
Marattia, 7; ‘salicina, 


ae silea mucronata, 65, 66 
Mason, Charles T., he Report of Audit- 
ing onuthittes: ee 


Matteuccia, 170, 7; pensylvanica, 9; 
Struthiopteris, 92, 94, 95, 156, 220 
wy Kanse peor ae ald L. Ferns and Allies in 


Meco: fou issum, 259;  dilatatum, 
359. P aabalainens age rarum, 259; 
sanguinolentum, 25 263; scabrum, 

Mehra Bir. Cytological 


P..N- and 8: 8. 

Ohanweiona on the Himalayan Species 

of Athyrium and Comments on_ the 
coh ili cawigael dt Status a e Genus, 276 

oe multifidum 

a ma, 7 

Yr, E. Hybrids in the Genus As- 

wena Found in Mabel apea sia and 
e 13 


tral Europe, 
Microchlaena, 98, : yunnanensis, 104 
Microgramma lycopodioides, 230 
Microlepia speluncae, 231 
Microsorium, 241-245; alternifolium, 
244; commutatum, 243, 244; diversifo- 
lium, 241 43, 259; multijugatum, 
244; novae-zea diae, 243; Parksi 
244; punctatum 5, 230; seandens, 


241, 243; Seolopendria, 85, 243, 244; 
sibomense, 243, sylvaticum, 243, 


Microstegia 194 
Mohria, 
Mohlenbr. oe Hy, dre Tsoet 
piclanopods in’ cities Illinois, 181 
Morton, ©. servations on Cultivated 
uu nm 


Fern ns, vr “$s Geran rently Known 
as Rumohra, 145; S ern Books 
for Different Regions of the United 
States, 169; ‘axonomic Notes 
Ferns, I, 275 

A Mule Gain Trip to Sierra Mohinora, 


Chihuahua, 66 

Multicellular and Branched Hairs on the 

n Gametophyte, 79 

Nayar K. The hag Pd eck and 
You ung Sporophyte of Athyrium eseu- 
lentum, 

Nishida, Makot A Synopsis of Scep- 
tridium in ea 127 


Nephrodium, 89, 


301 


275; crista tum £ 


acuminata, 80, 81; 
231; cordifolia, 53, 80, 
3 


ze 
= 
i 


w York Bot: 
A New Station tot Tric 
gia, 246 
ewell, Chauncey 


omanes in Geor- 


Jackson (Obituary), 

The Nomenclature of a Japanese Ophio- 
glossum, 124 

Notes on Pterido 
and New Caledonia 


phytes from Australasia 
, cele 


cote 158; schenborniana, 68; 
54, 162 calitocititn. 22, 158; 
159, t accessita 2, ssp. nigrescens, 
22; candida, , 167; cretacea, 159; 
Grayi, 68; incana, 75; Lemmonii, 165, 
166; limitanea, 166; mollis, 30, 31; 
Newberryi, 22; nivea, 54: peruviana, 
; sinuata, 162; Reine 159; 

61, 


sir Rape on Galavewsd Bo dd VI; 
Ferns Currently Kno as Ru- 


‘ich 
enropieds. 
Oleandra Tistebit, 280; Wallichiana, 80, 


On the Lava Fields of aon gitoto, 257 

On the Species of Dryopteris, subg. Pyc- 
nopteris 

Onychium, 158; densum, 19 

Onoclea sensibilis, 9, 64, 218, 220; ft: 


hemiphyllod 33, f. obtusilobata, 33 
Cpniososnus: : anugstatum, 125, 126; 
bucharicum, 125, 126; californicum, 
18; Engelmannii, 63, 64; japonicum, 
—126; lusitanicum ssp. californi- 
cum, 18; nipponicum, 124-126; nudi- 
caule, 54, 125, 6; palmatum, 48; 
petiolatum, Sans Savatieri, 125, 126 
thermale, 1 ucharicum, 126, v 
agi gat 126: ulpatica, B66 
ie 


219, 220, bucharicum, 125, 1 
xe 


la, 33; Claytoniana, 2 7 ; multi- 
fida, 130; regalis, 8, 41, 66, 220, 
spect is, 41; ternata, 131 

Osmundopteris, 12 

Paesia seaberula, 261 

Paltoni 


lanceolatum, 82, 85 

Papuapteris, 147 

balcinee _nllosuroides 68, 163; androme- 
difolia, 22, v. pubescens, cae atropur- 

63, 65, 219, 220, Bushii, 


pure v. occidentalis, 156; irackv iets, 


23: compacta, 23, 159; dealbata, 62, 
63; densa, 19; glabella, 64, 65, 220; 
mucronata, 23, v. californica, 23, 159; 
ornithopus. y. brachyptera, 23; ov vata, 
51; rafaelenis, 22; rotundifolia, 261; 
sagittata, 51, 163, v. cordata, 68, 163; 
Seemannii, 151, Fae Skinneri, 161, 
167; ternifolia. 70 


ot oe pate ra 82, 85 
gm rophlebia auriculata, 70; caryoti- 
Oia tobdiceus major, 9 
omer gee connectilis. 219, 220; elon- 


Pras Walter S. Report of Treasurer, 


Phie a diu 85 

aus | Seolopendrium, 41, 84, 139, v. 
am, 41, 55-62, 84 

teecinat 8, 2 scandens, 241; 


daca: , 248, 244; 
Scolopendria, 230 
, Rodolfo E. G. The Identi- 
ation 0 “As spidium distans Viviahi, 


Pik falaria americana, 66 

Pityrogramma eaten elanos X chrysophyl- 
la, 11; hybri vy. maxima, 11; tarta- 

bbe si ‘tria ng wilaxis v. palli da, 23, Vv: 

vis 28 

Plasiozs" ja & s ordata, 75, a 

Platycerium, 86: alcicorne, 82; 
lense, 230; grande, 82; stemaria, 230 

thy » 23 


ecos 
Pleceneinia conjugata, 83 
Pleopeltis hastata 5 
Pleurosorus, 276: rutifolia, 82 
Poelt, J. Dryopteris Borreri in Bota: 


114 
pop ae 7: caudata, 49; osmunda- 
9 
Besse: 75, 244, 245, 265, 266; 
adiantiform 147; amoenum, 245; 
ematitolni, 48: aristatum, 50— 
ee aur 163; iforni 23. 


chn 
1 creenitonin 
eae clato es: 152; 48; e 
tum, 103, 
ayy 68; faleara, 

32, v. Thane Ace fica, ee 


er 
cng 
ia 


AMERICAN FERN JOURNAL 


vee 38, 156, 220, 245, v. colum- 

75, v intermedium 

Polstichopsis 14 y gee © 8: raga | 

; chacrophylloides 15 
rida, 


5; Hasseltii, 
ee 5; ochropteroides, 155; 
pubes eis tes 
Polystichum, 13, , 145-149, 202, 250, 
4-266; acrostichoides, 64, 217, 220: 
culeatum alifornicum, 24, v. Dud- 
leyi, 24: adiantiforme, 146, 147; ama- 
bile, 153; aristatu «a Serger. 192; 
assamicum, 153; eali- 
fornicum, ba: capense, 146, aa ae 
vifolium, 153; coriaceum, 146, 147; 


distans, 135-137; Dudleyi, 24; Filix- 
reri, 135, 1 6; Hen- 
; = itum 


x ns, 
25, 42 ssp. udatum nudatum, 
24; inte aii 49; Bichard 2625 
speciosum, 154; Sta ndishii 145; ulgi 
nosum, nt 
Pond, Bremer Whidden (Obituary), 221 
Progress in tthe Study of Dryopteris Hy- 
brids, 87 
sitar: 7; nudum, 7, 269, 270 
Psilotum in Louisiana, 269 
Pteretis, 170 
Pteridium aquilinum, a 290; 281, Ve 
arachnoid 48, languinosum, 
eis v. Dp * 
5 pubescens, 73, 
iene, 261: latiusculum vy. 
cens, 4 
Pteridrys australis, 
85% rndromeditoia, 22; atro- 
}. ‘ 


ges Hel 23 grandifolia, 48; 
macilenta, : on icata, 53; similis, 
231; spinulifera, 231; tremula, 261, 
263; vittata, 47, 92, 9 


2, 94— 
Sieboldii, 98 
nsis, ee lanceolata, 230; 
“lingua 85; serpens, 259 
— William D. Peilothan in Louisiana, 
9 


gd rs of Audi fing Hage ee Bil; 
of Judge 


’Ve nent 

153: a teste 

see ee Be 
46, 147: 

; ‘chgerophso des 

52; 154; 


nipponi 4 
duadripinnata, sino-miqueliana, 
: ls seciosa, Standishii, 152: 
chii. 153 
235; 


154: 


Sadle 
Hilebran 


232 234; 
dii 
mace cree 
138 130 


2— myatheoides, 
234 

32, 127-132; daucifoliu 

j aoieog ee 129, 130; he 

sect. Multifida, 130, 181; 


INDEX TO VOLUME 50 303 


multifidum, 129, 130, 132, vy. robustum, subtriphylla, 83 
130; 231; obliquum, 129: robustum, Thelypteris, 71, 158; Biolleyi, 82, 84; 
131; ternatum, 130-132, vy. nipponi- hexagonoptera, 64; macrotis, 49; ne- 
cum, 130, 132 vadensis, 159; noveboracensis, 220; 
Schizaea, palustris, 64, 220; pilosa, 75; pu- 
Schizolepton, 109-112; cordatum, 110 berula, 159; Rosei, 54 
Schizoloma, 109115; Billardier Ke 109; ne Mary D. Notes on Pteridophytes 
cordatum, 109, 0 112; Guerinianum, m Australia and bg Caledonia, as 
109 “os Vein Patterns in Microsori 
Scolopendrium, 264, 26 ad Soh and Its ‘Allies, 2. 
Seudder, Anna E. Iecport of 1959 Ver- ig veered aerugineum, 229; diversi- 
mont Field-t i. Pane fro pe Hostmannianum, 49; Pe- 
Selaginella, 1 Ar- tarsi 24 radicans, 75 
senei, 163; asprella, 16; “Bigelow 16; Griancrla tnifoliat a, 47 
Bolanderi, 17; bry winsch cine- Tryon, “et a. The Ecology of Peruvian 
rascens, 17 Hansen iy as cee. Ferns, 
oides, a7 myosurus, 231; novoleonen- Two dg Scores Stations for Lycopo- 
sis, 167; pallescens, 70, 165; abi dium complanatum var. flabelliforme, 
ana, 54; pestris, 64, 68, 24 
Bolanderi, 17, v. Hansenii, 17; Sar ich Problem-species: Schizoloma corda- 
torii, 167; ee 76; Wallacei, m Gaud. and Syngramma pinnata J. 
pe Wrightii sm nith, 1 me 
ane caes ee var. densifolia, 8 Men Patte in Microsorium scandens 
e Smooth porte Rush and 2 Com- nd Its ‘Allies, 241 
fcniireae Vittaria, TIL 112; guineensis 230 
Some Fern eh for Different Regions Wagner, Warren H., Jr. Evergreen 
of the United States, 169 Grapeferns and the Meaning of Infra- 
Stokey, Alma G.  Multicellular and specific Categories as Used in North 
Bra ot Hairs on the Fern Gameto- American Pteridophytes, 32; Report of 
phyt Curator and Librarian, 2 
Sacviesi i Sahl gh Fern in Cen- What Is the Role of Spores in Fern 
tral New York, Taxonomy ?, 6 
Syngram = 113; atismitolia 110, 113; Wherry, Edgar T. re I Became In 
pinnata, 109-11 11 terested in Ferns, 225; Progress a the 
A Synopsis eo Roxpuiaiatn = Japan, 127 Study o Nrinoshietg iybrids, 87 
Taenitis, 109-113; blechnoides, 110-113; Wiggins, Ira L. Report of President, 204 
ordatum, 112 Woodsia, 75, 157, 215; Cathcartiana, 
Tagawa, Motozi and Kunio Iwatsuki. On 216; glabella, 216, 220, X_ ilvensis, 
Hs Species of Dryopteris, Subgenus 43; ilvensis, 133, 136, 218, v. alpina, 
yenopteris, 98 43; mexicana, 68, 76; mollis, 70, 163; 
ienger Mrs Charles XY Stat ee 222 montevidensis, 53-55; obtusa, 63, 64, 
Cremona, s on Ferns, 66, , 220; scopulina, 216 
Tectaria, 202! egy belong ane ee iffithii, Woodwardia virginica f. fertilis, 33 
33; incisa, 49, 83; irregularis, 83; Xiphopteris delitescens 82, 86 
ERRATA 


Page 4, uate line 4: For ‘‘supposted,’’ read ‘‘supported.’’ 
»b m line: Trans oe to become bottom line of p. 7. 
7: jhuen 20 and 22: Fo ‘‘eompliment, ’’ read “complement. ee 
Page 11, line 30: For ‘‘anomolous,’’ read anor alous.’ 
pig | Vax feet.’ 


a) 
so 9 
gg 99 
@ 
Hoy 
é 
a 


T, 
Page 66, line 25; For ‘ singe 7? read “pidest 
eos oo on map: Change ‘‘One inch = 12 sir 2? to ‘‘One inch = 16 


Page ry “Transfer legend under map to become legend under Plate 8, p. 74. 
Page 74: Transfer legend under Plate 8 to become legend under map, p. 69. 
i e of pape . Stokey: “piss 


: UBIFORME 
Page 112, line 19: For “cordatum,” read “cordata. 
Page 114, line 7: For ‘‘funtioning,’’ read “funetioning’ 
Page 136, line 23: For ‘‘distant,’’ read ‘‘dis 
Page 144: Interchange lines 1 and 2 


HENRY TRIPP 
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and cryptogamic botany 


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JUST PUBLISHED! 
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ALso In Stock 


FERNS OF MALAYA, by R. E. Holttum (Kew). 642 pp., illus. cloth 
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“THE PREPARATION OF 
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M. Johnston, of the Arnold Avboreien, Harvard a Pg has 


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Help for the amateur botanist, and hints for the professional collector, 
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American Fern Yournal 


A QUARTERLY DEVOTED TO FERNS 


Published by the 


AMERICAN FERN SOCIETY 


EDITORS 
RA L. WIGGINS 
Cc. V. MORTON ROLLA M. TRYON, JR. 
JOHN H. THOMAS 


VOLUME 51 
£901 


MONUMENTAL PRINTING COMPANY, BALTIMORE, MARYLAND 


Conients 


* VotumeE 51, NumBeEr 1, Paces 1-64, ISSUED Aprit 20, 1961 
Florida Rarities 2 Thomas Darling, JY. 
Hybrids in an ‘Asda Gymnocarpiums 

Elizabeth Eichstedt Root 
The Problems of Variation in North American aout 


nald Britton 

Some New see on the Vernation Differences of Sn dissectum 

end tem W. H. Wagner, Jr. 

Supntnentary Neel on Dryopteris Hybrids —-- Edgar et Wherry 
w Forms in Botrychium virginianum -------------_- . C. Co 

pediuas Noelis on poms: bl ee ee C. ues Morton 


Some Records of Michigan “es and Fern Allies ——. Jarl K, Hiltunen 
Shorter Notes: Recent Fern Discoveries in Western Pennsylvania ; 
Fern Monster?; Cystopteris tennesseensis Shaver ; Pellaea atropur- 


pure 
Sah Fern Literature 
Notes and aa Annual Meeting; aicinte, Dead: Botrychium 
Transplan 
American sid Society: Report of President; Rep. of Secretary ; Rep. 
f Treasurer; Rep. of Auditing ones Report of Judge of 
Elections; Rep. of 1960 Arkansas For 


VouuME 51, NumpeEr 2, Pages 65-112, Issurp JUNE 29, 1961 
My Lath House for Ferns 32252 Fay MacF adden 
A Parcel of Ferns Mee ‘ee mrp Geo 
Gladstone W. pest ell aor Franklin D. Snyder 


Another ss eine rom Kentue 
Dale M. Smith, Truman . Weed and Donald E. Tate 
C. 


Some Forms of Polypodium californicum —..-------- Horton 
On the Relative os ek of the Fertile Segments in Botrychium dis- 

sectum: and B. oneidense 22 W. H. Wagner, Jr. 
Another Genus of Ferns New to the United States .._C. V. Morton 


Iséétes echinospora Durieu in North America ——-- perwaihd Boivin 
A Hawaiian Thelypteroid Fern with Peltate Indusia Kunio Iwatsuki 
Observations on Microsporocarpic Material of Azolla caroliana 
Tey, Se W. Reinert, ne Richard D. Houk 
Adaxial Sori in Polypodium hesperium — mily L. Hartman 
Heat Resistance of Sporocarps of Marsitea aie 
William W. Bloom 


Peltapteris in Costa Ric Clyde F. Reed 
wihcing! of Isoétes BTN: 8. CL Verma 
rature 


es Fern Sei ociety: Report of Spore Exchange; Constitution of 
10 


the American Vern Society, anew. a 


VOLU: » Number 3, Pages 113-160, IssuED OcToBEr 4, 1961 
Growing eat Heats Spo Thorleif Fliflet 
Some Chromosome aban of Icelandic Ferns and Fern-alli 


ell Love and re Love 

Horns for-9 Lath House Gov oo Fay MacFadden 
A Resume of the oe Reorganization of a errant Subgenus 
ippochaete, I . Hauke 

An Lutebecting Ecological Niche Involving Salvinia ees Willd.. 
homas A, Hutto 

The esnesdoane! a Mecodium wrightii in Canada _____. Kunio Iwatsuki 
Notes on ee R. Proctor 


Concerning Azolla imbricata gee ene Yu-Feng Shen 


Shorter Notes: An Addition to the List of Ferns Growing Naturally 
15 


in hee eee Pe pions Alberto Chiaragi, 1901-1960 —. 
Notes and N A Fern Project Wins a Science Fair e wa oe Society 
of Economic Cae A Mass Collection vi Polystichum; Onoclea 
Spores rene able 
American Fern Society 


VoLuME 51, Numser 4, Paces 161-200, Issuzp DECEMBER 30, 1961 


The Fern Valley at the United States National Arboretum — 


Edith Bittinger 
Ferns in Cultivation, IV. Some Wall Ferns Sylvia Leatherman 
Southern Distribution of Botrychium ee a and B. multifidum 
F. R. Fosberg 

Dryopteris + Tavelii in the Valley of Chamonix, Frane 
Andr . Anne-Marie Ps e Collaris 
Basket Ferns for Southern California _ Fa = MacF adden 

Shorter Notes: Dennstaedtia eee (Poir.) 


Hieron. in Tex 
Pteridological Troglodytes; Marsilea Soseitien L. in fidlana as 
1 


Notes and News: Letter on Fluorescent Tubes 
sie Fern Society: Report on the Babetland National Forest 
For -: mace ae 


ie o Volume 51 


97 
99 
105 


113 


al 
Vou. 51 JANUARY-MarcH, 1961 No. 1 


American Fern Journal 


EDITORS 
Cc. V. MORTON 
ROLLA M. TRYON, JR. IRA L. WIGGINS 
: CONTENTS 
Florida Rarities Tomas DaRLine, JR. 1 > 


Hybrids in North American Gymnocarpiums 
ELizaBeTH Eicustept Root 15: 


The Problems of Variation in North American saa he. cori 
NALD BRITTON 23 : 


Some New Data on the — Differences ~ nas hium 


dissectum and B. ternatum___._________ . WAGNER, JR. 31> 
Supplementary Note on Dryopteris Hybrids. Epear T. WHERRY 33 ‘ 
New Forms in Botrychium virginianum ___________ W.J. Copy 36 + 
Taxonomic Notes on Ferns, II C. V. Morton 37 + 


Some Records of Michigan Ferns and Fern — 
arL K. Hintunen 40 . 


Shorter —— Recent Fern Discoveries in Western Pennsylvania; - 
Fern mster?; Cystopteris tennesseensis Shaver; Pellaea 
een 42 


Recent Fern Literature 45 


Notes and News: Annual Meeting; Reprints Desired; Botrychium 


a Bg) Society: Report of President; Rep. of Secretary; 
Han f Treasurer; tg of Auditing Commi mumittee ; i aonwt of 
Judge of Elections; Rep. of 1960 go twee Foray_ 49 


The American Fern Hociety 


Council for 1961 
OFFICERS FOR THE YEAR 


Cam A. Brown, Department of Botany, Louisiana State oer Baton 
Rouge, Louisiana sident 
Marcet RayMonp, Montreal Botanical Garden, 4101 East sherbrocko fcuk 
Montreal, Canada Vice-President 
DonaLp Hurrizston, Longwood Gardens, Kennett Square, Penna, 


Secretary 
— . Pures, Department of Botany, University of —_ Tucson, 
rer 

Cc. ¥. riety Smithsonian Institution, Washington 25, D. C. 
Editor-in-Chief 

OFFICIAL ORGAN 
American Fern Journal 
EDITORS 

C. V. Morton.__________Smithsonian Institution, a 25, D. C. 
Roiia M. "TRvo 22 Divinity Avenue, Cambridge, Mass. 
Ina L, Wiaeerns. "aie Herbarium, Stanford University, Stanford, Calif. 


An illustrated quarterly devoted to the general study of ferns, pub: ublished 
quarterly by te sap pn ern Society, a Elm Ave., Baltimore 11, Md. 
Second-class postage at Baltimore. 

Matter for publication A should be pot endl to C. V. Morton, Smithsonian 
Institution, Wa 

Subscription $2.50, pete nFt of agency handling ng sent free to mem- 
bers of the erican Fern Soci ity (anual duos, $2.50 mem- 
bership, $5. ne O; ite membership, $50.00). veh prennd St if ordered in 
advance, W. il be furnished authors at cost. a they should be ordered when 


is returned. 
Back volumes $2.00 each; single back numbers 50 cents each; Cumulative 
Index to vols. 125, 25 cents. Ten per cent discount on go of six volumes 
or more. 
Changes of address should be sent to the Secretary: Dr. Donald agen 

, Longwood Gardens, Kennett Square, Pennsylvania — and sub- 
seribers should allow two months for es to take effec 

pplications for membership, st orders for mack numbers, and 
hee geese communications should be addressed to the Treasurer: Dr. 
Walter 8. Phillips, Department of Botany, University of Arizona, Tucson, 


LIBRARIAN AND CURATOR OF THE HERBARIUM 
Dr. W. H. Wagner, University of se Gpaeh Ann Arbor, Mich. 

A regular loan department is maintained in connection with the library 
and herbarium. Members may borrow books on ~pevesnig be Pree! time, 
the borrower paying all postal or express cha: vn The pages of the J ournal 
ag are open to members who wish to arrange exchanges; a oaabarie list 

ublished at intervals, to assist those maauited in obtaining 
pb different localities. 


American Fern Journal 


Vou. 51 JaNnuary-Marcn, 1961 No. 1 


Florida Rarities 
THOMAS Daruina, JR. 


In April, 1941, I made my first trip to Florida, combining some 
casual fern hunting with other activities, mostly sightseeing. 
Since this was the period when the St. John brothers (Edward 
and Robert) were making their extensive studies of the fern 
flora of peninsular Florida, it is not surprising that I made a 
point of visiting them at Floral City, north of Tampa. More 
Surprising, however, was. the fact that Maurice Broun and his 
wife, having just returned from the West Indies, were visiting 
the St. Johns at that particular time. Maurice, noted ornitholo- 
gist and curator of Hawk Mountain Sanctuary (Pennsylvania), 
and also known in the botanical field as the author of “Index to 
North American Ferns,’’ had previously. introduced me to.some 
of the rarest ferns in Vermont when he was ‘superintendent of 
nature activities at Long Trail Lodge, home of the Green 
Mountain Club, at Sherburne Pass. 

The first day of my visit Maurice and I accompanied Edward 
and Robert St. John to certain areas near Floral City noted for 
unusual ferns. As a visitor familiar with only northern flora, 
this expedition was a liberal education. At the end of the day 
my mind was reeling with countless unfamiliar names. Surely 
Florida leads all other states in the number of ferns to be found 
within its boundaries. Because of the protective nature of lime 
sink formations, ferns are here to be found which otherwise are 
known only from the West Indies and other tropical regions. 

Although I encountered many ferns new to me during that 
memorable field trip, as I now look back on the experience I find 
that we discovered few actual rarities with the exception of 
Ophioglossum nudicaule (tenerum), tiniest of the Adder’s- 
tongues, near Homosassa Springs, and Blechnum occidentale in 


2 AMERICAN FERN JOURNAL 


the vicinity of Brooksville. According to Dr. Small, the former 
is probably more overlooked than uncommon, but he lists the 
latter fern from only two hammocks in Florida. Following our 
return that evening an unexpected tragedy, the sudden death of 
Edward St. John’s wife, cancelled our proposed field trips. The 
following day we had planned to visit Maynard’s Cave at Le- 
canto to see three of the rarest spleenworts, Asplenium plenum, 
A. scalifolium, and A. subtile, believed to be Florida endemic 
species by the St. Johns, but more probably hybrids; they are 
still a subject of controversy. Unfortunately, I was not to see 
the St. John brothers again, nor to revisit Florida for many 
years. 

Continuing to Miami along the Tamiami Trail, east of Mon- 
roe I stopped at dusk to look for Asplenium serratum, the 
Bird’s-nest Fern. The Everglades swamps are most forbidding 
as darkness approaches. Weird bird calls mingle with the splash 
of unseen and unidentified animals and reptiles. Wading waist 
deep into the morass along the highway I was rewarded by find- 
ing A. serratum growing luxuriantly on mossy logs and vege- 
tation together with the giant Strap Fern, Campyloneuron phyl- 
litidis. 

The late Professor Buswell, of the University of Miami, in- 
vited me to join him on an expedition along the Loop Road in 
the Everglades south of Monroe. Here, in addition to interest- 
ing epiphytic orchids, he showed me a fine display of the Vine 
Fern, Phymatodes heterophyllum (Craspedaria serpens) rooted 
in humus on the ground and climbing by rootstocks on the near- 
by shrubs and trees. 

The next day we journeyed southward to Castellow Hammock, 
near Homestead, where we visited the amazing lime sinks of that 
region. Here I first encountered that fiendish subtropical vine 
known as Devil’s Claws or Hold-back Vine, Pisonia aculeata, 
which literally can (and does) rip one’s clothing to shreds. At 
Castellow Dr. Buswell showed me Trichomanes punctatum, one 
of the rare filmy ferns, along with Tectaria minima (Small Hal- 
berd Fern) and Asplenium verecundum (Lacey Spleenwort). He 


FLorIpA RARITIES 3 


did not mention Tectaria Amesiana and T. coriandrifolia, two ex- 
tremely rare ferns known from but a single location (Hattie 
Bauer Hammock) about three miles distant from Castellow 
Hammock. Whether these rarities were still to be found at 
Hattie Bauer as of that date is a matter of conjecture. Also at 
Castellow I saw the beautiful South Florida ‘‘tree fern,’’ 
Ctenitis ampla. 

On my return northward via Route 1 along the east coast, I 
stopped at Vero Beach to look for the rare Hand Fern, Cheiro- 
glossa palmata, which grows as an epiphyte in humus at the 
bases of palmetto leaf-stubs higher than a man’s head. Although 
Maurice Broun had given me a sketch map showing the approxi- 
mate location one mile south of McKee J ungle Gardens, it took 
considerable searching before I finally discovered the prize in an 
area of canals and scattered cabbage palms. Although I never 
returned to the area, I understand that this unusual fern has 
been introduced into the Jungle Garden and may now be seen on 
trees there. It has also been reported from a few other localities 
including (surprisingly enough) along a busy highway south of 
Miami! 


I did not visit Florida again until December 1955, when I 
flew down to Sarasota to stay with relatives on Longboat Key. 
Since then I have returned every autumn around Thanksgiving, 
and each time have included some fern hunting in the overall 
program. 

Tn 1955, not having a car at my disposal, my activities were 
somewhat restricted. Fortunately John Beckner, a member 0 
the American Fern Society from St. Petersburg, invited me to 
join him one Sunday in a search for rare Aspleniums. 

First we drove to Maynard’s Cave at Lecanto, which I had 
hoped to visit many yea ss with the St. John 
brothers. Climbing down an iron ladder at the entrance, we 
found that a recent rock fall had aeganns part of the cave and 
made exploring hazardous. Asplenium verecundum was in evi- 
dence in places, and at one point John thought he detected 
A. Curtissii too far out of reach to be identified. Mature fruit- 


4 AMERICAN FERN JOURNAL 


ing fronds of the three rarities A. plenum, A. scalifolium, and 
A. subtile have not been seen for many years. Tiny sterile speci- 
mens of what might have been immature forms of A. scalifolium 
(once termed ‘‘A. swave’’ by Edward St. John) were scattered 
along the cave wall. One sterile plant resembled A. plenum, but 
otherwise the cave was barren of interesting spleenworts. 

In a lime rock area on high ground not far from the cave we 
found a limited occurrence of one of the rarer lip-ferns, Chet- 
lanthes microphylla. Near Brooksville, we saw Asplenium ab- 
scissum and A. heterochroum, neither of them rare, but both new 
to me, as well as the relatively uncommon and highly decorative 
Pteris cretica var. albolineata. 

The final object of our search was the epiphytic spleenwort 
Asplenium auritum, recently rediscovered by the Garrett broth- 
ers of Tampa. It grows southwest of Zephyrhills, in the Hills- 
borough River State Park area, usually appearing ‘‘on the north 
side of dead live-oaks leaning south.’’ And before dusk, after 
considerable search and in just such a location, we came upon a 
thriving colony of this rare and intriguing fern. 

Several days later I rented a car and drove to Lake Pana- 
soffkee from Sarasota (about 300 miles round trip) in an attempt 
to find the strictly local Hemlock Spleenwort, Asplenium cris- 
tatum, known in the United States only in this general region. 
En route I stopped at a point south of Tampa to see the unique 
Floating Fern, Ceratopteris pteridoides, not uncommon in open 
ditches along the road. I had been told that at Lake Panasoffkee 
A. cristatum grew sparingly just south of the outlet on low 
boulders in an elevated portion of the hammock, but though I 
searched carefully for several hours until dark I could not find 
it. The long trip from Sarasota had proved a wild goose chase, 
and I would have to try again the following year. 

In November, 1956, I again went to Florida by plane, this 
time going directly to Gainesville to explore that interesting roc 
formation west of the city known as Buzzards’ Roost. Renting @ 
car and consulting a sketch map given me by Dr. Erdman West 
of the University, I located this area without difficulty. At one 


Fiorwa Rarities’ 5 


time there was a profusion of rare spleenworts and other ferns 
on and around this cluster of lime rock ledges. Masses of 
Asplenium Curtissii, a Florida endemic formerly confused with 
A, myriophyllum of the West Indies, were described as forming 
dense mats or iridescent green cascading over the faces of these 
cliffs. Now, however, this fern has disappeared altogether from 
Buzzard’s Roost, and even A. verecundum (Lacy Spleenwort) 
and the Creeping Fern (Thelypteris reptans), which also used 
to carpet these steep walls, have dwindled to an alarming extent. 
The damage has apparently been caused by a gradual leaching 
out of the lime, followed by an extensive drying and crumbling 
of the rock. Furthermore, cattle have been invading ‘‘The 
Roost’’ from pasture land above, and browse around at will. 
There has been talk recently of preserving this area as a nature 
sanctuary before it is too late. 

Ithough unable to find Asplenium Curtissii, I was interested 
to renew my acquaintance with A. verecundum and A. abscis- 
sum, but the highlight of my visit was locating A. pumilum 
(Dwarf Spleenwort) on the very topmost ledges. This tropical 
fern is known from only a few stations in Florida. 

The next use to which I put my rented car was another trip 
to Lake Panasoffkee, south of Ocala, where I had failed to locate 
Asplenium cristatum (Hemlock Spleenwort) the preceding 
year. This time, with the aid of a sketch map by John Beckner, 
I had better luck, finding a few plants on low boulders in that 
elevated part of the hammock south of the outlet. The fern was 
So scarce, however, it is not surprising that my ‘‘wild goose 
chase’? from Sarasota in 1955 was doomed to failure. It is about 
this beautiful fern that Dr. Small wrote in his ‘‘Ferns of the 
Southeast,’’ ‘‘ As it is an attractive and rare fern, it might be the . 
unfortunate victim of attempts to exploit it commercially.’ 

Looking up John Beckner in St. Petersburg, I again joined 
him in an interesting expedition, this time to the Pineola grottoes 
near Istachatta. Along with Buzzards’ Roost, this is one of the 
few localities once famous for a lavish display of Asplenium 
Curtissii, but here again the fern has practically disappeared, 


6 AMERICAN FERN JOURNAL 


due largely to extensive quarrying operations. After many 
hours of search, we finally found one tiny plant halfway up a 
vertical rock wall, not at all representative of the species. But 
there were other interesting and beautiful ferns in the area in- 
cluding the luxuriant Brittle Maidenhair (Adiantum tenerum), 
the giant Bristly Shield Fern (Thelypteris setigera), the dimor- 
phie Florida Shield Fern (Dryopteris ludoviciana), and the rela- 
tively uncommon Thelypteris tetragona, with its dark green 
fronds and pinna-like apical lobe of the leaf blade. 

Following a brief visit with relatives near Sarasota, I flew 
back to Washington, stopping en route at Jacksonville to see 
Mrs. W. D. Diddell, a life member of the American Fern Society. 
There was time for a brief drive to Fort George Island, where 
I saw the odd-looking fern ally Psilotum nudum (Whisk Fern) 
growing beneath cedar trees near Rollins Sanctuary. The walls 
of the old slave houses on the island were covered with the tube- 
rous form of the Sword Fern, Nephrolepis cordifolia. 

In November 1957, instead of flying I decided to drive to 
Florida, to be more independent as far as fern exploration was 
concerned. Stopping at Jacksonville en route, I looked up Mrs. 
Diddell again and obtained information regarding rare Florida 
ferns, especially those in the Miami and Homestead areas. 

At Gold Head Branch State Park I found the relatively un- 
common Hypolepis repens (Spring, or Beaded, Fern) locally 
plentiful along the erystal clear stream at the bottom of the 
ravine. 

Continuing to Gainesville, I spent a rainy morning examining 
herbarium sheets at the University of Florida, which proved to 
be time well spent, especially in locating the St. Johns’ locality 

_ for ferns at Indian Fields Ledges in Sumter County. 

At Orange Lake, south of Gainesville, I looked up Don Me- 
Kay, who showed me the interesting Twin Sinks where formerly 
there was a fine display of Asplenium Curtissii. With the aid of 
ropes we ventured into the depths of these vertical shafts, and 
here for the first time I saw A. Curtissii approaching its typi- 
eal form, although the plants were extremely scarce. 


Forma Rarities 7 


The next day, with the help of a local guide, I found my way 
to the Indian Field Ledges, north of Wahoo in Sumter County. 
Ilere on low boulders and ledges in an elevated portion of the 
hammock I encountered a profusion of ferns of the genus As- 
plemum. A. abscissum, A. verecundum and A. heterochroum 
were plentiful in this area, and at one point I felt confident that 
I had finally stumbled across a typical station for A. Curtissii, 
since the St. Johns had reported the latter from this locality. 
Later study of the specimens collected indicated that they were 
not typical Curtissii, but that the roa sgn consisted of 
two apparent hybrid Aspleniums. One more finely cut of 
the two, closely resembled A. Curtissii. se sere less finely cut, 
resembled (rather surprisingly) the extremely rare A. plenum 
found many years ago at Lecanto Cave by Edward St. John. 

At the time of my 1957 visit I was not aware of this distine- 
tion and supposed the entire lot to be A. Curtissii. Not until I 
revisited Indian Field Ledges in 1958 and collected more speci- 
mens of these ferns did I suspect that two different species of 
Asplenium were represented, John Beckner concurring in this 
supposition. On my return to Washington, I showed these plants 

. V. Morton at the Smithsonian Institution. Comparing the 
less finely cut form with the single specimen of A. plenum in the 
National Herbarium, he agreed that there was a resemblance. 
Sample specimens of both ferns were then sent to Dr. Warren 
H. Wagner at the University of Michigan and to Mrs. Charles 
W. Crane at Summit, N. J., for microscopic spore study. Their 
investigations and reports to date have indicated that both ferns 
reflect a condition of allopolyploidy and are probably hybrids. 
What parent ferns are represented presently remains a mystery, 
although a parent must almost surely be A. verecundum. This em- 
phasizes the fact that at present little is known about Florida 
Asplenium hybrids, and this situation must prevail until a seien- 
tifie cytological study can be made of living plants of all related 
species 

At Tampa, Ralph Garrett, local fern collector, one of two 
brothers who have discovered many new stations for Florida 


8 AMERICAN FERN JOURNAL 


ferns, directed me by a sketch map to one of his brother Ray’s 
recent ‘‘finds’’—Dicranopteris flexuosa (Net Fern) about 15 
miles southeast of the city. Although Dr. Small listed only one 
locality for this fern in the United States (Mon Louis Island, 
Alabama, where it was discovered in 1913)," Maurice Broun 
states that it was subsequently extirpated at this spot.? Miss 
Mary L. Singletary reported the first discovery from Florida (in 
1947) at a point near Kissimmee, Osceola County. These plants 
were in poor condition and the station was described as a pre- 
carious one, exposed to storms and high water.’ I found little 
difficulty in locating the colony of Dicranopteris flexuosa south- 
east of Tampa, where it forms a luxuriant thicket in a strictly 
local area.t Alongside it in profusion grows the handsome Stag- 
horn Club Moss, Lycopodium cernuum, resembling a miniature 
Christmas tree. 

Continuing southward, I traversed the Tamiami Trail to Mi- 
ami to search for the rarer ferns of south tropical Florida. At 
Miami my efforts to find Aspleniwm biscayneanum in the vicin- 
ity of Brickell Hammock were unsuccessful. The entire area is 
now built up as an integral part of the city, and this rare fern 
(a probable hybrid between A. dentatum and A. verecundum) 
has disappeared from the lime rock formation along the sea wall 
where once it was found in limited quantities. 

Along the Coral Gables Canal I found the grasslike Sphe- 
nomeris clavata locally plentiful under the banks of the south 
side, with Thelypteris serra (Parchment Fern) occasional, 
though rare, in the vicinity. 

At Matheson Hammock, south of Miami, now a state park, I 
saw Asplenium dentatum growing sparingly along the sides of 
open lime sinks. 


In the vicinity of Naranja, near Homestead, Mr. Fred Fuchs 
showed me some interesting ferns in Sykes Hammock, of which 
1 Ferns of the Southeast; 329, 330. 1938. 


2 Index to North American Ferns, 60. 1938. 
Tuts JournAL, 40: 176. 1950. 


Forma Rarities 9 


he is the owner. Here I saw for the first time the decorative 
wavy-edged Filmy Fern, Trichomanes Krausii. 

With Fred Fuchs, Jr., I explored Timms Hammock nearby in 
an effort to find the extremely rare Stenochlaena Kunzeana 
(Holly Fern). His teenage son asked for a description and 
promised to keep a sharp lookout inside the walls of the open 
lime sinks which honeycomb the area. Rougher going could 


clutching at you from all angles and giant spider webs whip- 
ping across your face at frequent intervals, the hammock floor 
would suddenly give way without warning and down you would 
go up to your armpits in a treacherous semi-covered lime sink. 
Although Fred had found the Holly Fern at Timms Hammock 
several years previously, no sign of it appeared after hours of 
search. We were about to give up when the bright-eyed young- 
ster fii pilantly called us over to a small sink where he had 
spotted half a dozen specimens of the prize! 

With the same father and son as guides, I was shown dense 
masses of Sphenomeris clavata along the edges of lime sinks near 
the Homestead Air Base. We concluded the day searching for 
(and finding) a real rarity, Trismeria trifoliata, discovered as‘a 
genus new to the United States by Ray Garrett of Tampa in 
1953. Along the Card Sound Road southeast of Florida City 
this fern grows luxuriantly in close proximity with Blechnum 
ylang, which the sterile fronds of Trismeria somewhat 
resemble. 

Before leaving the Homestead area I tried to unravel the mys- 
tery of the ‘‘Rare Tectarias,’’ without success. This concerns 
the discovery many years ago, in Hattie Bauer Hammock, of the 
extremely rare Tectaria Amesiana and T. coriandrifolia. Dr. 
Wherry had written me that the lime sinks where these ferns 
used to grow had been taken over by the Fennell Orchid Jungle, 
and that the ferns had long since disappeared. John Beckner 
confirmed this, saying that the west end of the hammock, where 


* THIs Journan 45: 52. 1955. 


10 AMERICAN FERN JOURNAL 


the rare Tectarias were formerly found in a large sink, is now 
part of the Orchid Jungle, and that the ferns may have vanished 
in the 1930’s following the effects of a long continued drought. 
This supposition, however, regarding the date of their disap- 
pearance is not strictly accurate, since there are on record (at 
Smithsonian) herbarium sheets of both Tectarias collected in 
November, 1940, by the late F. N. Irving of Washington, D.C) 
As far as I can ascertain, however, this is the last known col- 
lection. Mr. Irving once described to me his experience in col- 
lecting these ferns ‘‘in the middle of a dense jungle’’ and how 
he had tried to rediscover the locality at a later date, without 
suecess. Roy Woodbury of the University of Miami, now in 
Puerto Rico, wrote me that he had never been able to find these 
ferns after many years of search. Maurice Broun reported 
thriving colonies of both species in February 1935, of which he 
collected some fine representatives. 

While visiting relatives on Longboat Key near Sarasota, Ai 
discovered gigantic specimens of Psilotum nudum (Whisk Fern) 
epiphytic on trees at the northern end of the island. Some of 
these plants measured approximately twenty inches long, greater 
than the upper limit given by Small and much larger than the 
terrestrial specimens I had seen on Fort George Island near 
Jacksonville. 

On my return trip I had planned to join the Garrett brothers 
of Tampa for a full day of exploration. Unfortunately, heavy 
tropical storms earlier in the day prevented our leaving until 
afternoon. But even in this brief interval I had the pleasure of 
being shown at least two rare ferns by these amateur botanists 
who have made such amazing discoveries of new locations for 
Florida ferns. On a rocky abutment to a railroad bridge along 
the Hillsborough River near Zephyrhills I saw Asplenium pum- 
lum, one of Ralph Garrett’s surprising discoveries while looking 
for additional stations of A. auritum, Then, west of Dade City, 
in a swampy hardwood hammock we visited a typical location 


5 Tuts Journa, 45: 10,11. 1955. 


Forma Rarities a 


for the graceful Thelypteris resinifera (T. panamensis), on 
of the most beautiful of marsh ferns, which Dr. Small has 
termed one of Florida’s rarer plants. The Garrett brothers 
have found a number of new locations for this uncommon fern. 
Recently, in November 1958, I drove down to Florida once 
again. En route to Gainesville I stopped at Gold Head Branch 
State Park, finding the xerophytic Selaginella arenicola locally 
plentiful in high sandy locations up ties Sheeler Lake. 

At Gainesville Dr. E. S. Ford, of the University of Florida, 
joined me in searching for a spot Nye of High Springs known 
to the St. John brothers as ‘‘ Fern Cave,’’ where they had found 
some rare Aspleniums identical (or at least similar) to those 
at Lecanto. A few specimens from this locality may be seen in 
the University herbarium. Stopping off at Buzzards’ Roost 
found the ferns there in poor condition due to the sedation 
previously described. In this area Dr Ford pointed out the dec- 
orative Pteris multifida, the so-called Huguenot Fern, which I 
had overlooked in 1956. Our search for Fern Cave proved un- 
successful. Although we followed a number of leads and clues 
in the supposed general region, no local residents recognized a 
cave by that name, nor could we find any such formation after 
several hours of search. 

Continuing southward I revisited Indian Field Ledges in 
Sumter County and found the unidentifiable Aspleniums still 
in evidence on the low boulders. This expediton was not with- 
out its hazards. A narrow wood road led to the spot, but recent 
rains caused soggy depressions in places where I nearly got mired 

own. At one point a herd of semi-wild bulls roaming the 
hammock land did not add to my peace of mind. Fortunately 
T was still driving, as I would not have cared to face these ani- 
mals on foot. After I had left the bulls, however, the road be- 
came so impassable I had to travel the last part of the distance 
on foot. Arriving at Indian Field before dusk, I found a good- 
sized coral snake guarding the very ledges I wanted to examine. 
This is the poisonous reptile of the southland whose venom is 
similar to that of the cobra, affecting the nervous system and 


Lys AMERICAN FERN JOURNAL 


usually causing death within 24 hours. But these snakes are for 
the most part rather shy, and after this particular one had dis- 
appeared in a hole, I felt free to continue my search for Aspleni- 
ums with the results mentioned at the time of my 1957 visit. 

Spending the night at Lake Panasoffkee, I arose early in the 
morning to renew my acquaintance with Asplenium cristatum, 
the Hemlock Spleenwort. A new road south of the outlet came 
perilously close to the limited stand of this fern which I had 

succeeded in finding after so much difficulty in 1956. It now 
appeared as if this station had been adversely affected by the 
road construction and felling of trees. In the vicinity a large 
cottonmouth moccasin raced across my path. Oddly enough, this 
was only the second poisonous snake which I had encountered 
ee all my wanderings in Florida—two poisonous snakes in 
wo successive days. I have yet to see the dreaded diamond- 
irae: 
_ Next y pace to explore the area north of the outlet, hay- 
ing heard just recently that A. cristatum had been found here 
also. Making my way across open shaded pasture land with 
masses of low boulders, I found nothing of interest for some 
distance. Suddenly I noticed A. abscisswm on the rocks, and 
then, much to my amazement, A. cristatum too began to appear 
in considerable abundance. Sometimes these two ferns carpeted 
the same rocks, even occupying identical crevices on. occasion, 
but after a careful search I could detect no sign of a hybrid. 
Le is one place where A. cristatum appears to be locally plenti- 
ul. 

At St. Petersburg John Beckner joined me on an all day trip 
halfway across the state in a search for Meniscium at Arbuckle 
Creek, Highlands County, northeast of Avon Park. After 
several hours of exploration along the low swampy areas border- 
ing the sluggish stream bed, we encountered a considerable 
number of immature plants of M. serratum, but none of the 
strikingly conspicuous fertile fronds. Many of the ferns were 
broken and deformed, giving the appearance of having been 
eaten by cattle, thus possibly explaining the lack of mature 


FLoripa Rarities 13 


plants. JF inally, however, we did manage to find two typical 
fertile fronds of this rare and interesting fern. Once common in 
the Lake Okeechobee area, it has long since disappeared with 
the drainage of the swamps. All along the mud flats of Ar- 
buckle Creek we saw many small fertile leaves of the Floating 
Fern, Ceratopteris pteridoides, sprouting out-of the alluvial 
soil, the spores having been deposited when the waters receded 
after flood stage. 

: Before leaving St. Potenbure I paid a visit to the site about 
25 miles north of the city where John Beckner in 1952 found 
a few tiny stems of the tropical Curly Grass, Schizaea (Actino- 
stachys) Germani.® My efforts to uncover this ‘‘needle in a hay- 
stack’? were to no avail. Although Beckner returned to this 
spot a number of times folowing his original discovery, he was. 
able to find only one additional plant. 

Again I journeyed to Miami via ‘‘The Trail.”’ This | time, 
with directions from John Beckner, I located the small hammock 
near Rickenbacker Causeway in the very center of the city where 
he had found several plants of the almost extinct Asplenium 
biscayneanum, Sure enough, in one of the small sink holes in 
the dense woods on the flat limestone surface I saw a single 
specimen of this’ probable hybrid growing in company with A. 
dentatum. 

On the grounds of the University of Miami, Dr. Taylor Alex- 
ander, head of the Department of Botany, showed me several 
thriving plants of Pteris grandifolid, a gigantic tropical bracken, 
found by Roy Woodbury near “Cutler in’ 1952. Mr. Woodbury 
had written me about’ this discovery, mentioning that it was a 
fern new to the United States, but that it had not yet been offi- 
cially reported or published. 

Continuing to the Naranja area near Homestead, I once more 
looked up Mr. Fred Fuchs, Sr., who showed me~ Thelypteris 
(Dryopteris) sclerophylla in yhien Hammock, a dark green 
leathery fern found in the United States only in this ‘hammock.? 
In a nearby lime sink I saw two filmy ferns .in close proximity 


6 Tu1s JourNAL, 43: 124,125. 1953. 


14 AMERICAN FERN JOURNAL 


—Trichomanes punctatum and T. Krausii—the former on the 
vertical rock wall, the latter on decayed logs at the edges. In 
his greenhouse, Mr. Fuchs showed me two of the rare epiphytic 
Strap Ferns, Campyloneuron angustifolium, originally from 
Timms Hammock close by, where it may now be extinct, and 
C. costatum, from the Big Cypress, near Deep Lake, in Collier 
County. 

In the Naranja region I heard a rumor that someone had 
accidentally encountered a single plant of the Fragrant Maid- 
enhair, Adiantum melanoleucum, in the Everglades National 
Park while searching for other plants. This rare maidenhair 
was believed to have become extinct in Florida many years ago. 
There was not sufficient time for me, however, to explore that 
distant area. 


One of the rarest filmy ferns, Trichomanes lineolatum, has 


Hammocks. With specific directions for the Ross Hammock 
location from Roy Woodbury in Puerto Rico, I carefully searched 
the area to no avail, nor did I see filmy ferns of any kind in this 
region. 

On the return trip I visited the Deering Hammock near Cut- 
ler, south of Miami, having obtained permission to explore this 
interesting place, The head caretaker showed me about the vast 
wooded estate, with its many roads and trails. From a pterido- 
logical standpoint it is famous chiefly for the lavish and abun- 
dant display of Asplenium dentatum on almost every lime out- 
crop and lining the walls of the sinks. This protected sanctuary 
remains, except for Matheson Hammock and a few other re- 
stricted areas to a much lesser degree, the one place in the United 
States where this Asplenium still grows in abundance. 

In a moist ravine at the Deering Estate, I happened to spot 
a few young and immature plants of the Giant Bracken, Pterts 
(Litobrochia) tripartita, the first time I had seen this fern in its 
native habitat. Although once common in many parts of Florida, 


7 TuIs Journau, 41: 86,87. 1951. 


Hyprips IN GYMNOCARPIUM 15 


it now appears extremely scarce. Previously, in Mrs. Y. C. 
Lott’s garden in Miami I had seen a fully developed fruiting 
specimen of this handsome plant, taller than a man. 

In conclusion, the above represents the story of random ex- 
plorations by a northern visitor with limited time available. 
Local fern students are probably familiar with other places 
where some of these rarities may be found. Any information 
regarding these would be much appreciated by the author. With 
the advance of civilization, and all that it entails, many of the 
rarest Florida ferns may be doomed to extinction unless in some 
way they can be preserved for future generations to see and 


enjoy. 
382 37th Pu. S. E., WasHineTon, D. C. 


Hybrids in North American Gymnocarpiums 
EvizasetH Eicustept Roor 

The relationship between the two North American species 
of Gymnocarpium, G. Dryopteris and G. Robertianum, has long 
remained a topic of speculation. 

These two cireumboreal species occur together in many 
northern localities, though G. Dryopteris is often found alone at 
somewhat lower latitudes and in a greater variety of habitats. 
The less tolerant @. Robertianum most commonly grows in a limy 
situation, on talus slopes or in rocky woods, but it has also been 
collected in swamps, where there are no rocks. In some habitats, 
populations including both species frequently oceur; it has been 
previously reported that within these populations are forms that 
are difficult to place with either species. Se 

Lawalrée (1950) maintains that the presence of an intermedi- 
ate form supports his view that G@. Robertianum represents a 
variety of G. Dryopteris, being adapted to a calcareous habitat. 
He cites Koltz (1877-78) who, without proof according to La- 
walrée, stated that the intermediate results from hybridization. 
Intermediates have also been turned up in North America. The 
intermediate characters found have concerned glandularity and 
leaf shape; G. Dryopteris has a very slightly glandular rachis 


POR Eee 
oF FB ee 
~ 8 Pago gs See 


Hysrips IN GYMNOCARPIUM ae, 


and a broadly triangular frond, whereas G. Robertianum is 
densely glandular on the rachis and lower leaf surface, and has 
a narrowly triangular frond. Hultén (1941) reports: ‘‘Some 
of the specimens of G. Robertianum referred to are very slightly 
glandular, and at the same time have a general appearance close- 
ly resembling tht of G. Dryopteris, while others are quite typical 
*. Robertianum. One specimen, Anderson 2609, from Circle 
Hot Springs, is very doubtful. The form of the frond ap- 
proaches strongly that of G. Dryopteris, the first pair of pinnae 
being much larger than the rest. It has, however, scattered 
glands on the stipes and on the lower surface of the leaves as 
well as the dark green color of G. Robertianum ...’’ The plant 
recognized by Tryon (1939) as Dryopteris Linnaeana forma 
glandulosa is more or less intermediate in glandularity also. 

A few mixed populations of the Gymnocarpium complex have 
been found in Michigan, at the tip of the Lower Peninsula and, 
more commonly, in the Upper Peninsula. In July, 1957, such a 
population was found by Dr. W. H. Wagner, Jr., in Marquette 
Co. The plants were growing in woods on rock cliffs about six 
miles northwest of Ishpeming. A collection was made by the 
discoverer in company with E. G. Voss and D. J. Hagenah, and 
these plants were turned over to me for study. Of these twenty- 
six specimens, several, on the basis of frond shape, eould not 
be placed with either species. Upon microscopic examination 
of these specimens, it was found that the degree of glandularity 
also varied according to frond shape. Further study of the 
variation was made possible through the help of C. V. Morton, of 


Microprojection drawings of upper epidermal cells of cleared aoe 
showing variations in size and anape a-f, GYMNOCARPIUM DRYOPTER 


M=p, Marquette Co., Mich.; 4q, Fernald, Long, & Fogg 1130, 
Sieecune > Novena (US); r. Fernald, Long § Fogg eG Lord 
nd Lady Cove, Newfoundland (US). 


18 AMERICAN FERN JOURNAL 


the United States National Herbarium, in lending to me ad- 
ditional materials and by his helpful suggestions 

The principal characters used to distinguish these species from 
one another have been those already mentioned of frond cutting 
and degree of glandularity. It is generally stated (Fernald, 
1950, Morton in Gleason, 1952) that the fronds and rachises of 
G. Dryopteris range from glabrous to rarely very slightly glan- 
dular. However, upon close examination of specimens from 
various localities it was found that every individual of this spe- 
cies examined had glands. Though few, they were always pres- 
ent, at least in the axils of the pinnae. The degree of glandular- 
ity still remains a good distinguishing characteristic, however, 
as the fronds and rachises of G. Robertianum are much more 


A B c 
* 
Le 
7 
are = 
Le 
“s 
a 
‘a - 
” EUG ie 
he 
+ 
x o are 
+ Pet fe 
te 
+ 
= 
te 
¥ le 
+ 
“ 
-7 
ey 
+ 
Le - 
-° 
my “ 
ak 
q 
= 
+ 
Le 
3 lo 
ag ~ 
+ 
E~a te 
+ &b t 
a + 
- + 
sd SEE 


7 
Fia. 1. Diagram Suowine THE SILHOUETTE OF A RACHIS SEGMENT BETWEEN 
A 


N); B: M 
Co., Micniaan) ; C: G. Dryopteris oon 4708. epoca Ae MICHIGAN ) 


Hysrips IN GYMNOCARPIUM 19 


densely glandular than those of G. Dryopteris. (Fig. 1) The 
difference in the frond cutting is due mainly to differences in the 
size and shape of the lowermost divisions of the blade. In G. 
Dryopteris, these are nearly as long as the terminal part of the 
blade; they are asymmetrical, the lower side having elongate 
pinnules. In comparison, those of G. Robertianum are about 
half as long as the terminal part, and they are more nearly 
symmetrical. As was mentioned earlier, specimens intermediate 
in these two characters were included in the Michigan collection, 
indicating the possibility of hybridization between these plants. 
Additional characters were required in order to substantiate 
this possibility. Therefore, a study of spores and epidermal cells 
was made. e Michigan specimens were divided into three 
groups, representing the two species and the intermediate. Mi- 
croscope slides were made of the spores and cleared pinnules of 
each group. The latter were prepared by clearing in sodium 
hydroxide and then staining with tannic acid and ferric chloride. 

Examination of the spores revealed only a subtle difference in 
structure, the perispore of G. Robertianum being a little more 
roughly sculptured than that of G. Dryopteris. Of major im- 
portance in the spore study is the fact that the majority of the 
spores of all intermediate specimens were found to be abortive. 
Subtle dissimilarities were noted also in the shape of the epi- 
dermal cells of the two species, those of @. Robertianum being 
longer, on the average, and with less sinuate margins than those 
of G. Dryopteris. Those of the intermediate were found to be 
large like G. Robertianum, but with the more sinuate margins of 
G. Dryopteris (Pl. 1). 


DISCUSSION 
The fact that the spores of intermediate forms are abortive 
indicates that the two presumed parental plants are distinct 
species. If they are mere varieties, as has been claimed, their hy- 
brids would most likely be fertile. The more roughly sculptured 
perispore of G. Robertianum provides a trait by which to sepa- 
rate the species. 


T I 7 5 LATE 2 
AMERICAN FERN JOURNAL VoLUME 51, PLATE 2 


as 


MERE aes 


ha 


¥ 


, GYMNOCARPIUNE Dryopreris X G. ROBERTIANUM, 


PROBABLE Hyprip 
MARQUETTE Co., MicHigan, Voss 4707 


Hyprips In GyMNocARPIUM vA 


The intermediate character of the leaf cutting, glandularity, 
and epidermal cell shape and spore abortion provide a sound 
argument for the hybrid origin of these intermediate specimens. 
A summary of the intermediate characters is presented in 
Table I. 

No other collections that I have seen from North America are 
clearly of the hybrid type. Several possible intermediates were 


nacana forma glandulosa Tryon. These bore a resemblance in 


TABLE I 
Character G. Dryopteris Intermediate G. Robertianum 
Frond Broadly triangular: Two lower pinnae Narrowly triangular: 
Shape two lower pinnae about one-half to two lower pinnae 
about three-fourths three-quarters as about one-half as 
as long as remainder | as remain- long as remainder of 
of frond or longer. der of frond frond. 
Indument Very sparse Moderate Very dense 
of frond 
an 
rachis 
Size of 100 u 122 uw 122» 
epidermal 
cells 
(average 
greatest 
length) 
a Margins deeply Margins moder- Margins shallowly 
epidermal _ sinuate. ately sinuate. sinuate. 
ells | 
(average) 
Per cent About 1 48-85 About 5 
abortion 


of spores 


22 AMERICAN FERN JOURNAL 


leaf-cutting and glandularity to the Michigan hybrids, but the 
spores are immature, and the epidermal cells were not examined, 
and so the true position of this plant in the Gymnocarpium 
complex remains unknown. 

There is a need for the investigation of additional collections 
of these species, in order to determine the frequency of hybridi- 
zation between them. A more detailed analysis of the habitats 
of each may also shed further light on the problem. It is sug- 
gested that observers in regions where G. Dryopteris and G. Ro- 
bertianum co-exist remain watchful for other hybrid populations. 

I wish to express my appreciation to Dr. Warren H. Wagner, 
Jr., for his guidance in this work and for his generous extension 
of laboratory facilities.| My thanks go also to Mr. G. M. Christ- 
man for his helpful advice on the illustrations. 


LITERATURE CITED 
FERNALD, M. L. 1950. Gray’s Manual of Botany. American Book Company. 
Boston, Mass. 
Morton in Gueason, Henry A. 1952. The New Britton and Brown Illus- 
trated Flora oe choca aan United States and Adjacent Canada. Lan- 


er, Pa. 
Huurén, Eric. 1941. Flora of Alaska and Yukon. Bot. Mus. Lund, Sweden. 
LAWALREE, ANDRE. 1950. Pteridophytes, in Flore Générale de Belgique. 
Jardin Botanique de . ’Etat, Brussels. 
TRYON, oF 1939. Notes on the Ferns of Wisconsin. Amer. Fern 
Journ. 29: 


e Mrs. Root’s study, Olga Lakela and I have found ee ——— 


ay of an intermediate Gy sae ian in June 18, 1959, on shaded ¢ 
below the observation tower a sh Riv St. Douis jill gets eae 
(Wagner 9034.5a). The leaf shape and pilin pa appear to be inter- 


mediate as deseribed by Mrs. Root, and the spores are conspicuously irreg- 

ular in form. 

: It might be mentioned that even though these plants are sagglan judg- 

ing by spore abortion, they can still form large populations by vege etative 
on ace 


reproduction, ount of their long, horizontal, undergr oa rhizomes, 
similar to those formed rosia inte ealens and Equisetw x 
trachyodon. O other hand plants with only approximate offsets, h 
as Polystichum acrostich Lon and most Dryopteris h prids, 
orm only one or a few suall eid 1 e orig 
inal living plant of ostichoic oe has been transplanted to 
his garé Hagenah because it was in dange ahaha 


Mr. Dale 
by eattle and sheep ; it will now be are available for cytological stu 
W. H. WAGNE 


VARIATION IN DRYOPTERIS 23 


The Problems of Variation in North American Dryopteris! 
Donaup Brirton 


Recent cyto-taxonomic work has helped us considerably to 
understand the evolution and classification of the North Ameri- 
can species of Dryopteris. The initial impetus for these studies 
came from Manton (1950). Later studies by Britton (1953), 
Manton and Walker (1953), Wagner (1955), and Walker (1959) 
have furthered our knowledge of this genus. It is now possible 
to exclude such species as Thelypteris palustris, T. Phegopteris, 
and T. hexagonoptera from Dryopteris, as is done in the New 
Illustr. Flora of Britton and Brown, both on the basis of their 
chromosome numbers departing from the basic number of 41 
in the genus and also on their morphology (See Table). Having 
done this, it is then possible to focus attention on Dryopteris. 


The novice soon learns to ane Dryopteris marginalis, D. 
Goldiana, D. cristata, and D. Clintoniana, although together 


cation. The truly difficult plants to identify are those that be- 
long to the spinulosa complex, and especially those forms which 
apparently have hybridized both within and without this com- 
plex. Examining the chromosome numbers of these entities 
(See Table) we find that marginalis, Goldiana, and intermedia 
all have 41 chromosomes, which is apparently the basic or 
monoploid number for the genus. These species all have a regu- 
lar and normal meiosis and apparently reproduce sexually as 
one would expect. Dryopteris spinulosa var. spinulosa is in 
part a sexually reproducing tetraploid (n = a2). as is D. cris- 
tata, and D. Clintoniana is a hexaploid (n = 123). All these 
Species have a regular meiosis with normal pairing of bivalents 
at meiosis I. 


1The author wishes to acknowledge Siesta! oe ree 
by an award from the Sigma Xi RESA research grants, and to thank Dr. 
Rolla M. Tryon, Jr., and Prof. F, H. Montgomery for their interest and 
help in lasdnomie matters. 


24 AMERICAN FERN JOURNAL 


For more than six years now, Dr. Stanley Walker, of the 
University of Liverpool, has been studying the cytology of this 
group and has been synthesizing hybrids between the different 
entities. The proof of the hypothesis that D. Clintoniana is an 
allopolyploid between D. cristata and D. Goldiana will have to 
await the publication of his results. aah ee hybridization ex- 
periments together with some genome analysis will show how 
many basic genomes or sets of chromosomes are involved in the 
species of Dryopteris. The entities Dr. Walker has studied have 
been shipped to him from North America and hence are removed 
from their native habitat, and their natural morphology may 
have changed even if they are grown by experts under ideal 
conditions. Walker has not had the opportunity to examine his 
material growing in large numbers at any one location or over 
its natural range. 

My own work has taken full cognizance of Walker’s prolonged 
study of this group. I have attempted to examine and collect 
large numbers of these entities in an effort to delimit the genetic 
and ecological variation. Fortunately, there are many woods 

swamps near Guelph that abound in Dryopteris, so I am 
well situated to study the variation in the genus. 


One should make a distinction between occasional rare hybrids 
such as the one between fragrans and intermedia reported by 
Tryon (1942) and some of those entities that make up a sub- 
stantial percentage of our fern flora. The latter group are 
extremely successful and are even ‘‘end-points’’ from an evolu- 
tionary standpoint, whereas the former tend to be curiosities 
such as some horticultural hybrids. The former can be plotte 
only as occurrences, whereas the latter may have a definite 
range. 


The first observation that might be made is that although D. 
marginalis (n = 41) is one of our more common species and is 
usually found in association with other Dryopteris species 
(especially intermedia), it apparently has hybridized with the 
others very infrequently. Although six different Dryopteris 
hybrids involving D. marginalis are noted by Chandler (1948), 


VARIATION IN DRYOPTERIS 25 


this is not an indication that marginalis commonly hybridizes 
with other species of Dryopteris. More probably, it is an out- 
come of the broad distribution of marginalis, together with its 
distinctive phenotype which is expressed in a modified but 
recognizable form in its hybrids 


The second observation is the rather astonishing frequency of 
triploids that one encounters in the field. This is more apparent 
if one considers that each triploid has arisen from a separate 
fertilization and is a final product with no means of sexual re- 
production. At one location, I collected seven specimens for 
cytological study; the three that looked like intermedia were 
diploid and the other four were triploids. At another location 
Six specimens were collected; three were diploids and three 
triploids, which shows how frequently triploids may be encoun- 
tered in this area. 


Since typical intermedia is apparently a sexually reproducing 
diploid species with a normal meiosis, it would seem that if one 
looked at a very large number of entities that approached inter- 
media in morphology it might be possible to determine the ex- 
tent of variation in this species. That is, instead of trying to 
key specimens down to intermedia, it should be possible to use 
diploidy as a basic criterion and group all diploids having an 
intermedia-like appearance as intermedia, since the only other 
diploids one encounters locally are marginalis and Goldiana, 
which will not be confused with intermedia. I have attempted to 
do this with a large number of plants and the only difficulty that 
I can visualize arising is that I may be putting another diploid 
species, e.g., a diploid var. americana (or campyloptera), in this 
grouping. However, I believe this possibility will be eliminated 
when all the specimens are examined critically and compared 
with one another. 

Having conveniently lumped the diploids into intermedia for 
the taxonomists to determine the limits of variability, one is left 
with a large group of triploids and a large group of tetraploids. 
Considering the tetraploids next, these are apparently true 
spinulosa. The lower inner pinnules next to the rachis are 


26 AmemrICAN FerN JOURNAL 


longer than the next pinnules and, most important, the indusium 
is glabrous. None of the 29 local specimens for which I have 
determined the chromosome number could be considered to be 
var. americana (or campyloptera) on the basis of the spacing of 
the basal pinnules. It will be necessary to try to establish the 
limits of variation of this group also. 


The last group are the triploids (including var. fructuosa) 
which as one would expect, are quite variable, although many 
approach spinulosa, except for the fact that the indusium is 
glandular. The chromosome numbers of 30 different specimens 
of these have been determined. As yet, it is too early to say 
whether the variability is due to reciprocal crosses exhibiting 
matrocliny, or of more than just the two entities spinulosa and 
intermedia being involved, as suggested by Manton and Walker 
(1953). From field studies, the hypothesis that only two paren- 
which, as one would expect, are quite variable, although many 
locations there is a preponderance of only one of the putative 
parents. In the dryer locations, especially rich deciduous woods 
in which Adiantum pedatum is present, intermedia and the 
triploid predominate, whereas in the wetter swamps spinulosa 
and the triploid are more frequent. 

In some of the local swamps Boottii has been collected and, as 
Manton and Walker (1953) reported, this is a triploid hybrid 
with little or no pairing of the chromosomes. Presumably it has 
two sets of chromosomes from cristata (4x) and one set from 
intermedia (2x). Confirmation of this will have to await Dr. 
Walker’s findings. Also found occasionally in these same 
swamps are some intermediates between cristata (4x) and Clin- 
toniana (6x) which are pentaploid (5x) as one would: expect. 

Out of the hundreds of marginalis specimens looked at in the 
field only one putative hybrid was located between this and 
intermedia. As one would expect from such a cross, 82 unpaired 
chromosomes were seen at meiosis, 41 from intermedia and 41 
from marginalis. 

As far as possible in this study I have attempted to study the 
plants in the field as well as in the laboratory. To this end, I 


VARIATION IN DRYOPTERIS par 


have placed Hartley Metal Labels with appropriate numbers in 

the ground beside certain selected plants so that I may return 

to the same plant another year if need be. Also, it will be possi- 

ble to obtain four or five fronds in the Fall for herbarium speci- 
ted i 


their natural Sonar seats The chief difficulty will 


in inaccessible eaiea 


It would seem that var. americana (or D. campyloptera) is 
not present in the localities that this author has investigated so 
far, and that it has not been a factor in the production of the 
triploids. Both the author and Prof. F. H. Montgomery have 
made extended search for this entity. 


Of the various characters used in keys for determining speci- 
mens, it would seem that if the innermost lower pinnules are 
shorter than the next succeeding pinnules that one is moving 
towards intermedia. The other good key character is whether 
the indusium is glandular or not. Further elaboration on these 
points will be required by competent taxonomists. 


Chromosome numbers of 110 different plants have been de- 
termined. The onset of meiosis was on May 29 in this area and 
specimens suitable for cytological study were collected for the 
following two weeks. Studies on the variation in the genus were 
made both before and dfter suitable meiotic material was col- 
lected, and approximately 100 additional collections were made. 
The longer a student studies the different plants in the field the 


his material. Accordingly, the sample collected at any one loca- 
tion may be reduced to five or six plants although the investiga- 
tor has looked at say a hundred intermedias at that one loca- 
tion. He rejects 94 of the intermedias as obviously falling into 
this species on the basis of his previous collections and previous 
chromosome number determinations, and consequently can con- 


28 AMERICAN FERN JOURNAL 


chromosome number determinations, and consequently can con- 
centrate on the remaining six plants. 

Sinee it is unlikely that triploid hybrids growing many yards 
apart, each arising from a separate fertilization, will arise in 
thick woods far removed from their parents, it is often necessary 
to return to the same locality to attempt to find the missing 
putative parent. If it cannot be found, as happened at one loca- 
tion, it would seem logical to assume that it has become extinct, 
although the age and longevity of the triploid hybrids is quite 
unknown to this investigator. Although unable to reproduce 
sexually the triploids can reproduce by vegetative means, but 
the speed of this vegetative colonization is unknown. 

Another observation that might be made is that some plants 
that have only one or two fronds, in contrast to strongly grow- 
ing plants with five or six fronds, show more variation in rela- 
tion to the spacing of the innermost lower pinnules than do the 
latter in the same locality. There is also variation in different 
fronds of the same plant. Some intermedias will have a frond 
with the innermost lowest pinnules as long as, or longer than, 
the next succeeding pinnules, but this will not be true for all the 
fronds of the same plant. The cutting of the frond and the 
over all lacy appearance are certainly of help in field indentifica- 
tion. 


The most variable character used in keys seems to be the 
spacing of the inner lower pinnules. For this general area, it 
would seem that the more remote spacing (over 0.5 em.) is found 
on the diploids rather than the tetraploids. Some sterile fronds 
exhibit more variation in this respect than do the fertile fronds 
of the same plant; occasional sterile plants have wide spacing, 
but unfortunately their chromosome number is not known. 

Further work is planned on these problems next year. It will 
be necessary to decide if all the diploids can be assigned to 
intermedia, and it will also be necessary to continue to search 
for var. americana. With the present specimens for reference, 


VARIATION IN DRYOPTERIS 


further collections can be made more and more critically. 


1. D. Thelypteris var. pubescens 


Thelypteris palustris 


2. D. simulata 
Thelypteris simulata 


oe 


co on 


BU Pe Ss Se See 


iD; veboracens 
Thely pteris nov cbariesaatl 


D. disju 

jcebapeerater ts Dryopteris 
D. Robertiana 
Gymnoearpium Robertianum 
D. Phegopteris 

Thelypteris Phegopteris 

D. hexagonopte 

Thelypteris ecnpapouties 
. Spinulosa 

austriaca var. spinulosa 


spinulosa var. 


spinulosa var. fructuosa 
austriaca var. fructuosa 


spinulosa var. americana 
austriaca var. austriaca 


X< Boottii 


D. cristata 


oo 


i. hala | Clintoniana 
. Clintoniana 


. intermedia 
austriaca var. intermedia 


spinulosa var. coneordiana 
austriaca var. concordiana 


35 
35 


41 
41 


pairs and 
singles 


unpaired 


CuromosomMe NuMBERS OF EASTERN Nor?TH AMERICAN DRYOPTERIS 


GAME RO- 
Geo cca PLoIDy 


AUTHORITY 


Britton, 1953 
Wagner, 1955 


Britton, 1953 
Wagner, 1955 


Britton, 1953 
Manton, 19505 


Manton, 1950° 
Britton, 1953 
Wagner, 1955 
Manton & 
Walker, 1953 
Britton, 1960 
M. & W., 1953 
Britton, 19603 
M. & W., 1953 
Britton, 1960 
Walker, 1959 


Britton, 19534 
: ., 1953 
Britton, 1960 


M. & W., 1953 
Britton, 1960 


M. & W., 1953 
Britton, 1960 


30 AMERICAN FERN JOURNAL 


11. D. Filix-mas 82 4x M. & W., 19545 
Doepp, iin 
12. D. celsa 82 4X Walker, 1959 
D. Goldiana subsp. celsa 
13. D. Goldiana 41 2X M. & W., 1953 
14. D. marginalis 41 2X M. & W., 1953 
41 2X Britton, 1953, 
and 1960 
15. D. fragrans var. remotiuscula 


2Number and name of each ce Sebel! Fernald, in Gray’ s Manual, 

tain edition apie ; alternative names are those of Mo rton, in Gleason, 
Ne ritton Pati! wn Illustr: alas ew qd 952). 

“ count n = ne Britton, 1953, should be rejected; the material 
was aE typical Satbec 

4The determination of the specimen is doubtful, and the cytological anal- 
ysis was incomple e. This was presented merely to show some of the problems 
that arise in a first survey of this kind. 

5Counts made from European plants, included here for interest. 


REFERENCES 


Brirton, Donald M. 1953. Chromosome studies on ferns. Amer. Journ. 
Bot. 40: 575-583. 


CHANDLER, A. 1948. Dryopteris hybrids. Amer. Midland Naturalist 40: 
763-773. 


OEPP, W. 1955. ee erzeugte “acta zwischen Dryopteris 
sro -mas (l.) Schott. und D. paleacea (Sw.) C . Planta 46: 70-91. 
ANTON, Irene. 1950. Problems of Cytology and Evolution in the Pteri- 
dophyta. Univ. Press., Cambridge, En sei nd. 


1954. Induced apogamy in Dryopteris aitatat gars aes Gra, 
: its 
tion of the two species. Ann. Bot., N.S., 18 
Tryon, R. M., Jr. 1942. A new Dryopteris hybrid. Amer. Fern. Jour. 32: 
81— 
Waener, Warren H., Jr. 1955. Cytotaxonomie observations on North 
American Ferns. Bieler 57: 219-240. 
Waker, S. 1959. Sonera va studies of some American species of 
pteris. Amer. Fern 4-112 
DEPARTMENT OF Bonasy, Ontario AGRICULTURAL COLLEGE, 
GUELPH, CANADA 


VERNATION OF BoTRYCHIUM 31 


Some New Data on the Vernation Differences of Botrychium 
dissectum and B. ternatum 


W. H. WAGNER, JR. 


No evidence has appeared thus far to indicate that the Ameri- 
can grapefern that resembles Botrychiwm ternatum (Thunb.) 
Swartz of the Old World! is other than that species, so this name 
has been adopted for this plant. It has been discovered that the 
four northeastern North American species of evergreen grape- 
ferns differ significantly from each other in periodicity of leaf 
development and sporangial maturation when growing together 
in the same habitat.2 The first species to start growing in the 
spring (the last week of April and first week of May) is 
Botrychium multifidum (Gmel.) Rupr. and the last species to 
start is B. dissectum Spreng. (the first half of June). Botrychium 
ternatum and B. oneidense (Gilb.) House are intermediate, the 
former preceding the latter to some extent. The same periodicity 
differences between the species are also reflected in sporangial 
maturation in the late summer and fall. So far as we have been 
able to determine, these differences are found wherever two or 
more of the species co-exist. 

Most of the data we have on time of early leaf development in 
these plants were obtained in 1959 during the period June 11-26 
in various counties of southern Michigan. The differences in ver- 
nation between B. dissectum and B. oneidense were also exam- 
ined on July 3, 1959, but no comparisons in the month of July 
of that year were made of B. dissectum and B. ternatum. Ac- 
cordingly, the following year, on July 14, 1960, a now well 
known locality in Midland County, Michigan, where the last 


1 American Grapeferns Resembling Botrychium ternatum: A Preliminary 
Report. sind JouRNAL, 49: 97-103. 

iodi and Pigme sponse in es th Subg. Sceptridium in the 
etthogaices: Tedted States. B Bull. y Bot. Club 87: 303-3 25, 1960. This 
work has been ¢éontinued as a ae periodicity differences between 
closely related species as a par obs . » National Science Foundation Research 
Project. 


32 AMERICAN FERN JOURNAL 


two species grow intimately together was visited (R.2E, T. 16N, 
Sect. 29, NW14, south side of road, along edge of more or less 
open, sandy field with scattered trees and shrubs). In order to 
assure as accurate as possible a comparison, 22 pairs of plants 
were collected, each pair including one each of the two species 
in question, these taken anywhere from 0.3 to 5.0 feet apart. 
They were taken no further than 5.0 feet apart so that the 
habitat conditions were as nearly the same as we could get them. 

The relative development, as in the earlier work cited above, 
was measured by the ratio of the length (to the nearest milli- 
meter) of the 1960 leaf primordium to the length of the 1959 
previous year’s leaf, expressed as percentage. The measurements 
were made by Miss Virginia M. Morzenti. The results, as shown 
in the table, confirmed the previous conclusion, namely that 
B. ternatum develops significantly earlier than B. dissectum. In 
paired specimens of the two species, B. dissectum averaged only 
55 per cent as far in development as B. ternatum. 

Botrychium multifidum oceurs scattered in the same habitat, 
associated here and there with the other two species. (No speci- 
mens at all of B. oneidense were found in 1960 at this locality, 
and only one was found in 1959.) The five complete specimens 
of B. multifidum (Wagner 9257) that were found on this date 
and in this habitat (though not in definite spatial relationship 
to the other two species) gave an average figure of leaf develop- 
ment of 109.8 per cent, and were thus much further developed 
than either B. dissectum or B. ternatum. This was entirely ex- 
pected, however, as all previous studies have shown that B. mul- 
tifidum matures its leaves considerably earlier than the other 
three species. 

These investigations have been most interesting in supplying 
additional data for the interpretation of inter-relationships of 
these controversial species of evergreen grapeferns. We must 
credit Professor W. L. Dix? for calling attention to the need of 


3 Observed seyret of Botrychium multifidum var. oneidense, THIS 
JOURNAL, 35: 37-39, 1945. 


DrYOPTERIS HYBRIDS 33 


making comparisons of subtle characteristics of these plants in 
the same habitats. It would be very desirable if our colleagues 
in Japan would make similar studies of B. multifidum and 
B. ternatum in habitats where these species grow together there. 


B. dissectum  B. ternatum  B, multifidum 


Collection no. 9256 9258 9257 
Sample size 22* 22* 5 
Average (%) 37.4 67.6 109.8 
Standard Deviation (%) 10.0 18.8 9.3 
Range (%) 21.7-58.7 40.1-93.4 102.2-124.2 


* Each specimen taken five feet or less from a specimen of the other 
asterisked species. 


Supplementary Note on Dryopteris Hybrids 
Epa@ar T. WHERRY 


In a recent article on Dryopteris hybrids,’ only five parent 
species were taken into account; two others are here discussed. 


be hexaploid, its ability to cross freely with tetraploid and dip- 
loid ones is somewhat surprising, yet evidence from morpho- 
logic characters indicates that it does so. The following have been 
reported 

D. CLINTONIANA X CRISTATA. This was published? as Aspi- 
dium cristatum < A. cristatum var. clintonianum. The fre- 
quently expressed view that D. clintoniana and D. cristata in- 
tergrade and are therefore only varietally distinct is manifestly 
based on the manner in which this hybrid combines their fea- 
tures. It should prove on cytologic study to be pentaploid. 

. CLINTONIANA X GoLDIANA. In reporting the existence of 
this hybrid, Dowell® considered it to represent D. goldiana ssp. 
celsa W. Palmer. Although this view has been widely accepted, 
eytologie study by Dr. S. Walker* has shown it to be incorrect. 


: ha JourNAL 50: 87-92. 

2 Bull. Vt. Agr. Exp. Sta. i87: 85. 1915. 
2 all Torr. Club 35: 137. 18 

4TxH1s JournaL 49: 104-112. 1959. 


34 AMERICAN FERN JOURNAL 


D. CLINTONIANA X INTERMEDIA = X D. dowellii (Farw.) 
Wherry, comb. nov. Reported by Dowell® and named Filix [x] 
dowellii by Farwell.® 

D. CLINTONIANA X MARGINALIS. Reported by Davenport’ in 
1902. 

D. CLINTONIANA X SPINULOSA = D. X benedictii (Farw.) 
Wherry, comb. nov. Reported by Benedict® and named Flix 
[X<] benedictii by Farwell.® 

Dryopreris ceLsa. While as above noted this taxon has often 
been considered a hybrid, the cytologic study by Dr. S. Walker 
(op. cit.) has shown it to be a tetraploid species. It is known 
from widely scattered stations from Dade County, Georgia, to 
Berks County, Pennsylvania, and is represented in many 
herbaria by specimens distributed as Plantae Exsiccatae Gray- 
anae No. 1003 from one of its Virginia localities. It is here- 
with suggested to be one parent of three hybrids: 

D. celsa x intermedia = D. X separabilis, new interpreta- 
tion. This taxon was named as a species by Small! and inferred 
to be D. goldiana & intermedia by the writer ;!! as it grows in 
association with D. celsa, however, this is herewith proposed as 
one parent. 

D. celsa  ludoviciana = D. X australis, new interpretation. 
The taxon named D. clintoniana var. australis by the writer” 
was raised to species status by Small.1# That its imperfect 
spores indicate it to be a hybrid was pointed out by Brown and 
Correll,"* and the cytologic study by Walker (op. cit.) favors 


> Bull. Torr. Club 35: 136. 1908. 

6 Pap. Mich. Acad. Sei. 2: 14. 1923. 
7 Rhodora 4: 10. 1902. 

8 Bull. Torr. Club ~~ ree 1909. 

9 Pap. Mich. Acad woes 15, 1998, 
° Ferns SE. foe beh 1938. 

Guide East. Ferns, ed. ss 163. 1942. 
THIS JOURNAL 27: 2. 

ties SE. epee 279, Pe 

Ferns. . .of La., 2 


DRYOPTERIS HYBRIDS 35: 


the view of the origin proposed herewith. 

D.cetsa X ? = D. X atropalustris. This taxon, named as 
a species by Small," is known only from the inadequate type 
specimen, other occurrences listed by Small being doubtful. 
Until it is rediscovered, its parentage remains dubious. 

In conclusion, attention may be called to the surprisingly close 
resemblance between some of the taxa discussed in this and the 
preceding article. In several cases distinction can be based only 
on spore studies, which Mrs. C. W. Crane has generously made 
for me. For example, while D. clintoniana normally has its 
lowest pinnae broadest at base, the material of it distributed in the 
Plantae Exsiccatae Grayanae series has them narrowed at base 
like D. celsa; this came, however, from New York state far north 
of the range of the latter, and its spores prove typical of D. 
clintoniana. On the other hand occasional plants of D. celsa 
have fairly broad-based lower pinnae, and Mrs. Crane finds these 
to have markedly different spore sculpture. The workers who 
have considered D. clintoniana X goldiana identical with D. 
celsa are not to be criticized, for the resemblance is striking, and 
only through the finding of imperfect spores can individual col- 
lections be identified as this hybrid. D. clintoniana X margina- 
lis presents the same problem, as does indeed also D. goldiana 
X marginalis (D. X leedsvi). : 

Two years ago Mr. Robert H. Gaede guided Mrs. Crane and 
me to a swamp in northern New Jersey that was being destroyed 
by the construction of a super-highway, where there grew hun- 
dreds of hybrid Dryopteris plants with no sign of any parents 
nearby. These showed combinations of the characters of various 
Dryopteris species, including D. celsa, which is not known to 
grow that far northeast, although it may formerly have done so. 
Mr. Gaede was fortunately able to rescue a considerable number 
of clumps. It is to be hoped that other similar occurrences will 
be discovered in places not endangered by human activities. 


UNIVERSITY OF PENNSYLVANIA. 


15 Ferns SE. States, 274. 1938. 


36 AMERICAN FERN JOURNAL 
New Forms in Botrychium virginianum!' 
W. J. Copy 


While conducting field work in preparation for the Southern 
Ontario Field Trip of the IX International Botanical Congress, 
the author came upon a Rattlesnake-fern (Botrychium vir- 
gimanum) which in addition to the fertile panicle bore sporan- 
gia on some of the pinnae of the otherwise sterile part of the 
frond. An examination of some 175 specimens of var. virgin- 
tanum in the herbarium of the Canada Department of Agricul- 
ture did not reveal any other specimens with sporangia disposed 
in this manner. There were, however, two specimens among 50 
sheets of var. ewropaeum which displayed this phenomenon. 

Although the occurrence of such forms has been known for a 
long time,?* they have apparently not been formally named. 
The following names are proposed: 

BorRYCHIUM VIRGINIANUM (L.) Swartz var. vIRGINIANUM f. 
anomalum, n.f. Ramus sterilis sporangia aliquot gerens. 

Ontario: Renfrew County, McNab Township, 214 miles west 
of Braeside, June 19, 1959, Cody & Dore 11133 (DAO, Tyre). 
Growing with the typical form (Cody 11134) in eut-over clear- 
ing in mixed Thuja occidentalis, Abies balsamea, Populus trem- 
uloides woods in rich black moist shallow soil over limestone. 

BoTRYCHIUM VIRGINIANUM (L.) Swartz var. EUROPAEUM Ang- 
strom f. heterodoxum, n.f. Ramus sterilis sporangia aliquot 
gerens. 


Rich woods, Saskatchewan, McKague, June 25, 1935, A. J. 


1 Contribution No. 106 from the Plant — Institute, Research 
Branch, Canada Department of Agriculture, Ott 

2 Chrysler, M. A. So nature of the fertile ie se in the Ophioglossaceae- 
Ann. Bot. 24:1-18, 19 


3 Clausen, R. T. A ean of the Ophioglossaceae. Mem. Torrey Bot. 
Club. 19;1— 177. 1938, 


Taxonomic Notes : 37 


Breitung sn. (DAO, Type); Bois tourbeux, Quebec, Vieux- 
Comptoir, 52° 37’ N, 78° 42’ W, E. Lepage 32,232 (DAO). 

The type specimen of f. anomalum has one of the lower pin- 
nules of each of the two outer segments of the ternate normally 
sterile portion of the frond almost completely fertile. In addi- 
tion five other pinnae scattered over the normally sterile por- 
tion of the frond also bear sporangia. The single isotype bears 
only a few sporangia on lower pinnules of the outer segments of 
the normally sterile portion. 

The type specimen of f. heterodoxum, in addition to the fertile 
panicle, bears scattered sporangia on the normally sterile portion 
of the frond. These sporangia are found mostly on the basal 
pinnules of the lower pinnae. The single paratype is one of two 
plants mounted on a sheet. One of the lower pinnules of a large 
outer segment of the ternate normally sterile portion of the frond 
is almost completely fertile. A few sporangia are also to be found 
on basal pinnules of the lower pinnae of the other two segments. 
The other specimen mounted on the sheet belongs to f. curo- 
paeum. 


Taxonomic Notes on Ferns, II 
C. V. Morton 


THELYPTERIS subg. THELYPTERIS sect. Glaphyropteris (Presl) 
Morton, comb. nov.! 

Some years ago I published a paper entitled ‘‘New South 
American Species of Dryopteris, section Glaphyropteris,’” in 
which four new species were described and one variety raised to 
specific rank. The section Glaphyropteris is extremely close to 
the subgenus Lastrea of Christensen’s Monograph of Dryopteris, 
differing only in the development of prominent aerophores at 


‘Glaphyropteris ie Abh. Boehm. Ges. Wiss. V, 5: 34, 1848. Typus: 
Polypodium decussatum L. Dr onene: te Glaphyropteris C. Chr., 
Arbeider Tileg ras = erat 80. 1911 eae ihe Glaphyropteris 
Alston, Jour. Wash. Acad. Sci. 48: 234. 

*Journ. Washington Acad. a . 28: 525- 530. 1938. 


38 AMERICAN FERN JOURNAL 


the base of the pinnae and also at the base of the costules in 
most species. This is a relative character, since some species 
classified as true Lastreas sometimes have minute aerophores, 
and doubtless is not of fundamental importance. Glaphyropteris 
can rank as no more than a section 

With the segregation of the large inclusive genus Dryopteris 
into smaller units, these plants fall in the genus Thelypteris. 
The following new combinations are necessary. Several species 
have already been transferred to Thelypteris—D. canadasii and 
D. mapiriensis by Alston and D. Thomsonii and D. decussata 
by Proctor. The others are: 


THELYPTERIS andina (Morton) Morton, comb. n 
Dryopteris andina Morton, Journ. Wachingtoh ‘Acad. Sci. 28: 
526. 1938. 


HELYPTERIS boliviensis (Morton) ipien comb. nov. 
Dryopteris boliviensis Morton, op. ¢ t. 527. 
THELYPTERIS comosa (Morton mn.) Moron comb. nov. 
Dryopteris comosa Morton, op. ¢ t. 528 
THELYpTeRIS Tatei (Maxon & fai Morton, comb. 
Beg? Tatei Maxon & Morton, Journ. Wachineton. ‘Acad. 
529. 19388. 


THELYPTeERIS brasiliensis (C. Chr.) Morton, comb. n 
Dryopteris metspe var. brasiliensis C. Chr. asi Vid. 
Selsk. Skr. VII. 10: 161. 1913. 
Dryopteris brasiliensis Morton, op. cit. 529. 
THELYPTERIS polyphlebia (C. Chr.) Morton, comb. no 
ch a polyphlebia ©. Chr. Dansk. Vid. Selsk. " gkr. Vil. 
1 19, 1913, 


THELYPTERIS macradenia (Sodiro) Morton, comb. no 
Nephrodium macradenium Sodiro, Ree. Crypt. ike Quit. 
1883. 


Dryopteris macradenia C. Chr. Ind. Fil. 276. 1905. 
BLECHNUM PENNA-MARINA (Poiret) Kuhn 

This attractive and characteristic species has been known 
from a number of widely separated localities—Australia, New 
Zealand, Chile and Patagonia, Tristan d’Acunha Island, Am- 


Taxonomic Notes 39 


sterdam Island, and Kerguelen Island. It is not reported from 
Madagascar in ©. Christensen’s Pteridophyta of Madagascar 
(1932), nor in Madame Tardieu’s recent treatment,’ but ap- 
parently it does grow there. Rather strangely, there are three 
sheets of the species in Christensen’s home institution, the Uni- 
versitetets Botaniske Museum, Copenhagen, that I discovered 
filed among the unidentified specimens of Polypodium. 

Only one of the specimens is adequately ticketed, this one 
reading ‘‘Polypodium. Commerson, Madagascar, ded. Dr. 
Thouin, Hb. Vahlii,’’ i.e. collected by Philibert Commerson in 


some mistake, but the other two collections also definitely say 
‘‘Madagascar,’’ and they are obviously from different collectors. 
One of them has no indication of the collector, but it is obviously 
an old sheet distributed from Paris. Some one will probabl 
able to recognize the handwriting, which is rather characteristic. 
The other sheet bears two initials indicating the collector, but 
these are not legible to me. [Can they refer to Pervillé?] It 
seems reasonable to conclude that B. penna-marina did grow in 
Madagascar 150 years ago and more. The absence of the species 
from recent collections is remarkable. 


ASPLENIUM VARIANS Hook. & Grev. 

. Although common in India and China, this species has not 
been reported from Malaya. In the herbarium of the Universi- 
tetetes Botaniske Museum, Copenhagen, there is a specimen 
collected by C. W. Franck at ‘‘Gunung Pulae,’’ Johore, April 
20, 1924, that I so identify. It does not represent the divided 
form that is commonest, but a form that is merely deeply 
pinnate-pinnatifid, quite similar to a specimen in the U S. 
National Herbarium from Yunnan, China (Maire 6108). This 
species has been known from Tonkin and Annam, and so its oc- 
currence in Johore is not unreasonable. 


1Jn Humbert, Flore de Madagascar, 5th Fam. 2: 1-19. 1960. 


40 AMERICAN FERN JOURNAL 


Some Records of Michigan Ferns and Fern Allies! 


Jaru K. Hintunen 


In the western end of Chippewa County, Michigan, adjacent 
to Highway M-28, about 1.5 miles west of Hulbert, occurs a 
plant community with an unusual variety of plants. This part 
of the county is a swampland with intermittent, low, broad, 
sandy ridges, which are probably old glacial lakeshore deposits. 
The swampland is dominated almost exclusively by black spruce 
(Picea mariana), tamarack (Larix laricina), and white cedar 
(Thuja occidentalis), while the ridees support scattered pine 


se 


In the summer of 1954, I discovered this habitat and noted 
that, among other plants, certain species of club-moss were es- 
tablished there. These were mostly Lycopodium clavatum, L. 
inundatum, and L. tristachyum, together with another species 
which T did not recognize. I was unable to collect specimens, 
however, until the summer of 1956. In September of that year 
I revisited the area to collect a sample of the unknown club- 
moss*. The plant was relocated along the margin of the moist 
sand-flat, where the latter meets a bank of dry sand, a remnant 
of a partly removed ridge. 

After a close examination, the plant was tentatively identified 


1 Contribution No, 54 from the Department of Biology, Wayne State Uni- 
versity, Detroit, Michigan, 


_ the author’s collections are in the Herbarium, Biology Department, 
Wayne State University, Detroit, Michigan, 


MicHIGAN FERNS 41 


as L. Selago. Mr. Dale J. Hagenah, Cranbrook Institute of Sei- 
ence, and Dr. W. H. Wagner, Jr., Department of Botany, Uni- 
versity of Michigan, agreed that it was this species or probably 
the variety patens. This proved to be an unusual find, for the 
only previous Michigan records of this species are from the west- 
ern and central parts of the Upper Peninsula (Isle Royale, 
Keweenaw Peninsula, and Schooleraft County). In July, 1957, 
however, I found the species in Ontario, along a margin of an 
upland lake, two or three feet from water’s edge. This latter 
station, which is approximately 30 miles north of Sault Ste. 
Marie, is in a region that is mountainous and unlike that in 
Chippewa County. Judging from the above, L. Selago appar- 
ently is established in scattered moist habitats around the east- 
ern end of Lake Superior. 

On a third visit to the moist, sand-flat area in Chippewa 
County, in July, 1957, I found a few individuals of a previously 
overlooked club-moss that had the appearance of L. tristachyum. 
Among them was a variant with only four or five branches aris- 
ing from one underground rhizome. Its specific determination 
was made more difficult by the fact that the plant was sterile. 
Further study showed, however, that the variant was not L. 
tristachyum, but that it appeared to be L. sitchense (L. sabin- 
ifolium var. sitchense). The specimen was sent to a student of 
the genus, Mrs. Joan H. Wilce, of the University of Michigan, 
who confirmed its identity. The only published Michigan rec- 
ords of this taxon are from Iron and Marquette Counties,* near 
the western end of the Upper Peninsula, so that this new record, 
as in the ease of L. Selago, establishes a notable eastward range 
extension in Michigan. 

Mr. Dale Hagenah recently visited the area described above 
and informs me that he has found the grape-fern Botrychium 
simplex in this habitat. The discovery of the grape-fern consti- 
tutes the third known record for the county, the first two being 


eT Cecil, Ferns of Michigan. Cranbrook Institute of Science, 
Bull, 1952. 


42 AMERICAN FERN JOURNAL 


those of Hagenah,* who found the species in a woodland habitat 
near Trout Lake. 

Since the discovery of this community, it has been Sound to 
bear other vascular plants worthy of note: Botrychium multi- 
fidum, B. virginianum, Carex flava, Gentiana rubricaulis, Habe- 
naria clavellata, Juncus articulatus, Epigaea repens, Equisetum 
palustre, E. variegatum, Ophioglossum vulgatum, Scirpus hud- 
sonianus, and Spiranthes cernua. 

AYNE Strate University, Detrorr, MICHIGAN. 


Shorter Notes 


Recent Fern Discovertes IN WESTERN PENNSYLVANIA.—AI- 
though western Pennsylvania is relatively well known botan- 
ically, two interesting ferns have been discovered here recently : 
Lyeoprum paLMATUM (Bernhardi) Swartz. Climbing Fern 

This fern has not previously, to our knowledge, been collected 
in western Pennsylvania, although it is known from northeastern 
Pennsylvania, southern Ohio, and West Virginia. In June, 1959, 
Mr. Robert Leberman, of Meadville, Pennsylvania, found a col- 
ony of Climbing Fern one mile north of Sugar Lake, Crawford 
County, in northwestern Pennsylvania. The plants were twining 
over cinnamon ferns and other vegetation in a mixed woods of 
red maple, yellow birch, and white pine. 

WoopwarpiA arEoLATA (Linnaeus) Moore. Netted Chain-Fern. 

Over fifty years ago, Dr. O. E. Jennings, Director Emeritus 
of the Carnegie Museum, collected this fern in the vicinity of 
Half-Moon Swamp, north of Mercer, in Mercer County, Penn- 
sylvania. Until 1959, this was the only known collection in west- 
ern Pennsylvania, although several efforts have been made to 
relocate this fern. Due to partial draining of the swamp, and 
other changes attendant on the passage of half a century, the 
plants could not be found and were suspected of being extinct. 
In June, 1959, however, the authors succeeded in finding a 
flourishing colony in a low wet woods in a part of the swamp not 


4 Hagenah’s collections are in the Herbarium of Cranbrook Institute of 
ein Bloomfield Hills, Michigan 


Suorter Nores 43 


previously investigated. With it grew such plants as wild calla 
and Virginia chain-fern—both rather rare in Pennsylvania so far 
south. Natives tell us that there was at one time a stand of 
American larch in this swamp.—l. K. Henry and W. E. BuKEr, 
Carnegie Museum Herbarium, Pittsburgh, Pennsylvania. 


Fern Monster?—Our member Mr. George R. Proctor has 
sent a clipping from The Star, Kingston, Jamaica, with the 
headline ‘‘Asked to Fight Lake Monster,’’ and the dateline 
September 20, 1960, Salisbury, Southern Rhodesia : 

“‘The Federal Government of Rhodesia and Nyasaland has 
asked the United Nations to help fight a battle. It is asking for 
scientists to help conquer the ‘Monster of Lake Kariba.’ 

‘‘Although Kariba is the largest man-made lake in the world 
—it was officially opened by the Queen Mother earlier this year 
—90 square miles of its 1160 square mile surface has been cov- 
ered by a fast-growing weed called Salvinia auriculata, known 
locally as ‘The Monster.’ 

‘‘Because the weed is so fast growing and cannot be con- 
trolled, the government is worried that before long it will cover 
the whole lake. This could bring the giant power supply engines 
to a standstill, partially crippling the Federation’s economy. 

As the problem is getting out of hand, the United Nation’s 
Food and Agricultural Organization has been asked to send a 
scientific team to Kariba to find a way of controlling the weed.”’ 

CYSTOPTERIS TENNESSEENSIS Shaver.—In October, 1958, I col- 
lected one live plant of Cystopteris tennesseensis from Shaver’s 
type station on the north-facing bluff below the quarry on 
Round Lick Creek near Highway US 70N. The station, I would 
say, is destroyed or nearly so by the quarry there, as I could 
only find one or two plants. 

This plant was considered a hybrid by Dr. Shaver—C. bulbi- 
fera X fragilis var. protrusa (Ferns of Tennessee, p. oat). I 
planted the spores in October on sand plus peat-moss. They 
were fertile, for I now have typical plants bearing both spores 
and bulbils. The bulbils grow rapidly into nice plants. I would 


44 AMERICAN FERN JOURNAL 


say that if Cystopteris tennesseensis is a hybrid, it is a fertile 
one and ean be classed as a spec 

I now have about fifty nae on the spores and will keep 
a colony growing, so that plants from the type station will con- 
tinue to be available—Ratpu H. Brnepict, 3106 Lapey Street, 
Rockford, Illinois. 

PELLAEA ATROPURPUREA.—The new edition of ‘‘Ferns and 
Fern Allies of Texas,’’ by Donovan §8. Correll, reviewed in an 
earlier volume of this journal,' gives on page 10 a schematic 
representation of the life cycle of a typical fern, using as an 
example Pellaea atropurpurea, as shown both by the drawing of 
the leaf and in the description of the figure. In the figure, P. 
atropurpurea is shown to have a sexual life cycle. This seems to 
be a noteworthy discovery, especially if sexual plants are as 
widespread as Mr. Correll indicates by choosing it to represent 
the prothallus of the entire fern flora of Texas. In fact, on page 
116 in his book he says that ao species “is perhaps the most 
widely distributed fern in Texas. 

I have been interested for some time in the prothalli of P. 
atropurpurea and have grown them from spores collected by 
myself in the Ozarks and from spores kindly sent me from the 
Missouri Botanical Gardens by Mrs. Alice Tryon (original col- 
lection from Gray Summit, Missouri). Prothalli from both of 
these sources are apogamous. 

That P. atropurpurea is apogamous has been known for some 
time. Reported first by Steil in 19112 and again in 1918,° it was 
studied in some detail by Manton‘ and discussed in her chapter 
on apogamy in ‘‘Problems of Cytology and Evolution in the 
Pteridophyta.’’ Miss Manton obtained spores (through the kind- 
ness of the late Mr. Alston of the British Museum) collected in 
the wild in California and found strong evidence of her material 


1 THs JouRNAL 47: 79-81. 1957, 
Pes il, W. N. Apogamy in Pellaea atropurpurea, Bot. Gaz, 52: 400. 

pees Beeb Studies of Some New Cases of Apogamy in Ferns, Bull. Torr. "Bot 
ub 


« Chiabriage University Press (England) 1950, chap. 10, pp. 184, 185. 


Recent Fern LITERATURE 45 

being triploid with a chromosome number’’ of about or exactly 

This same number was obtained by Tryon and Britton 

for Mrs. Tryon’s collection from Gray Summit, Missouri,> and 

for those who may be interested, my drawing of one of the 
apogamous prothalli appears in that paper. 


That material of P. atropurpurea critically studied to date 
has been found to be apogamous does not, of course, preclude 
the existence of sexual plants of this species. The interesting 
case of Asplenosorus ebenoides immediately comes to mind— 
readers will remember Dr. Wagner’s contribution in this jour- 
nal.® 


I should be happy to receive leaves of the Texan P. atropur- 
purea (in good fruiting condition, i.e., leaves dropping spores) 
in order to grow the prothalli for points of comparison with 
those from Arkansas and Missouri which I have already studied. 
—Lenerre R. Arkinson, 415 South Pleasant St., Amherst, Mass. 


Recent Fern Literature 


Fora Mauesiana, Ser. II. PrertpopHyta, vol. 1, pp. 1-64. 
1959, by R. E. Holttum.'—Professor Holttum, having finished 
his magnificent job on the ferns of Malaya, has now turned his 
attention to the ferns of Malesia, that fern paradise in the 
South Seas consisting of Java, Sumatra, Borneo, New Guinea, 
and hundreds of smaller islands. This first part contains a brief 
fore discussion of fern morphology, a key, admittedly tenta- 

0 


detailed, yet compact, in marked contrast to many contem- 
* This ‘‘evidence’? i 4 Flee because Pellaea atropurpurea does not occur 
“in the wild’’ in Cal a—C. V. 
°’ Tryon, Alice F. a "Donald Britton. Cytotaxonomic Studies on the 
5, 1s 


Pern Genus Pellaea. oes 12: 137-14! 
® Wagner, Warren H., A Cytological Study oe the Appalachian Spleen- 
1953. 


worts. THs JOURNAL 43: 109-114. 


‘Available from P, Noordhoff, Ltd., Groningen, Holland, $2.65. 


46 AMERICAN FERN JOURNAL 


porary works, too many of which seem to be just skeletons, con- 
taining only the bare bones, or at the opposite extreme diffuse 
discussions in which it is hard to locate any definite or satisfy- 
ing conclusions. Holttum’s works are always contributions to 
knowledge, because they are original, in both senses of the word 
—they are never just compilations, composed of parrotings of 
other people’s descriptions and ideas, and they always have new 
and stimulating ideas of their own; they are sure to remain 
indispensable to pteridologists. The many drawings deser 
special mention, for they are models of botanical illustration. 
One can only wish Dr. Holttum an unusually long life, for a 
treatment of all the ferns of Malesia on the same scale as this 
beginning is a lifetime job—C. V 

Recent publications of our Honorary Member GuALTERIO 
Looser that may be mentioned are: Los Helechos de la Isla de 
Pascua,’ an account of the ferns of Easter Island, that isolated 
island in the South Pacifie 2 ,000 miles west of Chile; and Clave 
de los Blechnum (Filicales) de Chile,? a eeomeets toa 
ous (1947) paper on Chilean Blechnums.—C. V. M 


New Poticres ror THE Brirish FERN GAZETTE.—The most 
recent issue of the British Fern Gazette (Vol. 9, No. 1) makes 
very interesting reading for American Fern Society members. 
In a foreword, the new editor, Mr. A. C. Jermy,? explains 
changes in policy adopted by the Council of the British Pterido- 
logical Society. He notes that “An interest in fern variation 
has been foremost in the Society from its beginning and this 
interest has in later years resulted in two factions. One is the 
fern grower who is anxious to find new and attractive varieties 
for his collection; the other is the taxonomist, equally interested 
in finding vihictien but who is interested also in the conditions 
under which the plant is growing and its possible relationships 


TRevista Universitaria (Santiago, Chile) XLII: ae ae fig. 1-16, 1958. 
*Revista Universitaria XLII: 123-128. fig. 1-17. 
Says ney army — succeeded to the post at the Britis in couse ~ = 


M ¥. Alst n, deceased. For the Gazette, he succeeds the 
Elliot, also dee ease ae 


Recent Fern LITERATURE 47 
to other varieties and other species.’’? The new policies of the 
Society provide for an extension of interest to fern allies and to 
pteridophytes from all parts of the world. It would appear that 
henceforth the Fern Gazette will have a much wider interest for 
fern students the world over and that the British Pteridological 
Society should find a greatly expanded membership. The ranks 
of British botanists, since the time of Bolton in 1795, have al- 
ways included leaders in the study of ferns, both as to classifi- 
fication, in the fundamental field of morphology and phylogeny 
(Bower and Lang), and in genetics and cytogenetics (Anderson- 
K6tto, Manton and Walker) 

This recent issue implements the new policy by its contents: A 
complete checklist of British pteridophytes carried down to 
counties and vice-counties by Mr. Jermy; a clearly and simply 
written account by R. E. Holttum of significant vegetative dif- 
ferences among common fern genera by which modern fern stu- 
dents supplement characteristics of fruiting structures formerly 
the chief reliance; and an article on the geographic affinities of 
New Zealand pteridophytes, by J. D. Lovis. There are shorter 
notes and also five pages which deal with the annual (1959) field 
excursion and other matters of ‘Society activities. 

A few comments on some of the articles follow. The Holttum 
discussion is a presentation by an expert taxonomist of data of 
real interest to amateurs, including beginners. I do not recall 
that the Fern Journal has ever published anything of this sort. 
The Jermy ‘‘Census’’ provides the best possible basis for a com- 
parison of British fern flora with that of our own. That the 
British Isles offer fern species not found elsewhere has been 
noted before. Of the 73 species listed, two only are noted as 
aliens which have had limited establishment, the sensitive fern 
and the Pacific holly fern, Cyrtomium faleatum. A dozen hybrids 
furnish additional distinctive taxa to the species list. Only one 
is also found in the U. S., Dryopteris uliginosa (D. cristata X 
spinulosa) Among the others are two crosses between the 
hart ’s-tongue, almost ubiquitous in many parts of Great Britain, 


48 AMERICAN FERN JOURNAL 


and two species of spleenwort; also several crosses involving 
the male fern and the mountain wood fern, D. dilatata. For the 
United States, I believe only one male fern cross has been re- 
ported, Winslow’s find of D. filix-mas & marginalis. With us, 
D. dilatata has a higher altitudinal distribution, which usually 
keeps it out of much contact with our lowland species, among 
which many hybrids have been reported. I recall one Adirondack 
station with a good growth of dilatata and D. intermedia in 
which there were indications of possible crossing. 

For the benefit of those who may wish to enroll as members 
of the British Pteridological Society, the following information 
is offered: The Society publishes at present one issue a year of 
the Gazette (the present issue is 32 pages). It conducts an an- 
nual summer excursion and holds one December meeting indoors. 
Members take part in plant exhibitions. The officers stand ready 
to provide information by correspondence. The dues are ten 
shillings per year and applications may be sent to the Secretary- 
Treasurer, Mr. J. W. Dyce, Esq., ‘‘Hilltop,’’ 46 Sedley Rise, 
Loughton, Essex, England —Raupu GC. BENEDICT. 


Notes and News 


AnNvuAL MEETING:—The annual meeting of the American 
Fern Society will be held during the last week of August at 
Purdue University, Lafayette, Indiana, in connection with the 
meetings of the A. I. B. S. Will members desirous of reading 
papers at this meeting please send the titles and approximate 
length immediately to the Secretary, Dr. Donald Huttleston, 
Longwood Gardens, Kennett Square, Pennsylvania. The annual 
field-trip will be held in Kentucky just prior to the meeting. 
Members and friends interested should write for information to 
Mr. Thomas McCoy, Catlettsburg City Schools, Catlettsburg, 
Kentucky, who has kindly agreed to act as leader. 

Reprints Destrep.—Our member Dr. R. Pichi-Sermolli 
(Instituto Botanico ‘‘Hanbury,’’ Via Balbi 5, Genova, Italy) is 
currently working on a new supplement to the Index Filieum. 
He asks that members of the American Fern Society, and other 


AMERICAN FERN Socrery 49 


pteridologists, send him reprints of their taxonomic papers, 
especially those issued since 1933, the closing date for Supple- 
ment III. Papers published in the American Fern Journal are 
not needed. 


BorrycHium TRANSPLANTING?—In volume 48, number 4, of 
the American Fern Journal, Dr. Benjamin R. Allison asked the 
question why it is that botrychiums are so hard to transplant. 
I have five plants of B. dissectum, two of B. multifidum, and also 
several of B. virginianum that I moved three years ago. The 
multifidums have increased in size, the dissectums this year grew 
two fertile and two sterile leaves to each plant, and the vir- 
ginianums have remained the same size. They are all in sandy 
soil on the north side of a building, well mulched with leaves at 
all times and kept well watered—RautpaH H. Benepicr, 3106 
Lapey Street, Rockford, IMlinois. 


American Fern Society 
Report of the President for 1960 


_ The American Fern Society celebrated the fiftieth anniversary 
of the establishment of the American Fern Journal by printing 
over 300 pages, a new record. Although the cost of these extra 
pages exceeded the original budget for the year, it was deemed 
worthwhile to draw upon our reserve funds for this expense, 
and some members made contributions also. L. R. Bolton, Presi- 
dent of the British Pteridological Society, sent hearty congratu- 
lations for the fiftieth year of publication of our Journal. 

Last May witnessed the gala opening of the Fern Valley in the 
U.S. National Arboretum. This project, initiated in 1953 and 
Supported by the National Capital Gardening League and others, 
has been under the supervision of one of our members, James 
W. Johnson. A full account will be published in a later number 
of the hinge Plan to visit it the next time you are in 
Washing 

Your 1 Pieanieit was able to accept the invitation of Biological 
Abstracts to attend its symposium and house warming in cele- 


50 AMERICAN FERN JOURNAL 


bration of its twenty-fifth anniversary and its move to new 
quarters. 

The successful operation of the American Fern Society falls 
upon the shoulders of the other officers and members of the 
Council, and I wish to express my appreciation to them for their 
willing cooperation. Conrad Morton, as Editor, has had a very 
active part in soliciting papers and in the time-consuming task 
of preparing the Journal. 

Walter S. Phillips, with the aid of his wife, has had the exact- 
ing fiscal duty of collecting membership dues and subscriptions 
to the Journal, and keeping the books, a task which has mate- 
rially absorbed much of his time, leaving him very little oppor- 
tunity for research or leisure. 

Dr. R. C. Benedict’s request to be relieved of his editorial 
duties was accepted with regret. The Society appreciates very 
much his interest and contributions over a period of 50 years. 
The resignation of Dr. A. C. Smith from the editorial board has 
also been accepted. Dr. Rolla M. Tryon, Jr., has accepted ap- 
pointment to the editorial board. 

Dr. Tom Cooperrider, Kent State University, Kent, Ohio, has 
been appointed Custodian of the back files of the Journal. At 
one time they were in the custody of Mr. E. J. Winslow; then 
transferred to Dr. Henry Svenson at the Brooklyn Botanical 
Garden; then in 1947 to Conrad Morton in Washington. 

Dr. A. C. Smith has been the Society’s representative on the 
AAAS Council for the past three years and has asked to be re- 
placed. Each year he has submitted an excellent report on the 
activities of the Council. 

Mrs. K. Boydson’s enthusiasm and work has made the spore 
exchange a success. 

Dr. Dwight W. Moore organized and led a highly successful 
field trip in northern Arkansas last August, the plea of which 
will appear elsewhere. 

Dr. Edgar T. Wherry kindly contributed the revenue derived 
from sales of his book ‘‘Guide to Eastern Ferns’’ to the Ameri- 


AMERICAN FERN Society 51 


can Fern Society, a gracious act showing his love and interest in 
the Society. 

Our member Dr. W. H. Hodge, Chairman of the Editorial 
Committee of the American Horticultural Society, wrote me that 
his organization is interested in devoting a special issue of the 
National Horticultural Magazine to ferns. The Horticultural 
Society will assume the cost of publication if members of the 
Fern Society will undertake preparation of the manuscript. The 
proposed publication is to stress the horticultural importance of 
ferns; their propagation ; how to grow them; kinds suitable for 
gardens in different sections of the United States; and kinds 
suitable for house plants. The exact scope is to be determined 
by an editorial committee which will have the problem of finding 
individuals who are willing to write on assigned topics. This 
project was discussed briefly at the meetings in Stillwater, and 
most of those present were in favor of it. It seems like a wonder- 
ful opportunity to prepare a much-needed publication. Several 
members have accepted appointment to this committee and are 
preparing lists of topics to be considered. After the scope of the 
publication has been approved, members will be asked if they 
will write on some phase in which they have had experience. 

There has been some dissatisfaction with the procedure fol- 
lowed in selecting candidates for offices in the annual election. 
Although the procedure followed is that specified in our Consti- 
tution, it seems more democratic to present a dual slate so that 
members may have a choice. The constitution will be examined 
and, if necessary, steps will be taken to — it if the members 
wish. I am ready for your suggestion: 

Mr. Thomas N. McCoy, spertaenl Kentucky, has kindly 
consented to lead the annual fern foray next August in the 
vicinity of Carter Caves, Natural Bridge, and Cumberland Falls, 
Kentucky. 

In conclusion, I would like to say that, on my part, I have 
enjoyed being your President for the past year. You are a 
pleasant and wonderfully congenial group. 


Respectfully submitted, Cuatr A. Brown, President 


52 AMERICAN FERN JOURNAL 


Report of the Secretary for 1960 

Due to the fact that a number of delinquent members were 
dropped from the rolls in July, the membership dropped for the 
first time in a number of years, from 797 to 749. By December, 
however, we had partially recovered from this drop and the 
present membership stands at 776. All are,.urged to persuade 
interested non-members to join so that we can,.resume our in- 
crease. 

I am sorry to report the deaths of six members of long stand- 
ing: Mrs. Walter Beck (1928), the Rev. E. A. Elliot (1939), Mr. 
F. N. Irving (1940), Dr. H. H. Bartlett (1944), Mr. H. B. Rust 
(1944), and Dr. S. F. Blake (1945). A note about Mr. Irving 
appeared in Volume 50, Number 2, 1960, of the Fern Journal... - 

The annual field trip, in the Ozarks of Arkansas, was well at- 
tended and is being reported upon elsewhere in the Journal. 

The annual meeting of the Society was held in connection with 
the A.I.B.S. on the campus of Oklahoma State University, Still- 
water, Oklahoma, on August 24, 1960. Since only seven papers 
were presented, there was only a morning session, but it was 
well attended by about 50 persons and considerable interest was 
shown. At the session, presided over by President C. A. Brown, 
the following papers were presented: ‘‘Ferns as Experimental 
Tools for the Study of Idioblast Differentiation—an Initial Re- 
port,’’ by George S. Dehnel; ‘‘Spore Studies in the Genus C'ys- 


(1679),’’ by Joseph Ewan (read by D. G. Huttleston) ; ‘‘Spore 
Studies in the Genus Anemia,’’ by John T. Mickel (read by 
Richard White) ; ‘‘The Species of Evergreen Grapeferns (Bo- 
trychium),’’ by Warren H. Wagner, Jr. ; ‘‘Fern Spore Studies,’’ 
by Clair A. Brown; and ‘‘Practical Checks to Microscopic Illu- 
sions,’’ by Clara S. Hires. Miss Hires also had an exhibit of 
her work in the Exhibit Hall. 

The luncheon at twelve o’clock was presided over by Dr. 
Brown and was attended by 35 members. At ‘a’brief meeting 
following the luncheon a suggestion by the American Horticul- 


AMERICAN FERN Socrery 53 


tural Society that the Fern Society write a handbook on culti- 
vated ferns, for future publication as a special issue of ‘‘The 
American Horticultural Magazine,’? was discussed and it was 
considered to be a worthwhile project. A committee to look into 
the feasibility of the undertaking is being appointed by Dr. 
Brewn. The Fern Society is grateful to Prof. U. T. Waterfall 
for acting as the Society’s local representative and for making 
the arrangements for the meeting rooms and luncheon. 

The annual meeting for 1961 will be held as usual with the 
A.I.B.S. meetings at Purdue University, Lafayette, Indiana, 
August 27-31. It is hoped that all who can will plan to attend 
and that a number will present papers. We are hoping to ar- 
range a field trip preceding the meetings. 

Respectfully submitted, ; 
G. Hurrieston, Secretary 
ort of Treasurer for 1960 

The balance on hand at the end of 1960 was much smaller 
than it has been for the last few years. However, most members 
will agree that the 1960 50th Anniversary numbers were well 
worth the extra cost, and in spite of the expense in printing the 
special number we still are in the black. 

Sale of back numbers continues to aid in our yearly receipts, 
but it is my opinion that the back numbers ought to be raised 
to be at least as high as the yearly annual subscription cost. They 
are now selling at 50¢ a number or $2.00 for the volume, which 
is 50¢ less than the yearly subscription price. Reprinting out-of- 
print numbers has proven expensive and more reprinting must 
be done soon. I suggest that $3.00 per volume or 75¢ per num- 
ber is a fair price and that old numbers be sold only when 
available as extras not needed for complete volumes. 

Correspondence seems to increase each year. Libraries, insti- 
tutions, both private and governmental, and especially foreign 
organizations are requiring extra invoices and receipts and usu- 
ally by air mail. The 10¢ extra for foreign postage hardly covers 
the cost of the business dealings necessary in most cases. 

Funds in reserve at the Green Point Savings Bank remain un- 


54 AMERICAN FERN JOURNAL 


touched and have shown yearly increases as the interest accumu- 
lates, but they do not show capital gains. Some of the funds 
ought to be invested in mutual funds so that capital gains can 
be appreciated. 

The Treasurer and his wife want to express appreciation to 
those who answer our annual dues billings promptly and hope 
more members will follow suit. Last year we mailed out 700 
first notices, 400 second notices, 100 miscellaneous back notices 
and answered some 250 miscellaneous requests for receipts, and 
so forth. 


Receipts 
Total 
$1,195.50 
ewals 
1961 Subscribers—Renewals (advance). 245.42 
1962 Subseribe sean ales (adVvalive yo 4.50 
Sale of Back MODOIS ce ee ee | 231.21 
Sale of Piakelbe PE AUEAEE Se 1 Steet ic) See oa anaes 404.32 
rn thee ee ee ee oe ae 50.00 
be SaboMahaoi et an US En Wa Ee orl eee 200.01 
x: : . S. subscription by member A.F.S, 1.0 
3,329.15 
4,524.65 
Disbursements 
A.F.J. Vol. 49, No. 4 eee 87.43 
A.F.J. Vol. 50, No. 1 1,521.86 
A.F.J. Vol. 50, No. 2 = 
A.F.J. Vol. 50, No. 3. RE SOOM TER Nomen ane te oC eae 459.48 
Reprints EE GOS 387.74 
Envelopes, ma ailing ‘a “postage. Uc Sse nan ea 202.55 


Shipping Back Number ese 26.77 


AMERICAN FERN SOCIETY 


A.IL.B.S. 1960 dues Peneto Fa ates 100.00 
A.LB.S. a8 Sub. by A.F.S. Member. 1.00 
Refund to Book Agent for Cancellation 2.50 
Editor Sgn, $25.00—1960, $1.37_-.._-----. 26.37 
Seeretary Expense—1959, $10.00—1960, a 86 Bea eel 14.86 
‘reasiiver Hxperse --f900 1 2 oo 63.49 
President’s Expense—1959 —.._---------- PSs he te 40.00 
Purchase of Piling ‘Cabinet 2 38.13 


Cash in Southern Arizona Bank, January 1, 1961. 


Statement December 31, 1960 
Assets 


Oasn in Southern Arizona Bank 22 
Cash in Green Point Savings Bank (Bissell Herbarium Fund) — 

Cash in Green Point Savings Bank (Life oar Pind): -¢ 
Cash in Green Point Savings Bank (Reserve Fund) —-—-- 
Cash in Green Point Savings ‘Bank (Una ee Veit ts) eure 
Inventory—American Fern Journal 
American Wern Society Library. 


: Liabilities 
Advance dues collected UU Sa RE ann ee Pe $348.77 
Accounts Payable (Vol. 50, No. a Se 613.99 


Fund Balances 
Bissell Herbarium Fund. 
Life lap acs Band ee 


Rese 
Una Weathety and. 
General Fun 


Respectfully saicai 
Watrter S. Pairs, Treasurer 


Report of the Auditing Committee 


55 


4,026.84 


$ 497.81 


10,716.69 


$10,716.69 


We hereby certify that we have seen the books and accounts 
of Dr. Walter 8. Phillips, Treasurer of the American Fern So- 


56 AMERICAN FERN JOURNAL 


ciety, Inc., and have obtained confirmation of the correctness of 
the Society’s balances on hand as set forth in detail in the ac- 
companying report of the Treasurer. 
CHARLEs T. MASON, JR. 
Ricwarp H. HEeviy 
Auditing Committee 


Report of the Judge of Elections 
Three hundred and fifty marked ballots were received. The 
tally of votes is as follows: 
For President : 


Clair A. Brown 346 
Mougiis Cobb 1 
R. Pichi-Sermolli ori ae 
Rolla M. Tryon,Jr. 1 
DY S. Correll: or 1 

350 

For Vice-President : 

Maton Mey 346 
Ald. Sharp 22 4 1 
Rolla M. Tryon, Jr. 1 
Tom Cooperrider, 4. 5. 2s 1 
K. U. Kran oo) ee 1 

350 


For Secretary: 
D. G. Huttleston _ 348 


Mrs, Geoffroy Atkinson. rt pi 1 
Mie ‘Tardieu Blot ee us 
350 


For Treasurer : 
Witter 8. Phillipe 349 
I therefore declare the following officers elected: Clair A. 
Brown, President; Marcel Raymond, Vice-President; D. G. 
Huttleston, Secretary; Walter 8. Phillips, Treasurer. 
Respectfully submitted, » 
Clark T. Rogerson, Judge of Elections 


AMERICAN FERN Society 57 


Report of the Curator and Librarian for 1960 

This has been an important year of organization and expan- 
sion of the herbarium. Much curating was done, and numerous 
contributions were made. In our herbarium, according to counts 
made during the summer, the total of mounted and filed speci- 
mens was 8,192. The Department of Botany of the University of 
Michigan contributed help to bring the collections up to date. 
Over a thousand specimens were mounted with plastic by Misses 
Patricia Deacon and Darleen Helmick, including a large backlog 
of unmounted materials plus 50 specimens contributed by Edgar 
T. Wherry and 31 specimens sent by Daniel B. Ward. A total 
of 1,051 specimens, mostly newly mounted on herbarium sheets, 
were inserted in the Herbarium. Mr. F. W. Hunnewell kindly 
contributed in addition three sets of specimens, these already 
beautifully mounted, from his collections, a total of 310 sheets 
altogether. The results of these contributions and the mounting 
and inserting program give the Society Herbarium a total of 
9,675 specimens as of the end of 1960. 

Also, during this year a number of books and pamphlets were 
contributed to the Society’s Library by E. T. Wherry, D. M. 
Britton, Barbara Joe Hoshizaki, John Thomas Howell, and 
Thomas Darling, Jr. The use of the Herbarium and Library, 
except by people in and around Ann Arbor, has been low again 
this year, with only three requests to borrow literature. 

espectfully submitted, 
Warren H. Wacner, Jr., Curator and Librarian 
Report on the 1960 Arkansas Foray 

American Fern Society members gathered at the dormitory of 
Arkansas College, Batesville, Arkansas, during the afternoon 
and evening of August 24, 1960. By 9 p.m. about 20 members had 
assembled and were briefed for the trip by Professor Emeritus 
Dwight M. Moore, of the University of Arkansas. A well-planned 
itinerary which also included geological profiles of the Buffalo 
and White Rivers and a check list of 41 species and varieties 
of ferns which we might expect to see was distributed to the 
members. 


58 AMERICAN FERN JOURNAL 


Our first stop on August 25 was at the Blanchard Springs 
Recreational Area some 50 miles northwest of Batesville. A size- 
able stream flowing out of and over the Everton limestone in a 
wooded valley provided the habitat for several ferns and other 
eryptogams such as Rhodobryum, Thuidium, Conocephalum, and 
Dumortiera. The latter was growing close to the spring falls 
where the water came from the rocks. By following the nearly 
empty stream bed above the spring we enjoyed the coolest and 
most unusual microclimate of the trip. A sinkhole from which 
issued cool moist air (ca. 64°F) allowed the rather lush growth 
of a number of ferns and sustained some flowering plants not 
otherwise found in Arkansas. Some of the ferns seen in this area 
were Adiantum pedatum, Asplenium resiliens, Botrychium vir- 
ginianum, Camptosorus rhizophyllus, Cystopteris fragilis, Dry- 
opteris hexagonoptera, Osmunda Claytoniana, Pellaea atropur- 
purea, Polystichum acrostichoides, Pteridium aquilinum var. 
latiusculum, and Woodsia obtusa. As we left the area a search 
was made along the roadside for Cheilanthes alabamensis, the 
smooth lip-fern, but it was not found. 

During lunch time at the Barkshed Recreational Area con- 
siderable time was allowed for exploring the area on our own. 
Though many of the group were not too active after topping off 
our sack lunches with watermelon which Clair Brown and his 
family provided, several of the members discovered a nice stand 
of Athyrium pycnocarpon at the base of a tall limestone cliff. 
Asplenium platyneuron was growing in the grass at our picnic 
area. 


had to be content with views from across the water by means of 
binoculars. Other ferns seen in this camp ground area were 


AMERICAN FERN SOCIETY 59 


Athyrium asplenioides and Cystopteris bulbifera. Near a small 
spring on the hillside there was an abundant growth of Selagi- 
nella apoda and a moss, Fissidens. The capture of a large copper- 
head snake which had been basking in the sun only a few feet 
from where most of the group had climbed a bank provided the 
excitement for the day. 

Returning to an area across the White River from Calico Rock, 
we stopped at City Rock Bluff on top of a sandstone flat some 
300 feet directly above the White River. Hanging several feet 
below the top of this sheer cliff were several plants of Woodwardia 
areolata protected somewhat in the crevice where they grew, but 
certainly out of their typical habitat in swamps 100 miles to the 
southeast. Besides this chain fern other ferns found here included 
Polypodium virginianum and P. polypodioides, Dennstaedtia 
punctilobula, rare in Arkansas, and Asplenium Bradleyi with 
small tapering fronds and stipes brown at base and green at tip. 
The mosses Climacium, Grimmia, and Sphagnum were conspic- 
uous near the edge of the cliff. Selaginella rupestris grew abun- 
dantly in the open areas on this sandstone flat. 

Descending to the White River our caravan crossed the river 
on a ferry which held 3 cars and was propelled back and forth 
by the current of the river. 

One disappointment on the trip came as we discovered that the 
Bull Shoals Park Lodge, where we had planned to stop for sup- 
per and stay overnight, was temporarily without a supply of 
water. However some water was provided to wash off some of 
the day’s accumulated dust and soon afterward a bounteous 
fried chicken dinner served family style restored us to amiable 
spirits. Mr. and Mrs. Charles Dear joined us during dinner and 
continued the rest of the trip with the group. The lodge manager 
called local motels and found rooms sufficient for the entire 
group. 

Friday, August 26, on the way to Harrison, we made a roadside 
stop along the highway to observe the differences between Juni- 
perus virginiana and Juniperus Ashei. Beyond Harrison the only 
car trouble which happened on the trip occurred. Dr. Jewel 


60 AMERICAN FERN JOURNAL 


Moore’s car overheated and caused her to miss part of the trip. 
On the way in to Big Bluff, the roughest road on the trip, a pro- 
truding rock broke the exhaust manifold of Mrs. Blanche Dean’s 
car. Mr. Dear was able to wire the parts approximately in place 
and Mrs. Dean went in to Fayetteville after lunch to have it 
repaired. 

At Big Bluff the fern of most prominence was Cheilanthes 
Feet growing in the cracks of the limestone bluff. The top of the 
bluff is composed of Boone chert which showed numerous flint 
deposits in the limestone. 

We ate lunch in a little private park adjacent to Scroggins 
Store, near Ponca. The store was a rather picturesque Ozark 
mountain general merehandise-grocery store where a variety of 
items could be purchased. 

Friday afternoon we visited Lost Valley, a secluded area off 
the highway between Ponea and Boxley. Up the valley of a small 
stream a high, overhanging cliff gave a shaded place for resting 
to the few hardy members who scrambled up the rocky stream 
bed. Along the stream were several ferns, some growing quite 
luxuriantly. Those seen at Lost Valley were Asplenium platy- 
neuron, Asplenium Trichomanes, Asplenium resiliens, Adiantum 
pedatum, Botrychium virginianum, Camptosorus rhizophyllus, 
Cystopteris bulbifera, Dryopteris marginalis, Dryopteris hexa- 
gonoptera, Polypodium virginianum, and Polystichum acro- 
stichoides. 

Before reaching Boxley we were shown an excellent display of 
Adiantum Capillus-veneris growing down the side of an under- 
cut roadside embankment. From between the closely clustered 
fronds of this fern hung long slender strands of a soft lime ma- 
terial from which water dropped to the ditch below. : 

On a side road a few miles from Marble we stopped beside a 
limestone cliff of the Boone formation. Here we were able to see 
Pellaea glabella and P. atropurpurea growing close together, af- 
fording a nice comparison. Cystopteris bulbifera and Asplenium 
Trichomanes also grew along the cliff, 

This was our last field stop for the day. A short while later 


AMERICAN FERN SOCIETY 61 


we checked in at the Chief Motel in Fayetteville, where swim- 
ming, eating, and talking were all enjoyed after we were cleaned 
up from a dusty road trip. 

Saturday morning the group congregated at the Union Build- 
ing of the University of Arkansas for pictures and to get ac- 
quainted with new members of the foray. Later that morning on 
a trip west from Fayetteville we turned off the highway and 
proceeded through Savoy and on to a steep east-facing hillside 
of Boone chert. Near the top of this hill we were afforded the 
rare opportunity of seeing and photographing three species of 
Cheilanthes growing together, C. alabamensis, C. lanosa, and C. 
tomentosa. Also Notholena dealbata, Pellaea atropurpurea, and 
Woodsia obtusa were seen at this location. Further on at Lake 
Wedington we were priviledged to see an abundant growth of 
Pilularia americana (pillwort) and were told an absorbing story 
by our leader, Dwight Moore, of the rediscovery of Pilularia in 
Arkansas and of its distribution in new artificial lakes which are 
being created in the state. 

On the way back to Fayetteville we investigated a poison ivy 
infested corner of a section north of Farmington. Most of us 
were turned back by the combination of Smilax and poison ivy. 
This poison ivy was the type that grew erect through the vegeta- 
tion to about head height as a relatively unbranched young tree 
and then branched out forming a canopy that was hard to avoid. 
The three or four hardy members who braved the thicket were 
rewarded by finding Botrychiwm dissectum and B. d. var. obli- 
quum, samples of which they brought out with them to show to 
the rest of the group. 

Saturday afternoon our trip took us north of Fayetteville to 
Martin’s Bluff. This was on a stream which was being dammed 
and the whole area was to be flooded. Collecting, therefore, was 
without limitation. Since many of us were heading toward Still- 
water and the AIBS meetings, however, there was relatively little 
collecting done. There had been high water in the area but a 
number of ferns and other eryptogams were conspicuous and in 
plentiful supply. Many of the ferns which had been previously 


62 AMERICAN FERN JOURNAL 


seen were found here also. In addition, we found Onoclea sensi- 
bilis and Osmunda cinnamomea, the latter peculiar in that it was 
growing as a cliff fern. A frond of Dryopteris marginalis found 
at this location was the smallest fertile marginal shield fern that 
I had ever seen. 

We were able to see at least 36 of the 41 ferns on our list. All 
of us are indebted to Dr. Moore for his very capable leadership 
and for arranging such a fine trip. 

Those attending the foray all or part of the time were Mr. and 
Mrs. Ralph H. Benedict, Mr. and Mrs. Clair A. Brown, Sarah 
Brown, Alan G. Cazart and daughters Cissy and Jeanie, Maxine 
Clark, Blanche E. Dean, Mr. and Mrs. Charles Dear, Elizabeth 
Eddy, Inez Hartsoe, Donald G. Huttleston, Sylvia B. Leather- 


mond, Eva Sobol, and Mary Walker.—Rosert C. LoOMMASSON. 


NEw MEMBERS 
Mrs. Elizabeth Arnold, Woodside Fernery, Route 2, Box 848, Jacksonville 
ida 


7, Flor 
Mr. M. Bange, 24 Rue Gr renette, Lyon 2, Frane 
Mr. Henry S. Bl ethen, 47 Federal St., Roan: M 


F a 
. R. Granville Curry, River Road, Rockville, Md. 
Angie Estill, 153 Navarre Drive, Miami Springs, Florida 
Mrs. Brooks Evert, 430 Thomas Avenue, Riverton, New Jersey 
: Ph, On e 


ss. 

Mrs. Edward I, Gr iffith, 135 Athania Place, Metairie, Louis 

Mr. Edward N. Hallman, 2308 Coventry Avenue, Lakeland, “Florida 

Mrs. David E. Harris, 713 E, Orange St., Lakeland, Florida 

Mr. N. Keith Harrison, Dept. of Hetaay. and Bacteriology, Montana State 
College, Bozeman, Mont. 

Mr. David’ W. Hutchings, Science Department, Cazenovia Junior College, 
Cazenovia, New Yor 

Miss Alice C. Jones, 218 High St., Mt. Holly, New Jersey 

Dr. Alan Johnston, Route 6, Martinsville, Indiana 


AMERICAN FERN SOCIETY 6: 


. E. W. Johnson, 15 Elinor estes Mill Valley, Calif. 
Mrs John N. Keers, Sr., 4290 Ristow Drive, La Canada, Cali 
Miss Ethel E. King, 120 South ae eg Drive, Dunedin, Florida 
vas Helen B. Krech sane pang sse 
r. Charles leu i Wes h St., Heian 8, Texas 
Mra J. L. Linkenhoger, 45 ene Trail, Houston 24, Texas 
Mr. D. A. MacInnes, Rockefeller Institute, New York 21, N. Y. 
Mr. Harry F. Malone, P.O. Box 1333, Cocoa, Florida 
Miss Norma A. Maloney, 1600 N. Rodney St., Wilmington 6, Siinerene 
ts. Lee B rsha 


Mrs. James H. Mason, Box 251A, Route 1, West Terre ye Indiana 

Mr. Frank F. ges 830 Sheridan Road, easton, Tllin 

Mrs. Martha H. Mosier, 490 West Washington St., eae India 

Mr. SOE poe Botany Department, Univesity of Washi me Se- 
attle 5, shington 


g 
W. P. Murray, 249 Seaview Avenue, San Rafael, Calif. 
k: 


Mr. James D. Perry, Dept. of Botany, University of Georgia, Athens, Ga. 
Mr. George Postma, 847 Courtney St. N.W., Grand Rapids 4, Michigan 
Mr. L i 


rL.,G; dale 
Mrs. Arno Reddel, Harbert, Michigan 


Mr. Donald E. Tate, 314 E. Garland St., West Frankfort, . 
Mr. William Carl Taylor, 8914 Francis Court, St. Louis 23, Misso 
r. Edmund A. Turnau, 20329 Lakeshore Road, Baie ean pre te 


Can: 
Mrs, P. = tecia: P.O. Box 516, Brazoria, Tex 
Mrs. E. O. Williams, 2601 Acacia Court, Fort AG Matte Florida 
Mr. David L. Woolsey, Route 1, Box 140, Ione, Calif. 


CHANGES OF ADDRESS 
Dr. Ralph D. Ade, 2805 26th St., Moline, Illinois 
Mrs. Lenore Alexander, 2104 Grant Ave., Redondo Beach, Calif. 


2 . n, Conn. 
Mr. Osear J. Byers, Jr., 563 28th St., Oakland 9, Calif. 
Mr. Thomas O. Cie, ne Peachtree St., N.E., Atlanta 9, Georgia 


64 AMERICAN FERN JOURNAL 


Mrs. Ruth W. Cotton, 13 High St., Beverly Farms, = 

Miss Tone Covert, 1105 Elizabeth St., So ees Norfolk 

Mrs. Lewis L. De afield, Piermont Road, Hewlett, oo York x 
Dr. eee E. DeVol, 65 Wan Shen ce are ath se Taiwan, Free 


Mr. meme K. Eosefow, 83 Salem St., Andover, Mas: 
Mr. acme M. Evans, Dept. of Botany, University of gat Ann Arbor, 


Mrs. eden T. Filer, 12665 Highview Lane, Redlands, Calif. 
Mrs. George N. Gardiner, Pikes Hill Ro: a Weston, Vermont 


Ohi 
Mr. Ray MeAdams, Apt. E-4, Harold Apts., 1130 Woodward Heights Blyd., 
Hazel Park, Michigan 
Mrs. Juanita S. McCoy, Biology Dept., Salem College, Salem, West. Virginia 
i 


ott 
Dr. Howard F. L. Rock, Dept. of Bi iology, cao ui pie Nash- 


ville 5, Ten 
Mr. George A. eS 13611 Wheeler Place, Santa Ana, Calif. 
Mrs. Maxine B. Sites, P.O. Box 562, Sweet Hom e, Orego 

Mrs. John Stephan, Route 2, Box 744, Pinson, Alabama 


vi 
Mrs. Carl Sybertz, South Millie Road, Rests 1, Box 298, Valrico, Florida 
Albert R. Tenney, Box 45, R. D. 1, Tor ronto, Ohi 
Dr: Stanley Walker, Dept. of Genetics, University of peo’ Liverpool 3, 
nd 


Mrs. Pride Wertman, 16632 Eagle Ridge Drive, Hopkins, Minn. 


HENRY TRIPP 
Successor to Henry George Fiedler 


Scientific Books and Periodicals 


Large stock of books on ferns 
and cryptogamic botany 


31 East 10th Street, New York 3, N. Y. 


JUST PUBLISHED! 
THE FERNS AND FERN-ALLIES OF WEST TROPICAL AFRICA, 
by A. H. G. Alston (British Museum). Keys and deseriptive text, line 
drawings. 1959. Paperbound, octavo, 89 pp. $1.65, postpaid 


Atso In SrTock 


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bound, 1954, $10.24, postpaid 
EDWIN A. MENNINGER 


The Flowering Tree Man 
Stuart, Florida 


FERNS 
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CAMBOSCO. i ag Mb Sl COMPANY 
37 ANTWERP STR c BRIGHTON STATION 
BOSTON, MASS. U.S.A. 


Vou. 51 APRIL-JUNE, 1961 


No. 2 


American Fern Journal 


A QUARTERLY DEVOTED TO FERNS 
Published by the 


ERICAN FERN SOCIETY 
is 


RYON, JR. IRA L. WIGGINS 


ts 
CONTENTS 


My Lath House for Ferns Fay MacFappen 


A Parcel of Ferns from Stephens County, Georgia 
GLADSTONE W. McDowWELL AND FRANKLIN D. SNYDER 


Another er emg Hybrid from Kentucky 
M. Suita, Truman R. Bryant, AND Donatp E, Tate 


Some Forms of Polypodium californicum C. V. Morton 


On aed a _—— — > Fertile —- in Bot 
and B. oneidense_.____-_____. Ae: fone et 


hea Genus of Ferns New to the United States___.C. V. Morton 
Isoétes echinospora Durieu in North America BERNARD BOIVIN 


A Hawaiian Thelypteroid Fern with Peltate Indusia 
Kunio IwatsvkKi 


Observations on a material of Azolla — 
R. K. Goprrey, Grapy W. REINert, AND RIc . HouKk 


Adaxial Sori in Polypodium hesperium_....___Emity L. Hartman 


Heat Resistance of Sporocarps of Marsilea Ap IS 
ILLIAM W. BLOOM 


Feltapteris in Costa Riea Ciype F, REep 
Cytology of Isoétes coromandelina._____-__- S. C. Verma 
meee Fern ENerwturn 2. 
—— Fern Society: Report of tg ——* Constitution 


f the American ye Gute: Inc 1 


a 
a 


The American Fern Hocietyp 


Council for 1961 
OFFICERS FOR THE YEAR 

Cuair A. Brown, Department of Botany, Louisiana State University, Baton 
Rouge, Louisiana President 
MagceL RayMonpD, oo Botanical Garden, 4101 East — Street, 
Montreal, Cana e-President 

DONALD Sen Longwood Gardens, Kennett Square, pace 
Secretary 
WALTER 2 — Department of Botany, University of a oe 
Ariz asurer 

Cc. V. iecena. Smithsonian Institution, Washington 25, D. C. 

Editor-in-Chief 


OFFICIAL ORGAN 
American Fern Journal 


EDITORS 
C. V. Morton. Smithsonian ome — blige “es 25, D. 
Rota M. Tryon, 22 Divinity Avenue, Cambridge, Mas 


Ira L. Wicerns____. one Herbarium, santora P eiareansity, Stanford, Calif, 


An illustrated quarterly gs to ay general study of ferns, published 
ei by the American Fern Society, 3110 Elm Ave., Baltimore 11, Md. 
ond-class postage paid Be “ocanc 
“Matter for publication should be addressed to C. V. Morton, Smithsonian 
Institution, Washington 25, 
ery oy $2.50, exclusive of agency handling fees; sent free to oor 


bers the erican Fern Society ogee dues, $2.50; sustainin 
Seven, $5. vat bare bet arapt Yours - do $50.00). Extracted reprints, if par ore in 


Be is 
ack volumes $2.00 each; single back numbers 50 cents each; Cumulative 
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Changes of address should be sent to the Secret tary: Dr. Donald Huttle- 
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LIBRARIAN AND CURATOR OF THE HERBARIUM 
Dr. W. H. Wagner, University of Michigan, Ann Arbor, Mich. 


egular loan department is maintained in connection with the library 
and herbari bers may bor books and s D: any time, 
t rrower paying all postal or ag oa —- The pages of the Ji ournal 


American Fern Journal 


Vou. 51 APRIL-JUNE, 1961 ' No. 2 


My Lath House for Ferns 
Fay MacFappen 


My early married years were spent in British Columbia, where 
I first became interested in ferns. Later we moved to Los An- 
geles. When in 1935 we eventually found a house on two lots, a 
lath house started to go up almost before we got settled. The 
first one was not large, only 16 by 20 feet with a roof eight feet 
high, but built so that it could grow in time. The uprights, red- 
wood four by fours, were cemented into the ground, and the 
framework was constructed of two by fours, with laths for the 
roof. It is better to have a small lath house well conditioned 
than a larger one poorly cared for. After the small one is estab- 
lished, you can always add to it. 

This part of Los Angeles had once been swampland and salt 
grass grew everywhere. Salt grass roots are ropelike and go very 
deep, forming a sort of impenetrable mass of roots that some- 
times completely cuts off drainage. Storm drains had not yet 
come and the water table was high in those years and the soil 
full of salt. Such a situation was mine, and so I eventually dug 
up the soil on the two lots two to four feet deep. I had a little 
grass and a mountain of roots to dispose of. But it was neces- 
sary to establish good drainage. 

A foundation of six inches of rocks was laid down. Every 
time we went for a drive, the old Hupmobile came home groan- 
ing with the weight of rocks. And we made many trips to the 
mountains for leaf mold. Our home was not far from peat beds 
at the foot of Baldwin Hills which used to catch fire and burn 
for months. Eventually the city controlled the fires and used the 
spot as a dumping ground for city sweepings. Over the years a 


Volume 51, No. 1, of the JouRNAL, pp. 1-64, was issued Apr. 20, 1961. 


66 AMERICAN FERN JOURNAL 


great quantity of leaves were deposited there and a bed of leaf 
mold several feet thick had been formed. The city was letting 
builders dump earth there so that it could be reclaimed ; and so 
we got a truck and sacks and anything that would carry leaf 
mold and worked feverishly to get quantities before the big 
earth-loaded trucks began to roll. I feel that much of my success 
with ferns is due to the leaf mold that I acquired at that time. 

The leaf mold was mixed with peat moss, not the finely ground 
kind that is now sold and which is soon gone but the fibrous peat 
that used to come baled from Europe and which we can not get 
now; the fibrous peat really built the soil and lasted for years. 
Ser the years I’ve used many ‘bales of peat moss. If you are 
buying moss at most nurseries in Los Angeles they will try to 
sell you Oregon green moss and tell you it is the same as sphag- 
num. However, this is not so. The green moss can be used for 
lining baskets but must have soil put in to make a growing me- 
dium. However, there are a few nurseries that will get baled 
sphagnum for you, and this can be used as the growing medium. 
It holds about seven times its weight in water. It is much cheap- 
er than Oregon green moss. 

In the lath house I planted several small Japanese maple trees. 
One of these, a four-pronged tree, is huge now. Due to its de- 
ciduous habit it allows sun in the winter and shade in the sum- 
mer. It comes into leaf in early April. The leaves fall in No- 
vember and I have them to mulch the ferns that grow under- 
neath. In spite of the network of tree roots, the ferns thrive. 
However, if one wants to move a fern it has to be cut out, and 
unfortunately some ferns have to have their location changed 
many times before the right spot is found for them. 

Between the trees we built a fish pond, which is ornamental 
and also provides moisture for the atmosphere. Basket ferns 
hang from the maple trees over the water. In a few years we 
added.to the lath house and eventually had 2000 square feet un- 
der lath, except that now we do not use lath but old fluorescent 
tubes. These are to be had free. They provide a light shade, 
have the advantage of lasting indefinitely, and are also orna- 


FERNS FROM GEORGIA 67 


mental. The back end of the lath house is a stucco wall, now cov- 
ered with ivy, where originally there was a beautiful old pepper 
tree. In recent years, with the construction of storm drains, the 
water table has gone down and after several drought years all 
the old and beautiful pepper trees in the neighborhood died, in- 
cluding ours. 

In the completed lath house we laid out brick walks and con- 
structed a little ravine where some rock ferns could be grown. I 
also planted an English yew tree, to satisfy my desire for the 
woods. In the center of the area I planted a Woodwardia radi- 
cans at the time the house was enlarged. It is tremendous now. 
The arching fronds produce buds which root and form new plants 
in the winter and late spring. These I cut away and hope I can 
find someone who wants them. The central area has several tree 
ferns also. 

The design for a lath house planting, or any other kind of gar- 
den for that matter, is a personal matter, like painting a picture: 
Two can’t do it. Really, if you want a fern garden, it is because 
you love ferns. It is the gardener who hasn’t much money to 
spend but has lots of love to give his plants who has the best 
luck with ferns, for every fern must be studied and loved for 
itself alone. 

In future articles I shall write about the kinds of ferns suit- 
able for a lath house in the Los Angeles area and about some of 
the problems to be solved in growing them. 

5450 Carin St., Los ANGELES 16, CALIFORNIA. 


A Parcel of Ferns from Stephens County, Georgia 

GuapstonE W. McDoweELL AND FRANKLIN D. SNYDER 
Recent explorations in the valleys of two streams tributary 
to the Tugaloo River in Stephens and Habersham Counties, 
northeast Georgia, have resulted in the finding of several ferns 
previously unreported in that region or which are rare in 
Georgia. The streams, Panther and Toccoa Creeks and their trib- 
utaries, lie partly in deep gorges that are not easily accessible. 
Perhaps the more spectacular gorge of the Tallulah River has 


68 AMERICAN FERN JOURNAL 


diverted the attention of botanists from these gorges which are 
only a few miles to the sou 

Several years ago we became interested in the area south of 
the Tallulah River because of published reports of limestone out- 
crops in the Brevard schist through which Panther Creek has cut 
deeply. It was not until 1957, however, that any field work was 
done. In January of that year the authors searched part of 
Panther Creek. The precipitous sides of the gorge made walking 
difficult, and at times it was necessary to wade the creek. Sheer 
cliffs rose from the water at our stopping place, and a sharp 
bend prevented a view farther upstream. A few species of ferns 
were seen, but none that would indicate the presence of limestone. 
One was Asplenium montanum, a first record for Stephens 
County. The most unusual plant observed was Carex plantaginea, 
a striking sedge reported previously in Georgia only from Rabun 
County. 

Subsequent trips were confined mostly to the portions of the 
streams in Stephens County, and the ferns mentioned hereafter 
are from that county. The second trip to Panther Creek, which 
was made in January, 1958, a year later, resulted in the finding 
of Asplenium resiliens on a small outcrop of limestone. This fern 
was new to northeast Georgia, the nearest station in Georgia 
being about 60 miles to the west. 

On March 22, 1959, larger exposures of limestone were dis- 
covered in the Panthes Creek gorge, where we found Cheilanthes 
alabamensis, Pellaea atropurpurea, and Camptosorus rhizophyl- 
lus. The Alabama lip-fern seems to be rare in Georgia and had 
been reported only in the Appalachian Valley of northwest 
Georgia. Also, C. tomentosa was collected in the gorge, and C. 
lanosa, though not seen there, occurs elsewhere in Stephens 
County. The walking fern was abundant, and forma auriculata 

. Hoftm. was collected. Some of the leaves of this form had only 
the right or left lobe elongated. 

The glade fern, Athyriwm pycnocarpon, and the brittle fern, 
Cystopteris fragilis, were found on May 16, 1959. The glade fern 
was abundant in one of the numerous small ravines tributary to 


FERNS FROM GEORGIA 69 


the gorge. This collection added a third county in which it has 
been reported. The brittle fern, which fits the description of the 
variety protrusa Weath., grew with the glade fern and in several 
other places on cool, north-facing slopes. It has been reported 
from five other counties, only one of which is south of Stephens 
County. 

In the summer of 1958, Mr. Henry W. Bookout found the first 
station for the walking fern in Stephens County in the deep 
ravine of Cedar Creek, a tributary of Toccoa Creek. A few days 
after that discovery, he and Snyder collected Trichomanes 
Petersti at the same locality. In October, 1958, Snyder collected 
Adiantum Capillus-veneris in the ravine. The southern maiden- 
hair has previously been known in north Georgia only from one 
station in Habersham County. 

On March 21, 1959, we visited Cedar Creek to see T. Petersu 
and to look for additional specimens of the southern maidenhair. 
None was found, but a thorough search was prevented by bad 
weather. One additional fern for Stephens County, Asplenum 
Trichomanes, was found on cliffs in the ravine. On June 21, 1959, 
McDowell made another visit to Cedar Creek and found a num- 
ber of plants of the southern maidenhair growing among the 
more abundant northern maidenhair. 

Much of the whole area remains to be explored, and we do not 
yet know what lies beyond the sharp bend in Panther Creek 
where the cliffs rose from the water. The two streams rise on the 
divide between the Gulf and Atlantic drainage systems formed 
by the Chattahoochee Ridge and the Tallulah Mountains. The 
maximum elevation reached is about 2,800 feet. All the ferns 
mentioned were found at elevations of about 900 feet. The pre- 
vious records given in this paper are based on Pyron and Me- 
Vaugh’s Ferns of Georgia and a mimeographed list of additional 
records prepared in 1957 by Dr. Wilbur H. Duncan, of the Uni- 
versity of Georgia. 


328 Hearp Srreer, ELBERTON, GEORGIA, AND 198 VALLEY Roan, 
Toccoa, Grorata 


70 AMERICAN FERN JOURNAL 


Another Asplenium Hybrid from Kentucky 
Date M. Suiru, Truman R. Bryant, and Donat E. TATE 


Intensive collection of Aspleniums in the Cumberland Plateau 

region of eastern Kentucky during the past two years has yielded 
much material of the species and hybrids comprising the ‘‘ Appa- 
lachian Spleenwort complex’? (Wherry, 1925, Wagner, 1954). 
All the basic species are present in this area; the most commonly 
encountered hybrids are A. X trudellii Wherry, and A. X 
gravesii Maxon (Smith, Bryant and Tate, 1961). It seems quite 
likely that all the possible hybrid combinations in the complex 
may eventually be found in this limited area. 
- One of the most elusive hybrids in the complex is that involv- 
ing A. Bradleyi D. C. Eaton, and A. montanum Willd. The first 
record of this hybrid was that of Wherry in 1935, based upon a 
specimen collected by him in New Jersey. This was followed by 
the publication of a line drawing of two lower pinnae of a rather 
large frond with the statement, ‘‘Exactly intermediate between 
parents’’ (Wherry, 1937). No additional information about this 
hybrid was given by Wherry in 1942, nor was mention made of 
this hybrid by either Fernald (1950), or Morton (in Gleason, 
1952). Wagner (1954) mentioned the hybrid briefly and pre- 
dicted that the plant would be a triploid. Evidently no addi- 
tional material of the hybrid has been detected since Wherry’s 
original discovery. 

It was our good fortune to find two plants of this hybrid in @ 
dry sandstone crevice, along with both putative parents, near 
Bear Track Lookout Tower, Lee County, Kentucky. These plants 
were quite small individuals, and were unfortunately overlooked 
as possible hybrids until the pressed material was studied later. 
Consequently, it was impossible to verify Wagner’s prediction 
of the triploid chromosome number for these plants, but the 
spores were abortive as in many other Asplenium hybrids. 

In consideration of the paucity of information about this hy- 
brid it seems well to give it a binomial and a more complete de- 
seription. It is a pleasure to name this fern in honor of one of 


AMERICAN FERN JOURNAL VoLUME 51, PLATE 3 


é 


a % 


Fig. 1. ASPLENIUM X WHERRYI, WHOLE PLANT FROM Lee Co., Ken- 
TUCKY; Fig, 2. A. X WHERRYI, SINGLE FROND FROM TYPE SPECIMEN FROM 


a 


Ss. 3- 
FROM Sure Rock CrevicE, LEE Co., Kentucky; Fic. 3. A MONTANUM; 
I - X WHerryI; Fig. 5. A. Brapiey|. ALL NaTuRAL SIZE. 


12 AMERICAN FERN JOURNAL 


the foremost students of American ferns. Dr. Edgar T. Wherry, 
of Philadelphia. 

AspLentum X Wherryi, hybr. nov. (A. bradleyi * montanum). 
Stipitibus — usque ad 2 em. longis, laminis foliorum semper- 
virentibus, lanceolatis, basi bipinnatis, sursum bipinnatifidis, 

apice pinn a pinnis 10-16, alternis, rachibus viridibus, 
do tak a otis 

Typ Beanie cliff 4.5 miles northwest of Blairstown, War- 
ren County New Jersey, Edgar T. Wherry, September 1, 1935, 
no. 725,2: PH 

This ene probably triploid hybrid is, as pointed out by 
Wherry, essentially intermediate between the putative parents 
in several A aiaahataas The stipe as well as much of the rachis 
is darkened in radleyi, but only the stipe bases are dark- 
ened in A, Sain In A. X Wherryi, the darkening extends 
only to the base of the frond blade, the rachis being entirely 
green. The overall shape of the frond in A. Bradleyi is typically 
narrowly lanceolate, in A. montanum decidedly triangular, often 
nearly deltoid, and in A. X Wherry lanceolate. The extent to 
which the fronds of both parental species are cleft is quite vari- 
able, but in A. Bradleyi only the basal pinnae are regularly deep- 
ly cleft. In A. montanwm it is not uncommon to find bipinnate 
fronds, or those which are occasionally subdivided to an even 
greater extent. The type specimen of A. X Wherryi is bipin- 
nate in the lower half, grading gradually near the tip to bipin- 
natifid, and at the apex it is pinnatifid. 

In eastern Kentucky, and perhaps elsewhere, A. Bradleyi and 
A. montanum occur rather abundantly in crevices of sandstone, 
often in contiguous clusters. Consequently, it is surprising that 
more specimens of A. X Wherryi have not been found. If eco- 
logical differences exist they are slight, with A. Bradleyt occu- 
pying slightly drier crevices, and A. montanum occurring in 
more moist or shaded places. More often than not the two par- 
ental species occur together, and perhaps with more careful 
searching additional hybrid individuals will be found. 


POLYPODIUM CALIFORNICUM 13 


LITERATURE CITED 

FERNALD, M. L., 1950. Gray’s Manual of Botany, 8th Ed. New Yor 

Morton, C, V., 1959. In Gleason, The New Britton and Brown Tihstratet 
Flora, ed. 3. New Yor 

Situ, D. M., TRUMAN R. BRYANT, AND D. E. Tate, 1961. Asplenium 
Gravesiti in Hentnsky: Brittonia 13: 69-72. 

Waener, W. H., Jr., 1954, Reticulate Evolution in the Appalachian Asplen- 
iums. Evol. 8: 103-118. 

WHERRY, ae T., 1925. The Appalachian Aspleniums. Amer, Fern Journ. 
5: 47-— 


aes Fern Field Notes, 1935. Amer. Fern. Journ, 25: 123-126, 


——____—_,, 194 
Universrry oF ILLINOIS, URBANA, ILLINOI 


Some Forms of Polypodium californicum 
C. V. Morton 
Mr. Donald Branscomb, of Willits, California, an amateu 
member of the American Fern Society, has been enthusiastically 
studying and growing the ferns of northern California. He has 
recently sent me a frond of a Polypodium californicum that he 
found in Humboldt County in which the lower segments are 
deeply lobed rather than merely serrate as they usually are. This 
is the sort of variation that is to be expected in species of the 
vulgare alliance, of which many forms have been described in 
Europe. No such form has ever been described in P. californi- 


shows. It is perhaps worth while giving this form a name: 
PoLypopium CALIFORNIcUM Kaulf, forma Bescascombil Morton, f. 
nov. 

A forma typica segmentis inferioribus perspicue lobatis lobis 
latis obtusis serratis 3—8 utrinque latere differt. 

Type in the U. S. National Herbarium, no. 2,257,987, collected 
on the Redwood Highway, 2.8 miles south of Trinidad, Hum- 
County, California, December 25, 1958, by Donald Brans- 


ds looking through the material of Polypodium californt- 


74 AMERICAN FERN JOURNAL 


cum in the National Herbarium, I came across another peculiar 
form which has lain undescribed in the collections for the last 
64 years. It is a form with the segments deeply laciniate and is 
a dead ringer for P. vulgare f. cambricum, the well-known Eng- 
lish fern, except this plant is surely P. californicum. It was 
found in one of the canyons coming down from Mount Tamal- 
pais, in Marin County, California, in 1895, by Miss Mary Eliza- 
beth Parsons. Miss Parsons sent additional material to Dr. Max- 

on eight years later, writing as follows: 
San Rafael, California 
September 14, 1903 

Dear Dr. Maxon: 

Several years ago I found in one of our canyons a Se 


The ileus year I found it again in the same locality but have 
never seen it anywhere else. 
I bro 


for more, and at any rate there would be none now at this dry 
season. Perhaps next spring I may be able to examine the spot 
again 

Will you please name and publish this form if you think it 
worthy of doing so? If you do publish it, I should be glad to 
know of it. The wig hg where this fern was found is on the 
property of Mrs. A. E. Kent (my aunt), Kentfield Sorae - 
oe North Shore Railway about two or three miles north of t 


by _one of the little streams making down from Mount Tamal- 
pais. The altitude was probably not over 100 feet, if that. 


Yours truly 
Mary ELIzABETH PARSONS 


1 Actually P. californicum. C. V. M. 


FERTILE SEGMENTS IN BotrycHIUM 1D 


PoLyPopIuM CALIFORNICUM Kaulf. forma Parsonsiae Morton, f. 
nov. 
rma typica lamina bipinnatifida, segmentis, saltem in- 
Dae ae fere usque ad costulam partitis differt. 

Type in the U. S. National Herbarium, no. 434,989, from a 
plant cultivated in a greenhouse in California: collected by Mary 
Elizabeth Parsons, September 14, 1903, originally transplanted 
from a canyon near Kentfield, Marin County, California. A par- 
atype in the U. 8S. National Herbarium, no. 469,766, is a similar 
specimen collected from the wild at Kentfield by Miss Parsons, in 
March, 1895. 


On the Relative Development of the Fertile Segments 
in Botrychium dissectum and B. oneidense 


W. H. Waener, JR.? 


The taxonomically controversial Botrychium oneidense (Gilb.) 
House differs from B. dissectum Spreng. in a number of char- 
acteristics, including the average shape and size of the segments, 
the pigmentation and periodicity of the primordial and mature 
leaves, the root diameter, and the geographical range. It has 
been ia a distinct species, a variety of B. dissectum, a 
variety of B. multifidum, or, possibly, the hybrid of B. dissectum 
and pee iy Botrychium oneidense is said to produce fertile 
segments less commonly than B. dissectum, and this has been ad- 
duced as an argument for the-theory that B. oneidense may be 
an interspecific hybrid of the two species with which it has been 
associated. The present paper adds new data on the relative de- 
velopment of the fertile segments, and discusses briefly some of 
the implications. 

Some years ago it was pointed out by Robert T. Clausen? that 


1 Re search supported by a grant from the H. H. Rackham School, Uni- 
versity of Michigan 

2 On the status of Botrychium dissectum var. oneidense. THIS JOURNAL, 
34: 55-60. 1944. 


76 AMERICAN FERN JOURNAL 


“In central New York var. oneidense is fairly common. Around 
Utica it apparently fruits as freely as do any of the other vari- 
eties of B. dissectum. In the section around Ithaca and the south- 
ern tier of counties in New York this is definitely not the case, 
for the plants develop fertile panicles only rarely.’’ His survey 
of his collections showed that only 15% of a total of 72 speci- 
mens of oneidense possessed these structures; but in ‘‘obliquum’’ 
(B. dissectum) 54% of a total of 148 had fertile segments, i. e., 
fertile segments were, relatively, over three and a half times as 
common. Clausen did not say, however, whether or not his ma- 
terials were comparable, i.e., from the same localities. In the 
Cornell University herbarium, his count showed that 66% of a 
total of 72 oneidense were fertile; and 88% of 217 specimens of 
obliquum were fertile. He suggested the hypothesis ‘‘that var. 
oneidense may be an interspecific hybrid, only more or less fer- 
tile,’’ on the basis of ‘‘a marked tendency toward lower fertility”’ 
plus some morphological and geographical observations. It is 
only with the ‘‘marked tendency toward lower fertility’’ that I 
am concerned here. There are two questions involved: (a) If 
B. oneidense were truly an interspecific hybrid, would this nee- 
essarily lead to the expectation that formation of fertile leaf parts 
would be influenced by its hybrid nature?; and (b) Are the data 
used for demonstrating the lower fertility of B. oneidense reli- 
able? The latter question will be discussed first. 

Comparative materials of different taxa of evergreen grape- 
ferns should be taken in the same localities, where soil conditions, 
climate, and associated plants are alike. It would be best if we 
could grow these plants together in routine greenhouse culture, 
but grapeferns are notoriously difficult to grow in pots and they 
usually die. The data reported here are from mass collections 
made by the author, assisted by one to five other collectors, and 
from each locality every leaf, young and old, seen during the 
time available was taken. The counts were made by Misses V. M. 
Morzenti and Therese Signaigo. The leaves were plucked at 
ground level and pressed. There is the possibility of occasional 
errors in rare cases where the leaf separated above the insertion 


FERTILE SEGMENTS IN BorrycHIUM E 77 


of the fertile segment or the fertile segment was completely de- 
stroyed by some natural agent. 

All of the 14 localities from which the samples were taken are 
in southeastern Michigan, roughly 350 miles west of the areas in 
which Clausen found such sharp differences in the relative pro- 
duction of fertile spikes. Table 1° gives a comparison of suec- 
cessive mass collections from three of our best localities for B. 
oneidense. Locality 2 showed a consistently low percentage of 
fertile spikes for both species in the years 1958 and 1959, not 
more than 5% of either species bearing fertile segments either 

year. However, localities 1 and 3 had higher percentages: 
1958, locality 1 produced approximately half the percentage of 
Portits leaves that it did in 1954 and 1959. Locality 3 had a great- 
er percentage of fertile leaves in B. dissectum than in B. onei- 
dense in 1958, but in the following year the proportion was re- 
versed. These data suggest that the number of fertile leaves per 
Species may vary from locality to locality and from year to year. 

Table 2 compares the averages of localities 1, 2, and 3 with 
mass collections from 11 additional localities. One point will be 
immediately evident to persons familiar with B. dissectum, as it 
grows, for example, in meadows and fields—in this table B. di. 
sectum shows a surprisingly low percentage of fertile spikes, 
and, with one exception, considerably lower than the 54% re- 
ported by Clausen. This may be explained, I believe, by the fact 
that the localities listed in Table 2 are all low, swampy, acid 
woods, the usual habitat of B. oneidense, with very rare excep- 
tions. But Botrychiwm dissectum is a much more common and 
generally distributed plant, with indications of a much wider 
amplitude of habitat and range tolerances, for it grows not only 
in localities where B. oneidense occurs, but in widely different 
habitats, often in much more open and exposed situations where 
it becomes much more freely sporangiferous. 

The average percentages of fertile segments in all 14 locali- 


3 All collection numbers of the writer’s will be reported with locality on 
Da’ nah in the near future as part nes a report on the g 


by le J... Hi 
graphical distribution of Ophioglossaceae in Michi igal 


78 AMERICAN FERN JOURNAL 


ties show that there is indeed a difference between B. dissectwm 
and B. oneidense, the average of B. dissectum being greater. 
However, seven out of 14 localities gave a difference of less than 
5% in the production of fertile leaves, and, in fact, in four lo- 
calities, B. oneidense exceeded dissectum in the percentage of 
fertile leaves. I would not call this a ‘‘marked tendency toward 
lower fertility.”” The data suggest that in Michigan the condi- 
tion is like that stated to be true around Utica, New York, «.e., 
B. oneidense fruits almost as freely as the varieties of B. dissec- 
tum. Studies should be made to determine why B. oneidense is 
more sterile in the area around Cornell University. 


TABLE I. epee oF sphap eouleg — COLLECTIONS 
FROM 


E Loca 
B. sy a B, ONEIDENSE 

No. Fert. o. Fert. 
Coll. lvs. lvs. % Coll. lvs. lvs. % 
4-29-54a (1954) 90 9 10.00 429-54b 25 4 16.00 
Loe. 1 8521 (1958) 29 2 6.89 8522 50 3 6.00 
9109 (1959) 154 21 13.63 9110 50 7 14.00 
av. 10.17 av. 12.00 
8599 (1958) 28 1 3.57 8600 156 5 3.21 
Loe. 2 9124 (1959) 60 1 1.66 9125 114 4 (3.52 
av. 2.62 av. 3.37 
8631 °-(1958) "70 | "20-2857 8630 132. 18 13.64 
Loe. 3 9059 (1959) 55 11 ~ 20.00 9061 60 17___ 28.33 
av. 24.29 av. 20.99 


W. L. Dix pointed out in 1945¢ that ‘‘var. oneidense [which 
he considered a variety of B. multifidum|] is a plant of the woods 
and is seldom found in fields’? and that ‘‘insufficient sunlight 
is a common cause of sterility among most plants.’’ He wrote 
further that, according to his evidence, when B. dissectum occurs 
in shady places (‘‘woods and thickets’’) it too has reduced fer- 
tility. The data reported here from southern Michigan tend to 
support Professor Dix’s idea that the ‘‘theoretical sterility of 


4 Observed Characteristics of Botrychium multifidum var. oneidense. 
THIS Joueeis, 35: 37-39. 


FERTILE SEGMENTS IN BotrRYcHIUM 79 


- . . oneidense may be due 2 environment rather than to hybrid- 
ity or a juvenile condition.’ 

If Botrychium oneidense were an interspecifie hybrid, which I 
do not believe it is, might we expect that the formation of fertile 
parts of the leaf would be arrested or curtailed by factors associ- 
ated with hybridity? Among the many sterile interspecific hy- 
brids known among ferns, there is no reason known to me why 

leaf axes or parts of leaves that bear sporangia should become 
aborted. The leaf blade and its fertile parts tend to develop nor- 
mally in sterile hybrids. The sporangia themselves also develop 
in the usual way after the fertile axes or fertile laminae mature. 


TABLE II. COMPARISON OF VARIOUS COLLECTIONS FROM 14 LOCALITIES 


B. DISSECTUM B. ONEIDENSE 
No. Fert. No. Fert. 

Coll. lvs. lvs. % Coll 1 lvs % 
See Table 1 nae An ailes 10.17 ae eos wi 12.00 
2 See Table 1 Ngee 2.62 ee Soe YA 
See Table 1 LF areas 4.29 Lae Cal eae 20.99 
8374 114. ih 13.15 8373 79 BS > 1648 
8394 4 4 0.00 8393 18 2 Ait 
€ 8403 48 15°.°31L.25 8401 63 8 12,88 
8519 10 vi 6.60 8517 68 0 .00 
8631 7 20 28.57 8630 132 18 13.64 
8641 7 6 8.21 8640 108 14 96 
1 8648 5 16° 28.07 8647 122 15.45 
11 8676 4 6 14.00 8675 53 4 D5 
12 9068 56 3 5.35 9069 28 2 oF 
13 9101 252 171, - 67.85 9102 4 31 48.44 
14 9119 46 2 6.08 9120 Bi ey Gace © § 3.67 
av, 19.73 av. 13.26 


Only at the time of meiosis, when spore production takes place 
do irregularities in chromosome distribution and other factors 
become apparent that tend to produce abortive spores. Often the 
failure of the spores to develop within their sporangia is so severe 
and early that the sporangia themselves become more or less 


uld be interesting to transplant B. oneidense from its ordinary 
woodland ‘habitata to open fields and meadows where B. dissectum (as well 
as B. multifidum and B. ternatum) tend to be highly fertile, and thus de- 


t fata, whether the woodland species ca made to react as does is- 
sectum and produce more fertile structures. Such pine are planned by the 
present author; it is possible, however, tha ense wi Cas narrow 


habitat requirements may not tolerate or survive pen condition 


80 AMERICAN FERN JOURNAL 


arrested, wes they may even fail to undergo the normal dehis- 
cence pro 

The fertile alcibienton of Botrychium pass through an extensive 
growth period prior to the maturation of the spore-mother cells 
prior to meiosis. How any irregularities in meiosis could affect 
the ontogeny of the fertile part of the leaf, a process taking at 
least several years, is by no means clear to me. The only plant 
J have thus far found in this genus that seemed to be an inter- 
specific hybrid (B. matricariifolium < simplex) had an entire- 
ly normal fertile structure; the spores, however, were extremely 
irregular, mostly dwarfed and distorted but some of them very 
large, suggesting the failure of cytokinesis. The idea that leaf 
axes or parts of a leaf that bear sporangia would be aborted in 
a hybrid in addition to the spores themselves lacks any basis in 
experience, to the best of my knowledge. 

That the average low percentage of production of fertile struc- 
tures in B. oneidense could be a result of abortion associated with 
hybridity finds even less support in a study of the meiotie proc- 
ess itself in this species. As I reported several years ago® cyto- 
logical studies of materials of B. oneidense from three localities 
showed that there was no evidence of hybridity, either in irregu- 
lar pairing or polyploidy. 

In conclusion, then, the relative development of fertile seg- 
ments in plants of Botrychium dissectum and B. oneidense grow- 
ing in the same localities tends to show what appear to this au- 
thor as only minor differences. The percentage of fertile leaves 
evidently varies from locality to locality and from year to year; 
it is imperative, therefore, in making comparisons to take ma- 
terials from the same habitats and at the same times. The area 
around Ithaca and the southern tier of counties of New York, 
where very profound differences have been found in production 
of fertile spikes between the two species, should be examined eare- 
fully. However, even if there are areas where there are marked 
differences in the relative development of fertile structures in 


se pemnenonerets Observations on North American Ferns. Rhodora 57: 


Fern New to UNITED STATES 81 


these two species, I do not believe that this can, without further 
knowledge, be taken as evidence in support of an idea that B. 
oneidense originated as the hybrid B. dissectum & multifidum. 
UNIVERSITY or MicuigAN, ANN ArzBor, MICHIGAN. 


Another Genus of Ferns New to the United States 
C. V. Morton 


Mr. Joseph Monachino, of the New York Botanical Garden, 
has been working recently in curating some of the collections of 
Dr. J. K. Small, with the assistance of Mr. George Cooley. He 
has sent me two sheets of a fern collected in Deering Hammock, 
Florida, in December, 1921, which was left unnamed by Dr. 
Small. It represents. Maxonia apiifolia (Swartz) C. Chr., a 
genus not hitherto known from the United States. It occurs in 
Cuba (rarely) and is not uncommon in Jamaica. Mr. Monachino 
has suggested that this specimen may have been from a cultivated 
plant, and this is certainly possible. Still, this species has never 
been known in cultivation. Very probably it is a natural oceur- 
rence but a recent introduction. It seems that the prevalence of 
hurricanes is having an effect, and quite a number of West In- 
dian species are being blown across to Florida, where some of 
them become acclimatized. The leaf collected by Dr. Small is 
from a large, vigorous specimen, and must have been from a 
well-established plant several years old. Still, it may not have 
been persistent, since no one else has found it in the last 38 years. 
One factor against its survival is that it is an epiphyte, and even 
in southern Florida the climate is not altogether the best one for 
large epiphytes. 

The genus Mazonia, named in honor of the late William R. 
Maxon, cannot be regarded as a very ““strong’’ genus, which is 
to say that it has no characters that are absolutely different from 
allied genera, unless it is the peculiar method of development of 
the indusium, as described by Christensen. It belongs to the gen- 
eral group of Dryopteris and Polystichum, and it has been re- 
ferred to both genera. The indusium which is large, vaulted, and 
conspicuous, is apparently peltate, but it is actually of the reni- 


82 AMERICAN FERN JOURNAL 


form type, being laterally attached in the early stages; in devel- 
opment, some large basal lobes grow out and overlap, thus making 
the point of attachment apparently medial. The type is that of 
Dryopteris rather than Polystichum. 

The rhizome provides perhaps the most distinctive generic 
character. It is elongate and climbing, unlike that of species of 

ryopteris, which are terrestrial. It is thick and cordlike and 
with its dense covering of bright brown scales suggests strongly 
the genus Polybotrya. The general shape of the blades is also 
rather like Polybotrya, and there may be a real relationship. 
However, in Polybotrya the sporangia are ‘‘acrostichoid,’’ i.e. 
distributed all over the fertile segments, without an indusium and 
not grouped into sori. The closest relationship of Mazonia is 
with some of the species grouped by Christensen in Dryopteris 
sect. Polystichopsis. The relationships remain to be worked out. 
The rhizome character and epiphytic habit can be considered 
characteristic for the present. Another, probably less important, 
character, is that in Maxonia the fronds are somewhat dimorphic. 
Sometimes the fertile blades are strongly contracted, with re- 
duced leaf tissue, and then resemble Polybotrya; again, only 
certain pinnae are contracted and fertile. Polystichopsis species 
have fertile and sterile fronds that are essentially uniform. 

The synonymy is as follows: 
Maxonia C. Chr., Smiths. Mise. Coll. 669: 3. 1916. Type: M. apiifolia 

(Swartz) ©. Chr 
a subg. Deiguciudan Maxon, Contr. U. 8. Nat Herb. 13: 39. 
1909. 


Maxonia aptirota (Swartz) ©. Chr., Smiths, Mise. Coll. 669: 3. 1916. 
Dicksonia apiifolia Swartz, Journ. Bot. Schrad. 18002: 91. 1801. 
Dr. 


Polystichum apiifolium C. Chr. Ind. Fil. 578. 

LORIDA: Deering Hammock, Cutler soul 1921, John K. Small, 
George K. Small, John B. DeWinkeler, ‘cell NX); 

Cusa: Batabané, Prov. Habana, in patch of royal palms, Dec. 10, 1920, 
Ekman 11599 (US). Moist woods between Ojo de Agua del Bafio and La 
guna de Piedra, Pinar del Rio, Mareh, 1937, Leén 16817 (US). In palm 
grove, Caibarien, Las Villas, Apr. 10, 1921;‘H, Fernandé:592 (US). In wet 


ISsOETES ECHINOSPORA 83 


palm jungle in swamp at end of road due north of Hotel Isla de Pinos, 

Nueva Gerona, Isla de Pinos, Feb. 18, 1955, Killip 44829 (US); ibid., Feb. 

6-10, 1956, Morton § Killip 9971 (US). Without special locality, De la 

Sagra ee cited by C. Chr. 

AMAICA: Vicinity of Troy, Hollymount, Moneague, Tiddenham, and else- 
where, various collectors. 

MAXONIA APIIFOLIA var. DUALIS (Donn. Smith) C. Chr., op. cit. 4. 
Nephrodium duale Donn. Smith, Bot. Gaz. 15: 20. pl. 4. 1890. 
GUATEMALA: Pansamalé, Dept. Alta Verapaz, von Tuerckheim 1408 

typus). Dense wet limestone forest near Chirriacté, Dept. Alta Verapaz, alt. 

900 m., Apr 9, 1941, Standle = seh 
PANAMA: Barro Golerade't ad; Cs%  piukaley 31401 (US), Kenoyer 22 

(US), Taylor 1330 (US). pig ae Rive er, C. Z., Killip 2610 (US). Trail 

from Campana to Chica, Cerro Campana, Prov. Panama, 600- oa m., Allen 

2658 (US) 

UADOR: Vicinity of Quininde, alt. 50-150 m., Holdridge 1640 (US). 
This is the first report of this species in Ecuador, in fact the first record of 
the genus in South America at all 

SMITHSONIAN INSTITUTION, WASHINGTON, D. C. 


Isoetes echinospora Durieu in North America! 
BERNARD BOIVIN 


The North American variants of Isoétes echinospora Durieu 
are generally distinguished by the presence of stomata on the 
eaves, whereas typical European plants lack stomata. This char- 
acter is scarcely a practical one, but in view of its geographical 
value it may be retained to separate the American and European 
phases of this species. As the presence or absence of stomata does 
not appear to be linked with any other character, specific rank 
for the American phases appears to be hardly justifiable, but our 
plants may be conveniently regarded as a subspecies as follows: 
Isorres ECHINOSPORA Dur. subsp. muricata (Dur.) Boivin, stat. 


nov. 
Isoétes muricata Dur., Bull. Soe. Bot. France 11: 100,101. 1864. 
The type locality of I. muricata is Woburn, Massachusetts. 


1 Contribution No. 33 from the Plant Research Institute, Research Branch, 
Canada Department of Agriculture, Ottawa, Canada. 
2 THIS JOURNAL 35: 84. 1945. 


84 AMERICAN FERN JOURNAL 


Live specimens collected by Boott were sent to Durieu and eul- 
tivated in the botanical garden at Bordeaux. 


ISsoETES ECHINOSPORA Dur. subsp. MURICATA var. MURICATA. 
Leaves rather long, 15-40 em. long, flaccid and often twisted. 


Specimens have been examined from Nova Scotia, New Bruns- 
wick, Maine, New Hampshire, Massachusetts, and New York; 
also reported from New Jersey. The New Hampshire plants 
have been referred to var. robusta, to which they are somewhat 
intermediate, but they appear to be closer to var. muricata. 


IsoETES ECHINOsPoRA Dur. subsp. MURICATA var. ROBUSTA En- 
gelm. Trans. St. Louis Acad. Sci. 4: 380. 1882. 


Somewhat intermediate between var. muricata and_ var. 
Braunii. Plants stouter with leaves numerous, usually 20 or 
more, and 15-25 em. long as in var. muricata, but more rigid and 
mostly strongly arched as in var. Brawnii. Specimens of this 


they are roughly distributed around the old Champlain Sea, they 
may be retained as a geographical variety. Specimens have been 
examined from Ontario (Marmora, Gillett 6765; Deux-Riviéres, 
Dore 10235), Quebec (Wakefield, Calder, Cody & Gillett 1746; 
Templeton, Calder 1222; La Tuque, Marie-Anselme ; Saint-Jean, 
Raymond 1762), and Vertuant (Isle LaMotte, Pringle). The 
type is from Isle La Motte, Lake Champlain, Vermont 


IsorrEs ECHINOSPORA Dur. subsp. MuRIcATA var. Braunii (Dur.) 
Engelm. ex Gray, Manual, ed. 5, 676. 
Isoétes Braunii Dur., Bull. Soe.. Bot. ieee 11: 101,2. 1864. 
‘ hapa echinospora var. Boottii Engelm. ex Gray, Manual, ed. 
Tsoétes maritind Underw., Bot. Gaz. 18: 94. 1888. 
Isoétes echinospora var. eanais Eaton ex Gilbert, List N. A. 
Pter. 10, 27. 1901. 


Plants smaller, with 5 to 20 leaves, these 3-15 em. long, rigid, 
at first straight, soon becoming arched. nats about 0.5 
millimeter across, varying from 400 » to 600 p» 


ISOETES ECHINOSPORA 85 


This plant is disjunct-transcontinental, ranging from Nova 
Scotia and the northeastern United States to southern Mackenzie, 
then along the Pacific Coast from the Aleutian Islands to Van- 
couver Island. The western specimens have been segregated on 
the basis of their blunt spines on the megaspores, but this char- 
acter does not appear to be convincing. Type in Berlin, from 
Lake Winnipiseogee, New Hampshire. 

I have not seen any material of I. muricata var. hesperia Reed,” 
but from the original description this variety should belong here. 
eek ECHINOSPORA Dur. subsp. MuRICATA var. Savilei Boivin, 


ar. 
eal foliis 5-10 (15) in planta, (2) 5-10 (12) em. longis, 
rigidis, strictis vel saepius arcuatis; megasporae 300-4 
e spores are smaller than in any of the other aia istoak: 
ly averaging about 14 mm. in diameter 

GREENLAND: Tjgrn ved Ivigtut, Avert 17, 1937, J. Grontved 
606 (DAO 

QUEBEC: Great Whale River, 1.5 miles northeast of Post, Un- 
gava, 55°17'N, 77°47’W, in rock pool with black organic mud 
bottom, numerous in this pool, not in adjacent ones, September 
8, 1949, D. B. O. Savile 792 (DAO, type) ; same eon — 
553 & 738 (DAO); Fort Chimo, Ungava, 58°07’N, 68°23’W. 
muddy-bottomed stream between 2 lakes about 3 a8 witit of 
base, common locally, rare in area, August 17, 1948, J. A. Calder 
2668 & 2669 (DAO) 

OnTario: Shores of Lake Attawapiskat, Patricia, 52°14’, 87°53’, 
August 28, 1952, Dutilly, Lepage & Duman 30,714 (DAO). 

In all the varieties enumerated above the size of the spor- 
angium varies concurrently with the size of the leaf. For a more 
complete synonymy of the varieties listed above, see C. F. Reed, 
Amer. Fern Journ. 35: 77-86. 1945. Incidentally I may mention 
that among the material at hand is a collection of W. J. Eyerdam 
(No. 3373), collected in 1939 at Prince William Sound, Alaska, 
and distributed as I. Braunii var. maritima and as I. Nuttallii 
A. Braun. This specimen apparently belongs to I. asiatica 
Makino, new to North America. 


86 AMERICAN FERN JOURNAL 


A Hawaiian Thelypteroid Fern with Peltate Indusia 
Kunio [wAtsvuKI 


Natural groups of species are defined by a combination of 
characters. However, some of these diagnostic characters are 
sometimes overestimated, so as to result in unnatural groups, for 
there is no single feature that can cireumscribe the higher taxo- 
nomic groups. 

Aspipium Boyptag,! an interesting but little known Hawaiian 
species described by D. C. Eaton, has long been considered as a 
representative of the genus Cyrtomium because of the presence 
of a peltate indusia, though its affinity has also been thought to 
be with Aspidium (Cyclosorus) cyatheoides. In A. Boydiae, the 
indusia are round and peltate; the fronds are impari-pinnate, 
with narrowly triangular-lanceolate, papyraceous pinnae with 
reticulate venation. On account of these features, this species 
was placed in Cyrtomium. A peltate indusium is a convenient 
key character to define the polystichoid ferns within the dryop- 
teroid group of genera. Still, the similarity of such a character 
may be relatively unimportant, and in this case the most inti- 
mate alliance of A. Boydiae is found in A. cyatheoides, a species 
undoubtedly thelypteroid. Because of its interesting character- 
isties, A. Boydiae has been investigated from the phylogenetic 
point of view, comparisons having been made especially wit 
Cyrtomium, a genus of the polystichoid series, and with the 
thelypteroid series of ferns. 

The rhizome of A. Boydiae is ascending and radially con- 
structed, as in both the series. In A. Boydiae, the stipes are 
terete, stramineous, and subpubescent. The scales are very sparse 
near the base of stipe only, as is usual in almost all of the 
thelypteroid ferns. In Cyrtomium, the stipes are often densely 

1 Aspidium Boydiae D. C. Eaton, Bull. Torrey Bot. Club 6: 361. 1879; 
mre from Oahu, Hawaiian Islands. 
Aspidium cya athe oides var. depauperatum Hilleb. Fl. Haw. Isl. 572. 1888. 

Dryopteris cyatheoides var. tetris e C. Chr, Ind. Fil. 66. 1905. 


Cyrtomium Boydiae W. Robinso Bull. Torrey Bot. Club 40: 204, pl. 10. 
ee Ind. Fil. Suppl. II: 11. 1917; Bull. B. P. Bishop Mus. 25: 10, 


HAwaAlAN THELYPTEROID FERN 87 


covered, with the scales lacerate-fimbriate on the margin. The in- 
ternal structure of the stipe of A. Boydiae indicates more evident- 
ly its thelypteroid nature; two vascular strands are present in 
the base of stipe, which are united upwards to a single strand, 
U-shaped in eross section, a feature typical of the thelypteroid 
and athyrioid ferns. In the polystichoid ferns, on the contrary, 
the stipe contains several separate vascular strands. 


YW» 


pees — 


DING 
4 NSN cies 


A 


Fies. A-C, Aspiplum Boyptar, DRAWN FROM Faurie£ 352. A, A LATERAL 
Pinna, SHOWING THE VENATION AND THE POSITION oF Sorl, X 1.5; B, A 
Sporz, X 300; C, a ScaLe aT BASE or Stipr, X 7.5. 


The form of frond varies in both series. Impari-pinnate blades 
are also found in both of them, as observed in most species of 
Cyrtomium and of Abacopteris. There is, however, a distinct 
difference in the anastomosis of the veins between these two series. 
In the polystichoid species having reticulate venation, the veins 
anastomose to form irregular areoles, which usually contain one 
or more veinlets. On the contrary the venation of the thelypteroid 
ferns is, when anastomosing, typically goniopteroid or meniscioid, 
and a sinus-membrane usually develops. The venation of A. Boy- 
diae is goniopteroid in pattern, but somewhat irregular, a sinus- 


88 AMERICAN FERN JOURNAL 


membrane being also found. On the abaxial surface of the fronds 
of A. Boydiae oceur sparsely some setiform hairs, and no scales 
are found on the laminar surface, both characters of thelypteroid 
ferns. In the polystichoid ferns, scales are usually present on 
laminar parts and no setiform hairs. 

With the exception of the indusia, the sori of the polystichoid 
and thelypteroid ferns exhibit no fundamental differences. The 
indusia of A. Boydiae are, as mentioned above, round-peltate in 
construction. Except for this aberrant feature, A. Boydiae may 
be considered as a member of the thelypteroid group on the basis 
of the brief discussion given above. In the thelypteroid ferns in 
general the indusia are reniform to round-reniform, rarely some- 
what asymmetrical, but peltate indusia or ones intermediate 
between reniform and peltate have been recorded. The peltate 
indusia of A. Boydiae are so constant in their construction that 
there is no clue as to their probable origin. At present, only the 
morphological presumption that the peltate indusia of A. Boy- 
diae have been derived from the basifixed reniform ones may be 
admitted. This is supported by the generally accepted assump- 
tion that the various types of indusia found in the athyrioid 
ferns have been derived from a reniform type. 

The thelypteroid relationship of A. Boydiae may be assumed 
from the facts mentioned above, but its final taxonomic position 
is still in question. According to the current delimitation of the 
thelypteroid genera, this species might be included in Cyclosorus, 
if only the indusia were not peltate. The similarity between A. 
Boydiae and Cyclosorus cyatheoides is apparent, but the differ- 
ence in their indusia seems to be evidence that the position of A. 
Boydiac is fairly distant from Cyclosorus proper. It appears to 
be necessary not to include this species in Cyclosorus, but, as 
Cyclosorus itself is not yet completely studied, no new taxon 
or combination is proposed here. The proper systematic position 
of A. Boydiae may be clarified when Cyclosorus is better known. 


DEPARTMENT OF Botany, University or Kyoto, JAPAN. 


AZOLLA CAROLINIANA 89 


2 aa hal agin on Microsporocarpic Material 
Azolla caroliniana 


R. K. Goprrey, Grapy W. RerNert, and RicHarp D. Hovx! 


Svenson (1944), in reference to Azolla caroliniana Willd., 
wrote that diligent search by him revealed but two herbarium 
collections in which microsporocarps were present and implied 

he saw no megasporocarps. For several years the senior 
author has been collecting and examining plants of this species 
in the hope of obtaining ‘‘fruiting’’ material. Only recently has 
this met with limited success. On October 8, 1960, about 6 miles 
south of Perry, Taylor County, Florida, a small collection was 
made, subsequent examination of which revealed the presence 
of microsporocarps. Since no megasporocarps could be found 
amongst the meagre amount of material collected, we revisited 
the locality and procured a larger quantity. In this, although 
microsporocarps were bountifully present, stil} no megasporo- 
carps were seen. 

Neither Svenson (ibid.) nor Mason (1957) appear to be overly 
confident about the characters by which the species of Azolla are 
to be distinguished, particularly vegetatively. Both authors use 
the septation, or lack of it, of the glochidia of the microsporangic 
massulae as primary characters. Svenson indicates that the char- 
acter of the surfaces of the megaspores is distinctive for three of 
‘the four species with which he was concerned, the megaspores of 
A. caroliniana being unknown. 

Azolla caroliniana is the only species of the genus reputedly 
occurring in Florida. Plants of the collection here discussed 
appear to be not unlike those that the senior author has observed 
during his years of botanical work in the southeastern states and 
it is assumed that they represent this species. 

Microscopic examination of the microsporangic massulae of our 


1 This investigation was supported (in part) by a PHS research grant, 
RG-6305, to the senior author from the Division of Genesl ort il Sciences, 
Publie Health Service. Illustrations were prepared by Mr. 


AMERICAN FERN JOURNAL VouuME 51, Pate 4 


AZOLLA CAROLINIANA 91 


recent collection shows that the glochidia of a given massula are 
both septate and nonseptate without exception. A single instance 
was observed in which a glochidium was branched, one branch 
being septate, the other nonseptate. According to Svenson the 
glochidia of Azolla caroliniana are not septate; those of A. filicu- 
loides are not septate or rarely have 1 or 2 septae at their apices; 
A. mexicana and A. microphylla, on the other hand, have many- 
septate glochidia. We are not suggesting that A. caroliniana and 

liculoides are not, therefore, distinguishable for we have in- 
sufficient material for comparison. We simply question the use 
of the nature of the glochidia as distinguishing characters for 
these two plants. And we encourage collectors to keep a weather 
eye out for sporocarpie material of Azolla, particularly A. caro- 
liniana, so that in time there may be more ample and more suit- 
able material for comparative study. 

It is difficult in our material to ascertain the number of spor- 
angia in each sporocarp because some sporangia apparently do 
not develop at all and gradually disintegrate. Of those which do 
mature, not all mature at the same rate. Svenson gives the num- 
ber per microsporocarp of Azolla caroliniana as 8 to 40. For the 


record, counting all microsporangia in a given sporocarp which 
could be distinguished regardless of stage of development, these 
are the counts fo ty-four pate aman 23, : 


50, 60, 66, 44, 29, 64, 60, 23, 24, 45, 24, 19, 45, 24, 50, 60, 44, 29, 
64, 60;-19, 32,30; 89,16, 49, 52,22, 19, 46. 

Svenson, in his key to the species, gives the number of massu- 
lae per microsporangium as 4 to 6 in Azolla filiculoides, and four 
in A. mexicana. In discussion of A. microphylla he reports an 
earlier investigator’s count as six for this species. He gives no 
count for A. caroliniana. The number of massulae in virtually 
all of the numerous microsporangia examined in our material 


Fig. 1. AZOLLA CAROLINIANA: A, Hapit, Upper SuRFACE OF STERILE 
PLANT; B, LowER SuRFACE, PLANT WITH MICROSPOROCARPS ; Ms MIcROSPORO- 
CARP; D, MICROSPORANGIUM WITH MassuLA BEING DISCHARGED; E, GLO- 
CHIDA TYPES FROM A SINGLE MASSULA; F, THE ONLY BRANCHED GLOCHIDIUM 
OBSERVED; G, TIP OF A GLOCHIDIUM AS SEEN UNDER OIL IMMERSION; H, THE 
Two-Losep LEAF 


92 AMERICAN FERN JOURNAL 


was three; only one of those examined contained four. 

SAcimens of our microsporocarpous material have been dis- 
tributed to some herbaria as Godfrey 60394 and Godfrey & Houk 
60474. Some material has been preserved in F A A and is avail- 
able should anyone wish to see some. 


REFERENCES CITED 


Svenson, H. K., 1944. The New World Species of Azolla. Amer. Fern Journ. 
34: 69-84. 


Mason, Hersert L., 1957. A Flora of the Marshes of California. Berkeley 
and Los Angeles. University of California Press 

DEPARTMENT OF BroLoaicaL SCIENCES, 

Fioripa State University, TALLAHASSEE, FLORIDA. 


Adaxial Sori in Polypodium hesperium 
Eminy L. Hartman 

While collecting ferns and fern allies in the Gunnison Na- 
tional Forest, in central Colorado, in June, 1959, two colonies of 
a Polypodium were discovered which were unique in that the 
fronds had distinet sori on both the adaxial and abaxial surfaces 
(Fig. 1). The plants readily keyed to Polypodium hesperium 
Maxon, the taxonomic status of which is somewhat uncertain as 
evidenced by the fact that Maxon in Abrams! and Harrington” 
regard it as a distinct species, whereas Broun® listed it as a 
variety of Polypodium vulgare L. and Fernald‘ considered it a 
synonym of Polypodium vulgare L. var. columbianum Gilbert. 
Without attempting to decide the taxonomic status discrepancy, 
the collections here reported will be referred to as P. hesperium. 
. The two colonies were growing in the erevices of rocks on 
mountain slopes covered with a pifion-juniper association at an 
altitude a little over 7680 feet. Although the habitats were quite 
similar, they were separated geographically by over 30 miles. 


1 - itaelpey Flora of the Pacifie States ie oy, 1926. 
2 Manual of the Plants of Colorado 15. 
31 Thdek t sor th American Ferns 144 1908. 
4 Polypodien virginianum and P, vulgare, Rhodora 24: 125-142. 1922. 


AMERICAN FERN JOURNAL VOLUME 51, Puate 5 
I ; ! 


t 


PoLYPoDIUM HESPERIUM WITH ADAXIAL Sori 
(CoLoRADO, HARTMAN 1653) 


94 AMERICAN FERN JOURNAL 


The presence of adaxial and abaxial sori was widespread 
throughout both colonies, occurring on more than 50% of all the 
plants present. In general, a single plant would have this tend- 
ency developed to some extent on all of its fronds. The plants 
of one colony were about twice the size of those in the second 
colony, but within each of the colonies the individual plant size 
was fairly uniform. The larger fronds averaged 15.0 em. in total 
length and there was some occurrence of arrested pinna-devel- 
opment, in which case the pinnae were either reduced to slight 
protuberances, mere laminal flanges of the rachis, or were totally 
absent. ere was no apparent distributional pattern of these 
abnormalities. Mohlenbrock,® in reporting an unusual form of 
Asplenium pinnatifidum Nutt., mentions abaxial sori being vis- 
ible from the adaxial side. This occurred on three specimens of 

opulation, and in each case there was extreme suppression of 
laminal development. 

he abaxial sori were large and located about one-half way 
between the margin and the midrib or submarginal. The adaxial 


superficial examination of the pinnae would lead one to this 
conclusion, microscopic study of microtomed sections showed 
that there were two distinet sites of sori initiation, one definitely 
abaxial and one definitely adaxial. Thus what appears to be @ 
confluency of sori is merely due to the bushy development of 
sporangia within a sorus. It should also be pointed out that 
there was not an adaxial counterpart of every abaxial sorus. 
The ratio between viable and non-viable spores was essentially 
the same in both sori. Approximately 15-20% of the adaxial 
and abaxial spores were non-viable. Viable spore size was also 
essentially the same, ranging from 60-65 » in length X 38 p in 
width 

A specimen is on deposit in the herbarium of the U. S. Na- 
tional Museum. Specific collection data is as follows: Crevices 


BS as Unusual Form of Asplenium pinnatifidum, THIs JOURNAL, 46: 91. 


Heat RESISTANCE OF MARSILEA 95 


of large rocks on pifion-juniper covered mountain slopes, Spring 
Creek Camp Ground, Gunnison National Forest, 26 miles north- 
east of Gunnison, Gunnison County, Colorado, June 28, 1959, 
elev. over 7,680 feet, Hartman 1653 

DEPARTMENT OF Brotoay, Kansas Stare TEACHERS COLLEGE, 
Emporis, Kansas. 


Heat Resistance of Sporocarps of Marsilea quadrifolia 
Wiu1am W. Broom 


In an earlier paper! the author reported on the comparable 
viability of sporocarps of Marsilea quadrifolia L. in relation to 
age. The same paper summarized some of the reports in the 
literature on the unusual resistance of sporocarps of the Marsi- 
leaceae to aging, drying, alcohol, and herbarium poisons. The 
present paper deals with the unusual resistance to moist heat 
of sporocarps of M. quadrifolia. 

On November 21, 1953, sporocarps of M. quadrifolia were col- 
lected at the botany pond of Eastern Illinois State College. A 
Similar collection was made on November 3, 1953, by Dr. W. N. 
Stewart at the Kikapoo State Park in Illinois and forwarded to 
Dr. Paul D. Voth, of the University of Chicago, under whom the 
author was working at the time. 

When attempts were made to germinate sporocarps from each 
lot soon after collection, the results were poor. Few of the 
Sporocarps opened typically, and relatively few spores were re- 
leased. Few sporophytes were produced. However, when sporo- 
carps were placed in a 65°C oven in a study of their moisture 
content, excellent germination results were obtained after two 
days of drying. Both lots of sporocarps were stored in a steam- 
heated room in the Barnes Laboratory of Botany at the Univer- 
sity of Chicago during the fall and winter. These sporocarps 
showed normal germination behavior by early spring. 


om, William W. hig coke My Viability of Sporocarps of Marsilea 
Bestel. in Relation to Age: Ill. Acad. Sei Trans. 47: 72-76. 1955. 


96 AMERICAN FERN JOURNAL 


Early in the spring of 1954, when the normal sporocarps were 
giving good germination results, it was decided to check their 
resistance to boiling. Sporocarps boiled for two hours showed 
normal germination. Numerous gametophytes developed and 
later sporophytes were produced. 

The writer thought it would be desirable to repeat the experi- 
ment in the fall of 1957 and extend it by determining whether 
the spores within the sporocarps could withstand autoclaving at 
15 lbs. pressure for 15 minutes as well as boiling. The results of 
this repetition were somewhat surprising. At this time the spores 
within the sporocarps were unable to resist even the five minutes 
of boiling. 

Since the sporocarps might have taken up considerable mois- 
ture during the humid summer of 1957, selected sporocarps were 
subjected to drying at 65°C to a constant weight. The weight of 
100 sporocarps decreased from 1.895 grams to 1.745 grams for a 
loss of 0.15 grams or 7.9%. The weight loss of a control lot was 
negligible during the same period. 

Sporocarps from the dried lot and the control lot were boiled 
for 15, 30, 90 and 120 minutes. Similar numbers of sporocarps 
were autoclaved at 15 pounds pressure for 15 minutes. Follow- 
ing this heat treatment, sporocarps were scarified with a file and 
placed in individual vials containing about 20 ml of tap water 
each. The dried and boiled sporocarps that were germinated 
contained viable spores which developed into good gametophytes 
and later sporophytes developed, similar to those of a set of con- 
trols. The spores from undried sporocarps that were boiled failed 
to develop into gametophytes. The undried sporocarps that were 
autoclaved also lacked viable spores. Some of the sporocarps 
that were dried and autoclaved contained no viable spores, some 
had viable megaspores, and some had both viable megaspores 
and microspores which resulted in the production of sporophytes. 
In a series of 20 dried and autoclaved sporocarps, 1 did not open, 
9 contained no viable spores, 7 had viable megaspores only, and 
3 had viable megaspores and microspores which resulted in 
sporophyte development. 


PELTAPTERIS IN Costa RIcA 97 


It is interesting to note that the microspores were much more 
susceptible to the autoclaving, as they were to aging, as was 
pointed out in the earlier paper cited previously. This greater 
resistance of the megaspores to autoclaving is a handy tool for 
hybridizing attempts with the Marsileaceae. Since megaspores 
are readily visible, it is possible to mix microspores from one 
source with megaspores of another, making it possible to work 
with thousands of megaspores with a minimum of labor as com- 
pared to that required for the manual separation of megaspores 
and microspores. 


VALPARAISO UNIVERSITY, VALPARAISO, INDIANA. 


Peltapteris in Costa Rica 
CuyprE F.. REED 


Peltapteris Link is a small genus of wide-creeping, epiphytic 
ferns, with slender-branching rhizomes and dimorphic fronds. 
The more or less dissected, flabellate sterile fronds tend to sep- 
arate this genus from Elaphoglossum, to which it is nearly re- 
lated. The fertile fronds are smaller and roundish, cordate, 
crenate-dentate or lobed, with the sporangia covering the fertile 
surfaces. 

Morton! made transfers of the species and varieties to Peltap- 
teris Link from Rhipidopteris Schott. There are two species, each 
with one or more varieties or forms, found in Costa Rica. 

In addition to the many specimens of the following species and 
forms in the United States National Museum, collections in the 
Reed Herbarium, purchased through grants to H. EB. Stork and 
C. K. Horich, give an idea of the distribution of the genus Pel- 
tapteris in Costa Rica. The general distribution for each will 
also be given based on the specimens in the United States Na- 
tional Museum, I wish to thank C. V. Morton for helping in the 
identification of the specimens in the Reed Herbarium. 

ELTAPTERIS FLABELLATA (H. & B.) Morton. Atlantic rainforest 


1THIs JourRNAL 45: 11-14. 1955. 


98 AMERICAN FERN JOURNAL 


of Tapanti, dense jungles at base of the northern Cordillera de 
Talamanca, along the upper headwater area of Rio Reventazon 
and Rio Macho, south of Orosi, alt. 1100-1200 m., Horich 193 
(Reed Herb. 33487-88) ; Santa Clara de Cartago, alt. 1950 m., 
Maxon & Harvey 8238 and Lankester 662 (US) ; vic. of El Gen- 
eral, Prov. San José, alt. 1680 m., Skutch 3043 (US). Also Pan- 
ama, Colombia, Peru, and Ecuador. 

PELTAPTERIS FLABELLATA forma STANDLEYI (Maxon) Morton. 
Creeping in moss on tree in wet forest, vic. of Tilaran, Prov. 
Guanacaste, alt. 500-600 m., Standley & J. Valerio 44454 (US) ; 
also collections in US from Los Ayotes, La Palma, Pejivalle, Que- 
brada Serena, La Tejona, and El Silencio. This form is usually 
at lower elevations than the typical one. Also Panama, Vene- 
zuela, and Ecuador. 

PELTAPTFRIS PELTATA (Swartz) Morton forma PeLTATA. Peralta, 
Stork 493 (Reed Herb. 19936). West Indies, Mexico to Panama. 

PELTAPTERIS PELTATA forma FOENICULACEA (Hook & Grev.) 
Morton. Highest Carpintera, alt. 6000 ft., Stork 1372 (Reed Herb. 
19937) ; Carpintera, cloud forests, sichakni near San Ramén de 
Tres-Rios, alt. 1700-1880 m., Horich 20 (Reed Herb. 32809). 
This is the most divided form, with the sterile fronds up to 6- 
times divided and the ultimate segments filiform, making them 
about one-half as wide as those in the typical form. Also south 
to Venezuela and Eeuador. 

PELTAPTERIS PELTATA forma potentillifolia (Christ) Reed, 
comb. nov. Based on Acrostichum (Rhipidopteris) peltatum 
Swartz var. potentillaefolium Christ, Bull. Soc. Belg. 35: 242. 
1896. La Palma. Costa Rica, alt. 1550 m., Dee. 18, 1888, Pittier 
704 (Isotype, US 833,938). Cloud forests of Montafia del Cedral, 
south of San Antonio de Escazti, crest section and ridges at 2000- 
2400 m. alt., Nov. 1959-Jan. 1960, C. K. Horich 52 (Reed Herb. 
33039-40). The lamina is several times dichotomous, with 6 to 8 
segments which are decussate-dentate at their apices and from 
0.3-0.5 em. wide, making the segments about twice as wide as 
those of the typical form. Only known from Costa Rica. 


Reep Hersarium, 10105 Harrorp Roap, Bautrore 34, Mp. 


Cyrotoey or IsorrEs 99 
Cytology of Isoetes coromandelina 
S. C. VERMA 


Isoétes coromandelina L. is a native of India (ef. Reed, 1953) 
and is most commonly met with in South Indian localities along 
with three other species, namely: J. dixitei Shend., I. sahyadru 
Mahabale and I. sampathkumaranii Rao. Apart from South 
India, I. coromandelina has been reported to occur in Serampur 
(Bengal, cf. Ekambaram and Venkatanathan, 1933), Bombay 
(McCann, 1934), Banaras (Bhardwaja, 1935) and Baroda 
(Gaekwad and Deshmukh, 1956) from where a new variety has 
been described. Recently Bhambie (1957) has reported its wild 
occurrence at Meerut (N. India), the exact location being about 
four to five miles north of Meerut, on the Meerut-Mowana road, 
where it is fairly common on field margins during the monsoons 
and dries up in winter. The present cytological observations 
concern only the Meerut gatherings. The morphology of the 
genus as a whole is being dealt with by Dr. S. N. Bhambie ; the ey- 
tology was investigated by the writer in August and September 
1956, 1957, and 1958 with the kind permission of Prof. V. Puri. 
Almost all the fixations of the wild as well as the material kept 
in the greenhouse were made in 1:3 acetic-alcohol by Dr. S. N. 
Bhambie, to whom the writer is very grateful. 

Extensive examination of the material has revealed the pres- 
ence of only megasporangiate plants at Meerut. Curiously 
enough, Bhambie (1957) too, who has studied many more indi- 
viduals, has only recorded the megasporangiate material thus 
far. As Prof. Puri tells me, microsporangia have so far not been 
observed in any of the individuals studied at Meerut. Therefore, 
the present account is limited to only the megaspore-mother-cells. 
The number of megaspore-mother-cells per sporangium is Lone 
ally very small, which evidently renders this difficult cytological 
material. The usual acetocarmine squashes (cf. Manton, 1950) 
have been obtained and made permanent by McClintock’s (1929) 
technique. 


AMERICAN FERN JOURNAL VoutuME 51, PuaTE 6 


CyvToLoey or IsoETES 101 


The sequence of meiosis in the megaspore-mother-cells is in- 
teresting. Early prophase stages have not yet been critically 
followed, but the evidence seems to be convincing that mostly 
the chromosomes remain unpaired during prophase. In a very 
few mother-cells, the univalent chromosomes are associated in 
groups of three or more. There is no regular division in sue 
cells and depending upon the groups micronuclei are organized, 
or in some cases there is one mitotic division of the chromosomes 
followed by an irregular anaphase separation, which results in 
abortive spores. Rarely the entire contents, first distributed in 
groups, get included into a single large nucleus and result in a 
““monad.’’ However, in 95% of the cells examined there is com- 
plete asynapsis and at the first meiotic phase 33 univalents and a 
small fragment (marked ‘f’) are clearly discernible (Fig. 1). 
Three to five chromosomes are usually observed to be associated 
with the nucleolus where some of them may show end to end 
associations, observed only in a few preparations). The small 
fragment has invariably been noticed and is therefore believed 
to be centric. It is further supported by a similar count in two 
Squashes of root tips which were pre-treated with 8-hydroxyqui- 
noline (cf. Tjio and Levan, 1950) There is an equational division 
of the chromosomes with a regular anaphase separation and sub- 
sequent organization of two nuclei with the unreduced chromo- 
some number (Figs. 2 and 3). There is no indication of a second 
meiotic division. Tetrad nuclei are therefore absent and the 
dyads are the end products of ‘‘meiosis.’’ This type of meiotic 
behavior is in general agreement with hybrid species where 
almost all the chromosomes appear at metaphase I as univalents 
(ef. Darlington, 1958). Complete details of the process will ap- 
pear elsewhere. 


E Total asynapsis in a megaspore-mother-cell of Isoétes pa 
lina uke showing 33 + 1 frag. chromosomes (all univalents) at the first 
phase, 


loti 430; Fia. m ore-m ell = hase I, 
43 aspore-mother-cell showing dyad nuclei, X 430; Fig. 
4 Tetrahedrally partitioned megaspore her-cell showi two nucleate 
i two seneput ha pores, the nucleate ones with csbgprticel nucleoli; X 430; 


5. Tetra hearaliy partitioned spore-mother-cell showing two nucleate 
ane two enucleate spores, X 430. 


102 AMERICAN FERN JOURNAL 


The dyads with the unreduced nuclei now undergo cytokinesis 
in a tetrahedral manner, like the ferns with tetrahedral spores, 
resulting in two nucleate and two enucleate spores (Figs. 4 and 
5). The latter shrivel and ultimately abort. Mature megaspores 
are invariably tetrahedral, which supports the present observa- 
tions and also confirms that this type of wall formation is the 
rule in the Meerut plants. Evidently the suppression of one 
meiotic division has not affected the cytokinetic behavior, 
which occurs here as it would take place in a normal tetrad. It 
may be pointed out that in other ferns such as Ophioglossum 
vulgatum L. (forma) (Verma, 1956) and Trichomanes insigne 
(v.d.B.) Bedd. var. B (Mehra and Singh, 1957) where dyad 
formation has been reported, there is only one wall laid down to 
separate the dyads. Sometimes, however, here in [soétes coro- 
mandelina (in Meerut) incomplete walls may be laid and the 
whole mass containing two nuclei remains a single structure with 
the impression of walls on it. This part of the study and the 
probable explanation of the events has appeared separately 
(Verma, 1960). It may be pointed out here that to the writer’s 
knowledge there have been no previous reports of such a type of 
cytokinesis: The laying down of more than one wall when dyads 
are developed either by the suppression or failure of one meiotic 
division. This appears to be the first report in plants. The con- 
clusion well supports the general view expressed by Swanson 
(1958), that ‘‘the two processes—karyokinesis and cytokinesis 
are distinct and have arisen independently in evolution.’’ 

Only the nucleate spores seem to be capable of germination. 
Furthermore, in the absence of any microsporangiate plants or 
an accessory mode of reproduction, it is inferred that the unre- 
duced megaspores develop parthenogenetically (apogamously )- 
Spore germination studies are in progress. 

Abraham and Ninan (1958) have recently reported on the 
cytology of two of the South Indian species, [soétes sampath- 
kumaranii and I. coromandelina. The latter was studied from 
Kovalam, Veli, Quilon, and Crangannore, in Kerala state and 
from Waltair, in Andhra Pradesh. All these gatherings were 


CytoLogey or IsorTEs 103 


shown to be diploid asynaptie with 22 + 1 frag. chromosomes 
at meiosis in the megaspore-mother-cell and also in root-tip 
squashes. One of the plants collected from Kovalam was, how- 
ever, a triploid, 33 + 1 frag. chromosomes having been observed 
in its root-tips. The Meerut populations of I. coromandelina are, 
in contrast, all triploids, and are thus of perhaps some cyto- 
geographic interest. Like the South Indian material, the Meerut 
plants also possess an additional (centric) fragment. Abraham 


South Indian material. Furthermore, since Ekambaram and 
Venkatanathan (1933) observed bivalents and regular meiosis in 
oth mega- and microsporangia in Coromandel coast as well as 

Madras material, the search for a sexual population is likely to 
be rewarded in South India which may perhaps give a clue to 
the origin of the triploid taxon. Such a probability is further 
indicated by Abraham and Ninan’s statement that in the diploid 
race “‘asynapsis is probably the result of accumulation of struc- 
tural hybridity rather than the consequence of hybridization,”’ 
which means its origin from a previously sexual species. 

Abraham and Ninan’s results corroborate the earlier findings 
of the lowest monoploid number in Isoétes echinospora Dur. 
(n = 11, Ekstrand, 1920) and I. asiatica Makino (2n = 22, 
Takamine, 1921). The chromosome number in the Meerut popu- 
lation of I. coromandelina lends additional support to the mono- 
ploid number of 11. Polyploid series based on 11 have earlier 
been reported in I. japonica A.Br. (2n = 43-45, Takamine, l.c.; 
n = 33, Yuasa, 1935) and I. lacustris L. (n = 54-56, Manton, 
1950). It is believed that the rest of the numbers in Isoétes re- 
ported by various authors (ef. Delay, 1953) and also in I. coro- 
mandelina in all probability may have been derived from the 
number 11 

The writer wishes to express his sincere thanks to Prof. P. N. 
Mehra for suggestions and encouragement, to C. V. Morton for 
some literature, to Prof. V. Puri (Meerut) for allowing him to 
work out the cytology and for laboratory facilities, and to Dr. 
S. N. Bhambie for the fixation of the material. 


104 AMERICAN FERN JOURNAL 


LITERATURE CITED 
ApraHam, A. and ©. A. NinaN, 1958. Cytology of Isoétes. Curr. Sci. 27: 
60, 6 


Buampsigr, S. N., 1957. Studies in igh ee I, Mee shoot apex of 


IsoZtes coromandelina L. Journ. Ind. Bot : oGt 
Buarpwasa, Y., 1935. The oecurrence of a in aie Cart. Sci. 3: 
300. 


DaRineton, C. D., 1958. Evolution of Genetic Systems. London 

Dezay, C., 1953. Nombres chromosomiques chez les ceypiiiniaae (1938- 
1953). Rey. de Cyt. et Biol. Veg. 14: 59-107. 

EKAMBARAM, T. and T. N. VE Sa in, Rene Studies on Isoétes 
coromandelina LL. Journ. Ind. Bot. 12: 191-225. 

“EKSTRAND, H., 1920. Ueber die sche rein’ von Isoétes echinospora. 
Svensk. Bot. Tidsk. 14: 312-318. 

Garkwap, L. K. and Y. §. DesHmuKH, 1956, Oceurrence of Isoétes at 
Baroda in poe from Bombay state. Sci. and Cult. 22: 346 

McCann, C., 4. Oceurrence of Isoétes coromandelina L. Seare. Bomb. 

i 01, 


2. 


19 

nent. Stain Tech. 

Manton, I., 1950. Proteins of Cytology and Evolution in the Pteridophyta. 
Camb. se v. Pre 


MenrRa, P. N. and € heat te 1957. Cytology of Hymenophyllaceae. Journ. 
Geneties ae: 379-3 

Reep, C. F., 1953. ‘odes Isoétales. Bolet. Soe. . 27: 5-72. 

SWANSON, C. P., 1958. Cytology and Cytoge ebm ondon., 

TAKAMINE, N., 1921. Some observations in the life as a Isoétes. Bot 


4-90, 
. H. and 4 ‘AN, 1950. The use of oxyquinoline in chromosome 
analysis. Ann. Estae. Exper. de Aula 2: 21-64. 
ERMA, S. C., 1956. Cytology of Ophioglossuwm. Ourr. Sci. 25: 398, 399. 
—————. 1960. Enucleate spores in Isoétes peice Caryologia 
13: 274-284. 
Yuasa, A., 1935. Studies in the cytology of Pteridophyta vii. Reduction 
division in Isoétes japonica A. Br. Bot. Mag. Tokyo 49: 31 


Borany DEPARTMENT, PANJAB UNIVERSITY, CHANDIGARH, INDIA. 


REcENT FERN LITERATURE 105 


Recent Fern Literature 


Tue FEerRNs AND OTHER PrERDOPHYTES OF Iowa, by Tom S. 
Cooperrider! is the latest addition to state fern floras, and a 


a specimen in the National Herbarium—Thelypteris novebora- 
censis, which was collected in a ravine in Tom Range,? Iowa, 
July 22, 1921, by B. Shimek (distributed by the University of 
Texas with the number 152). The other species mentioned 
should be searched for; the Adiantum would hardly have been 
misidentified, and probably not the Dryopteris, but the Equise- 
tum and Woodsia could have been errors of determination. This 
Thelypteris brings the number of Iowa pteridophytes to 55; and 
Equisetum scirpoides, first collected in Towa in 1959, in Winne- 
shiek and Allamakee Counties by Thomas mpi according to 
a letter from Cooperrider, makes the total 56—C. V. Morton. 
THE GOLDBACK FERNS OF CALIFORNIA.—To those feiniline with 
the California flora it is perhaps a wonder that the conspicuous 
and variable goldback ferns, Pityrogramma triangularis sensu 
lato, have not long ago been investigated by biosystematists. 
Karen §S. Alt and Verne Grant of Rancho Santa Ana Botanic 
Garden at Claremont have now given us the first investigation 
of this complex using modern cytogenetic techniques.’ Although 
the study is not yet complete, the results are of substantial in- 
terest both to pteridologists and general vascular-plant taxono- 
mists. 


1State University of Iowa Studies in Natural History, vol. 20, 1, 
pp. 1-62. Undated, but Pg gem July 22, 1959, according to a settee: ps 
be oe September 

*Thus far, I have eles gene to identify this locality, which is unknown 
to Dr. idee define er also. 

’Cytotaxonomie observations on the goldback fern. Brittonia 12 (no. 
3): 153-170, figs. 1-9, tab. 1-3. 1960. 


106 AMERICAN FERN JOURNAL 


Pityrogramma triangularis was treated by Cc. A. Weatherby 
in 1920 as comprising, in addition to the widespread typical 
variety that ranges from Baja California to the southeastern 
edge of Canada, several other varieties, namely pallida, viscosa, 
and Mazonii, each with much more local and southern distribu- 
tion patterns. The results of Alt and Grant show that ‘‘typical’’ 
P. triangularis itself comprises at least three population sys- 
tems—two diploids (a large, coarse one with long segments, 
‘“A.’’ and a small, more herbaceous one with short segments, 
“B’’) and a tetraploid with n — 60 chromosomes. They con- 
sider ‘A’? and ‘‘B’’ to be best regarded as poorly separated 
‘‘semispecies’’ because of the many intermediates that exist. 
But they interpret the tetraploid populations as representing 
possibly a distinct, sibling species. 

The ‘‘varieties’’ pallida (of the Sierra foothills of Central 
California) and viscosa (of maritime southern region) are 
sympatric with P. triangularis. They retain their distinctness 
where they overlap, in spite of evidences of some hybridization 
in certain localities, and the authors therefore propose that 
pallida and viscosa be treated as species coordinate with triangu- 
laris. The primarily desert-inhabiting var. Mazonii is still so 
little known that the authors have made no suggestions in re- 
gard to its status. 

These cytotaxonomic studies have revealed, according to Alt 
and Grant, ‘‘an unsuspected heterogeneity within this small 
assemblage,’’ and they interpret the traditional taxonomic fern 
species Pityrogramma triangularis as a complex of biological 
species, not unlike a number of taxonomic species of Californian 
flowering plants such as Artemisia tridentata, Elymus glauca, 
and others which are similarly constituted —W. H. WaGNeER, JR. 


AMERICAN FERN SOCIETY 107 


American Fern Society 
Report of the Spore Exchange, 1960 


The activities of the spore exchange, consisting of receiving 
and filing contributions of spores and of filling requests for them, 
seem to follow a seasonal pattern, the peak load coming in the 
fall when spores in most of the country have recently ripened 
and folks are returning from summer vacations. The next busiest 
time is in early winter—no doubt when things are being cleared 
away and put in order after the Christmas rush is over. The 
period of least activity is in the spring, although people in warm 
places and those who have greenhouses may request or send in 
spores at any time. Letters and replies go on more or less all the 
year. In 1960 there were many more of these than in the previous 
year, considerably over a hundred. 

As to spores, 146 packets were received from 27 kind members 
and of these about 40 were species new to our list, already num- 
bering 239 kinds available to members. During the year, 1,173 
packets were sent to 48 members requesting them. 

At the present time, our supply of mimeographed lists of avail- 
able spores is exhausted. A new revised and up-to-date list is 
planned for the fall of 1961; it will be sent to any who in the 
past two years have shown interest in the exchange either by in- 
quiries or by contributions or requests for spores. It will be 
sent also to any member of the Society requesting one. 

In order to make this new list as inclusive as the last, an urgent 
request is made to all members to send in spores or fertile fronds 
by September 10 or sooner. Even if the ferns you have or the 
ferns you see on your trips seem to you too common, remember 
they are not common in another part of the country or the world. 
We have a good many requests, now, from other countries. Also, 
even if you have sent the same ones before, remember the sup- 
plies must be kept fresh each year and many species with spores 
that are now two years old will have to be dropped from the 
present list. 

Preparing a new spore list is a task requiring a considerable 


108 AMERICAN FERN JOURNAL 


amount of time, and so if it is to reach you in the fall, spores 
must be on hand in earliest September. Your cooperation in this 
will be sincerely appreciated. 

Before closing this report a more or less personal note should 
be added. Because of other full time commitments from October 
15 through Christmas, it will be impossible for letters to be an- 
swered or spore requests filled during that time. It was a source 
of real concern when this proved true 1960, and many letters 
of apolog : had to be written in Janua 

So, please, your spore cceib titi hotars September 10 and 
your spore ievats before October 15. Thank you. 

Respectfully submitted, 
Katuryn E. BoypDsTon, 
Fernwood, Route 3, Niles, Michigan 


AMERICAN FERN SOCIETY 109 


CONSTITUTION OF THE AMERICAN FERN 
SOCIETY, INC.* 


ARTICLE I. NAME 


Section 1. The name of this society shall be THE AMERICAN FERN Society, 
INCORPORATED. 


ARTICLE II. OsBJEcts 


Section 1. The objects of the Society shall be to affiliate those who are in- 
terested in the study of ferns and allied plants, to foster such an in- 
terest, to encourage correspondence and exchange of specimens between 
members, and the publication of matter pertaining to this group of 

ants. 


ARTICLE III. MEMBERSHIP 

Section 1. Any person interested in the objects of the Society shall be 
eligible to membership. 

Section 2. Application for ena beed accompanied by the required fee of 
two dollars and fifty cents may be made at any time to the Secretary, 
and when so receiv vel approved by two members of the Council, and 
acknowledged, the eer shall be considered a member for the 
current year 

Section 3. The admission fee shall be two dollars and fifty cents payable 
when application for membership is made. This fee shall also consti- 
tute the dues for the current year. 

Section 4. The annual dues shall be two dollars and fifty cents, payable on 
January first of each year. Sustaining membership is credited to any 
person upon the annual payment of five dollars. 


Section 5. Any eligible person may bee mber on payment, at 
any one time, of a fee of fifty ny oF 258 thereafter be subject 
to no dues nor assessments. All such fees shall be held and invested 


vote of the Council. Contributions for the purpose and other available 
moneys may be added to this fund at the discretion of the Council. 
Section 6. Honorary members may be chosen when unanimously nominated 
y the Council, and their names submitted to the members at the next 
succeeding annual election. Three-fourths of the votes cast on the 
question shall be required for election, and the total number cast must 


1 As amended by the membership at the annual elections of 1935, 1936, 
1940, 1947, 1949, 1954, and 1958. 


110 AMERICAN FERN JOURNAL 


be at least twenty. Honorary members shall be entitled to all the 
privileges of the Society without payment of dues. The number of such 
members shall not exceed five at any one time. 

aes a Every member in good standing is entitled to all the privileges 

Pate including its publications. 
Section 8. Members one year in arrears for dues who have been twice noti- 
fied of their in eee shall be considered not in good standing and 

shall forfeit all privileges of the Society including its publications. Any 
such member may be reinstated at any time durng the succeeding year 
by the payment of arrears to the Treasurer. If at the expiration of this 
second cs and without justifying cause his dues shall remain unpa aid, 
he shall cease to be a member of the Society, provided, however, that 
the Coghett” pail ae the power to remit any dues for reasons which it 
considers sufficie 


ARTICLE TV. OFFICERS 
Section 1. The officers of the Society shall be a President, Vice-President, 


Secretary, and Treasurer. Their term of office shall begin January first 
they shall serve for one year, or until their successors are duly 


Section 2. The President shall be in immediate charge of the general in- 
terests of the Society; he shall appoint all committees not otherwise pro- 


the treasurer for that year and who shall report to him as soon after 
the close of the year as possible. 
Section 3. The Vice-President shall act in the absence or disability of the 


Section 4. The wasp shall keep the records of the Society, including 

he official list o ers, and conduct the correspondence pertaining 

to his office. He aaa turn over to the Treasurer all money received a2. 
shall report annually to the Society. 

Section 5. The Treasurer shall receive and hold all moneys belonging te 
the Society subject to the direction of the Council 
bills when approved in the manner prescribed by the Council, make an 
annual report to the Society, and at the end of his term of office shalt 
deliver to his successor all money and other property of the Society in 
his possession 

At such inion as the Council shall direct, he shall farnish the Councit 
with a statement showing his financial transactions since the date of his 
previous report, any outstanding indebtedness, the cash balance in hand, 


AMERICAN FERN SOCIETY 111 


and such other simple facts as shall enable the Council to know clearly 
the financial condition of the Society at the time. He shall close his 
accounts for the year promptly as of December 31 of each year, and as 


so! 

such records, vouchers, ete., as shall make possible a proper auditirg 
of his accounts, 

Section 6. The unexpired term or vacancy in any office shall be filled until 
the ensuing election by appointment by the Council. 

ARTICLE V. COUNCIL 

Section 1. The President, Vice-President, Secretary, Treasurer, and Editor- 
in-Chief shall constitute a standing committee to be known as the 
*“Council.’’ 

Section 2. The Council shall have general charge of the affairs of the So- 
ciety; of its publications and property; shall have power to expend the 
Society’s money and to act upon all questions not requiring a vote of 

the Society. 


ARTICLE VI. ELECTIONS 
Section 1. Before the first day of September of each year, the President, 


with the approval of the Council, shall appoint a nominating committee, 
consisting of a chairman and two other members, none of whom shall be 
y 


t. 
and received by the Secretary not later than October fifteenth, shall 
incorporated in the ballot for that year. 
Section 3. The President shall immediately thereafter appoint some mem- 
er not a candidate for office to act as Judge of Elections, and shall 


= 
Pa 


ll send his ballo 
be 
Immediately after election the Judge of Elections shall send to the 


to the chairman of the nominating committee. The aorred receiving 
the largest number of votes shall be declared oe shall be noti 
fied of his election by the Secretary. In case of a dhs nominating 
committee shall cast the deciding vote and shall ais the Secretary of 
its action 


112 AMERICAN FERN JOURNAL 


ARTICLE VIJ. AMENDMENTS 


Section 1. Proposed ainendments to this Constitution must be presented to 


v 
on the question of its adoption, the amendment shall be declared adopted. 


HENRY TRIPP 
, Successor to Henry George Fiedler 


Scientific Books and Periodicals 


Large stock of books on ferns 
and cryptogamic botany 


31 East 10th Street, New York 3, N. Y. 


CASTANEA 
Published by the 
Southern Appalachian 

Botanical Club 
Devoted to the botany of the interesting Southern Appalachians. 
Published Quarterly. Yearly subscription, including membership 

in the Club, $4.00. 
Address 


Dr. Earu L. Coreg, Editor 
West Virginia University 
Morgantown, West Virginia 


FERNS 
SPECIALISTS IN TROPICAL AND SUBTROPICAL FERNS 
We ship Price List Available 


LEATHERMAN’S GARDENS 
2637 North Lee Avenue South El Monte, Calif. 


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Treetise: 
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“THE PREPARATION OF 
BOTANICAL SPECIMENS 
FOR THE HERBARIUM” 


r. I. M. Johnston, of the Arnold Arboretum, Harvard University, has 
pi Ba shared the benefits of his long and successful experience in the 
collection and preparation of plant noodmans. 


Help for the amateur poco and hints for the —— collector, 
abound in this thirty-six a kerggetioa which D —_— 
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One copy of the treatise is yours for the asking. With it will b 
eluded samples of Botanical Papers (Driers, Mounting Sheets and het 

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Just ask for ‘‘a copy of the Johnston treatise.’ 


CAMBOSCO te ae gait COMPANY 
7 ANTWERP STREET GHTON STATION 
BOSTON, MASS. U.S.A. 


Vou. 51 JULY-SEPTEMBER, 1961 No. 3 


American Pern FJournal 


A QUARTERLY DEVOTED TO FERNS 
Published by the 


RICAN FERN SOCIETY 
is 


IRA L. WIGGINS 


CONTENTS 


Growing Ferns from Spores THORLEIP FLIFLET 


Some Chromosome Numbers of Icelandic Ferns and Fern-allies 
AsKELL Love anp Doris Live 


Ferns for a Lath House Fay MacFappsen 


A Resume of the Taxonomic Reorganization of Equisetum, 
Subgenus Hippochaete, I Ricwarp L. HavKE 


An Interesting oe Niche Involving Salvinia 
Seat Tomas A. Hurto 


The Occurrence of Mecodium wrightii in Canada Kunio IwatsvxK1I 
Notes on Polystick Grorce R. Procter 


The Genus Grammitis in Japan, with Description of a New Species 
Moroz Tagawa AND Kunio IWATSUKI 


Concerning Azolla imbricata. EUGENE Yu-Fene SHEN 


Shorter Notes: An Addition to the List of Ferns Found Growing - 


ally in Tennessee; Obituary: Alberto Chiarugi, 1901- 


Notes and News: A Fern Project Wins a Science Fair Award; 
iety of Economic Botany; A Mass Collection of Poly- 
stichum; Onoclea Spores Available__.____-__-__. 


American Fern Society 


ROLLA M. TRYON, JR. 


131, 


156 


157 


The American Fern Hociety 


Council for 1961 
OFFICERS FOR THE YEAR 
Cram A. Brown, Department of Botany, Louisiana State ious — 
Rouge, Louisiana 

Marce, RaYMOND, — Botanical Garden, 4101 East shernooke pr 
Montreal, Cana resident 

DonaLD oa Longwood Gardens, Kennett Square, Pte 
Secretary 
WALTER = Pures, Department of Botany, University of aati Tucson, 
Ariz asurer 
Tra L. ‘Wueine, Dudley Herbarium, Stanford University, ————e Calif. 
Editor- in-Chief 


OFFICIAL ORGAN 
American Fern Journal 


EDITORS 
get i. eo Herbarium, Stanford University, Stanford, a 
C. V, ____ Smithsonian Institution, Washington 25, D. 
Roa x pal oN 
Gra: rbarium, Harvard University, Cambridge 38, 
Joun H. Tides Dasleg Herbarium, Stanford University, Stanford, Calif 


An illustrated quarterly devoted to ‘to the — study of ferns, pub: blished 
quarterly by the American Fern Society, 3110 Elm Ave., Baltimore 11, Md. 
Second-elass postage paid ai sae imore, Md. 

Matter for publication nea id be addressed to Ira L. Wiggins, Natural 
History Museum, Stanford University, Stanford, Sno 

SS $2.50, exclusive of agency handling f mpl sent biti to poor 


bers of the erican Fern Society (annual dues, $2.50 

bership, 00; life membership, $50. 00). Extracted po Hg if v ordered in 
— e furnished authors at cost. They should be ordered when 
proo 


Back volumes $2.00 each; single back numbers 50 cents each; Cumulative 
Index to vols. 1-25, 25 cents, Ten per cent discount on orders of six volumes 
or more. 

Changes of address should be sent to the Secretary: Dr. Donald Huttle- 


ston, panned Gardens, Kennett Square, Pennsylvania a bers and sub- 
seribers should allow two months for changes 

Applications for mem Pp, subscriptions, orders tee rent numbers, and 
other business communications should be addressed to the Treasurer: Dr. 


Arizo’ 


LIBRARIAN AND CURATOR OF THE HERBARIUM 
Dr. W. H. Wagner, University of Michigan, Ann Arbor, Mich. 
oan department is 


di ed mnection with the library 
and herbarium. Mem! ret a har ne time, 
the borrower paying all postal or meres & — The — of the Journal 


also ied + 
is published at intervals, to assist th e exchang € 
from different 1 ties. ‘ose ge in obtaining specimens 


American Fern Journal 


Vou. 51 JULY-SEPTEMBER, 1961 No. 3 


Growing Ferns from Spores 
THORLEIF FLIFLET 


INTRODUCTION 
Some years ago I acquired a small wooded area next to my 
garden and to make it more interesting, I began planting ferns 
there. At the same time a friend, Mr. Thomas Carlson, who is a 
local pteridologist, introduced me to the American Fern Society, 
and I was all set for a new hobby. Later I became acquainted 
with the late Mr. M. D. Mann, Jr., then Treasurer of the Ameri- 
can Fern Society. Many will remember his interest in the So- 
ciety and its members, his beautiful fern garden and his ‘“‘give 
away’’ department. Being a conservationist rather than a bot- 
anist, I began growing ferns from spores, doing some experi- 
menting in order to find the most practical methods for the ama- 

teur fern grower. Of course, I did some reading, too. 
This review was originally intended only to cover the develop- 


which, in the case of viable spores, requires only that the spores 
be surrounded by an atmosphere at or close to the point of satu- 
ration (of moisture), while resting on a moist surface and sub- 
ject to favorable light and temperature conditions. However, the 
amateur who sows fern spores does so in order ultimately to pro- 
duce ferns for his garden. So it appeared logical to pursue the 
subsequent development of the tiny sporophytes or fernlets 
through transplanting, and so forth. This article covers only the 
growing of ferns by the amateur not possessing a greenhouse. 
Commercial methods are not considered. 


Volume 51, No. 2, of the JourNat, pp. 65-112, was issued June 29, 1961. 


114 AMERICAN FERN JOURNAL 


GENERAL DESCRIPTION OF METHODS FOR GROWING FERNS FROM SPORES 

The amateur gardener who wishes to grow ferns from spores 
has two simple methods available to him, the flowerpot method 
and the method using clear plastic containers. Both methods 
employ organic soil for the cultures and require only easy means 
of sterilization. Methods involving a high degree of steriliza- 
tion or the use of crocks, stones, peat, or agar will not be con- 
sidered. However, reference is made in the appendix to articles 
in the AMERICAN FERN JOURNAL covering this subject and gen- 
erally employing more highly sterile conditions. 

The process of growing a mature fern from a spore involves 
the following steps: 

a. Collecting and sowing of the spores. 

b. Germination of the spores as indicated by the green emerg- 
ing prothallia. 

ec. Period of growth of the prothallium (first stage) during 
which it attaches itself to the surface of the medium and de- 
velops its characteristic shape and size. 

d. Fertilization, a process which depends on the waterborne 
movement of the sperm cells and their fusion with egg cells. 
However, many species of ferns reproduce without recourse to 
the fertilization process. 

e. Development of the sporophyte or fernlet by the grower 
until mature enough to be planted in the garden. 

The entire process, commencing with spores, involves one and 
one half to two and one half years. 

Assuming that the spores are sown in the fall, the miniature 
ferns must be transplanted from the cultures in the following 
spring and kept indoors another year enclosed in a propagation 
box, ‘‘window greenhouse,’’ or a terrarium. By then the ferns 
are one and one half years old (counting from sowing of the 
spores) and may still be too fragile for the garden, in which case 
they should go into a cold frame or possibly into a lathhouse. 
The cold frame will require attention as to shading, ventilation, 
and watering in the warm season. Protection against freezing 


GrowIna Ferns rrom Spores 115 


during the following winter should be provided so far as this 
is practical. After the third winter, when the ferns are two and 
a half years old, they are set out in the garden, where they 
usually make rapid growth. All transplanting is done with the 
soil adhering to the roots and vigorous watering should be de- 
layed until they are well anchored so as to avoid disturbing the 
root system. 

The time schedule given above will be affected to some extent 
by the temperature and light available by the method employed, 
the propagation box described below resulting in a faster growth 
than the other method. The rate of growth also varies with the 
species of fern. The above period of two and a half years is 
based on planting young ferns outdoors in the spring only, 
avoiding fall planting. However, in some instances the grower 
may find that a year and a half period may be sufficient. 


THE FLOWER Por MrTHop ror GROWING PROTHALLIA 

. Equipment. Unglazed earthenware flower pots, 3” to 5” 
Size with drainhole, each provided with a piece of windowglass 
on top and a saucer underneath. Several pots may be placed 
in an aluminum or tinned iron pan. The pots should have a 
Piece of crock in the bottom. 

- Material. Water, which must be boiled. Drainage materi- 
al, preferably vermiculite, which should be washed go as to re- 
move soluble salts. Soil, principally or entirely humus, screened 
through one sixteenth inch mesh. 

¢. Procedure. The washed pots are half filled with vermicu- 
lite and soil is then added up to three-fourths inch below the 
top and pressed slightly down to a smooth surface. The pots and 
the covers are then heated in an oven (the pots without the covers 
on) for an hour or so at 250°F together with the saucers and the 
pan. After cooling the covers are put on and the pots partly sub- 
merged in water to moisten the contents. When the surface of 
the soil appears moist (not muddy) and the pots cooled to room 
temperature, the spores are sown without touching them with 
the fingers. The cultures are then placed in a warm room, near 


116 AMERICAN FERN JOURNAL 


a window but well shaded against direct sunlight. The pan or 
saucer will require watering once or twice a week to replace the 
loss from evaporation. A slight loss of moist air from the pots 
will also occur. The cultures should not be exposed to frost. 

The prothallium obtains and utilizes the soil water through its 
hairlike rhizoids. After fertilization and the emergence of the 
sporophyte, the prothallium disappears. The organic soil con- 
tains the necessary nutrients, although the microbial life has 
been temporarily destroyed by the sterilization process. Con- 
densation of water on the glass creates somewhat of a problem 
as large drops of water falling down in the pot may wash spores 
down into the soil. It is advisable to remove condensation 0c- 
easionally, for instance by tilting the whole culture. After 
germination, however, drops falling on the prothallia may even 
be of some advantage in facilitating the movement of the sperm 
cells in the case of sexual species that follow the normal process 
of reproduction. 


VARIATIONS OF THE FLOWER Pot METHOD 

There are, of course, many variations of this method as re- 
gards use of drainage material and sterilization of the soil. 
Some growers sterilize a flower pot in an oven, and then put in 
their own soil mixture, with or without drainage material. Next 
the soil is packed down and saturated with boiling water, and 
then a thin layer of charcoal and ground crocks passed through 
a screen and heated in an oven, is placed on top. The surface 
is then moistened by a fine spray. This arrangement corre- 
sponds to the use of sand over a seedflat, and is intended to 
avoid attacks by ‘‘damping off’’ fungi. The use of such an 
inert surface is said to hasten the germination of the spores by 
providing a medium of neutral reaction. Other growers, again, 
may sterilize the soil to some extent by holding a bag of soil in 
a pail and pouring boiling water over it; the weight of the water 
used must be at least twice the combined weight of the pail and 
the soil. However, there is no doubt that a higher degree of 
sterilization is obtained by heating the soil in an oven as de- 
scribed above. 


GROWING FERNS FROM SporEs 117 


PLASTIC CONTAINER METHOD 

The use of plastic containers already has been discussed by 
Dr. Benedict (1). I have used plastic jars for several years and 
believe they are well suited to the amateur. They are four 
inches in diameter and two and three-fourths inches high and 
made of clear plastic with clear plastic covers. They do not al- 
ways withstand boiling water so I wash them in water not hot- 
ter than my hands can stand. After rinsing them I dry them 
with paper towels. Mild solutions of some household disinfect- 
ant, such as Chlorox, unquestionably would be of some help in 
the cleaning of the jars. 

I fill the jars an inch and a half deep with washed vermiculite 
which is still wet, add some water (about an inch and a half 
above the bettom) and then three-fourths inch of woods soil 
(leafmold) which has been screened through a sixteenth inch 
mesh mosquito netting or kitchen strainer and then heated. 
The heating of the soil can be done outdoors over an open fire or 
in the kitchen oven. The pan containing the soil should have 
Some water in the bottom and, if an open fire is used, the soil 
must be stirred to avoid carbonization. The oven should be set 
at 250°F and the pan of soil kept at this temperature for an 
hour or so, resulting in a dry powder. 

Considering now the jars with vermiculite, water, and soil, 
the surface of the soil is leveled off and the jars are left to stand 
for a while with covers on until the excess water which may have 
floated some of the vermiculite has sunk down sufficiently to per- 
mit the surface of the soil to be pressed gently down to the 
proper density and made smooth. Due to its dryness the oven- 
heated soil has difficulty in absorbing water by capillary action 
and it may take a day’s time before the surface is moist and 
ready for the sowing of the spores. Dry spots in the surface 
must be avoided. Builders sand could be used instead of vermi- 
culite; it must be washed and then rinsed in boiling water. 
However, sand is much heavier and absorbs only half as much 
water as the vermiculite does. 


118 AMERICAN F'ERN JOURNAL 


Plastic jars are much handier to deal with than are regular 
flower pots and their appearance does not usually conflict with 
the decor of a room. The covers are sufficiently tight to obviate 
any further watering. As wild growth (mosses and fungi) does 
not occur to any serious extent, I have found the method satis- 
factory. Occasionally ‘‘damping off’’ fungi have appeared, some- 
times introduced perhaps with the spores themselves. Airborne 
spores of such fungi and of mosses may enter the cultures later 
in the process. 

THE WARDIAN CASE (TERRARIUM) 

In 1836, Dr. N. B. Ward, a Londoner, invented the case that 
bears his name, a glass-walled and -covered container provided 
with a layer of moist soil over drainage materials on the bottom 
for growing plants. His invention was considered important 
enough at the time to be made a subject of a lecture by the great 
physicist Michael Faraday at the Royal Institution in April, 
1838, and thereafter the invention became generally known (2). 

The Wardian case was at one time much used for ferneries in 
the home but was probably of greater value in transporting 
living plants to botanie gardens. Today, the case is most often 
found in the form of a plant terrarium. The principle involved 
is that moisture-loving plants are protected against dry air and 
that only infrequent watering is required. There are various 
forms of the Wardian case, an interesting one being the ‘‘ Won- 
derglobe’’ described in ‘‘Wildflowers for Your Garden’’ by 
Helen S. Hull. 

It should be noted that ordinarily the Wardian case is not 
airtight around the cover, so that some air leaks in and out. 
The air leaking out is moist and the air coming in is compara- 
tively dry, and accordingly there is a loss of water which, in the 
case of a terrarium with a volume of five gallons amounts to a 
few teaspoonfuls a month. 


GROWING FERNS FROM Spores 119 


METHODS FoR GROWING PROTHALLIA AND FERNS IN THE WarDIAN CASE 
a. FLOWER Pots IN THE TERRARIUM 

The equipment for this method is easily obtainable. <A five- 
gallon aquarium with a glass cover, used as a terrarium, pro- 
vides space for eight three-inch flower pots. Such a terrarium 
may be built in a home workshop, using glass for the two sides 
and the cover and one by ten inch lumber for the ends and the 
bottom. The bottom should be covered with heavy roofing 
paper bent up a couple of inches along the edges to form a 
watertight bottom. The boards and bottom should be painted 
with white lead. The dimensions of the case can vary; I like 
them ten inches high and 24 inches long. For cultures use three- 
or four-inch flower pots without glass covers or saucers. Spread 
one inch of vermiculite on the bottom of the case and begin with 
a half-inch depth of water. This case will give room for 12 
three-inch pots, each of which may contain four small ferns 
after transplanting from the cultures. However, a two-foot 
Square box with ten-inch sides and with glass only used for the 
cover may be easier to make and is twice as roomy. The glass 
pane should be divided in the middle and be double thick sheet 
glass (one-eighth inch). 

b. FLowER Pors IN PropaGaTIoON Box WITH FLUORESCENT LIGHT 

The various forms of the Wardian case described above can be 
used where daylight is available but they may also be placed 
under a fluorescent light and kept out of way, say in the cellar. 
However, if greater capacity is desired, a larger propagation 
case made from transite boards is recommended. The case used 
So successfully for spore cultures and small ferns by Mr. Mann 
has been described and illustrated by Dr. Benedict (1). This 
method gives results more quickly than other methods since the 
illumination and temperature are under full control. 

This box may be 30 inches wide, 60 inches long, and 30 inches 
high, with two doors forming the front. Two 48 inch fluorescent 
lights in a reflector are placed in a recess in the top of the box. 
At the ends of this recess should be a few ventilating holes. 
The bottom of the box is covered with an inch of vermiculite, kept 


120 AMERICAN FERN JOURNAL 


wet by an inverted bottle of water such as is used for supplying 
water to a small trough in a chicken-house. Additional space 
is available by a ten-inch shelf placed 20 inches above the bottom. 
The total capacity is probably 130 three-inch pots. A soil- 
heating cable and a breeder thermostat may be provided as well 
as a time switch for the electric light, which may be kept on at 
least 20 hours a day. Cultures are kept in this box as well as 
plants, although the most common practice is to keep cultures in 
subdued light until after germination. Mr. Mann used washed 
vermiculite as drainage material in the pots with a soil two-thirds 
‘*hyperhumus’’ and one-third screened peatmoss. He sterilized 
only the pots. He recommended a temperature of 70 to 75°F 
and also that the cultures be sprayed with water when the pro- 
thallia were well developed 

Mr. William S. Johnston employs a similar box and method 
except that he does not require the heating cable. His soil mix- 
ture is ‘‘hyperhumus,”’ sand, and garden soil. He also referred 
me to an English writer, A. J. Macself, who recommended tem- 
peratures in the 60’s so as to get hardier plants, although this 
would result in much slower growth. The above propagation box 
if requiring heat beyond that furnished by the fluorescent lamps 
could be supplied with one or two ordinary light bulbs rather 
than a soil-heating cable. 
¢. PLASTIC CONTAINERS IN ‘‘ WINDOW-GREENHOUSE’’ 

A “window-greenhouse’’ consists of a case holding a set of 
shelves with clear plastic cloth in the back and provided with a 
removable cover in front also covered with clear plastic cloth. 


Wardian case built in several stories. This ‘‘greenhouse’’ de- 
pends on room heat, but as the nights may be quite cold in the 
ordinary dwelling, the growth will be much slower than in dex 
case of a propagation box. The advantage of the ‘‘greenhouse”’ 
is that everything is visible; on the other hand, the capacity is 
rather limited. 

The frame and the shelves are built of one- by four-inch 
boards for holding the four-inch plastie jars or alternatively 


GRowING FERNS FROM SPoRES I2i 


three-inch flower pots with their saucers. Two intermediate 
shelves are placed eight to ten inches apart making the height 
of the case 24 to 30 inches. The plastic cloth is held by half-inch 
halfrounds (such as used for holding the sereencloth to the 
frame of a window screen) against the rear edge of the boards. 
The front cover is made up by a frame of one- by two-inch 
boards set on edge, braced at the corners, and provided with a 
vertical board in the middle to hold the plastic cloth better. The 
length of the case could be equal to the width of an ordinary 
window. Plants should be kept away until the paint solvents 
have evaporated. The case can be supported from the floor by 
extending the sides downward to provide legs which can be 
secured to the floor by a small angle iron and serews. For stabil- 
ity, the top of the case should be braced to the window casing 
and, accordingly, the sides should be continued upward to per- 
mit headroom under the braces. Placing the case 18 inches away 
from the wall will permit access to the window. The case could 
be supported from the windowsill and braced higher up but this 
would make the window inaccessible. A 24 by 36-inch wooden 
kitchen table can also be used to support the case. With such an 


and screws, The table-top should be at the same level as the 
window-sill. The capacity of this “table model”’ is 21 four-inch 
containers, but it will hold 28 if made of 12-inch lumber, the 
case then being 30 inches high but with only one intermediate 
shelf, assuming that the case is to be 36 inches long. 

This “‘greenhouse’’ provides a safe place for keeping the sea- 
son’s supply of cultures with their covers on, but it is principally 
built to hold the small ferns after they are transplanted into 
four-inch diameter and 3-inch high plastie jars. These will have 
an inch and a half of wet vermiculite on the bottom and an inch 


122 AMERICAN FERN JOURNAL 


of soil. Four plants may be used in a Jar. Using containers that 
have no drainhole may seem unusual but when properly used 
there will be no stagnant water in the soil and v vatering will only 
be required at intervals of several weeks. Paper may be used on 
the shelves. Some condensation will oceur on the woodwork and 
it will be subject to mildew in the long run. Empty shelves may 
have jars filled with just water to help moisten the air. The case 
may also hold three-inch flower pots with their saucers, but the 
additional weight may have to be considered. 


MISCELLANEOUS COMMENTS 
a. DRAINAGE MATERIAL. 

In fern literature one often sees references to drainage and 
drainage material. As is well known, the root system of a fern 
must have access to oxygen and soil in order to make nutrients 
available to the plant. At the same time, water must be supplied 
at the required rate. Considering the ordinary flower pot the 
common practice is either to place broken pieces of crock in the 
bottom or gravel covered with a layer of dead sphagnum moss. 
The watering is done from above and a saucer collects the excess 
water. One finds at times, an expression of the idea of supplying 
air to the roots by the lugs that are sometimes found on special 
saucers or flower pots. 

In nature, plants receive their water as rain from above or 
through capillary action from below. Ferns have shallow roots 
and may be planted in flower pots which are filled one-third with 
vermiculite. The pots should be placed inside a container reach- 
ing half way up the pot. Such an arrangement is suitable for 
houseplants requiring much water. Watering can then be done 
from below and at greater intervals as a storage of water is 
provided. Submergence of the roots in water is easily avoided 
and proper aeration of the roots is obtained. The use of the 
plastic jars for small ferns in the ‘‘window greenhouse’” as 
described above also necessitates the same drainage material 
for water storage; watering is done from the top, and care must 
be taken to keep the water level below the roots. 


GROWING FrRNS FROM SporsEs 123 


b. Som Usep For QCuLrurEs. 

The use of organic soil as a medium for the growing of spores 
appeals to the amateur as well as to the commercial grower, as 
transplanting is easily done. Each grower has his own prefer- 
ence as to a soil mixture, which generally is made up from good 
loamy garden soil, humus, and builder’s-sand in equal amounts 
or with the humus predominating. Soil recently manured must 
be avoided. After the soil has been screened it is sterilized. 

Soil taken from gardens or woods contains insects, insects’ 
eggs, earthworms, and so forth and it is also alive with spores of 
fungi and mosses. Accordingly, the soil must be sterilized, which 
will also destroy the nematodes that cause browning of the leaves. 
c. SPOREs. 

If a fertile blade of fern is taken at the time of maturity and 
pressed between two sheets of absorbent paper, an outline of the 
blade will appear. This configuration is made up of empty spore- 
cases, free spores, and in certain cases also pieces of indusia. 
Unopened sporangia also may be present. By depositing this 
powder on a black surface and removing the larger pieces of de- 
bris a brown colored powder remains, part of which consists of 
pieces which are visible to the naked eye. The other, only visible 
as a fine dust, is comprised mostly of spores. By applying a 10 
magnifying glass, the spores become individually visible. It is 
not necessary to separate the spores from the empty or full 
Sporecases, but the mixture should be examined to judge the 
number of spores available for sowing. It is important to scrape 
the spores off from the paper of the envelope where they have 
been kept, because an inspection will often reveal a surprisingly 
small number of free spores. Sometime it is possible to see the 
sporangia exploding while resting on a black surface, scattering 
the spores around; some of these are carried away by the air- 
currents. As a sporangium contains normally 64 spores, the in- 
dividual spore is so small that it is practically inyisible to the 
naked eye, which led to the belief, in folklore that the spores 
were actually invisible and so useful in witcheraft. 


124 AMERICAN FERN JOURNAL 


The first problem confronting the fern grower is the collecting 
of spores in the woods or in the garden, which requires a close 
inspection of the fronds with at least a 10x hand lens (3). When 
collecting spores from small ferns an entire frond may be 
taken, but in the case of a large fern only a few pinnae need to 
be gathered. The harvest is taken home, dried to release the 
spores and finally cleaned of such debris as is easily removable. 
A frond in a bag or envelope of ordinary paper will dry to a 
degree depending on the temperature and the amount of mois- 
ture in the air in the bag. It is a slow process; the paper soaks 
up moisture on one side and lets it evaporate on the outside, 
assisted by the movement of air through any opening available. 
After the house heating system starts up in the fall, the drying 
process speeds up. However, if the frond has been picked at the 
' proper time the spores will fall off in a few days. 

In my own experience the collecting of spores is the weakest 
link in the process of fern growing by amateurs. The owner of 
a greenhouse is able to catch the spores as soon as they begin to 
‘ripen under his watchful eye, and he may enclose the frond with 
a paper bag before he removes it. 

The second problem concerning spores is that of their viabil- 
ity. Most viable spores will germinate visibly to the naked eye 
within a period of a few days or weeks depending on the species 
and the degree of dryness. The temperature of the culture is 
also a factor. In most of the species dealt with by the amateur 
it is probably a matter of a couple of months. The life of a 
spore kept at ordinary temperatures will in most cases span 
several months or even years. Spores of the Osmundaceae are 
quite short-lived, however, unless kept in cold storage (4). Spores 
of some hybrid ferns are practically sterile and may not germi- 
nate, but the amateur will only meet such cases very rarely. 

The third problem when growing ferns from spores involves 
germination of the spore and fertilization of the prothallium 
which leads to the development of the sporophyte and the young 


GROWING FERNS From Spores 125 


fern. There are at the present no data available as to the per- 
centage of germination of fern spores. Before germination, very 
little can happen to the culture except that spores may have 
been washed down by drops of condensation. After germination 
prothallia growing too closely together will have to be thinned 
out in order to provide space for further growth. Also, close 
spacing appears to interfere with the fertilization process. 
The prothallium will keep growing and reach the stage when 
fertilization is to take place. At this point some growers ree- 
ommend the use of free water, either as a fine spray or by sub- 
merging the prothallia for a short time so as to encourage the 
movement of the waterborne sperm cells. In nature, there is a 
great difference between day and night temperatures, resulting 
in dew, a phenomenon similar to the condensation taking place in 
the spore cultures when they are subjected to cooling. At a uni- 
form temperature no condensation occurs, but it is most probable 
that the sperm-cells will find their way through the water-film 
covering the prothallia and the surface of the soil without re- 
course to spraying. Cultures are generally subject to a daily var- 
iation in temperatures resulting in condensation on the culture. 
Through the fertilization of the prothallium the second genera- 
tion, the sporophyte, is initiated, which thereupon develops into 
a fern. Ina culture, however, there may be a long interval of 
time between the appearance of the first and the last sporophyte 
and my patience runs out when a dozen are ready to be trans- 
planted and the remaining prothallia are then abandoned. 


Now, a word of encouragement to the amateur fern grower. 
He should know that the spores he takes home and sows have a 
much greater chance of producing ferns than if he had left 
them on the plant, and, also that the professional grower only 
grows ferns which propagate very readily and, in many eases, 
asexually, which is the easy way for a prothallium to turn into 
a sporophyte. Readers interested in the scientific study of 
Spores will find much material published in the AmerTcAN FERN 
JOURNAL, 


126 AMERICAN FERN JOURNAL 


d. ILLUMINATION. 

In nature there is a daily cycle of light and darkness during 
which time plants grow continuously. In an article describing 
the forcing of ferns in a greenhouse for the 1938 International 
Flower Show, Dr. and the late Mrs. E. D. Thurston, Jr., made 
the observation that the growth was greater during the night 
than during the daylight hours (5). This observation has been 
confirmed by growers of flowering plants under natural light. 

r. R. G. Benedict (1), in discussing the use of fluorescent light 
by Mr. M. D. Mann, refers to the period of light as 20 hours a 
day and also mentions some cases of continuous illumination. 
His discussion dealt with spore cultures rather than ferns how- 
ever. In greenhouses, some flowering plants may have the elec- 
tric light on during the night while other plants may be kept in 
continuous darkness for a time to accelerate the time of flower- 
ing. Information regarding artificial illumination for erowing 
ferns in a propagation box is incomplete. 
CONCLUSION 

he above article is rather lengthy. It has been written on 
the assumption that a reader interested in a hobby is able and 
willing’ to make an effort to explore the basic elements of his 
hobby. In Great Britain the study and growing of ferns has 
always been popular, especially among people living in retire- 
ment but who continue to seek happiness in their surroundings, 
gardens, or woodlands nearby. bese oyna to indoor living 
may well enjoy a ‘‘window-: Tt is hoped articles 
in the American Fern Jontnal for the use of ue layman will 
stimulate the interest of American garden-lovers in ferns, na- 
tive and foreign. Although most fern gardeners will appreciate 
the botanical features of their ferns, there is pleasure also in 
developing attractive landscape effects. 


LITERATURE CITED 


. Beneptcr, R. C. Aids to Spore Culture. Amer. Fern Journ. 45: 60-64. 


2. Smirn, Jonn. Ferns: British and Moreign. London, 1866, (Chapter IV, 
Cultivation in Ward’s Cases.) 


Curomosome NuMBERS or Icenanpic Ferns 127, 


3. KLEINSCHMIDT, W. F. Growing Ferns from Spores. Amer, Fern Journ. 
42: 117 1952. 

Stoxey, A. G. Viability of Spores. Amer. Fern Journ, 41: 111-115. 

1951 


a 


Oo 


THursToN, BE. D. Foreing Native Ferns for Exhibition. Amer. Fern 
Journ, 29: 85-94, 1939 
ADDITIONAL REFERENCES 
BENepict, R. C. Fern oe Amer. Fern Journ. 38: 148-154. 1948. 
(Ripening of spore 
ass eae A. Wincie with Fern Spores. Amer. Fern Journ. 38: 12-16. 
. (Cultures on stones.) 
Kise , W. F.. A. Met ue of Preparing Spores for Fern Cultures. 
Am aes Journ. 47: 95-98. 1957. 
ig B. Te ape te A the Porat, of Spores of Ferns for 
Artificial rane Amer. Fern Journ 161-163. 1958. 
CLARKE, HERBERT M, ie Fern Prothallia. Amer. Biol. Teacher 16: 
214, 215. 1954. 
WAGNER, W. , JR. Asplenium ebenoides x platyneuron, a New Triploid 
pals Rees under Artificial Conditions. Amer. Fern Journ. 46: 
1956. 


aes Growing Fern Prothallia in Liquid Culture. Amer. Fern 
Journ. mm er 161. 1955. 


128 Kenttworta Roap, Mountain Lakes, N. J. 


Some Chromosome Numbers of Icelandic Ferns and Fern-allies 
AskeLL Live anp Doris Live 

In connection with the collection of material for a cyto- 
taxonomic review of the higher plants of Iceland (Live & Love, 
1956), the present writers also made fixations of Icelandic 
pteridophytes. The results obtained from the Lycopsids have 
already been reported (Live & Live, 1958), but chromosome 
numbers determined from other groups still remain unpublished. 
More exhaustive investigations based on this material are not 
possible in the near future. Since the observations already made 
have a certain interest, there seems to be justification in making 
them available without further comments. All the counts here 
reported were made on somatic material fixed and treated in the 


128 AMERICAN FERN JOURNAL 


same way as the root tips of the higher plants studied by Love & 
Love (1956). For previous reports on the same species and 
races, the list of European chromosome numbers by Love & Love 
(1961) can be consulted. If no exact locality is given below, 
the material originated from several places in the regions men- 
tioned. The authors wish to thank Dr. M. S. Chennaveeraiah 
for assistance with some of the chromosome number determina- 
tions. 


SPECIES 2n LOCALITY 

Siscneetiia SELAGINOIDES (L.) Link 18 SW. & N. Iceland. 
EQUISETUM AR - NSE L. 

ssp. ARVENS 216 SW. Iceland. 

ssp. BOREAL ethene) Love 216 SW. & N. Iceland. 
EquiIseTUM nee NSE Ehrh. 216 SW. Iceland. 
EQUISETUM FLUVIATILE 216 SW. Teeland 

IPPOCHAETE HYEMALIS (L.) Borner 216 N. Iceland. 
pe atoms VARIEGATA, (Schleich. ) 216 SW. Teeland. 

orner 


ne 
OPHIOGLOSSUM VULGATUM L. 
OS! 


var. ISLANDICUM - éve & Love 480 SW. & W. Icela 
BOTRYCHIUM BOREALE (F'r.) Milde 90 Eyjafjordur, = Toot. 
BoTRYCHIUM LANCEOLATUM (Gmel.) 90 Hengill, SW. Ice 

Angstr. 
Borrycuium Lunaria (L.) Swartz 90 SW. & N. Iceland. 
PHEGOPTERIS CONNECTILIS (Michx.) 90 SW. Iceland 

Watt 
Gymnocarpium Dryopreris (L.) Newm. 160 SW. & W. Iceland. 
ASPLENIUM VIRIDE Huds. 2 Fagurhélsmsri, 8. Iceland. 
ATHYRIUM FILIX-FEMINA (L.) Roth 80 SW. Iceland. 
ATHYRIUM DISTENTIFOLIUM * ac aue 80 Hafnarfjérdur, SW. Iceland. 
WoopsIA ILVENSIS (L. 82 SW. Iceland. 
Woopsia ALPINA (Bolton) Ss: B. bits 164 abla deme W. Iceland. 
CYSTOPTERIS FRAGILIS (L.) Ber 168 SW. Ice 
PotysticHtumM Loncuitis (L.) Roth 82 apalluhrae®. aah besitos 

v , Ww. 


DryopTeris DILATATA (Hoffm.) A. Gray 164 SW. & N. Iecelar i 
BLECHNUM SPICANT ue ae E. Smith 68 Kaldarsel, SW. ap pan 
POLYPODIUM VULGAR 148 Hvalfjérdur, SW. Iceland. 
LITERATURE CITED 
Love, A. & D. Lévr, 1956 Cytotaxonomical Conspeectus of the Teelandic 
Flora. Acta Hort. Gotob. 20: 65-2 
—1958 Cytotaxonomy me Classification of Lyeopods. Nucleus I: 
—1961 Chromosome Numbers of moshing and Northwest European Plant 
Species. Opera Botanica 5 (in presi 
Institut Boranigue, eioneains pE MonrtréaL, Montriat, 
CANADA. 


FERNS ror a Lata House 129 


Ferns for a Lath House 
Fay MacFappren 

A great many kinds of ferns ean be grown in small lath houses 
in Los Angeles, California. Many prefer sun in their native 
haunts, but most of them prefer shade in southern California, 
at least the kind of light shade provided by a lath house of the 
kind described in my last article in this Journal. Very near 
the coast, where the air is moister, the sun-loving ferns will 
stand lots of sun, but inland they want less sun; in the interior 
valley, where conditions approximate desert conditions, lots 0 
shade and artificial humidity is a necessity. In southern Cali- 
fornia all ferns look better in some shade. There are plenty of 
other kinds of sun-loving plants for full sun. 

When T started growing ferns in Los Angeles in the early 
thirties, Mr. Baldwin had been growing ferns from spores for 
the wholesale and retail trade, although only a few nurseries sold 
them. My first ferns from the Baldwin Nursery were species 
of Pteris and some of the cultivars of Pteris cretica. Most of 
these I still have, such as P. dentata, P. quadriaurita, and P. 
cretica ev. Drinkwateri and Childsii. 1 also got P. tremula and 

- quadriaurita ev. argyraea, both beautiful for two or three 
years, but which have to be replaced frequently. Most Pteris 
ferns are rugged and compact, and are among the easier subjects 
for cultivation. : 

One of the interesting ferns sold by Baldwin was listed as 
‘‘Nephrodium K. O. Sessions, ’’ evidently introduced by the hor- 
ticulturist Miss Kate 0. Sessions, but the botanical indentity of 
which has been in doubt until C. V. Morton recently indentified 
it as the Japanese species Thelypteris acuminata. It is an orna- 
mental fern, so well adapted to conditions in southern California’ 
that it has escaped from cultivation at least casually. 

Among the larger ferns suitable for a lath house are the 
Microlepias. In a corner M. platyphylla is good; it is tall-grow- 
ing and rather coarse. Microlepia strigosa, mistakenly sold as 
M. spel wneae, is even more ornamental, but is perhaps better out- 


130 AMERICAN FERN JOURNAL 


side if the lath house is small, because it eventually takes con- 
siderable space. A peculiar cultivated form that I have grown 
for many years, probably obtained originally from Baldwin, has 
been described as M. strigosa forma MacFaddemiae. Wood- 
wardia radicans is another suitable large, decorative fern for 
the lath house. These Mierolepias and Woodwardias do well 
outside under trees or on the north or east sides of houses, where 
they get some protection from sun and wind. Another fern that 
may eventually get too large is the leather fern, Rumohra adi- 
antiformis, but it is good for several years. 

Some large ferns should be grown with caution. Hypolepis 
punctata and the other species are a constant care in keeping 
the old fronds cut. The common bracken, Pteridium aquilinum, 
has long, running rhizomes that will make such a jungle that it 
will be difficult to get rid of; it should probably never be 
planted, except perhaps outdoors in places where space is no 
problem. 

The native southern maidenhair fern, Adiantum Capillus-ven- 
eris, is easily grown when once started, and in fact it became a 
weed in my lath house, but many people have a difficult time in 
starting it. It likes lime and if it is put next to a limestone 
rock or the foundation of a house it starts with ease; good soil 
and oyster shells help. After it is started it seems to grow any- 
where. On the other hand, the northern maidenhair, Adiantum 
pedantum, only grows outside or in lath houses in southern Cali- 
fornia for a short time; although it may be possible to keep it 
alive for a few years its beauty is soon gone. It grows naturally 
along streams and in deep moist woods; it is too hot and dry in 
southern California. However, it does well in a greenhouse, if 
given good soil, oyster shell, good drainage, and lots of water. 
The other maidenhairs such as A. hispidulum, A. formosum, and 
the cultivars of A. euncatum T have been unable to establish 
outdoors due to drying winds and low humidity. 

I grow successfully a great many other attractive ferns. 
Among these are Rumohra aristata, Asplenium bulbiferum, Poly- 


Resume or Suscenus HippocHarrr 131 


stichum Dudleyi, Blechnum occidentale, Dennstaedtia rubiginosa, 
and D. glauca. 

Of course, a fern lath house need not contain ferns exclu- 
sively. Fuchsias, begonias, and azaleas like the same growing 
conditions and give needed color. Camellias are good, although 
they eventually get too large for the average lath house, and they 
may get too much shade. Helxine is beautiful, of course, but it 
is better not to be planted, as it eventually gets in the roots of 
all your ferns, and is one thing that makes fern growing difficult. 

Once ferns are established, they are the easiest kind of garden- 
ing. Let a hand-lens be your constant companion in the garden 
and you will be surprised at all the things you are going to find 
out about your ferns. Growing ferns is very rewarding. To 
watch a fern develop from spore into a beautiful plant, even 
though this takes a long time, does something to the gardener 
spiritually. 


5450 CARLIN Street, Los ANGELES, CALIFORNIA. 


A Resume of the Taxonomic Reorganization of 
Equisetum, Subgenus Hippochaete, I. 
Ricuarp L. Hauke 

In a recently completed study of Equisetum, subgenus Hippo- 
chaete (the scouring rushes, snake grasses, or joint grasses), I 
developed a taxonomic system that varies from that presently 
accepted for this group of plants. I wish to present a résumé of 
this system in the American Fern Journal that it may be read 
criticized, and possibly used by people working with the vascular 
cryptograms. 

The study was monographie in scope, involving field experi- 
mentation, garden culture, gametophyte culture, and study of 
over 6,000 herbarium specimens, and including the statistical 
analysis of measurements from more than 2,000 specimens and 
observations of microscopic anatomy of over 400 specimens. I 
concluded that because of frequent interspecific hybridization, 


132 AMERICAN FERN JOURNAL 


clinal variation, and extensive morphological variability result- 
ing from traumatic injury or environmental stress, an ex- 
cessive number of infraspecific taxa have been described and 
named. Most of these do not warrant formal taxonomic recog- 
nition. 

Perhaps the best known work on Equisetwm is Milde’s Mono- 
graphia Equisetorum (1867). In this, Hippochaete is regarded as 
a separate genus, containing the following infrageneric groups: 


A. Equiseta pleiosticha! E. debile 
a. Equiseta planifolia b. Equiseta mexicana 
E. xylochaetum E, myriochaetum 
E. martii E. mexicanum 
hb. Equiseta angulata c. Equiseta hiemalia 
iganteum i 
E. pyramidale E. robustum 
. schaffneri E. laevigatum 
B. Equiseta ambigua d. Equiseta trachyodonta 
E. ramosissimum E. trachyodon 
E. sieboldii E. variegatum 
C. Equiseta monosticha E. seirpoides 


a. Equiseta debilia 


Milde never made any specifie combinations under Hippo- 
chaete. Baker (1887) called Hippochaete a subgenus, and listed 
E. myriochaetum, E. mexicanum, and E. schaffneri as synonyms 
of E. giganteum, but maintained E. rylochaetum as a separate 
species. He never mentioned E. marti or E. pyramidale. Boer- 
ner (1912) and Farwell (1916) independently recognized Hip- 
pochaete as a separate genus. Farwell divided it into two sec- 
tions, sect. Euhippochaete, containing H. hyemalis, H. variegata, 
H. scirpoides, and H. prealta, and sect. Ambigua, containing 
H. laevigata and H. nelsonii. Rothmaler (1944) followed Boer- 
ner in treating Hippochaete as a genus, and recognized the fol- 
lowing subdivisions: 


“Equiseta pleiosticha,” and similar group-names, being in binomial 
torte must be considered as descriptive phrases, and are not validly pub- 
lished subgeneric, sectional, or subsectional names, The same applies to 
Schaffner’s groupings such as ‘‘Equiseta Primitiva.’’ 


Resume or Susaenus HippocHarre 133 


Sect. Stichopora (A. Br.) Rothm. 
Subsect. Hiemalia (A. Br.) Rothm. 
A, hiemalis 
H, trachyodon 
Subsect. Homocormia (Pfitzer) Rothm. 
H. variegata 
Sect. Univaginata (Pfitzer) Rothm. 
HA, scirpoides. 


Schaffner (1921, 1925, 1930a, 1930b) developed a radically 
different classification of the genus Equisetum. He did not ac- 
cept Hippochaete as a taxon, but rather put the species of Hip- 
pochaete into four groups: 


I. Equiseta Primitiva 

E. xylochaetum, E. giganteum. 
Il. Equiseta Hiberna 

‘H mytioetiaet E. , E. debile, E. laevigatum, E. preal- 


tum, E. hiemale. : 
TIT. Equiseta Ambigua 

!. kansanum, E. funstonii, 
IV. Equiseta Pusilla 

E. nelsonii, E. trachyodon, E. variegatum, E. scirpoides. 


Although not a comprehensive review of the taxonomic treat- 
ments of the scouring rushes, this should illustrate the disagree- 
ment about the proper classification of this group of plants. It 
Should also serve as a background against which I can place the 
System of categories I believe most accurately reflects the rela- 
tionship of the various taxa of Hippochaete. This system follows: 


Equisetum, subgenus Hippochaete 
A. Seetion Incunabula 
1. Hquisetum giganteum 
B. Seetion Ambigua 


2. Equisetum ramosissimum 
2a. Subspecies ramosissimum 
2b. Subspecies debile 

3. Equisetum myriochaetum 


Cay 


Equisetum laevigatum 


134 AMERICAN FERN JOURNAL | 


C. Section Hippocha 
Ca. Subsection oe nnantia 
5. Equisetum hyemale 
- ar. hyemale 
. Var. vem 
Cb. Subsection Homocormia 
quisetum raricatom 
6a. Var. iegatum 
6b. Var. save num 
7. Equisetum scirpoides 


ybrids 
Noliecti um X schaffneri (E. aang < myriochaetum) 

Equisetum hyemale * myriochaetum 

Equisetum X* moorei (EF. hyem bos 4 fusca 

Equisetum X ferrissii (BE. hyemale < laevigatum) 

Equisetum X trachyodon (E. hyemale < eae a 

Equisetum X nelsonii (E. laevigatum X variegatum) 

Equisetum ramosissimum X variegatum 
Descriptions of all taxa will be included in this résumé but 
oniy the most important synonyms will be given. Complete lists 
of synonyms and citations of specimens examined can be ob- 
tained by writing University Microfilms Inc., Ann Arbor, Michi- 
gan, for a microfilm of the thesis, entitled A taronomic mono- 
graph of the genus Equisetum, subgenus Hippochaete. 

In the descriptions that follow reference is made to the col- 
lenchyma and the endodermal patterns. These are determined 
from a cross-section of an internode from below the middle of the 
stem. The collenchyma is the supporting tissue under the epi- 
dermis; it oceurs in thickened strands under the ridge (carinal 
eollenchyma) and under the groove (vallecular collenchyma). 

he endodermis is a distinctive layer of cells in the vicinity of 
the vascular bundles, and it occurs in three patterns. If each 
vascular bundle is encircled by an endodermis it has an indi- 
vidual pattern; if there is one endodermis surrounding the ring 
of vascular bundles, it is called the outer common endodermis ; 
and if there is one endodermis surrounding the ring of vascular 
bundles and another internal to the ring of vascular bundles, 
that is called the double common endodermis. 


Resume or Susgenus HierocHarre 135 


Subgenus Hippocuarre (Milde) Baker, Fern Allies 3. 1887. 
Equisetum Il Homophyadica A. Braun, Flora 22:308. 1839 (pro parte). 
Equisetum II Sclerocaulon Déll, Flora des Grossherz. Baden 1:65. 1857. 
“‘Equisetum II Equiseta eryptopora’’ Milde, Abh. Schles, Ges. 2:138. 
1861. (Invalid.) 
Equisetum subg. Cryptostoma Milde, Verh. Zool.Bot. Ges. Wilen 14:526. 
18 Pro syn 
Hippochaete Milde, Bot. Zeit. 23:297. 1865. 
Equisetum sect. 2. Hippochaete Milde, Fil. Eur. 230. 1867. 
Hippochaete sect. Stichopora Rothm. Repert. Sp. Nov. Fedde 54:81. 1944. 
Equisetum sect. Cryptopora Milde ex Grinteseu in Salvulesecu, Flora 
Republicae Popularis Romanicae 1:67. 1952. [Sectional name may have 
an earlier valid publication! | 
Although I am aware of a body of opinion by botanists past 
and present that Hippochaete should be maintained as a genus 
Separate from Equisetum, I shall nevertheless continue to treat 
it as a subgenus. There are, indeed, absolute and apparently in- 
violable differences between the two groups in stomatal structure, 
the chromosomes have some distinctions, so far as they are known 
(Manton, 1959), and hybrids between these groups are probably 
never formed. These differences, however, are few when com- 
pared to the great similarities in general morphology, anatomy, 
and reproductive structure. The contrasts of deciduous stems 
with blunt cones in Equisetum vs. evergreen stems with apicu- 
late cones in Hippochacte, though usually distinctive, are vio- 
lated by E. laevigatum and E. ramosissimum subsp. ramosissi- 
mum. The stomatal arrangement of E. arvense is matched by 
that of FZ. giganteum. Thus the differences between Equisetum 
and Hippochaete are far outweighed by the similarities between 
them. To reflect this taxonomically, they should be included 
within a single genus. Since a few distinct differences do exist 
between all of the species in one group and all of those within 
the other group, the groups may be maintained as subgenera. 
The type for this subgenus is Equisetum hyemale, 


136 AMERICAN FERN JOURNAL 


Diagnostic Kry To SEcTIONS, SUBSECTIONS, SPECIES, SUBSPECIES AND 
VARIETIES OF SuBG. HIPPOCHAETE 
Stems with regular verticils of branches; stomata in bands of tw 
more lines; individual endodermises in stem — 


1. E. giganteum 
Stems with regular verticils of branches, irregularly branched, or un- 
branched; stomata in single 
more lines; individual, outer 
mis in ste 
b 


nae sometimes in bands of 
mon, or double common endoder- 
em. Characters never in combination found in sect. Incu 
abuia. 
Stems branched and evergreen or deciduous, or unbranched and de- 
ciduous; stomata in one to several lines; ridges ¢ 
Sect. Ambigua 
Stem with individual endodermises; stomata in single lines; teeth 
regularly breaking off; stems branched and evergreen. 
2b. E. ramosissimum subsp. debile 
Stem with double common weeny stoma 
lines; teeth persisting or breaki 
bcdnchda. evergreen idu 


a in one to several 
g off; stems branched or un- 
Stomata in one to several lines; teeth persistent; lower sheaths 
ually’ tans Sos 2a. E. ramosissimum subsp. ramosissimum 
Stomata always in oné line; teeth breaking off 
gree 


; lower sheaths 


Stems evergreen, with regular verticils of branches. 


3. E. m brava ee 
Stems deciduous, rl eka mana 4. E. laevigatum 
Stems unbranched, ever stomata shin i in one sages idge: 
arely convex, iby acme ue oe ee t. Hippockuate 
Rhizome with individual endodermises; sheaths with black girdles, 
feeth articulated =. 7 Subsect. Perennantia 
Teeth persistent to falling; ridges convex to bituberculate; 
lants of North America and easternmost Asia. 
. E. hyemale var.affine 
Teeth soon falling; edges always bituberculate; plants o ur- 
pts 1 Wadia bag) .... Ba. EB. hyemale var. hyemale 
Rhizome with an outer or double common endodermis; sheaths 
reen, rarely black above; teeth not ue ce. always re- 
tained : _. .. Subsect. Homocormia 
Sheath with four or more © teeth; stem and rhizome with double 
common endodermis 


ReEsuME or Susgenus HippocHarre 1o7 


Teeth incurved, with narrow white margins or all black. 
E. variegatum var. alaskanum 
Teeth straight, with wide white margins. 
6a. E. variegatum var. variegatum 
Sheath with three teeth; stem and rhizome with outer endoder- 
mis only _ .......1. EH. scirpoides 
N. B.: du teriielaies haprean 1 ane 3, ga anda 5a, 2a and 6a, 
3 and 5b, 4 and 5b, 4 and 6a, and 5a or 5b and 6a are probably 
hybrids, especially if they have abortive spores. Seeming inter- 
mediates are sometimes produced under unusual growth condi- 
tions or as a result of injury. 


LITERATURE CITED 

Baker, J. S., 1887. Handbook of the Fern Allies. London. 

Boerner, C., 1912. Eine Flora fiir das deutsche Volk. Leipzig. 

FarweEu, C. A., 1916. The Genus Hippochaete in North America, North 
of Mexico. Mem. New York Bot. Gard. 6:461—472. 

Manon, I., 1950. Problems of Cytology and Evolution in the Pteridophyta. 
Cambridge University Press 

Mitpr, J., 1867. Monographia Equisetorum. Nova Acta Acad. Leop.-Carol. 
32(2) 


RoTHMALER, W., 1944. Pteridophyten-Studien I. Repert. Spec. Nov. 54: 
2. 


Mexico. Amer. Fern Journ. 1 
1925. Main Lines of Evolution in Equisetum. Ibid. 15:8-12, 


Scnarrner, J. H., 1921. North aii Species of Equisetum North of 
1:65-75. 


w 
a 


——————. 3930a. Diagnostic Analysis and site aes Relationships of 
the Main Groups of ana Tbid. 20:11-18. 
=>. 1990b. Geo: pani Distribution of the Species of Equisetum 
in Relation to Their ste bid. 20:89-106. 
(To oe joi aS 
DepaRTMENT or Botany, UNIVERSITY OF Ruope IsLanp, Krn@s- 
TON, RHopE TSLAND, 


138 AMERICAN FERN JOURNAL 
An Interesting Ecological Niche Involving Salvinia rotundi- 
fo i 


Tuomas A. Hutro 


Early in February of this year I accompanied Dr. Wilbur 
H. Duncan and a group of students from his advanced taxonomy 
class at the University of Georgia on a field trip to Sapelo Island 
off the Georgia coast. When Dr. Dunean extended an invitation 
to take this trip I readily accepted because Sapelo Island is 
privately owned by Mr. R. J. Reynolds and is not open to the 
public. The University of Georgia Institute of Marine Biology, 
which is financed by Mr. Reynolds, is located on the island. 

Sapelo is one of a chain of barrier islands just off the coast 
of Georgia. It is bounded on the east by the Atlantic Ocean and 
on the west by an extensive salt marsh. The island is in view of 
the famous ‘‘Marshes of Glynn’? about which Georgia poet Sid- 
ney Lanier wrote. The vegetation of the area is interesting and 
varied, and so Dr. Duncan is preparing a flora of it. 

During the course of our trip the group visited a large, fresh- 
water lake at the north end of the island to see the water fern 
Salvinia rotundifolia Willd., which Dr. Dunean had collected in 
1956 (Duncan 20,111). The plants were abundant in the water 
near the edge of the lake and I collected a quantity in a plastic 
bag. 

After returning to Athens, I placed the Salvinia in a con- 
tainer of water in a window. It was then that I noticed that 
one plants were covered with green objects which appeared to be 
insects. With a microscope I saw that they were aphids which 
were abundant along the veins of the leaves; they had pene- 
trated the veins with their mouthparts, and were so firmly 
attached that attempts to remove them with forceps proved 
difficult. Cast skins of the aphids were scattered about the sur- 
face of the leaves. 


AMER y FERN IRN 5 
ICAN FerRN JouRNAL VoLuME 51, PLATE 7 


APHIDS FEEDING ON SALVINIA ROTUNDIFOLIA WILLD. 


140 AMERICAN FERN JOURNAL 


While observing the aphids feeding I found an unidentified 
insect of the order Hemiptera feeding on the aphids. This tiny 
predaceous bug apparently kept the prolific aphid population in 
check. Several of these bugs were located, all busily feeding on 
the aphids by piercing their soft bodies and sucking the fluids. 
After consuming an aphid a bug would turn to another aphid 
on the same plant or would stride across the surface of the 
water to a different plant. I did not observe any of the aphids 

ing from plant to plant across the water surface and I 
assume that they were unable to do so. Apparently the aphids 
were at the mercy of the more maneuverable bugs. However, 
considering the rapidity with which aphids are known to repro- 
duce it was obvious that the bugs were quite busy keeping the 
population in check. 

Finding the aphids feeding on Salvinia aroused my curiosity, 
because, to my knowledge aphids do not occur on other ferns. 
For that matter, the ferns as a group seem to be generally free 
of insect pests, although Durand (1949) states that ferns grown 
as house plants may become infested with insects and Boydston 
(1958) mentions that ferns grown from spores may be attacked 
by thrips. It might prove interesting to make a future study of 
the insect pests of: the ferns. 

DEPARTMENT OF ‘Botany, UnIversiry or Groreta, ATHENS, 
GEORGIA. ¢ 


in 


BIBLIOGRAPHY 


Boypston, ey 1958. An Amateur Plants Fern Spores. THIS JOURNAL 
48: 1-18. 


Davipson, TREAT. 1961. Rose Aphids’: Cameras ded the Bizarre World 
of a Garden Pest. National Geographic Mag. 119: 59. 

Duranb, Herpert. 1949. Field Book of Common Ferns. G. P. Putnam’s 
Sons. New York, p. 212 


MercopruM WRIGHTII IN CANADA 141 


The Occurrence of Mecodium wrightii in Canada 
Kunio [watsukt 


I have recently received from Mr. C. V. Morton a notice of 
the occurrence of an interesting filmy fern in British Columbia. 
Complying with my request, he kindly lent me the specimen in 
the United States National Herbarium, and suggested that I 
should prepare a note on it. The specimen examined is labelled 
as Mecodium wrightii, an identification of E. B. Copeland, It 


ada in the 
1957, the locality Shine North end of Dawson Inlet, off Skide- 
a Chaantl, Graham Island, Gui Charlotte Islands, British 
Columbia, on west exposed shady vertical precipice, ca. 10 m.s.m 
There is no question that this filmy fern is Mecodium wrightii 
(v. d. Bosch) Copeland. 

Mecodium wrightii was originally described by van den Bosch? 
on the basis of a specimen collected by C. Wright at Hakodate, 
in Hokkaido. The known range of distribution has been from 
Saghalien? and the southern Kuriles* to Kyushu and southern 
Korea. The headquarters of its distribution lie in the central 
and northern districts of Honshu, usually at altitudes lower 
than 1500 meters. In Shikoku and Kyushu it is found, rarely, at 
500 to 1000 meters elevations. In the regions where M. wrightu 
is most abundantly found, it is usually growing on the bases of 
the trunks of large trees, especially of aged Cryptomeria 
japonica (Pl. 8, 9), in humid dense mountain forests. These 
substrata are in general moist and mossy. In northern Honshu 
and Hokkaido, it grows mostly on mossy rocks or cliffs in shade 
and usually in places with high humidity. Viewed in the vertical 
distribution, it grows, even in Hokkaido, at altitudes higher 
than 1000 meters, but it has never been collected in the alpine 
regions of central Honshu, where M. polyanthos is eokaibaaly 
found. Thus, M. wrighti has more limited habitats and distri- 


' Nederl. Kruidk. Arch. 4: 391. 
é Fomin, Fl. Sib. Or. Extr. 5: ear 1930: 
3 Kunashiri tatand: Kuriles, y. Matsumura (KYO). New to the Kuriles. 


yt {4 p 8 
AMERICAN FERN JOURNAL VOLUME 51, PLATE & 


FOREST OF OF PTOMERIA 
KYOTO, WITH THE SHAKADO TEMPLE AT THE BA 


JAPONICA AT SHAKADO ON Mount een NEAR 


bution than its close relative, M. polyanthos, of world-wide distri- 
bution. 

The collection in British Columbia is, therefore, a remarkable 
range extension. Moreover, it is a notable fact that this is the 
first record of any filmy fern in western North America. Among 
true ferns there is no known example of such a pattern of dis- 
tribution as the occurrence in northern Japan and in western 
Canada. But an example 


is seen in the hepatic Takakia 
lepidozioides S. Hatt. et H. 


Inoue’, an interesting liverwort 
occurring only in the alpine regions of central Honshu, Japan, 
and in British Columbia. Dr. 
British Columbia, 


Persson® first discovered it in 
the same station as Mecodium wrightir. I 
am not certain why they have such distinct gaps in their dis- 
tribution. Contrary to Takakia, which is taxonomically isolated 


4 Journ. Hattori ae foe 19: 137. 1958; 20: 296. 1958. 
» Bryologist 61: 3 


AMERICAN FERN JOURNAL VOLUME 51, PLATE 9 


ee 


= ; 
% vite 
oe 
oo ee 
2 


CLOSE-UP or COLONY OF MecopIUM WRIGHTIL SHOW 


144 AMERICAN F'ERN JOURNAL 


enough to merit a monotypic family or even an order, Mecodiwm 
wrightii is closely allied to the variable cosmopolitian fern M. 
polyanthos. The distinctive features between these two were 
briefly summarized by Copeland® and by Iwatsuki.. As M. 
wrightti is considered to be a direct northern derivative of 
M. polyanthos, it may be that this species became dispersed from 
the Far East through the north coast of the Pacific Ocean 
to western Canada. It may be growing now in the North Pacific 
Islands or along the north coast of the Pacific Ocean, but 
there has been no record as yet. 

The problem of generic delimitation in the Hymenophyllaceae 
has been questioned since the time that Copeland® reclassified 
the family after his extensive study of Old World species. I 
adopted his generic classification wholly in my investigation on 
the species of Japan and the neighbouring regions. This is only 
due to the fact that Trichomanes and Hymenophyllum in the 
older sense are not natural when numerous intermediate species 
groups and characters are taken into account. It is endorsed also 
by recent investigations by cytotaxonomists. However, as the 
taxonomic characters have not been precisely studied from the 
viewpoint of comparative morphology, the evaluation of charac- 
ters for giving the definition of the genera of filmy ferns depends 
chiefly upon the experience of the specialists. For the present, 
I follow conveniently Copeland’s classification, because his sys- 
tem seems to be the most natural one among those given until 
now. 

Tn closing, I wish to express my best thanks to Mr. C. V. Mor- 
ton for his kindness in lending me the Canadian material, and to 
Mr. N. Kitagawa, a specialist in hepatie taxonomy in our insti- 
tute, and Dr. M. Tagawa, who were kind enough to review my 
manuscript. 


DEPARTMENT or Borany, Universtry or Kyoro, Kyoro, JAPAN. 


6 Phil. Journ. Sci. 64: 15 
7 Acta. Phy ba ay Ge sey tH 
8 phil. ‘Journ, Sci. 67; 1, 1938; ily TiL 31, 1947. / 


Notes on POLYSTICHOPSIS 145 


Notes on Polystichopsis 
Grorar R. Procror 


In. a recent paper, C. V. Morton (1) has presented a dis- 
cussion of Ruwmohra and some related fern genera, including 
the small, chiefly West Indian one now to be known as Poly- 
stichopsis. The purpose of the present publication is to offer 
some additional data on this group. 

Mr. ton recognizes 4 species (P. pubescens, P. chaerophyl- 
loides, P. lurida, and P. ochropteroides) only the last of which 
occurs outside the West Indies. However, within the group 
there exist several other West Indian entities, including two 
which I consider valid species, not dealt with by Mr. Morton. 
A re-enumeration of the whole genus is herewith presented. 
POLYSTICHOPSIS PUBESCENS (L.) Morton 

The typical form occurs in Jamaica, Cuba, and Hispaniola. 
On the latter island also occurs a distinct variety, as follows: 
POLYSTICHOPSIS PUBESCENS var. haitiensis (C. Chr.) Proctor, 
comb. nov. 

Dryopteris pubescens var. haitiensis C. Chr. Monogr. Dry. 

II:105. 1920 (Dansk. Vid. Selsk. Skr. VIII. 6(1) :105). 

This variety was based on Picarda nos. 277 & 733. A dup- 
licate of No, 277 at the Institute of Jamaica shows a plant with 
very long-attenuate fronds, and sori bearing large, persistent, 
hairy indusia. The rhizome-scales, not mentioned by Christensen, 
are shorter, relatively broader, and more noticeably clathrate 
than those of typical pubescens from Jamaica. In other respects, 
a close relationship to P. pubescens is obvious. 

Another form of P. pubescens, with somewhat broader, more 
divided blades, occurs in Jamaica. This was confused by Christ- 
ensen (2) with the Lesser Antilles plant now recognized as P. 
muscosa, but it is my belief that they have nothing in common. 


1 Some of the data used in this paper were obtained through field work 
in the Lesser Antilles supported by Grant No. G-4441 from the National 
rise gs Foundation, in cooperation with Dr. R. A. Howard, Arnold Arbore- 

um, Harvard University. 


146 American Fern Journat 


The Jamaican form may not be taxomically distinguishable 

from typical P. pubescens. 

PoLysticHopsis muscosa (Vahl) Proctor, comb. nov. 
Polypodium muscosum Vahl, Eclog. Amer. 3: 1807. 
Aspidium muscosum (Vahl) Swartz, Adnot. Be 68. 1829. 
Phegopteris villosa Fée, Mém. Foug. 11: 53. 1866. 

Dryopteris pubescens var. muscosa (Vahl) C. Chr., Mon. Dry. 
TE 105. 1 
The type of Piliewndiuni muscosum was collected in Montser- 
rat by Ryan. The species has since also been found in Guade- 
loupe, Martinique, and St. Lucia (Proctor 21652) in the Lesser 

Antilles. Christensen cited specimens from Jamaica, Trinidad, 

and Margarita Island (Venezuela), but the Jamaican material, 

at least, is really a larger, more divided form of typical P: 

pubescens, as noted above. The Lesser Antillean population 

differs significantly from P. pubescens by having longer, paler 


with densely pale-villous stipes, and larger sporangia (averag- 
ing ca. 207 p in diameter vs. ca. 175 » for P. pubescens). P. 
muscosa is described as being without an indusium, and this 
seems to be true for much of the material, but some specimens 
from Guadeloupe (Proctor 20384) bear small, pale, suberect 
indusia which are fimbriate at the apex. These are quite dif- 
ferent from the brownish, more or less entire, orbicular-reniform 
indusia of P. pubescens. 
POLYSTICHOPSIS CHAEROPHYLLOIDES (Poir.) Morton 

Christensen recognized a variety sericea (first described as 
Phegopteris sericea in 1860 by Mettenius). Aside from slightly 
greater hairiness, this form seems to have no distinguishing 
characters by which it can be separated from typical chaerophyl- 
loides. The type of var. sericea came from Cuba (Wright 1054), 
and similar hairier forms have also been collected in Hispaniola. 
The type of P. chaerophylloides was collected in Puerto Rico by 
Ledru. This species is known to occur on all four of the Greater 
Antilles. 


Nores ON POLYSTICHOPNIS © 147 


PoLysticHopsis LuRIDA (Underw. & Maxon) Morton 
Typical P. lurida is localized in the Mt. Diablo area of central 
Jamaica, where it has been collected repeatedly. A slightly dif- 
ferent form, orginally described as a distinet species, occurs 
farther west, where it is rare. It can be recognized as a form: 
spar a LURIDA forma leucochaete (Slosson) Proctor, 
comb. n 
Sagara lencohaet Slosson, Bull. Torrey Club 40: 184, pl. 
35 fig. 2 LOLS: 
Dryopteri lurida f. leweochacte (Slosson) C. Chr., Mon. Dry. 
ee | 920. 


Thelyptens lurida f. leucochacte (Slosson) Proctor, Bull. Inst. 
Jam. Sci. Ser. 5: 61. 1953 

POLYSTICHOPSIS OCHROPTEROIDES (Baker) Morton 

No further comments are here offered on this seldom-collected 
species. The type is a Hart specimen from Jamaica 
PotysticHopsis argillicola Proctor, sp. nov. 

Rhizoma breviter repens ca. 0.5 em. erassum apice paleis 
lanceolato- attenuatis integris 0. 5-1 em. longis nitidis rufo-brun- 


deltoidea, plerumque 12— 20 em. longa, basi 9-12 em. lata, tripin- 
nata, chartacea bees) sube coriacea, supra 4 atrovirens, subtus palli- 


ior, t costis pubescentibus, pilis longis et brevibus, 
venulis ultimis Sent subtus sparse setosis. Forma pinnularum 
et segmentor chropteroidet similis. Sori Ee er 


reniformibus tenuibus marginibus subtiliter undulatis 

Rhizome short-creeping, about 0.5 cm. thick, noely ‘alothed 
at apex with glossy, red-brown, translucent, lance-attenuate, 
entire scales 0.5—1 em. long. Stipes 15-25 em. lone, densely soft- 

pilose toward base, otherwise cee and nearly or quite de- 
void of scales. Blades deltoid, “mostly 12-20 em. long, 9-12 em. 
broad at base, tripinnate, chartaceous or subcoriaceous, 
green above, paler beneath; rachis and costae pubescent with 


Spar sely setose beneath. Shape of pete and segments about 
as in P. ochropteroides. Sori covered by large, reniform, thin, 
glabrous indusia with finely undulate margins. 

TYPE; Jamaica; Parish of Portland, on ridge 2 2 miles northeast 


148 AMERICAN FERN JOURNAL 


of High Peak, above Murdocks Gap, on shaded clay banks at 
3900-4500 feet elevation, Proctor 5842, san Sie May 2, 1951. 


Differs from the somewhat similar P. apie SE with 
which it was included in the writer’s ‘‘Checklist’’ (3), by its 
very much smaller size, non-scaly stipes, setose ultimate costules, 
and non-ciliate indusia. 

It is interesting to note that five of the six known species of 
Polystichopsis occur in Jamaica, and that two of them are con- 
fined to that island. Although a relatively minor example, this 
can be added to the impressive list of genera for which Jamaica 
is a center of speciation. 


BLIOGRAPHY 
- Morton, C. V. (1960). Observations on cultivated ferns VI. The ferns 
currently known as Rumohra. Amer 0: 145-155. 
CHRISTENEEN, Cart (1920). A monograph of tie genus Dryopteris. Part 
eh 


a 


to 


= 


penh 
cTOR, GEORGE R, (1953). A discard checklist of vhudanke Pteri- 
Shae. . Bull. Inst. Jam. Sci. Ser ae 


THE INSTITUTE OF JAMAICA, Raneavens: JAMAICA. 
The Genus Grammitis in Japan, with Description of a New 
Species 
Moroz! Tagawa AnD Kunio IwaTsuKI 


The ferns of southern Japan and of the northern Ryukyu 
Islands have recently been studied rather minutely by the 
writers both in the field and by using herbarium specimens, 
including in the course of their investigation the genus Gram- 
maitis. 

In 1954, the senior author published a revision of grammitoid 
ferns in Japan, the Ryukyus, and Taiwan!, in which only one 
species of Grammitis, G. dorsipila? was credited to Japan. At 

1 Ta agawa, Acta Phytotax. Geobot. 15: 182-1 1954, 

Aotirciaeie i Pocgions (Christ) C. Chr. et. Tard, Not. Syst. 8: 179. 1933; 


be abalelie oahiié Ogata, Icon. Fil. Jap. 5: pl. 248, 1933, 


THE GENUS GRAMMITIS IN JAPAN 149 


that time, this species was known to occur rarely from the 
southern extremity of Kyushu and the northern Ryukyus and 
in South China and Cambodia. Since then several collections 
have been added to the herbarium of the University of Kyoto 
from the same regions, where G. dorsipila is fairly common on 
“mossy tree trunks or on mossy rocks near the very summit of 


Figs A-C, HoLtorypr or en puagigeon SP, Nov. A, A WHOLE PLANT, 
A 1D; B, A Spore, X 3 ; C, a Part or FRonD, 

higher mountains (alt. 500-1,000 meters) where there is constant 
moisture from the daily mists. These additional specimens are: 


Kyusnu: Mount Hoyoshi-dake, south of Kanoya-shi, Pref. Kagoshima, at 
900 meters elevation, Tagawa § Iwatsuki 1141. Ryukyu IsLanps: Mount 
Yuwan-dake, Amami-Oshima, at 690 meters elevation, Tagawa § Iwatsuki 
2919, ibid., T. Shin s.n.; Mount Inokawa-dake , Tokunoshima Island, at 640 
meters elevation, Tagawa g° Iwatsuki 2778, ‘Bi d., T, Shin 

In 1950 our associate, Mr. G. Murata, collected an interesting 
Grammitis at Nachi, in Wakayama Prefecture in southern 


150 AMERICAN F'ERN JOURNAL 


Honshu. Recently Prof. S. Hatusima, of Kagoshima University, 
sent a specimen of Grammitis to the writers for determination, 
which was collected by his student, Mr. K. Kawanabe, on the 
island of Yakushima. These two specimens are identical and so 
distinct as to warrant a description of a new species as shown 
below. It is distinet from @. dorsipila by its broader, harsher, 
more strigose lamina, with larger and subcostal sori. The dif- 
ference in the breadth of lamina makes a difference between 
the two species in the relative position of their sori. In addition 
of these features, the two species are ecologically distinct from 
each other, G. dorsipila being found only in a cloud zone near 
the summit of mountains, and the present new species, on the 


podium hirtellum Blume of western Malesia, which is, however, 
distinct from the two Japanese species by its larger lamina with 
more sparse hairs less than 1.5 mm long, and by the smaller 
rhizome scales. 
Grammitis nipponica Tagawa et Iwatsuki, sp. nov. Figs. A-C, 

G. dorsipilae affinis, frondibus ubique setis longioribus ad 
2 mm. longis densiore obsitis distincta ; frondibus 2-3.5 em. longis 
linearibus vel lineari-lanceolatis; venis fere ad marginem pro- 
longatis; soris costalibus, elliptioie in maturitate saepe plus 
minus confluentib 

Rhizome very short, ascending to. almost erect, usually im- 
mersed in mosses on rocks, densely covered with scales at the 
apex; rhizome scales narrowly lanceolate, up to 1.7 mm. long, 
0.7 mm. broad, acuminate to attenuate at the apex, entire, 
glabrous, pale brownish, subclathrate. Stipes terete, short, less 
than 1 em. long, ca. 0.3 mm. in diameter, stramineous, rather 
densely covered with shining, ebeneous-brown setiform hairs 
less than 0.5 mm. long. Leaves tufted, suberect; lamina linear 
or linear-lanceolate, gradually attenuate and long-decurrent 


8 e.g. Polypodium hirtellum sensu Makino, Bot. Mag. Tokyo 15: 60. 1901; 
Phan, Pter. Jap. Te. TM. 1: pl. 87, 1902, 


CONCERNING AZOLLA IMBRICATA 15k 


into the short stipe, (1.5) 2-3.5 (4.5) em. long, 3.5-4.5 (6) mm. 
broad, rounded to moderately acute at the apex, entire or very 
slightly undulate, subcoriaceous and rather thick in texture, 
both surfaces rather densely covered with ebeneous-brown, stiff 
hairs up to 2 mm. long, more densely so on the costa and the 
margin; veins entirely hidden, simple or furcate above the base 
in fertile condition, the acroscopie veinlet very short, the basi- 
scopic veinlet nearly reaching the margin. Sori elliptic, dorsal on 
simple veins or on anterior veinlets, arranged in a row close to 
costa, usually more or less confluent at maturity; sporangia 
naked or bearing a long seta; spores nearly spherical, trilete, 
minutely tuberculate. 

Honsuvu. San-no-taki Falls, at Nachi, Pref. Wakayama, on 
riverside mossy rocks in constant spray of a waterfall in dense 
mountain forests at about 400 meters elevation, G. Nakai 5052 
(type KYO), ibid., Iwatsuki 2510 (KYO). Kyusuu. Along the 
Suzu-kawa, Yakushima Island, in thickets at about 50 meters 
elevation, Kawanabe 4991(KAG). 

DEPARTMENT OF BoraNy, University or Kyoro, Kyoro, JAPAN. 


Concerning Azolla imbricata 
Eugene Yu-FENG SHEN 

Azolla is a very common fern in Eastern Asia. Diels (1901) 
listed the plants from Central China (Hupeh) that were col- 
lected by Henry as Azolla pinnata R. Brown, and Matthew 
(1911) in his Enumeration of Chinese Ferns used the same 
name. Dunn and Tutcher (1912) called the plant from Hong- 
kong Azolla caroliniana Willd. In 1925 Nakai suggested that the 
correct name for the fern that is found from India to Japan 
is Azolla imbricata (Roxb.) Nakai, stating that it is a species 
resembling Azolla africana, differing from that by the shorter 
papillae of the leaves and less pointed leaf-apex. Ching (1933) 
accepted Nakai’s name and used it in his list of the Pteridophyta 
of Kiangsu, as did DeVol (1945) in his Ferns and Fern Allies 
of Eastern Central China, and most botanists have been calling 
this fern by this name since that time. However, we have often 


AMERICAN FERN JOURNAL VoLtuME 51, PLATE 10 


Ia. 1, A Microsporocare (MI) AND MecasporocarP (MA); 2, MEGASPO 
CARP WITH WALL REMOVED, eyes FUNNEL- LIKE CILIA (cy, 'FLoats (F), 
AND psecieng vb (M); 3 3, MICROSPORANGIL 7, CONTAINING MASSULAE; 

ASSULA, VENTRAL View; 5, Massuua, Sipe View, witH VACUOLATED 
TRICHOMES CT) $6, Seovion or Mra ASPOROCARP, WITH Mr@asPor ROCARP WALL 
(C), ZONE BETWEEN MEGASPOROCARP WALL AND MeGasporancium (Z), 
Ouver beng OF MEGASPORANGIUM WALL (T), with VERMIFORM PAP Res 
(P), INNER, Nog: CELLULAR LAYER OF MEGASPORANGIUM WALL (N), ANP 
MEG capaalts Watt (8). 


CONCERNING AZOLLA IMBRICATA 1538 


wondered about the status of this species, for the real distine- 
tion between species of Azolla rests on the nature of the massulae 
and structure of the megasporangium and not merely on the 
papillae or shape of the leaves. So Charles DeVol suggested 
that we make a detailed study of the fruiting bodies of this 
aquatic fern with the object of finding out whether A. imbricata 
is really distinet from A. pinnata or not. 

Genkei Masamune (1936) in his ‘‘Short Flora of Formosa”’ 
had called this plant A. japonica Fr. et Sav., which was an 
error in determination, for true A. japonica has never been 
collected in Taiwan. In Masamune’s 1954 revision of his list of 
Taiwan plants he lists two species A. africana and A. imbricata. 
It has been thought that A. africana from Madagascar and many 
parts of Africa was distinct from A. pinnata of Australasia, but 
Christensen (1906) was correct in reducing it to a synonym. 
Azolla kiangsiensis is also no doubt the same fern. Specimens 
collected by L. H. Bailey in Kiukiang, Kiangsi, now in the U.S. 
National Herbarium are small sterile plants which appear the 


ul alu 
structure of the sporocarps of Azolla africana andl Azolla im- 
bricata have not been previously reported. 

Vegetative characters alone cannot be used as the sole criteria 
to distinguish the species of Azolla. It is more correct and reli- 
able to determine the species by using their fruiting structures. 
The morphology of the massula and its trichomes, the structure 
and the contents of megasporocarp, including the character of 
its ciliated apex, the number of floats, the presence or absence 
of a collar, and the structure of the megasporangial wall are 
all important. 

Last year the writer had a chance to get fruiting specimens 
of Azolla from Nanching, Chiayi Hsien. By carefully studying 
their morphological structures he found them to be similar to 
Strasburger’s drawings of Azolla pinnata. We do not know 
where Strasburger me his material but his drawing shows it to 
be from a plant like ours. Both have massulae with trichomes 


154 AMERICAN FERN JOURNAL 
attached on one surface and without glochidia. The anastomosing 


drawing of Azolla pinnata may be due to the over-lapping of 
some of the endings of the trichomes. The structures contained 
within the megasporocarp and the vermiform papillae on the 
wall of the megasporangium shown in the Strasburger’s drawing 
are also similar to ours, and the shape of the massulae appears 
similar to our species. Therefore, it seems safe to conclude that 
these taxa may be just a single species, and if so, then the correct 
name for the fern from Eastern Asia should be Azolla pinnata 
R. Brown. Specimens received from Australia sent by Ima G. 
Stone and from Singapore sent by Hsuan Keng have been 
examined and found to be similar to ours. 


A DESCRIPTION OF THE IMPORTANT STRUCTURES 

The plants found in Taiwan are triangular in shape. They 
fragment easily, and therefore vegetative multiplication is very 
rapid. The leaves are alternate and stand on the dorsal surface 
of the rhizome in two rows. They are each divided into a dorsal 
aerial and a ventral submersed lobe. The dorsal lobe is several 
cells thick in the central region and is photosynthetic, with 
papillae on the upper surface. The ventral lobe is thin, one cell 
layer thick through most of its extent, and is non-photosynthetic. 
The leaf-lobes are broader near the apex, being trapezoidal and 
1.2 to 1.4 mm long. Unbranched roots on the lower side of the 
plants extend a short distance into the water. They are enveloped 
with a sheath and cap when young. The sheath and cap are 
sloughed off and the root hairs spread out when the root is fully 
mature. 

The sporocarps containing microsporangia or megasporangia 
are different in size and shape. e microsporocarp is large and 
globular and the megasporocarp is smaller and ovoid in shape 
(Pl. 10, Fig. 1). The wall of the sporocarp, which is the indusium, 
is two cells in thickness with an opening at the apex. Within the 
microsporocarp are many long-stalked spherical microsporangia 
(Fig. 3) borne laterally on a receptacle. The order of develop- 
ment of the sporangia within the sorus is gradate. There are 


CONCERNING AZOLLA IMBRICATA 155 
four or more ‘‘massulae’’ in each microsporangium. The shape 
of the massulae of our species is almost like a hat with a dorsal 
and ventral face; it is not isodiametric (Figs. 4, 5). Three to 
eight trichomes hang from the ventral side, these not being 
glochidia-like. The contents of the trichomes are highly vacuolate 
and thus may appear to be septate, but they are not branching 
and not anastomosing. The microspores are 13-18 » in diameter. 

Embedded in the megasporocarp wall at the base is a spheri- 
cal megasporangium, above which are nine floats arranged in two 
tiers, with three floats above and six floats below. There is no 
collar between the spore and floats. Above the floats there is a 
funnel-like tuft of long cilia. A single large megaspore is formed 
within the megasporangium (Fig. 2). The wall of the mega- 
Sporangium of our species has two layers, just like that illus- 
strated by Strasburger (Strasburger, Fig. 103 b) ; the inner layer 
is non-cellular and appears homogeneous; the outer layer is 
finely tuberculate and covered with scattered vermiform papillae. 
The megaspore wall has close striations (Fig. 6). 


LITERATURE CITED 
CHING, ey C. (1933). The Pteridophytes of Kiangsu Province. Sinensia 
348, 


iaverece (1906). Index Filicum. Copenhag 

DEVo1, C. E 945). Ferns and Fern Allies of pape Central Chin 

DIELS, oe ce Die Flora von Central-China. Bot. Jahrb. Semel 29: 
169-6 


Dunn, ss T. tener W. J. (1912). Flora of Kwangtung and Hong 
Kong ous. Lond 
Masamung, G. (1936). Sad Flora of Formos 
Masamune, G. (1954). A List of Vascular Plant ts of Taiwan. 
Marrnew, C. G. (1911). Enumeration of Chinese Ferns, Journ, Linn, Soe. 
39: -393 
aioe ea (1925). Notes on Japanese Ferns, II. Bot. Mag. Tokyo 39: 
183-18 
SHEN, ts P, _ Anabaena azollae and its host Azolla pinnata, Tai- 
wania 7 
STRASBURGER, “ (1873). Ueber Azolla. Jena. 
DEPARTMENT OF Borany, NaTIonaL TAIWAN UNIVERSITY, 
Tarper, TaAIwan, CHINA 


156 AMERICAN FERN JOURNAL 


Shorter Notes 
_Awn AppITION To THE List oF FERNS Founp Grow1nG NATURAL- 
LY IN TENNESSEE.—During the Annual Wildflower Pilgrimage to 
the Great Smoky Mountains National Park, in the third week 
of April, 1957, a fern was discovered along the Huskey Gap 
Trail by Dr. Frank Barclay and the writer. This specimen 
could not be readily identified in the field, but upon further 
study proved to be Botrychium matricariifolium A. Br. An un- 
usual feature of this plant was that a few sporangia had de- 
veloped on the sterile frond. A specimen has been deposited in 
the Herbarium of the Great Smoky Mountain National Park. 

A detailed description of this plant is given in Gray’s Manual 
of Botany. Its range is given as extending from Newfoundland 
south to West Virginia. Westward, it is found in Michigan, 
Wisconsin, and Idaho. No mention of this species is to be found 
in any of the excellent works dealing with the flora of Tennessee, 
such as those of Anderson and Shaver. This find then represents 
a new addition to the ferns of Tennessee as well as to the unique 
flora of the Great Smokies. 

The southernmost stations for this plant previously known 
were in Maryland and West Virginia. Professor Ogden, of Alfred 
pepo New York, tells me that he has collected specimens 

n the mountains of West Virginia. — Herman O’DeELL, Hast 
Pnionis State College, Johnson City, Tennessee. 

OxitruaryY : ALBERTO CurARuGI, 1901-1960.—The botanical world 
suffered a loss with the death of Alberto Chiarugi on February 
25, 1960. Founder of the serial ‘‘Caryologia,’’ he directed its 
publication for 12 years. His wife Emilia is now undertaking 
the continuation of the publication. Botanists will be interested 
in perusing volume 13, no. 1, of ‘‘Caryologia’’ for a list of Dr. 
Chiarugi’s publications, which number 174 and attest to a long 
and active career. Of particular interest in this volume and 
number is a posthumous article on ferns entitled, “«Mavole 
Cromosomiche Delle Pteridophyta,” in which paper of 123 pages 
is discussed the cytological situation in the pteridophytes, with 
lists of all known chromosome numbers in this group. The list 


Norrs aNp News 157. 


of numbers is by far the most complete attempt existing, and 
lacks only the more recent papers (such as an article by Mehra 
and Loyal 1959 in which Marsilea brachypus is listed as having 
n=20 and a new number of n=20 is given for M. quadrifolia). 
Since accurate chromosome counts in ferns with numerous 
chromosomes is only possible with the squash technique, it is 
necessary to evaluate individually counts made prior to 1950. 
Dr. Chiarugi’s list will serve as an important reference point 
for many years to come.—Irvine W. Knosiocn, Michigan State 
Umiversity, East Lansing, Michigan. 


News and Notes 


A Fern Progect Wins a ScreNcE Farr Awarp.—In the 
Plainfield, New Jersey Courier-News for March 31, 1961, there 
was published a photograph of a young lady receiving a certifi- 
cate, with the caption: 

“Her Ferns Are Tops—Emily Anne Carver, a junior at the 
Watchung Hills Regional High School, in Warren Township, 
received an award last night for the best project submitted by 
a girl in the Central New Jersey Science Fair at Rutgers Uni- 
versity.’’ 

The text notes that as a grand prize winner she will enter 
the National Science Fair in Kansas City May 8 and 9. Miss 
Carver’s project consisted of a study of the ferns and fern 
allies of Somerset County, New Jersey, listing for each the 
known localities, the geological formation, the climate, the soil 
acidity, ete—E. T. W 

The Society for Economic Botany is interested in stimulating 
scientific studies of plants useful to man, by annual meetings, 
and the publication of ‘‘ Economic Botany,’’ the official journal 
of the Society, which publishes monographs, review articles and 
original studies on economic plants. The Society welcomes new 
members (annual dues, including subscription to journal, $7.50). 
Inquiries to Richard M. Klein, New York Botanical Garden, 
Bronx Park, New York 58, N. Y. 


158 AMERICAN FERN JOURNAL 


A Mass Couuection or PotysticHum.—TIwo members of the 
Fern Society, Mr. Harry K. Roberts and Mr. Donald L. Brans- 
comb, of Guerneville and Willits, California, respectively, have 
made mass collections of Polystichums native in the California 
Coast Ranges between Santa Cruz and the Oregon state line. 
Fronds were taken from populations of P. muwnitum, P. cali- 
fornicum, and P. Dudleyi. The specimens exhibit amazing varia- 
tions and intergradations, and suggest that these ‘‘species’’ may 
be a polymorhiec complex within which introgression occurs 
freely. 

Several hundred unmounted fronds, ranging from 3 deci- 
meters to 1.5 meters in length, are stored at the Dudley Her- 
barium, Stanford University. Anyone interested in studying 
this raaterial inquire of the Editor—I. L. W. 

OnocLea Spores AVAILABLE :—Dr. Ralph C. Benedict, Pilot 
Knob, New York, has a quantity of fresh spores of the sensitive 
fern, Onoclea sensibilis, available, if college teachers or others 
wish them for demonstrations or laboratory work. 


American Fern Society 


New MempBers 
Mrs. Paul Bartsch, Gunston Hall Road, Lorton, Va. 
Bruce H. Beeler, La Casa Verde, Rt. 1, Box 40, Homestead, Florida 


Marguerite Crookes, 6 Coronation Rd., Auckland, S. E. 3, New Zealand 
Mrs. Lois D. Cross, P. O. Box 1107, Washington 13, D. C. 

Mrs. Stuart Dunlap, 5618 Forest Blvd., Spokane 15, Wash. 

Mrs. Robert B. Elliott, 218 Pine St., Seaford, Del. 


AMERICAN FERN Society 159 


James Fiore, 1448 Joyce ape Seaford, N. Y. 
Mrs. Robert J. Fisher, Rt. 3, Box 206, Lokipnville, Ohi 
Dr. Chas. K. Gardner, M.D., Gis Charlotte Ave. | Nashville 9, Tenn. 


Mrs. Katherine Goodpasture, 3407 ins St., Nashville, Tenn 
T. Hammett, 1401 Rutledge Ave., Charlo 
Mrs. John R. Hart, 760-39 mond, Cali 


W. H. Hastings, 701 W. 7th Ave., Corsicana, Texas 
Teofilo Herrera, Instituto de a Ciudad Universitaria, Apartado postal 
29817, Mexico 18, D. F., Mex 


Mrs. tater E. House, 934 Scenic Deis , 8.W., Knoxville, Tenn 

Ralph S. Hudson, Shadies Lake, bisver, Uelowns County, Vermont 

Dr. Hirosi Ito, e/o Botanical Institute, Faculty of Science, Sa Univ. of 
Education, Otsuka-Kubomachi, Bunkyo-ku, Tokyo, Japan 

Mrs. J. E. Jackson, 108 Highland Ave., Bastrop, Louisiana 

Mrs. Dewey Janet, Sr., 701-89th Ave. W., St. Petersburg iy Florida 

George L. Kirk, 8 East Washington St., Rutland, ve 

Mrs. Johannas C. Klomp, R. D. 7, Saltsman Rd., Erie, a 

Mr. Dick Korda, Jr., 1640 E. 243 Street, Euclid 17, Ohio 

Dr. Satoru Kurata, c/o Inst. of Forest-Botany, Faculty of Agriculture, 


Dr. Thomas P. McCutcheon, Smoke-house — owningtown, given 
Thomas a 1501 N.W. 29 Ave., M 

Miss Cora Miller, 513 E. Allegheny Ane, opi Be ia ae 

Mrs, Vera .— Beech Tree Rd., Brookfield Center, om 
Mrs. Merril Niebel, R. F. D. No. 3, Napoleo: “Oh 

Mrs. R. Olivarez, 3024 Castro Street, San pa 12, Calif. 

W.S. Parks, Jr., 306 South Main, Pasadena, Tex 

Mrs. Paula Peck, 552 Riverside Drive, New ‘otk oe NOW 

Mrs. Harold O. Perkins, Dog Lane, Storrs, Con 

Miss Penny Pretzinger, 549 E. Cypress Ave., eens: Calif. 

Fred Rickson, 4554 Lemp Avenue, North on 4 Calif. 

Miss Jennie Riggs, 2005 Capers Ave, Nashville, n. 

Mrs. John Riker, Tabor Road, Morris Plains, ew Soles 

Mrs. Donald Russell, ae Otis Bl wr a anburg, South Carolina 
Stephen Spongberg, 1 East State Stet, Rockford, Illinois 
Mrs. C. C. Stevens, 14 tal liwood Road, Roslyn, N. 

Mrs. Dan J. Sullivan, 2556 Woodbour ne ve.,  boaibe, Kentucky 
Mrs. Olean T. Tatro, 7512 Elphick Seal, an Calif. 

Earle W. Tie 1904 Farris Ave., Fresn 

Mrs. Theodore Thompson, 935 Esplanade, ‘Plan, i ae ge 


160 AMERICAN FERN JOURNAL 


mehr Jesse T. Vogdes, Lemon St., No. of 5th St., Media, Penna. 
tuart W. Webb, Stonington, Conn 

eset R. Webe vee Dept. of Biology and Geology, Otterbein College, 
Westerville, Ohi 

Mrs. E. S. Webster, Hes Hill Road, Mt. Tabor, New Jersey 


Mr. Victor F. Wright, 7481-18th Ave., Burnaby 3, B. C., Canada 
Dr. M. R. Zingeser, 502 Portland Medical Center, Portland 5, Oregon 


CHANGES OF ADDRESS 
Don S. Caldwell, Jr., P. O. Box 1247, Odessa, Texas 
Mrs. E. M. Carroll, 1171 Belle Road, Paradise, Calif. 
Mrs. Kenneth L. eake a! Orangegrove, Orange, Calif. 
Mr. Orville W. wder, 2247, Washington 13, D. C. 
Dr. Pierre eae te Yoh Botanical Gistdien:” Bronx Park, New York 


Prof. Deda W. Dunlop, University of Wisconsin at Milwaukee, 3203 IN: 
Downer Ave., Milwaukee 11, Wis. 
Mr. Sidney K. Eastwood, 526 Madison St., New Orleans, Louisiana 


. ntu 
H. William Johansen, gee A. Woolsey, Berkeley 5, Calif. 
Walter F. Kleinschmidt, 1800 N. Dixboro Road, Ann Arbor, Mich. 
Hal M. Moseley, 6664 Avalon, Dallas 14, Tex: 
Maria T. Murillo Pulido, Apartado Numero sad Bogota, Colombia 
Ray A. Peters, 5271 Sereno Drive, Temple Cit rere 
awd 


Calioce, na Luis pee Calif 


Mr. Cedric Sidney, P. O. Box 53, Natalbany, os 

Dr. Harry N. Stoudt, Science Dept., Glassboro = 2 es Glassboro, N. J. 

Edmund T, Turnau, Department of Botany, University ef Malaya in Kuala 
Lumpur, Pantai Valley, Kuala Lumpur, Malaya 


HENRY TRIPP 
Successor to Henry George Fiedler 


Scientific Books and Periodicals 


Large stock of books on ferns 
and cryptogamic botany 


31 East 10th Street, New York 3, N. Y. 


CASTANEA 
Published by the 
Southern Appalachian 

Botanical Club 
Devoted to the botany of the interesting Southern Appalachians. 
Published pcm Gs arma subscription, including membership 

in the Club, $4.0: 
Address 


Dr. EARL L. Corr, Editor 
West Virginia University 
neg West Virginia 


FERNS 
SPECIALISTS IN TROPICAL AND SUBTROPICAL FERNS 
We ship Price List Available 


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2637 North Lee Avenue South El Monte, Calif. 


~vace € ad 
THE PREPARATION 
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of this 
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“THE PREPARATION OF 
BOTANICAL SPECIMENS 
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generously , oe e benefits of his long and successful experience in the 
collection and Srapacaiien of plant specimens. 


Help for the amateur — and Larges eo the Agee sgg =e» collector, 
abound in this thirty-six p: ted tri which Dr. Johnston 
describes time-tested roth arr for scabies, gp te een and mounting 
herbarium orn sagye Special — poietic pettnek for treatment of aquatic 
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One copy of the treatise is yours for the asking. With it will be in- 
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Vou, 51 OcToBER-DECEMBER, 1961 No. 4 


American Pern FJournal 
A QUARTERLY DEVOTED TO FERNS 


AMERICAN FERN SOCIETY 
ts 


IRA L. WIGGINS 
Cc. V. MORTON ROLLA M. TRYON, JR. 
JOHN H. THOMAS 


is 
CONTENTS 


The Fern Valley at the United National Arboretum 
EpitH BirTincer 161 


Ferns in Cultivation, IV. Some Wall Ferns ........ SyniviA LEATHERMAN 174 


Southern Distribution of Botrychium oneidense and B. multifidum 
F. R. Fossrere 175 


Dryopteris < Tavelii in ~ Valley of Chamonix, Fran 
André and Anne-Marie ae CoLLARIS 180 


Basket Ferns for Southern California .................++ Fay MacFappEn 186 
Shorter Notes: Dennstaedtia globulifera (Poir.) Hieron. in Texas; 


Pteridological Troglodytes; Marsilea quadrifolia L. in Indiana 
Again 188 


Notes and News: Letter on Fluorescent Tubes 191 


Eggo Fern Society: Report on the spaegenrese National Forest 
Fora 192 


A Rr “80; [Aa 
Index to Volume 51 Coens. Go Bad eg vss 196 


/ 
( JAN 2 9 1989 


The American Fern Society 


Council for 1961 
OFFICERS FOR THE YEAR 


Cua A. Brown, Department of Botany, Louisiana State nas Baton 
Rouge, Louisiana resident 
ARCEL RAYMOND, ee Botanical Garden, 4101 East SES Street, 


gg ty Cana Vice-Pr esident 
DonaLp cen Longwood Gardens, Kennett Square, Penna. 
Secretary 


Watrter 8. Pumutes, Department of Botany, University of — Tucson, 
Ariz pat 


Ina L. Wiacins, Dudley Herbarium, Stanford University, Stanford, . 
Editor- ae 


OFFICIAL ORGAN 
American Fern Journal 


EDITORS 
Tra L. a aes Dudley Herbarium, Stanford University, Stanford, Calif. 
©. V. Moeroy. .__Smithsonian Institution, Washington 25, D. C. 


Roa M. mao 
Gray Herbarium, Harvard University, Cambridge 38, Mass. 
Joun H. Saowan Dudley Herbarium, Stanford University, Stanford, Calif. 


An illustrated quarterly devoted to the general study of ferns, published 
quarterly by bus American Fern Socie age 3110 Elm Ave., Baltimore 11, Md. 
Second-class postage paid at Baltimore, 

Matter pg petieaton should be cies ssed to Ira L. Wiggins, Natural 
History Museum, Stanford Ragin Stanford, California. 

Subscription $2. ~ exclusive of agency handling fees; sent free to mem- 
bers of the ern ‘Becht (annual dues, $2.50; sustaining mem- 

ip, $5.00; life m embership, $50.00). —— eg a ordered in 


a 

Back volumes $2.00 each; single back numbers 50 cents each; Cumulative 
pongo to vols. 1-25, 25 cents. Ten per cent discount on orders of six volumes 

Changes of address should ec" sent to ~ Secretary: Dr. Donald Huttle- 
ston, Longwood Gardens, Kenn vapour Members and 
seribers should allow two om for cae te 

dg one for membership, subscriptions, orders for back. numbers, and 
other business communications should be addressed to the Treasurer: Dr. 
alee 8. Phillips, Department of Botany, University of Arizona, Tucson, 


LIBRARIAN AND CURATOR OF THE HERBARIUM 
Dr. W. H. Wagner, University of Michigan, Ann Arbor, Mich. 
rtment is maintained in ectio 


A regular loan department ed in conn m wil library 
and herbarium. Members may boo! specimens at any 
the borrower paying or express = The of the J sg 


American Fern Fournal 


Vou. 51 OctToBER-DECEMBER, 1961 No. 4 


The Fern Valley at the United States National Arboretum 
EpitH BirrinGeR 


“The U. 8S. National Arboretum was established by Act of Con- 
gress on March 4, 1927. The purpose of the Arboretum is to 
conduct research with woody plants, and to further public edu- 
cation with respect to trees and shrubs peed of cultivation 
in the climate prevailing in the Washington, D. 
strategie location of the Arboretum enables it ie serve aS a nas 
tional institution in a very real sense. It is in an intermediate 
Ea ceey nee enabling the cultivation of plants from a wide range 
of habita 

The rege consists of 400 acres in the northeast section 
of the District of Columbia.2 Before the government acquired it 
for an Arboretum it was rolling farm land with two hills. One 
hill, named Mount Hamilton, is the site of the Arboretum’s mag- 
nificent collection of azaleas. In the midst of the fields was a 
valley, undoubtedly called a gulch or a gully, of no use except 
as a dump for farm trash. It was a nuisance, a useless piece of 
land, so fortunately for its later use, the trees were not cut nor 
the gully filled. For this we are eternally grateful. This is Fern 


Fern Valley is 900 feet long and 150 feet wide, and has an 
area of about four acres. It runs east and west, and has a spring- 


1 Quoted from a folder on a nega Arboretum published by the United 


States Department of Agric 
2 Directions: From the Sone ecehoaat on Maryland Avenue to its termi- 
nus, the main entrance sr wp Arboretum. Open Monday through Friday 
a.m. ends mi ay ao - mid- es 10 a.m. to 7 p.m. 
and late Oetaber to ety pet 10 a o 5p 


Vol. 51, No. 3, of the JourNAL, pp. 113-160, was issued October 4, 1961. 


162 AMERICAN F'ERN JOURNAL 


fed rivulet winding down the center. It is pleasantly wooded 
with magnificent old tulip, beech, oak, hickory, and some serub- 
pine trees. The soil is a thick layer of leaf mold on a rather acid 
clay loam. Since it drains the surrounding area, the brooklet is 
subject to quick rises, sometimes as much as three feet during 
the flash summer rains, and storm debris was deposited above an 


US DeranrwanT oF AGRICULTURE 
NATIONAL ARBORETUM 
ey WARMNOTON, De 
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See 
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Fig. 1. Map or Tue Unrrep States NarionaL ARBORETUM 


unsightly six-foot sewer pipe which crosses the valley. At the 
lower end was a swampy area, where the ground was covered 
with Japanese honeysuckle intertwined with poison ivy. This 
was Fern Valley as I first saw it in 1957. 

In 1950 Dr. James W. Johnston, Jr., Professor of Physiology 
at Georgetown University, visited the Matoual Arboretum with 
Mr. Oliver M. Freeman, then Botanist of the Arboretum and 
author of “Annotated List of Plants Growing at the National 
Arboretum,” as a guide. At that time Mr. Freeman pointed out 
the valley. Dr. Johnston had been interested in ferns for many 
years, and with no place to plant them himself he thought of the 


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164 AMERICAN F'ERN JOURNAL 


Arboretum valley as a possible fern garden. In 1953 he met with 
Mr. Matthew Mann and Dr. Ralph C. Benedict, then Treasurer 
and President respectively of the American Fern Society, and 
suggested that the American Fern Society start a fern collection 
at the National Arboretum. As a result of this meeting Dr. Ben- 
edict authorized Dr. Johnston to seek a conference with Dr. 
Skinner, Director of the Arboretum, who approved of the pro- 
posed garden. Thus the Fern Society could make a start at plant- 
ing under a special committee with Dr. Johnston as chairman. 
I quote from Dr. Johnston’s notes: 

“May, 1953. Dr. Johnston discussed the project with Dr. Skin- 
ner and Dr. de Vos, Assistant Director of the Arboretum. They 
approved the use of the valley. ... It was planned initially as 
a series of pilot plantings since intensive garden culture was not 
possible then. .. . Three species of ferns occur naturally in the 
vall 


“ce 


Mann, Jr. Dr. R. C. Benedict and Dr. Edgar T. Wherry also 
gave ferns and advice. 

“October, 1953. Dr. Benedict wrote a letter to Dr. Skinner on 
behalf of the American Fern Society expressing appreciation for 
the cordial welcome given the fern project. 

“May, 1954. About 20 members of the American Fern Society 
and friends met at Fern Valley, where they were greeted by the 
Arboretum Staff and taken on a tour of the area by Dr. Johnston. 

“May 28, 1954. Mr. James Benedict led several local members 
of the Fern Society along the Virginia Side of the Potomac. The 
purpose was to collect the more abundant species for planting in 
the garden. 

“The pilot plantings at Fern Valley were five to ten plants 
that were put in prepared pockets. The adjacent weeds and 
honeysuckle were eliminated and the ferns entrusted to nature. 
Dr. Johnston built a non-caleareous rock wall for the smaller 
species. The late summers of 1953 and 1954 were very hot and 


FERN VALLEY 165 


dry, so some of the plantings failed. By 1955 it was clear that 
the initial phase was ended, and that what was needed was in- 
tensive garden culture and vigorous volunteer labor before any 
serious plantings were carried out. 

“Partial success was obtained, however, in the cases of the 
fine red-dotted shield ferns of Japan given by Matthew D. Mann, 
Jr., the goldie ferns from Cyrus Darling, and the Leedsii ferns 
from Dr. Edgar Wherry, which are still growing well where they 
have been for five or six years.? 


The list of Dr. Johnston’s ferns is as follows: 


10 Lady ferns 2 Ebony spleenworts 5 D. intermedia 

9 Hay-scented ferns 15 Walking ferns 1 D. cristata 

14 Maidenhair ferns 3 Cinnamon ferns 1 D. Boottii 

15 Polypodies 8 Oak ferns 1 D. spinulosa 

11 Resurrection ferns 6 New York fern ID; Seep sia 
ao 3 Dryopteris eaptaiesie 1 Ostrich fer 


During the winter of 1957 and 1958 the National Capital Gar- 
den Club League? sponsored a course in Tree Identification. This 
course was held at the National Arboretum with Dr. de Vos as 
lecturer, and the class walked all over the Arboretum on field- 
trip demonstrations. At the last lesson I asked, by chance, if 
there was a fern collection at the Arboretum, so Dr. de Vos too 
us into the bes they hoped to develop into a fern and wild 
flower sanctua 

T later waked Dr. Skinner and Dr. de Vos if they would like 
to have ferns sent in, and then learned of the previous planting 
undertaken by the American Fern Society, by that time inactive. 


3In the summer = 1959 I received a c. from Mrs. Donnald, our plant- 
3 Sear tien reporting progress. She e: “When I checke a = lower 
end of Fern Va ey yeste road : found That Dr. Johnston had been much 
oo "successful n he thou . I found over two dozen individual 
ferns growing ri 

ie National Capital eon Club League is to the greater ca araged 

a of Washington, D. C., t a State Federation of Garden Clubs is 

pen State. Thee, seo a mem Ree clubs, =, Sd one-third each sue 
Maryland, Virginia, and the District of Columbia 


166 AMERICAN FERN JOURNAL 


The Directors thought nothing should be attempted without con- 
sulting the Fern Society and gave me Dr. Benedict’s address. I 
wrote to him and quote from his reply: 

“May 26, 1958. I am delighted to know of your interest in 
developing representative fern plantings at the National Ar- 
boretum. The Fern Society made a start at such planting just 
about four years ago, after consultation with authorities. A 
special committee, with Prof. James Johnston, of Georgetown 
University, as chairman was appointed. . . . I recommend that 
you get in touch with Dr. Johnston and with Mr. James Bene- 
dict, Vice-President of the Fern Society at this time. . . . These 
suggestions are in no way intended to indicate any ior claim 
or rights in the activity for the Fern Society; rather to express 


will be ready to aid in further steps for expanding the plantings.” 

Dr. Benedict’s reply reached me after I had left Washington 
for the summer, and I was unable to contact Dr. Johnston and 
Mr. Benedict until autumn, but on the basis of Dr. Benedict’s 
letter, I began a summer project of collecting ferns for the Ar- 
boretum. My plan was to make a basic collection of our common 
native ferns and such rarer ones as I could find. Knowing from 
experience how inadequate a planting of only three or four ferns 
of a kind is, and also aware of the characteristics of the area to 
be planted, I decided on units of 25 specimens of a species if 
possible. To this basic collection more could be added as needed, 
and rare ferns provided later by gift, purchases, or field-work. 

I had much plasant cooperation in my project. My sister, Miss 
Mary Gay, a friend, Miss Irene Walker, and I, long addicted to 
fern-hunting, searched our own countryside, eastern Massa- 
chusetts, and made trips to western Massachusetts, New Hamp- 
shire, and Vermont. In the latter state we discovered, sadly but 
approvingly, that a law had just been passed prohibiting the 
removal of some rarer ferns. The late Mr. Donald Wyman, of 
Bay State Nurseries, allowed me his professional discount in 


FERN VALLEY 167 


purchasing ferns; Mr. Abbey, of Gardenside Nursery in Shel- 
burne, Vermont el for me two native Scolopendriums 


from a friend in New York; my own town Tree Warden, Mr. 


Roy Parks gave me a double handful of Lzbelia cardinals seeds, 
wich gave us beautiful blooms in the summer of 1960. 

In October, 1958, I sent to the Arboretum nearly 1,000 ferns 
of 88 species. I did not include any of our weedy ferns—bracken, 


hay-scented fern, and sensitive fern, and I made no effort to— 


collect Christmas ferns, Woodsia obtusa, and ebony spleenworts, 
which grow in quantity in the Washington area. My ferns were 
heeled in at the Arboretum as they arrived. A list of the species 
sent to the National Arboretum is: 


List or Ferns SENT TO ARBORETUM IN OCTOBER, 1958 


150 Adiantum pedat 25 Gymnocarpium gi amb 
25 Asplenium Trikina 25 Lygodium palm 

25 Athyrium — Femina (upland) 25 Osmunda Listens 

25 A. Filix-femina (lowland) 25 O. Claytoniana 

30 O. regalis 

25 A. thelypt 25 Pellaea atropurpurea 

2 ptosorus ences daa 15 Polystichum Braunii 

25 Cystopteris bulbifer 30 Polypodium Sela 

20 C. fragilis 25 P. polypodi 

25 Dryopteris Boottii 25 elaine ‘Strutniopteri 
25 D. Clintoniana 10 Scolop 

25 D. cristata ree 8 English) 
25 'D, Filix-mas 25 Thelypteris he cagonoptera 
25 D. Goldi 25 T. Phegopte 

25 D. intermedia 25 T. sept ad 

25 D. marginalis 25 

25 D. spinulosa 25 Woodsia ilve 

15 D. dilatata 25 Woodwardia « pp 


Up to:this time it was a “one man project,” but in the sum- 
mer of 1958 the Board of Trustees of the National Capital Gar- 
den Club League voted to make Fern Valley a League project.> 
To bring it to general attention, a fern screen for exhibition at 
garden shows, was made by Mrs. Donnald and Mrs. Grant. 

Dr. de Vos expressed a wish that a committee be appointed 
for the development of the valley. This was done with Mrs. 


>The League also maintains a guide service for the National Arboretum. 


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FERN VALLEY 169 


Morrill Donnald, Chairman for planting, Mrs. Frederick Lee, 
Mrs. Clifton Luce, Mrs. Benjamin Thoron, Mrs. Benjamin Powell, 
for wild flowers, Dr. James Johnston, for the American Fern So- 
ciety, and Mrs. Charles Bittinger, Chairman. 

As some money is essential for any kind of program, we gave 
a benefit and the proceeds were deposited with the “Friends of 
the National Arboretum.” Gifts have been added to this fund 
from time to time, from individuals and clubs. Some gifts were 
for specific purposes, for example the entrance planting with a 
stone bench and map shelter, ae a map of the valley and a 
list of workers and donors is display: 

In November, 1958, the Conintter unk at the Arboretum with 
the Directors to formulate plans. It was obvious that before any 
planting could be done much basie word had to be accomplished, 
and Dr. Skinner offered to arrange for this. The Arboretum has 
operated for many years on rather limited appropriations from 
Congress. With roads and bridges to build in order to open the 
various sections, there was little money or labor available for 
other projects. It was our good fortune that when the League 
began its plans the Arboretum was able to undertake a basic 
landscaping and clean-up task that the League would have been 
unable to accomplish or afford. 

It was agreed that the honeysuckle and poison ivy should be 
removed, that a main path five to six feet wide should encircle 
the valley, that the debris from flash floods should be cleared 
out, and that outlets from the main water system be installed. 

Under the able and artistic direction of Dr. de Vos work went 
on through the winter and summer of 1958 and 1959. The land- 
scaping was not finished but the results far exceeded our hopes. 
The ugly sewer pipe was camouflaged and made into a bridge 
across the stream, the rivulet was cleared, dammed, and widened, 
making miniature waterfalls, pools, swamp, and marsh areas; the 
honeysuckle was nearly eliminated. We, the League’s Fern Val. 
ley Committee, were ready to begin. 

Garden tools, hose, and soil testing equipment were bought, 
and five tons of lime soil and ten tons of lime rock were brought 


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from Leesburg, Virginia. Mrs. Donnald sent out a notice for 
workers and the planting began. It seemed to us that a lime- 
stone wall was essential for the lime-loving ferns. Once again 

r. Skinner came to our assistance by asking if he might super- 
vise its construction. Dr. de Vos knew of a limestone wall in 
Frederick, Maryland, part of the fortifications built by Braddock 
in the French and Indian war. Sixty tons of this we bought and 
trucked in. Our limestone “cliff” is ten feet high and sixty-five 
feet long, built against a dirt bank and filled with lime soil. It 
faces north but, following the contour of the bank, has some 
slight east and west exposures. The Arboretum has planted many 
hemlocks in this area. 

During the summer of 1959 Dr. de Vos asked if we could buy 
a collection of wild flowers. We agreed and he ordered and trans- 
ported 750 plants of 45 varieties which were planted by our 
workers. Mrs. Donnald collected the ferns and planted the bank 
which we had envisioned as “dripping with Christmas ferns” 
with 375 plants. The season ended with 3,000 ferns and 2,500 
wild flowers plante 

The winter of 1959 and 1960 saw the construction of two 
bridges completing the circuit of the valley, thus bringing the 
path back to the entrance. A small pine woods was added where 
Cypripedium acaule grows naturally ; a non-calcareous wall was 
built ; many native shrubs and trees, including two cypress trees, 
were planted by the Arboretum. 

e planned the formal opening of Fern Valley for May 24, 
1960. Early spring found every one at work, more ferns were 
brought in from the Washington area, more ferns were bought, 
and some rare ferns donated. By this time the original hope that 
rare ferns would be added to the basic planting was being ful- 
filled. Dr. Wherry sent ferns (Athyrium hybrids) and gave ad- 
vice. Dr. Laura Barnes brought Dryopteris chrysoloba and D. 
viridescens. 

At the opening ceremonies Mrs. Donnald reported 4,000 ferns, 
of 47 species, and 2,000 wild flowers representing 90 species had 
been planted. Seventy workers from 21 garden clubs had con- 


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FERN VALLEY 173 


tributed 326 man-days of work. Gifts of plants and money had 
come from 61 individuals and clubs. 

When we began all ferns were welcome, even the weedy ferns 
for restricted areas, but by the summer of 1960 it was decided 
to limit planting in Fern Valley to native ferns and wild flowers. 
We wished to have a group of foreign ferns for comparison and 
study, so a planting of exotic ferns was started in an adjoining 
ravine. Dr. Johnston was in charge of this section, and used 
exotics from his pilot planting as a nucleus. 

The exotic ferns transplanted to this section included Athyri- 
um Goeringianum var. pictum, an Athyrium hybrid (A. angus- 
tum X Goeringianum var. pictum?), Dryopteris chrysoloba, and 
D. viridescens. Dr. Meyer donated three each of Dryopteris 
Filix-mas ev. ‘Daedalea,’ ev. ‘Eroso-crenata, ev. ‘Furcans,’ ev. 
‘Polydactyla,’ and ev. ‘Subintegra, D. pseudomas ev. ‘Polydac- 
tyla, Athyrium Filix-femina ev. ‘Frizelliae, ev. ‘Montrosum,” 
ev. ‘Multidentatum,’ ev. ‘Pseudo-Victoriae, and some others. 

We still welcome gifts, for work continues on this long-term 
project. Anyone wishing to send ferns write to Mrs. Morrill 
Donnald, 3703 Jones Road, Chevy Chase 15, Maryland, and ask 
about species desired and for shipping instructions. 

I shall go ferning again, for the search is the spice of life, and 
the acquisition of new and rare ferns, in order that our National 
Arboretum may have as complete as possible representation of 
our native ferns in its climate, is our satisfaction. Our success 
to date is due to the interest, help and encouragement of the 
entire Arboretum staff, its Directors, Botanists, Plant Propaga- 
tors, Secretaries, Workmen and Truck-drivers; to Mrs. Don- 
nald and her husband, to the 75 women diggers and their hus- 
bands, but most of all to Mrs. Donnald, whose endless enthusi- 
asm, devotion, and hard work has made Fern Valley what it is. 

So here is Fern Valley today; a lovely shady valley with its 
beautiful old beech and tulip trees, sloping hillsides, rivulets, 
pools, ferns and flowers, a joy to all, a dream come true. 

3403 O Srreer, N. W., WasHrNneTon 7, D. C. 


174 AMERICAN FERN JOURNAL 


Ferns in Cultivation, IV. Some Wall Ferns 
Syivia LEATHERMAN 


A wall is a straight harsh line, which we all try to soften in 
order to have an attractive setting. Ferns are good for over- 
coming these straight lines. One I like to use is Microlepia platy- 
phylla, a common fern in cultivation which seems to lack a com- 
mon name. Here in southern California it will grow to seven 
or eight feet tall and is therefore an ideal background fern for 
landscaping along a wall that faces east or north. Good speci- 
mens with graceful arching fronds develop only with some sun. 
T have tried it in a shady corner, but the stipes become soft and 
do not have the sturdiness to hold up the large heavy fronds. 

Microlepia strigosa is another popular fern that can be used 
along a wall to create a soft, graceful effect. I have grown this 
one successfully where it receives the early morning sun or the 
late afternoon sun, and also where it receives filtered sunlight, 
or even in a location where it received no sun at all but had good 
light. It has been satisfactory in all these locations. In the sun 
the fronds are yellow-green and have a thick texture, whereas 
the ones in the shade with good light are greener and thinner. 

The Leatherleaf Fern, Rumohra adiantiformis (sometimes er- 
roneously called Polystichum capense or Aspidium capense), also 
called Iron Fern, is popular with flower arrangers, because the 
fronds will last as long as three weeks after being cut. To have 
good texture and fronds that will stand up, the ferns must have 
a good deal of sun. Inland where I live is not considered a good 
location for ferns, for we are far enough away from the ocean 
to have dry heat in the summer and frost in the winter. How- 
ever, Rumohra will grow well with me if given care. It desires 
morning sun until about eleven o’clock or the afternoon sun from 
three thirty p.m. on. Along the coast, where there is better hu- 
midity, it will taken even more sun. This fern is often called a 
“buffer” plant; when planted at the windy end of a wall it will 
break the wind and protect other ferns planted along the wall. 

2637 Norra Ler Avenur, Eu Monte, CALIrorNia. 


SouTHERN DISTRIBUTION OF BOTRYCHIUM 175 


Southern Distribution of Botrychium oneidense 
nd B. multifidum 


F. R. FosBere 


The resemblance of a plant collected for Botrychium dissectum 
in Greenbrier Cove, Sevier County, Tennessee (Fosberg 40332) 
to one found in Quebec (Fosberg 40154) a few weeks previously 
and called B. multifidum by botanists there directed my atten- 
tion to differences between this Smoky Mountains plant and two 
other collections from the Smokies near the Chimneys Camp- 

round, Sevier County (Fosberg 40334, 40336). The aberrant 
collection, representing a small colony, has short stubby ultimate 
segments, rounded to acute at the tips, but the fronds are not 
as dissected as those of plants of B. multifidum from Shenan- 
doah National Park. It corresponds well with the description 
of B. dissectum var. oneidense (Gilb.) Farw. as given by Ferna 
in the 8th edition of Gray’s Manual. When W. H. Wagner ex- 
pressed an interest in seeing this collection I sent it, along with 
a number of others of the Sceptridium group of Botrychium. He 


as a separate species. So far as we know this constitutes a new 
record for Tennessee and for the Great Smokies. Another collec- 
tion, Fosberg 23958b, from Little Hunting Creek just east of 
Mount Vernon, on the Potomac River, Fairfax County, Virginia, 
as well as the one from Quebec, three miles west of Napierville 
(Fosberg 40154), also proved to be B. oneidense. 

The Virginia and Tennessee records rather amplify the known 
distribution of B. oneidense in the southeastern United States. 
It has previously been reported from Rockingham County, Vir- 
ginia, by Clausen (1944) ; Yellow Creek, Graham County, North 
Carolina, by Clausen (1938); from Roan Mountain, Mitchell 
County, North Carolina, by Clausen (1943) ; and from Gatewood 
Switch, Pendleton County, West Virginia, by Strausgbaugh and 
Core (1952). The latter specimen has been verified by Dr. Wag- 
ner as B. oneidense (in litt. Aug. 7, 1960). 

The only localities south of Pennsylvania in eastern North 


176 AMERICAN FERN JOURNAL 


America in which B. multifidum grows are in the Virginia Blue 
Ridge. Besides the well-known colony at Big Meadows (see Wag- 
ner, 1946), two others have turned up not far away, one just to 
the north, at Old Rag View Overlook, on the Skyline Drive, 
Madison, County (Fosberg, 36135a), the other a little to the 
south, at Milam Gap, also on the Skyline Drive, Madison County 
(Fosberg, 41088). All of these Blue Ridge collections are from 
meadows resulting from the abandonment of ridge-top farms 
when the land was bought for the Shenandoah National Park. 
There was undoubtedly a swamp at Big Meadows, even in the 
farming days, and a surprising number of plants otherwise rare 
or absent in the area found refuge and persisted there. Among 
these are such otherwise northern species as Caltha palustris, 
Menyanthes trifoliata, and Betula populifolia. Botrychium mul- 
tifidum may well have existed there as a relict from a colder 
period, and with the new availability of the meadow habitats. 
may now have started to spread into them, north and south, ex- 
actly as has Betula populifolia. 

If there is anything to the hypothesis that Botrychiwm onet- 
dense is a hybrid between B. multifidum and B. dissectum (see 
Clausen 1944) it could be expected to turn up at Big Meadows 
and Old Rag View, where these species grow side by side. It has 
not been found there as yet, but it has not been specially sought. 
Furthermore, the existence of B. oneidense at five localities in 
the Appalachians and one on the Coastal Plain all well out of the 
present range of B. multifidum, make it seem unlikely that any 
such hybridization is taking place at present in this southern 
part of its range. 

On the other hand, though these three plants are distinguish- 
able, they seem pretty close together to be regarded as distinct 
species. The fact that these three, with their several other close 
relatives, all now being studied critically by Dr. Wagner, seem 
to require statistical methods to be discriminated with certainty, 
suggests that in Botrychiwm taxonomy we are almost getting to 
the field mouse stage. It may be that the suggestion (Clausen 
1944, p. 59) that B. multifidum and B. dissectum might be “only 


SoUTHERN DISTRIBUTION OF BOTRYCHIUM 177 


subspecies of a polytypie species” has merit and should not be 
abandoned altogether. 

I am familiar with Whgner's (1960) discussion of infra-spe- 
cific categories in this and other groups of pteridophytes, in 
which he maintains, among other points, that “if two taxa co- 
exist over a large range and maintain their characters, they 
should be interpreted as species.” I find this statement 4 little 
hard to reconcile with that which immediately follows it: “This 
means that not only do the taxa have diagnostic features be suffi- 
cient number and nature to be readily separated. . As 
read Wagner’s article it seems to be an attempt to pines the cate- 
gory “subspecies,” at least, on an objective basis. However, the 
words sufficient and readily in the above quotation place us right 
back where we were before, as these involve purely subjective 
judgments. It is evident that so long as we employ the present 
linear hierarchy of categories in which, because of the complex 
character of natural phenomena, we must include many kinds 
of taxa in each category (see Camp and Gilly 1943) any hope of 
placing these categories on a really objective basis is illusory. 

Since I have been interested for many years in the taxonomic 
treatment of infra-specific categories, I would like to comment 
a little more fully on Wagner’s discussion. Although it is of con- 
siderable value to consider the range of usage of given categories 
among students of pteridophytes as well as of other plants, it 
does not seem that we should allow preponderance of usage to 
influence our judgment unduly. We have a definite and limited 
series of categories available under the International Code of 
Nomenclature, and there seems to be no point in discarding any 
of them merely because other botanists have misused them. The 
purpose of having such a series is quite simply to indicate the 
degree of divergence between the discernible groups of plants, 
giving a number of levels at which they can be discussed and, 
hopefully, to indicate in some measure the evolutionary relation- 
ships of the populations concerned. The extremely diverse nature 
of the populations, at any one level of differentiation (or rank), 


178 AMERICAN FERN JOURNAL 


would seem to make a single objective definition of any category 
impossible. 

The category of forma should, in my opinion, be limited to 
what Wagner calls “genetic forms,” as there is certainly no place 
for non-genetic accidental or environmental modifications in the 
taxonomic system. Probably this rank should comprise both the 
sporadic but conspicuous genetically different individuals that 
occur among populations of ordinary individuals and the geo- 
graphically limited populations of individuals which differ only 
very slightly from the rest of the species. I cannot agree at all 

with Wagner’s opinion that “no trivial form deserves a botanical 
name.” Whether or not such entities should be named depends 
on whether we need to refer to them by name. 

Although admitting that variety has been variously misused 
T could not countenance dropping it altogether or even de-empha- 
sizing it. I expressed myself on this matter some years ago 
(1944) and have as yet found no reason to change my mind about 
it. My principal difference from Wagner’s conclusions on this 
is that I regard varietas as the ordinary geographical subdivision 
of a species, rather than assigning to this the term subspecies as 
is done by the zoologists. It is certainly very likely that varieties, 
in this sense, will intergrade with each other, especially where 
their ranges come together, but on the other hand, this inter- 
gradation does not seem to me a necessary criterion. There are 
sterility and other barriers, even between populations that are 
morphologically indistinguishable or almost so, that prevent any 
gene exchange. It still seems to me to be a matter for the judg- 
ment of the individual taxonomist, preferably based on experi- 
ence with many groups of plants, to decide how distinct these 
should be before they should be called varieties or species. The 
same is true of intergrading populations, some of which cer- 
tainly merit the rank of species, though most are of lower rank. 

The category subspecies should, I think, in general be reserved 
for groups of closely related varieties within species, but there 
seems no inherent reason why the term should not also be ap- 
plied to well separated single entities where this serves the pur- 


SouTHERN DISTRIBUTION OF BOTRYCHIUM hT9 


poses of clarity or emphasizes evolutionary diversity. 

It is easy to agree with Wagner that hybrids should not be 
regarded as varieties or subspecies of either of their parents so 
long as we are dealing with ordinary hybrids, either sterile F'; or 
the segregating “pleomorphic” hybrids whether sporadic. or 


In any event, it is obvious that the Sceptridium group of grape- 
ferns should be much more carefully studied in the field, and 


taining to the plants discussed in this note. 


LITERATURE CITED 

Camp, W. H. and C. GILLY, 1943. The Structure and Origin of Species. 
Brittonia 4: 323-385. 

Ciausen, R. T., 1938. A Monograph of the Ophioglossaceae. Mem. Torrey 
Bot. Club 19(2): 1-177. 

1942. Studies in the Ophioglossaceae: Botrychium, Subgenus 
Sceptridium. Amer. Fern Journ. 33: 11-27. 

Cpe BOE On tae Status of Botrychium dissectum var. oneidense. 
Amer, Fern Journ, 34: 55-60. 

FERNALD, M. L., 1950. Gray’s Manual of Botany, Eighth Edition. 1-1632, 
New York 


Fosserc, F. R., 1942. Subspecies and Variety. Rhodora 44: 153-157. 


180 AMERICAN FERN JOURNAL 


ee P. D. and se L. aca 1952. Flora of West Virginia (Part I). 


—————-,, 1960. Seren Grapeferns and the Meanings of Infraspecific 
Categories used in North American Pteridophytes. Amer. Fern Journ. 
50: 45, 


212 Houmes Run Roap, Fatus Cuurcn, VIRGINIA. 
Dryopteris x Tavelii in the Valley of Chamonix, France 
ANDRE AND ANNE-MARIE LAWALRE&E-COLLARIS 
In August, 1960, we spent several days collecting between 
Servoz and Chamonix, at the foot of Mont Blane, in the Depart- 


a rich flora. Dryopteris X Tavelii Rothmaler is abundant in 
this region, although the little work on the ferns of the environs 
of Mont Blane by Venance Payot! mentions only typical D. Filiz- 
mas. In 1959, the senior author in outlining the distribution 
in France of D. X Tavelii? had not at that time seen a single 
specimen from Haute-Savoie. Probably Christian Bange, of 
Lyon, was the first to observe this fern in this region. He wrote 
to us September 10, 1960, that he had found it in 1958 in the 
gorges of Diosaz, and later in several localities in Savoie and 
Haute-Savoie, among others in the environs of Chamonix. He 
believes that he has discovered D. Borreri Newm. also, on the 
left lateral moraine of the Glacier of Argentiéres. In collabora- 
tion with M. Berthet, M. Bange communicated his first discover- 
ies to the Societé Linnéenne de Lyon in June, 1960, and an 
account has now been published.? 

he specimens of D. X Tavelii from the valley of Chamonix 
differ from those of Belgium and Luxemburg. They have the 


Payot. Les dg res des environs du Mont Blanc, in Florule du 
Meee Blane, ou Guide du botaniste et du touriste sur les Alpes Pennines, 
deoxiene partie IT + 22 p., Trembley, Genéve. 1881. 

2A. Lawalrée. Répatition en ee nce et présence en Autriche et en You- 
goslavie de Dryopteris x Tavelii Rothm. Bull. Res. Council Israel, Sect. D., 
oe 7D: 181-183. 9 

3 P. Berthet and ©. Bange. Notules d’herborisations ptéridologiques: 
IV. Bull. Meus. Bok: Linn, Lyon 29: 227-231, 1960. 


DRYOPTERIS X TAVELIL IN FRANCE 181 


blades often narrower and more coriaceous, doubtless because of 
the more severe climate. Sometimes they have the pinnules al- 
most entire and separated by narrow sinuses, thus being so simi- 
lar to D. Borreri that we have consulted with Léopold Reichling, 
of Luxemburg, who after a full study has replied (10 October 
1960) : ‘‘Specimens of true Borreri such as those that I have 
collected here in Luxemburg and in Spain have characters and 
a general aspect that I have not found in any of your specimens 
from Haute Savoie. It will be necessary to establish by cytologi- 
eal research the nature of these forms of the Alps that leave us 
uadrpided before being able to judge their specific relation- 
hi 


We identify therefore all our plants as D. X Tavelii, although 
no. 11,120 is however very near to D. Borrert.t Moreover, Reich- 
ling has indicated that numbers 11,128, 11,137, and 11,141 are 
practically identical with two specimens (nos. 18,829 and 
18,830) collected in the Swiss cantons of Uri and Tessin by a col- 
laborator of F. von Tavel, E. Oberholzer, and determined by 
the latter as D. Borreri var. insubrica von Tavel.® It is true that 
von Tavel slassified his var. insubrica among his typical Bor- 
rert forms, but is not this plant related rather to D. x Tavelii? 

Poelt® speaks of his difficulties in sorting his Bavarian speci- 
mens into D. Borreri and D. * Tavelti, and our specimens pre- 
sent the same problem. Does D. X Tavelii behave in the valley 
of Chamonix like it does in Bavaria, on the northern slopes of 
the Alps, where Poelt studied it? Both regions have a very cold 
winter and a rather high humidity during the growing season. 
In the valley of Chamonix it grows on all exposures and far 
from being restricted to the bottom of the valley it climbs the 
slopes at least 400 meters higher near the Glacier of Bossons on 
western exposures and 200 meters near Vaudagne on northern 

4 Our collection numbers 11,113 to 11,142 of August, 1960, from Cha- 
monix are in the herbarium of the J. ardin hence de 1’Etat, Bruxelles. 

von Tavel. Dryopteris Borr ihr Formenkreis. Verh. 


SF, i New 
Schweiz. Naturf. Ges., 118 Jahresversamm! 153-154, 1937. 

6 J, Poelt, Dryopteris Borreri in Bavaria, Tus JourNaL 50: 114-117. 
1960, 


182 AMERICAN FERN JOURNAL 


exposures. One finds it in shade and in full sun, generally in a 
deep soil, although it sometimes accompanies Aspleniwm septen- 
trionale on almost vertical schistose rocks that very likely the 
snow does not cover in winter; but in this habitat the plants 
are weak, for example no. 11,139 (see no. 5 below). 


\(Brévene fy 


“af ' 2525 2 
x ZARLAVEYRON ge A ac pu sh 
: y 


LAigui tte 
po Ba 


\VAUDAGNE $ 


Con pe. La Fore! 
OUR 
“1556%.. 


\ 1007 


Map oF THE VALLEY OF CHAMONIX WITH THE LOCALITIES WHERE THE 
AUTHORS HAVE Founp D. & TAVELU, MARKED WITH A CROSS; ALTITUDES 
IN METERS, CHAMONIX Is JUST OFF THE MAP TO THE HAST. 


Here going from lower to higher elevations, are the places 
where we have observed D. & Tavelii: 

A. On the right bank of the Arve River: 

1. Gorges of the Lower Diosaz River: A little above the monu- 
ment to Frédéric Auguste Eschen, left bank (no. 11,122) ; under 
the falls of the Porte River on the left bank in a little side ravine 
(no, 11,121) ; steps above the Barme Rousse Bridge, left bank; 


DRYOPTERIS X TAVELI IN FRANCE 183 


under the falls of the Aigle River, left bank; at the Achille 
Cazin Bridge, right bank. 

2. Gorges of the Arve River: Abundant from the Pelissier 
Bridge to the viaduct of Sainte-Marie-aux-Houches, e.g. no. 
11,141. 

3. Along the Coupeau road, on wooded slopes at two places. 

4. Between Les Gaillands and Les Bossons, on wooded slopes 
200 meters above the railway bridge over the Arve River (no. 
11,116), two tufts in company with D. Filiz-mas, D. carthusiana, 
and Athyrium Filix-femina. 

B. On the left bank of the Arve River: 

5. Sides of National Highway 506: A little below La Fontaine 
(no. 11,139), on schistose rocks on the south side of the road. 
with Asplenium septentrionale; between La Fontaine and Pelis- 
sier Bridge, on the south side; at 100 meters above Pelissier 
Bridge, on the south side, on schistose rocks; at 500 meters above 
Pelissier Bridge (no. 11,133), on the north side, with Alnus 
tncana; higher, on both sides of the road, rather frequent up to 
the road to Les Houches (no. 11,134), for example in a wood of 
Picea Abies, with galls of Chortophila signata (Brischke) de 
Meijere. 

6. Along the road between Vaudagne and Highway 506: In 
a forest of Picea Abies and Fagus sylvatica, where the road 
winds frequently (no. 11,130) often with Pilystichum aculeatum 
(no. 11,131) ; on schistose rocks beside the road, with D. Filix- 
mas, Athyrium Filix-femina, Veronica latifolia L. em. Scop. 
(= V. urticifolia Jacq.), Hieracium gr. murorum L. em. Huds., 
Sazxifraga cunetfolia, Deschampsia flexuosa, Prenanthes pur- 
purea, Vaccinium Myrtillus, Melampyrum pratense, Epilobrwm 
montanum, Chaerophyllum hirsutum, Solidago Virgaurea, As- 
plenium Trichomanes, Polypodium vulgare, Thelypteris Phegop- 
teris, Rhabdoweisia fugaz, Bartramia pomiformis, Hypnum 
cupressiforme, Dicranum scoparium, Rhytidiadelphus triquetrus, 
Rhacomitrium aquaticum, R. heterostichum, Hylocomium splen- 
dens, Fissidens cristatus, Polytrichum juniperinum, Schistidium 
apocarpum, Tortella tortuosa, Diplophyllum albicans, Lophozia 


184 AMERICAN FERN JOURNAL 


quinquedentata,’ Peltigera horizontalis,’ and other species. 

7. Between Vaudagne and Les Chavants, frequent in hedges 
and talus (no. 11,120) and in the forest (nos. 11,128 and 11, 129). 
Here is a phytosociological record of 70 square meters of this 
light forest, 50 meters south of the Vaudagne-Les Chavants road, 
at about 100 meters elevation, on a 20% slope exposed to the east. 


Arboreal stratum, attaining 30 m. in height and a covering of 40%: 

Pi i Karst. va 4 

Shrubby stratum, attaining 4 m. in height and a covering of 30%: 
L. 


Corylus avellana gee. 
Alnus incana 4 Moench 1—2 
Populus tremula L aes) 
Herbaceous’ stratum, ahetatiig 1 m. in height and a 100% covering: 
tenuis Sibth 4 
Vaccinium Myrtillus L 4—4 
Melam ite pratense L. 3—3 
Baba 2—2 
i penibiia erecta (Iu.) Raeusch at 
Dryopteris Filix-mas (l.) Schott eae 
Dryopteris & Tavelii Rothm ee 
opteris dilatata (Hoffm.) Gray ee 
Athyrium Filiz-femina (L.) Roth? +2 
Blechnum Spicant (.) Roth ayers 
Thelypteris coiarati es (All.) H. P. Fuchs ae 
Carex pallescens 1. tol 
Prenanthes peat a L. oe 
iola sp. 4+—1 
Hieracium gr. murorum L. em. Huds. et 
Orchis maculata L amis 
Fragaria vesea L. ok 
oo odoratum lL. Bae 
oleu +—1 
Peni ek eee (L.) Crantz se 
Veronica officinalis L. - 1 
Prunella vulgaris L. Fo 


ambin 
gall of Chortopiile 4 sie oen “(Brischke) de Meijere is abundant in 
the eae on Athyrium Filix-femina. 


Dryopreris X TAVELIL IN FRANCE 185 


Ajuga reptans 1. +-—1 
Vaccinium Vitis-idaea L. +—2 
Mossy stratum covering about 80% 
Cirriphyllum piliferum (Heb) Grout 
pesintirey scoparium Hec 
todon Schreberi Gay Moenkem. 
is splendens (Hedw.) Bruch & Schimp. 
Ptilidium crista-castrensis (Hedw.) DeNot 


hy phus 
Lophocolea bidentata (L.) Dumort. 
LZ i r 
Plagiochila asplenioides (1.) Dumort. 


8. Les Houches, on the road to Les Aillouds (no. 11,126), 
wooded slopes facing west, with the following species: Alnus in- 
cana, Acer Pseudoplatanus, Sorbus aucuparia, Fraxinus excelsior 
Fagus sylvatica, Picea Abies, Larix decidua, Prenanthes pur- 
purea, Oxalis Acetosella, Deschampsia cespitosa, Rubus idaeus, 
Chaerophyllum in ssscil Solidago Virgaurea, Fragaria vesca, 


Maianthemum bifolium, Athyrium Filix-femina, Dryopteris 
Filix-mas, Polysti og OE eile vulgaris, and vari- 
ous bryophytes. 


Along the path to the Glacier des Bossons up to 1415 
meters elevation (no. 11,137), the highest altitude where we have 
observed D. X Tavelii, a little below the chalet of the Glacier des 


a sub-story characterized by an abundance of Prenanthes pur- 
purea, mixed with Veronica latifolia, Pasion limbosperma, 
T. Phegopteris, Dryopteris dilatata, D. Filix-mas, Athyrium 
Filix-femina, and other species. 

SAN MINIATO, 3 AVENUE VAN ELDEREN, BRUXELLES 16, BELGIUM. 


186 AMERICAN FERN JOURNAL 


Basket Ferns for Southern California 
Fay MacFAappENn 

In southern California ferns may be easily grown in baskets 
which may be hung from the branches of trees, from the rafters 
of a porch, or from the roofs of lath houses. 

The best medium for growing plants in baskets I have found 
to be pure sphagnum moss; it is light in weight when dry and 
can be easily moved. Baskets lined with Oregon green moss are 
good too, but then the baskets must be filled with a soil adapted 
to the growth of epiphytes; these are then heavy to lift. Each 
spring I tuck more sphagnum in the holes where the birds have 
stolen it to build nests, and around the edge and top of the basket 
Otherwise baskets require little care. They may be watered with 
a hose or by soaking them in a tub of water. 

Polypodium aurewm (also known as Phlebodium aureum) and 
some of its cultivars, such as ‘Mandaianum,’ make huge baskets. 
The fronds turn yellow and fall eventually. I cut them off as 
the new fronds appear. Like many other epiphytes they can be 
grown in soil, where they are beautiful when planted in good 
soil at the base of trees and posts. I usually plant them in sphag- 
num at first, but eventually they spread out to whatever soil 
they find. 

The most commonly grown basket-fern in southern California 
is Polypodium subauriculatum ev. ‘Knightiae’, which is tough 
and resistant. The old fronds are ready to drop off in April, 
when the new fronds begin to show, peeking through the basket, 
are black before they unroll. These fronds cover the whole 
basket and a well-grown plant is a beautiful sight. In this form 
the margins of the leaves are deeply and irregularly cut. Plants 
of the botanical species P. subauriculatum, with the margins 
uncut, are also grown in southern California, but they are very 
tender; they grow only during the summer months, and must 
be protected when the days get really cold. 

Another fern that may be grown in baskets is Polypodium 
vulgare and its varieties. They are at their best in early spring. 


Basket FERNS FoR SOUTHERN CALIFORNIA 187 


Among the most unusual ferns for baskets are the rabbit’s- 
foot-ferns, Davallia and Humata, which creep over logs and rocks 
in nature as well as growing as epiphytes in trees. Humata 
Tyermannu, a native of southeastern Asia, has long-creeping 
rhizomes, covered with white scales, which completely cover a 
basket. The faster-erowing Davallia trichomanoides, sold in the 
local trade erroneously as D. canariensis, is similar but has tawny 
rhizome scales. Both these ferns have deciduous fronds. Humata 
Tyermannii is beautiful when its fronds mature. These are quite 
thick; some time before they fall they turn various shades of 
yellow and eventually brown, but never dry until they fall. The 
fronds are jointed to the rhizome, and thus these ferns prune 
themselves, as do some species of Polypodium. An attempt to 
pull a frond off even a few hours before it is ready to fall is of 
no avail. Davallia trichomanoides, on the other hand, is incon- 
spicuous in its ripening process; the fronds when withering and 
drying up may be pulled loose easily without damaging any new 
growth. In the fall most fronds of the Humata usually fall 
off, leaving the basket a mass of rhizomes, and the same thing 
may happen at other times if the plant becomes too dry. No 
doubt this is nature’s way of carrying these ferns through peri- 
ods of drought. Late winter or early spring will bring new 
growth; the new fronds do not really develop well until summer 
arrives and the ripening takes place in late summer. 

Another rabbit’s-foot-fern, Davallia feejeensis, is a fern that 
maintains a good appearance all year, although it does not grow 
in the winter. Whether one is conscious of it or not, most ferns, 
even the tropical and subtropical ones, have a period of rest. 
Some may grow half-heartedly during part of the winter, some 
go completely dormant, like this Davallia, even though the fronds 
of the preceding summer remain green, and some disappear en- 
tirely, like Polypodium californicum, which sends forth fronds 
in winter and spring and which dies down completely in summer. 
The fern gardener shold take note of where such things are 
planted, to avoid digging around them when they have ceased to 
show. The fronds of D. feejeensis are even more ornamental than 


188 AMERICAN FERN JOURNAL 


those of the other Davallias, being finer cut and lacier. In sum- 
mer this species grows luxuriantly, but in winter it should be 
grown under glass. The frond usually withers and dies; the old 
stipe is left on and can be pulled out. I never wait for self- 
pruning in D. feejeensis, but remove the worst-looking fronds; 
still, the old fronds do protect the rhizomes somewhat. These 
rhizomes grow either upright or horizontal, but although they 
can cover the basket they do not wrap themselves around the 
basket in the complicated manner of D. trichomanoides. 

Suitable for baskets in the shade are the staghorn-ferns, Platy- 
cerium bifurcatum and other species, which form huge, exotic- 
looking plants. Other good basket ferns are Polypodium poly- 
carpon (sold as P. punctatum or P. irioides) and Pyrrosia lingua. 

5450 CarLIn Street, Los ANGELES 16, CALIFORNIA. 


Shorter Notes 


DENNSTAEDTIA GLOBULIFERA (Porr.) Hreron. in Texas (PI. 15). 
Recently the writer reported the occurrence of Dennstaedtia 
globulifera in Texas (Wricutia 2(2) :108-110, fig. 20. 1960). 
This species is not only new to Texas but also to the United 
States. In order to call the attention of members of the Fern 
Society to this discovery, who might otherwise not see the 
WricHtra report, Mr. Morton suggested a reprinting of the 
illustration and this note. 

Dennstaedtia globulifera was collected in Fern Cave, 18 miles 
north of Comstock in Val Verde County, Texas, by Larry Hoff- 
man (s.n.) on November 2, 1958. The plant or plants grew in a 
eave 50 feet in depth and had fronds which were said to be 13 
feet in length. 

This species is now known to occur throughout the West In- 
dies and from northern South America northward in the tropi- 
eal and subtropical regions of Central America and Mexico, with 
a disjunct station in Val Verde County, Texas. It is usually 
found in moist thickets and partially shaded places—DoNovaN 
S. Corretu, Texas Research Foundation, Renner, Texas. 


AMERICAN FERN JOURNAL 


VoLuME 51, Puate 15 


Wp 
CVF 
Sy 
OZ 
We di 
Se M4, 
ENG ag 
ne, 3B 
SNE Se 
Ne 
ANN PRE NNND) og 
QE SP 
SX Q SQV Assy” 
AS <i aWeP 
SESS QASSS 
a NG yay, @ Ss) R\n 
SEE eA 
aNETGNS GSS 
ole Q . O NN 
EGS” “ Ry : 
Sst abs = 
SH SK 
Pris 


EQ 
RED 


DENNSTAEDTIA GLOBULIFERA (PoIR.) Hreron.: 1, PINNULE, SHOWING SORI, 
ABouT X 3; 2, soRUS, X 20; 3, TWO PAIRS OF SECONDARY PINNAE, ~ 
ILLUSTRATION BY PHOEBEJANE HORNIN' 


190 AMERICAN FERN JOURNAL 


PrERIDOLOGICAL TrRoGLOpyTES.—It is well known that many 
ferns flourish in caves, or in limestone sink-holes, often well be- 
low the surface of the ground; the plants adjust to available nat- 
ural light. But ferns that depend almost entirely upon artificial 
light and survive unnatural conditions, as real troglodytes, are 
rare. 

For several years two clumps of ferns have been putting on a 
brave show deep in the limestone recesses of Crystal Cave, three 
miles west of Kutztown, Pennsylvania. The ferns were shown 
to me in early June, 1961, by Ralph B. Lutz, one of the Cave 
guides, who has taken a special interest in these misplaced mem- 
bers of the fern community. 

The plants, sadly depauperate specimens, barely exist without 
benefit of soil; they cling to thin, moist limestone ledges about 
200 feet back from the Cave entrance, and 110 feet below the 
surface of the ground, where the mean average annual temper- 
ature is 56°. Clump ¥1, a Christmas Fern, Polystichum acrosti- 
choides, had five fronds the longest of which was seven inches. It 
enjoys more ample light than its neighbor, an Ebony Spleenwort, 
Asplenium platynewron—just barely recognizable as such. t 
the time of my visit I added a generous handful of rich humus 
to the Christmas Fern. Two months later Mr. Lutz informed me 
that this clump developed two additional fronds, and that the 
plant had become healthier, more vigorous with the addition 
of soil. 

Illumination for these plants is derived from single 75-watt 
projection lamps, part of the lighting system of the Cave. In the 
case of the Christmas Fern, the lamp is 27 inches from the plant 
and, when illuminated, the temperature at the fern is 78°. The 
lights are on for 5 hours daily from mid-February to late May 
and from early September to December 1, and 8 hours daily 
from Memorial Day to early September—approximately 1,780 
hours of the year. 

The fern spores could easily have been introduced into the 
Cave via human visitors, or by bats; but it is remarkable that 
the ferns could develop under a light intensity of 8 foot-candles 


SHORTER NOTES 191 


an indicated by a photoelectric meter—MauvricE Broun, Hawk 
Mountain Sanctuary, Kempton, Pennsylvania. 


MARSILEA QUADRIFOLIA L. IN INDIANA AGAIN.—Recently while 
collecting ferns and fern-allies in Indiana, I came upon a large 
colony of Marsilea quadrifolia Li. in Cooley’s Pond, a farm pond 
given over to Nuphar advena and Marsilea. This pond is on a 
farm one and one-quarter miles south of Bordens on route 60 in 
Clark County. The pond varies from 150 to 200 feet in diameter. 
The Nuphar grows out in the center of the pond, while Marsilea 
grows from along the shore, where some plants are actually 
emersed, to ten feet into the pond where the water is at least 
two feet deep. One end of the pond is entirely covered by the 
Marsilea. 

Deam (in Flora of Indiana, p. 1021, 1940) excluded Marsilea 
quadrifolia Lu. from the flora of Indiana because he was unable 
to find the species in 1937 in the area reported by Grimes in 1911, 
that is, from an old mill pond on the south side of the Vandalia 
Railroad in Greencastle, Putnam County, Indiana. 

Therefore, finding Marsilea quadrifolia lL. in Cooley’s Pond 
near Bordens, in Clark County, reestablishes its existence in 
Indiana. Many specimens were collected from this locality Aug. 
16, 1961 (Reed 52412). Some plants have one sporocarp and 
some bear pairs of sporocarps. Representative specimens have 
been placed in the United States National Herbarium.—CLybE 
F. Reep, Reed Herbarium, Baltimore 34, Maryland. 


Notes and News 
July 18, 1961 
Dear Sir: 

In Volume 51 of the Journal, Mrs. MacFadden suggests the use 
of old fluorescent tubes in lath-house construction. Although 
modern tubes may be safe for such use, older tubes are danger- 
ous and should never be used. They contain various salts of 
beryllium in a fine, powdery form. Leaking and broken tubes 
permit escape of this material, which, if it gains entry into lung 


192 AMERICAN FERN JOURNAL 


or injured skin, is capable of producing unsightly, painful, and 
even fatal granulomatous lesions. 

I feel that readers of the Journal should be apprised of the 
very real danger involved in the proposed use of the tubes. If 
the age and contents of fluorescent tubes are in doubt they should 
not be used, but should be disposed of in the manner recommend- 
ed by the Public Health Department. 

Please refer to Flint, Thos. Emergency Treatment and Man- 
agement (Saunders, 1958), pp. 220-221. 

Sincerely, 
J. V. Gilkey, M. D. 


San Leandro, California 


American Fern Society 
Report on the Cumberland National Forest Foray 

Participants gathered for the foray at the Seven Gables Motel, 
Burnside, Kentucky, during the afternoon and evening of August 
23, 1961. A short briefing was given by Dr. Clair Brown, Presi- 
dent of the American Fern Society, and by Mr. Thomas McCoy, 
leader of the foray. 

During the morning of August 24th two stops were made in 
Cumberland Falls State Park. The first was above the Falls 
where the group walked up the river to observe the ferns on cliffs 
and talus slopes. The next stop was at the Falls, where we ate 
lunch and looked for ferns in the immediate vicinity. The no- 
table find at that station was Trichomanes radicans (T. boschi- 
anum) made by Conley Webster. He guided three different 
groups to the colony of this attractive little fern. 

The drive to Natural Bridge State Park, the only long one of 
the foray, took most of the afternoon. The night was spent at 
Hemlock Lodge in the Park. 

During the next morning and early afternoon we made several 
short trips in the area, visiting Natural Bridge, Nada Tunnel, 
Sky Bridge, and Devil’s Kitchen. Spectacular scenery furnished 
attractive backgrounds for the many ferns observed. In the late 


AMERICAN FERN Society 193 


afternoon we drove to Morehead College, where we stayed over 
night in the college dormitories. 

On August 26 we drove to Carter Cave State Park where we 
spent the forenoon and ate lunch. In addition to several ferns, 
the Crane-fly Orchid (Tipularia discolor) and the Three Birds 
Orchid (Triphora trianthophora), both in flower, attracted a 
great deal of attention although a few persons had seen both at 
Cumberland Falls. After lunch the group began to break up, 
but a few went on to explore Sandy Hook. 

The 33 persons who participated in all or part of the foray 
were indebted to Mr. McCoy, “the only McCoy who got away 
from the Hatfields,”’ for his careful planning, meticulous ar- 
rangements, and able leadership. 

Those attending the foray were Robert G. Aborn, Earl Bishop, 
Dr. R. F. Blasdell, Dr. and Mrs. Clair A. Brown, Mrs. Muriel P. 
Brown and daughter and grandson, Dr. and Mrs. Lewis Dick- 
inson, Frederick Dunlap, David L. Emory, Mr. and Mrs. Ed- 
ward G. Heinzelman, Dr. and Mrs. L. K. Henry and two sons, 
Dr. Donald G. Huttleston, Mr. and Mrs. J. W. Kunneke, Mr. and 
Mrs. John T. Laitsch, Mr. and Mrs. T. E. Landry, Thomas N. 
McCoy, Miss Lillian McKee, Mr. and Mrs. G. H. Peters, Dr. 
Elizabeth L. Sawyer, Miss Mabel Slack, Miss Eva Sobol, and 
Conley Webster. Special mention should be made of Earl Bishop 
who, although a 1961 high school graduate, was more familiar 
with the native ferns of the area than were most of the partici- 
pants, including this reporter. 

The following 46 species, varieties, and forms of ferns were 
seen during the foray: Adiantum pedatum, Asplenium bradleyi, 
A. cryptolepis, A. montanum, A. pinnatifidum, A. platyneuron, 
A, platyneuron var. incisum, A. resiliens, A. trichomanes, Athyri- 
um filix-femina var. asplenioides, A. filix-femina var. michauxn, 
A. pycnocarpon, A. thelypterioides, Botrychiwm dissectum f. ob- 
liquum, B. virginianum, Camptosorus rhizophyllus, Cystopteris 
bulbifera, C. fragilis var. protrusa, Dennstaedtia punctilobula, 
Dryopteris goldiana, D. hexagonoptera, D. marginalis, D. novebo- 
racensis, D. spinulosa var. intermedia, D. thelypteris var. pubes- 


194 AMERICAN FERN JOURNAL 


cens, Equisetum arvense, E. hyemale var. affine, Lycopodium 
co Sa ee var. flabelliforme, L. lucidulum, L. obscurum, L. 
porephilum, L. tristachyum, Lygodium palmatum, Onoclea sensi- 
bilis, Osmunda cinnamomea, O. claytoniana, O. regalis var. spec- 
tablis, Pellaea atropurpurea, P. glabella, Polypodium polypodio- 
ides, P. virginianum, Polystichwm acrostichoides, P. acrosticho- 
ides f. incisum, Pteridium aquilinum var. latiusculum, Selaginella 
apoda, and Trichomanes radicans. 

It was undoubtedly the aspleniums that received the greatest 
amount of attention, with a considerable bit of discussion—almost 
argument—over species and hybrids.—Donatp G. HuTTLESTON, 
Longwood Gardens, Kennett Square, Penn. 


NEw MEMBERS 
Mrs. Roy Anderson, 511 Lake Street, Lawrence, Kans 
i 


Mrs. Paul B. Barton, 4838 Rolling Hills Road, Pittsburgh 36, Penna. 
Miss Sara Olive hci Ce Route 1, Box 106, Anderson, South Carolina 
Miss Louise Brehm, Tal e, Nebraska 

Mrs. John H. Butler, doe oe erry Bottom Road, uke Ohio 

Mr. R. K. Ch 


: fashion 
Mrs. A. Dorvue elie: 701 Eagle Bend Rond,: Pee er 
Worth 12, Texas 


Mrs Ohio 
Mr. ‘Ralvh M. Evans, Eastman Kodak Co., Kodak Park Bldg. 65, Rochester 


Mr. Donald B. Foster, 220 Triphammer Road, Apt. 2B-3, Ithaca, N. Y. 
Mrs. Priscilla G. Fowler, 2845 Carlisle Road, Birmingham 13, Alabama 
rs. : i 


. Graves, 4606 Ortega Blvd., Jacksonville he Florida 


as 
Mrs. Don W. ines 113 W. How wx Street, Grint, North Carolina 
Mr. William §. Hart, P. 0. Box 583, Valdosta, Geo 

Mr. J. M. ee 3507 Eastis Drive, a seg Car olin. 

Mrs. Benjamin C. Howard, Amberley, R.F.D. 2, Annapolis, Magni 
Miss Se Hyland, aay Fie Tree Drive, Miami Beach 40, Florida 
Mrs. Claude E. Hynds, R. D. 2, Delanson x 

Miss Surjit Kaur, National Botanic Giiders, Lucknow, India 


. Mr. Gladstone W. McDowell, 6 Oak Ridge Road, Asheville, North Ca 


AMERICAN FERN SOCIETY 195 


Mrs. Dale R. King, Rt. 1, Box 61, Crescent City, Florida 
Mrs. Robert Kingston, 1420 West Sonoma, Stockton 4, Calif. 
Miss Margaret A. Koopmann, 2732 Latonia Blyd., Toledo 6, Ohio 


Mr. Curry J. Lacour, Box 154, Marksville, Louisiar 

Mrs. Philip D. Macbride, artis Farm, We sodinr ile Washington 
Miss Edna G. Merritt, Route 1, Box 423, Monticello, N. 

Mrs. illar, 178 Floral et Pahokee, Flori 


a a 

Mrs. Siowos S. Mitchell, 1150 Yew Street, San Mateo, Calif 
Mr. Philip W. Murray, 162 Mar Monte Court, grey Calif. 

. Lars Noren, Spinnrocksvigen 14", Bro on eden 
ey Hideaki Ohba, 157-6chome, Ojimacho, ub , Tokyo, Japan 

. H. G. Pack, 4308 Modoe Road, Santa a Cali 
rit eH: Patten, 2717 So. Hill Street, Los Angeles 7, Calif. 
Mr. Ray M. Petersime, a sburg, Ohio 

ard A. Pettis, 1120 River Road, AERA, Mass. 


at. 
Mrs. P. Blake Saunders, 58 Highland Avenue, Buffalo 22, N. Y. 
Dr. Elizabeth L. Sawyer, Department of Biology, Greensboro College, Greens- 
oro, North Carolina 
Mrs. E. K. Schlesinger, 1508 Rolling Road, tae 27, eee 
Mr. W. G. Shields, 8316 4th Avenue, N., Birmingh labam 
Mrs. George J. Skala, 3350 Green Road, Beachwood aus. Cleveland 22, Ohio 


Miss Helen E. Steyaert, 4436 Irvington Avenue, Fremont, Calif. 
Mrs. nb Van Orman, 83 Old Short Hills Road, me ‘Hills, N. J. 
Mrs. W. L. Whyte, 22482 Argus, Detroit 19, Michig 

Mrs. hae’ Wilburn, 1400 East Texas Avenue, Bay ‘a own, Texas 
Mrs. Marilyn Williams, 53 South Udall Street, Mesa, Arizona 
Mrs. L. B. Windham, Rt. 5, Box 110, Tyler, Texas 


CHANGES OF ADDRESSES 


Mr. Lee Andreas, Wisconsin State College, Stevens Point, ——— 


Mr. Boughton Cobb, 116 East 66th Street, New York 21, N. Y. 

Dr. Hans Peter Fuchs-Eckert, Clematislaan 61, Wassenaar (ZH), Netherlands 

Mr, Ralph S. Hudson, Route 2, Box 1205, Nokomis, Florida 

i rolina 

Dr. John T, Mickel, pee of Botany & Plant Pathology, Towa State 
University, Ames 


Miss Mary H. oa err Winlow Street, San Diego 5, Calif. 


196 


AMERICAN FERN JOURNAL 


Index to Volume 51 


Abacopteris, 87 

Abies balsamea, 36 

Acer Psi ov 185 
Acrostichum pe 
Actinostachys Corman 1 
Adaxial Sori in Poipteanie hesperium, 


Addition to the t of Ferns Growing 


es 
Adiantum capillus-veneris, 58 


105, 130; cuneatum, 130; Saas Py 

130; hispidulum, 130; melanoleucum, 

14; pedatum, 26, 58, 60, 130, 167, 
; tenerum, 

Srosue tenuis, 184 


reptans, 185 

Mets Chiarugi 1901-1950, 156 

Alnus incana, 183-185 

Another Asplenium Hybrid from Ken- 
tuc 70 


se of Ferns New to the 


odoratum, 184 


trichomanes, 60 3; 
Trudellii, 70; varians, 39; erecundum, 
2, 3, 5, 7, 8; viride, 128; x W erryi, 
Asplenosorum igen des, 45 
oo m, Pick angustatum Goe 
um, 73. Bapledioides, 59: 
distentifoliam, 127; Filix-fe 
5. A 


ngianum vy. pictu 
ay 58, 68, 167, 193; ‘hay. 
Asa” 167, 193 
eee oe tte R. Pellaea atropur- 
Azolla, — africana, 151 : hn i 
iana, 89- OL 26h. filicnloides, 91; 
bricata, 151, 153; japonica, ‘153: 
Hangar 153; age cae 91; micro- 
phylla, 91; pinnata, 151, 158, 154 


Bartramia pomiformis, 183 
Basket Ferns for Southern California, 


Benedict, Ralph H. Cystopteris tennesse- 
ensis Shaver, 43 
Betula populifolia, fa 4 
aes Edith. The Bien Valley at the 
ited States National mite nis 
iden 131 


ale ae 9: ceed 


Bloom ; William W. Heat Resistance of 
rept ied of Marsilea quadrifolia, 95 
Boi oo ard. Isoletes echinocarpa 
Du i g. Novi kon erie. 68 
Botrsehiua borew ue, 128; dissectum, 31 
33 


32, 75-81, 175, 176; simplex, 41; ter 
natum, 31-33, 79; virginianum, 36, 42, 
9, 58, , 298, i anum, 36, f 
anomalum, 36, v. europaeum, 36, 


37, f. het kecodactini, 36, 
Boydston, Kathryn EK, Report of Spore 
e, 1 


roble — on Vari 
teris, 23 


tion in North Am ue te: Dry 
Broun, Maurice Pastideingicat Troglo- 

dytes, 190 
Brown, Clair A. Report of President, 49 
Bryant, Truman R. See Smith, oo M. 
Buker, W. E. See Heat ry, Li. 
Caltha palustris, 176 

60, 68, 


Camptosorus rhizophyllus, 58, 
167, 193 


Camylon neuron a 14; costa- 
tum, 14; phyllitidis, 

Carex flava, 42; Sean. 184; plan- 
taginea, 


Ceratopteris pteridoides, 4, 
Chaerophyllum hirsutum rh “186 
eo alabamensis, 58, 61, 
60; lanosa, 61, 68; caorepayiie: 
greta entosa, ia 68 
Chebrotoass palm: 3 
Chortophila sign ore "183, 184 
Ceca | ee 185 
Climac 
Cody, W. J. 
virginianum, 36 
Concerning ‘Azolla imbricata, 151 
Conocephalum. is 
Correll, Donovan S. Den neimer oe globu- 
lifera (Poir.) “Hieron. in Texas, 188 
Corylus avellana, 


Ne ew Forms in Botrychium 


41 
ides, 88 
©. e, 86-88; faleatum, 47 
a ft herrea 59, 60, 167, 293, 
x fragilis v. protrusa, 43; fragilis, 


INDEX TO VOLUME 51 197 


ri 68, 128, mee YF Pitas ap 69,193" 
ennesseensis, 
Suasoene spa ag get Shaver, 43 
Cytology of Isoétes coromandelina, 99 


as, Jr. Florida Rarities, 1 

Davallia canariensis, 187; fijiensis, 187, 
188; trichomanoides, 187, 

Dennstaedtia glauca, 131;  globulifera, 
188; punctilobula, 59, 193; rubiginosa, 

Dennstaedtia gceniet (Poir.) Hieron. 
in Texas, 1 

Descha mpsia cexpitosy, 185; bi Sas 183 


38; aaa 82; 
onan fo lis. 


tris, 385 stralis, ; austriaca 
austriaca, 29, v. concordiana, 29, v 
ructuosa, 29, v. 29, v 


34; com 38; cristata, 23, 24, 
3 167 i 


rae’ x t m 
29; spinulosa, 93, 25, 26, 29, 165, 


7 5 concordiana, 29, 
fructuosa, 36, "20. v. Cpe cers 23. 
29 y. spinulosa, Tatei, 38; 
. Tavelii, 180-182, ave Lhervotae is 

pubescens, 29, 193; Thomsonii, 38; 
wigiates, 47; viridescens, 171, 173 


Dryopteris . Tavelii in the Valley of 
ce, 180 


altum, 133; pyramidal 13 ramo- 
3. Rap: aie, 188; 
133, us 


: e 
-134; c. ie 

boldii, 132; trachyodon, 132, 133; 

eae ngage 42, 182-134, ¥. alaskan num, 

134, v. variegatum, 134; xylochaetum, 

132, 133 

Fagus sylvatica, 183, 185 

Fern Monster, ?, 

Fern Valley at the United States National 
Arboretum 


ra tess 
Filix <X Dowellii, 3 34. x rcecul. 34 
Fissidens, 59; cristatus, 183 
Flifiet, Tho: rleif. Growing Ferns From 
Ae 


pores, 
rida Rar 
Lier posh qo 29 
Fosberg, F. R. Southern Distripation of 
B chium oneidense and B, multi- 
canes vs 
Fragaria vesea, 184, 185 


Prarbnis excelsior, 185 

Gentiana rubricaulis, 

Genus Grammitis in Japats with Descrip- 
tion of a New Species, 148 

Godfrey, R. K., Grady W. Reinert, and 
Richard D. Houk. Observations on 

rpic Material of Azolla 


ca 

ipearpita: 148-150; nipponica, 
eae 

Grim 559 

pide ‘ae "Ferns from Spor 

G i y 
ober tian ia. 17-19, Re 

Habenaria clavellata, 


Hartman, Emily L. “Adair Sori in Poly- 
podium hesperium, 


198 


Hauke, tee = _A Resume of the 
onomic Reo: ization oY, waeieen, 
enn s ainpethonts Bee SS 3 
awailian “gar ae roid Fern with Pel- 
tate Indusia, 
Heat Resistance oe ee of Mar- 
silea ~ “tig ifolia, 
De i ine, 1 
mry, XK. d W. E. Buker. Recent 
week Boscoveris in Western Pennsyl- 
vania, 
Wore cast murorum, 183, 18 
Hevly, Richard H. See Mason, TChiaivies 
ay OEe 
orties Jarl me Records of 
Michi; es ig ae Fern Alli ies, ee 


ae, D. See vsewpind R. K. 

Humata recto 18 

Huttleston, D. G. Report nm Cumberland 
paseast Forest Foray, 192; of Secre- 


Hut it, “Tho s A. An Interesting Ecolog- 
ical Niche. Tavehing Salvinia rotundi- 
folia Willd. 

ayerise in North American Gymnocarpi- 


leon um tego ge 183, 185 
Hymenophyllum, 

Hypnum Pcowse} ae 183 
Hypolepis punctata, 130; repens 6 


Interesting Ecological 

Salvinia 
Tsoétes 

America, 

serge. asiatica, 85, ul Braunii, 84, 

ritim acnaetag: 99, 

. oe 


Niche Involving 
rotundifolia Malo 138 
0. Durieu in North 


vi ba als 
84, v. Savilei, 8 
ata, 84 i 
maritima, 84; uricata, 83, 
ria, : Nuttallii, 85; sa) hyaidh, 
9; sampa athkumaranii, 99, 102 


“The ' Gen us Grammitis in 
Japan, ‘With temnerden of a New 
Species, 148 


Juncus articulatus 42 
Juniperus Ashei , BO; virginianum, 59 


Knobloch, Irving W. Obituary: Alberto 
Chiarugi, 1901- 1960, 156 


AMERICAN FERN JOURNAL 


decidua, 185; 


Larix Jaracina, 40 
tae, 37 


Lawalrée-Collaris, André, and Anne- 
Marie. Dryopteris Tavelii in the 
Valley of Chamonix, France, 

Leatherman, Sylvia. Ferns in B irtaghls 
tion, IV. Some Wall Ferns, 

Lithobrochia RN Maht 14 

Lom st rt C. Report on 1960 


ohan 
ine cocR: bidentate: 185 
Lophozia barbata, 185; quinquedentata, 


184 

Love, Askell, and Bing Léve. Some 
Chromosom Nw of Icelandic 
Ferns and Rech. ales, * 107 

Lveopadiumn cernu 8; clavatum 40; 
complan atum Sy AAUETROEIIC. 194; 
inundatum, 40; ulum, 194; o 
scurum, 194; porophilum, 194; sabini 
folium vy. sitchense, 41; Selago, 41, v. 
Aged 41; sitchense, 41; tristachyum, 


41, 194 
ies palmatum, 42, 167, 194 


MacFadden, Basket for 
Southern’ California 186; vere ers a 
aaa House, 129; My Lath House for 


Fer: 

Matanthinns oh bifolium. 

MeDowell, Vos stie mene we Frankl 
‘ans Parcel of "Pems ood 
taphonte “Gounty, Georgia. 

Marcing quadrifolia L., a ‘In 

gain, 191 
Marsilea, aig coe 157; quadrifolia, 95, 


chcae A- 


Moan Charles T. Jr., and Richard H. 
Hevly. Report of Aadi iting Committee, 

Matteuccia Str As tee ry 167 

Maxonia apiifolia 82 

Mecodium 0) 


Menyanthes trifoliate 76 

Microlepia “platyphyla, 10, 174; spelun- 
cae, ark ae raeet, 129, 174, f. Mac- 
Faddeniae. 

Morton, r Genus of Ferns 
New to the Gaited "states, 81; Some 
Forms of ie A eae californicum, 73; 
Taxonomic Not : 

My Lath House for Forn 

Nephrodium macradenium, 

Nephrolepis cordifolia, 6 

New ms in Botrychium virginianum, 


36 
Notes on Polystichopsis, 145 
Notholaena dealbata, 61 
Nuphar advena, 191 


Observations on Microsporocarpic Ma- 
terial i org lla caroliniana, 89 
a Mecodium wrightii in Can- 


} 


INDEX TO 


O'Dell, Herman. An Addition to bet ug 
of Ferns Found Growing Natur: in 
Tennessee, 156 

On the Relative Devel lopment of the Fer- 

tile Spey in Botrychium dissectum 
a B. oneidense, 75 
Onoclea eenainilia, 62, 158, 194 
pa ts cosa nudicaule, 1; tenerum, 1; 
vulga 42, 102, 128 


nd cinnamomea 62, 167, 194; 
Claytoniana, 58, 167, 194; regalis, 
167, ¥. ee a 194 
Oxalis’ acetos ella, 


A Parcel of Ferns from Stephens County, 
Georgia, de 
Pellaea atro rea, 44, 45, 58, 60, 61, 
68, 167, 194; slabella, 60, 194 
Peltapteris nanan ate f. Standleyi, 
98; peltata, 98 i: "foe iculacea, 98, f. 
peltata, oS £; potentillifolia, 98 
Peltapteris in ee i 
Sehaces. horizontalis, 
Phegopteris eonnanton. 128; lurida f. 
leucocha ete, 147; sericea 146; villosa, 


ca, 


Phillips, Walter S. Report of the Trea- 
surer, 
Phiebodiuse aureu 186 
Phymatodes heterophylium, 2 
Picea ahi. 183-185; mariana, 40 
i aI 


ityrogr: ane: triangularis, 105, 106, ¥. 

a a 106, PARE are oak 106, v. vis- 

; pallida, 106; viscosa, 106 
Plaplochatle’ asplenioides, 185 


Platycerium yey catum, 
Riishouver 
Polypodium, gs : ah < 8; 
aureum. 6, cv. Mandaianum, 186; 
187, Ss 


146; po! lyear- 
pon, 188; polgpod folded, 59, 167, 194; 
punctatum 6, 


bia. 
Polraenate 82; abla oo — 
hyllo: ides, 145, 46, 
146 , 145 
q 4 chroptero- 
ides, 145, 147, 148: ght iene 145, 
146, v. haitiensis, 1 
Polystichum, 81; agers age: 183, 185; 
acr i 0, 100; - 104, 2 
incisum, 194; apiifoliui, 82; Braunii, 
; 158; capense, 174; 


mente | 130, 158; lonchitis, 128; 
m 


, 85 
Th roblem of scitriation in North 
American Dryop 23 


VouuME 51 199 


Proctor, George R. Notes on Poiystichop- 
sis, 145 

Prunella vulgaris, 184, 185 

Psilotum nodum, 6, 10 

pets ida pov ibe 40, 130, v. latius- 


Pteridological be pe 190 

Pteris cretica, albolinea 
Childsii, 5g pos rink ad 2a 
dentata, 129; grandifolia, _ — 
— t quadriaurita, 12 


a, 
zs 9; Arg: 
129: tremula, 129: teipartiine vi 
Ptilidium crista- castrensis, 185 
Pyrrosia lingua, 188 
Recent Fern Discoveries in Western 

Pennsylvania, 
Reed, “Olvde KF. Marsilea quadrifolia fa 


in neg es 191; Peltapteris 

Costa Rica 

Bagi Grad e Godfrey, R. K. 
port of: A ‘oray, 193: Audit- 


if xonomic Re Soepaate bell 
of gta ty Bocas Hippochaete, 
81 
Rhabdoweisia ever: 183 
ern uaticum, 183;  hetero- 


ichum 18 

Rhipidoptoris, 97; peltatum v. potentil- 
pe prin 98 

Rhodobryu 58; roseum, 185 

Rhytidiadelpias loreus, 185; triquetrus, 
183, 18 

Rogerson, ant T. Report of Judge of 
Elections, 

Root, Elizabeth Eichstedt. Hybrids in 
North American EOS carpiums, 15 

Rubus, 184; idaeus, 

Rumohra, 145; edicontotecbe 130, 174; 
aristata, 13 


Salvinia auriculata, et oe lurcisaiiaey 138 
Saxifraga ener ete 
tridiu 179. 


ella apoda, 59, 194; arenicola, 11; 
pes 59; selaginoides 128 
Eugene Yu-Feng. Concerning Azolla 


'ruman R. Bryant, and 

" Another Asplenium 
Hybrid from aged 

Snyder, is nklin hg McDowell, 
1 vin aurea, 183, 185 

Soe hseus : ome Numbers of Icelandic 
Ferns pad ey rest allies, 127 

rms of Polypodium californicum, 


Some New Data on the Vernation Differ- 
pew bell Batryehium dissectum and B. 
terna’ 


200 AMERICAN FERN JOURNAL 


Some Records in Michigan Ferns and Thuidium, 58 
Fern Allies, Thuja occiden aa 36, 40 
Sorbus a avae 185 ip 
Southern Distribution of Botrychium Tortella SaPiea e 
oneiden nd B. multifidum, 175 ee tae 144; Boschianum, 
102; 


192; 
Sabacannn 5 Krausii, 9, 14; Tineolatum, 
Sphenomeris clavata, 8, 9 tiny “Pe eters, 69; punctatu Pee 
Spiranthes cernua, radic 192, 194 
Stenochlaena Kunzeana, 9 PB open tr rianthophora, 193 
sis RT ety Note on Dryopteris Hy- Trismeria trifoliata, 9 
rids, 33 


Genera 40; Myrtillus, 183, 184; 


Tagawa, Motozi, and Kunio Iwatsuki. itis-idaea, 185 
The Genus Gra msne in Japan, with ver rma, Ss. ne Cytology of Isoetes coro- 
Description of a New Species, 148 ndelin a 

Takakia lenideaioiden: 14 Veronica latifolia, 183, 185; cfficinalis, 
‘ate, Donald E. See Smith, Dale M. 184; urticifolia, 183 

Taxonomic Notes on Ferns, II, 37 Viola, 184 

Tee ia se er rege 3, 9; coriandrifolia, 
3, minim Wa Warren H., Jr. On the ep 

Thelypter: ee Moana. 129; andina, 38; Disclowniont: of the oe Segn 
boliviensis, 38; Drasiliensis, 38; comosa, Botrychium _— an a Ouaee 
38; hexagonoptera, 23, 29, 167; limbo- dense, 75; Report of Curator 
sperma, 184, 185; jurida f. leucochaete, Librarian, 57; Some new Data on the 


p Ff Where Edgar cppiementeey Note 
167, 183, 185; polyphlebia, 38; resini- yopteris Hybrids, 

11; reptans, 5; scler lla, 13; Woodsia alpina, 8; ilvensis, 128, 167; 
se seti; wii : Leo: obtusa, 58, 61; scopulina, 105 : 
Tatei, 38; teen agona, 6; subg. Seen areola ata, 42, 59; radicans, 
teris sect. Claphyrenteris: 37 7, 130; virginica, 167 

ERRATA 
Vol. 50, 
Page ae ae For “from Darjeeling and Mussoorie respectively,” 
read “from =P te rie and Dar. obi ling respectively.” 
Page 278, hace gy for figures 1, 2, 7, 8, 9, 10: For “500,” read 
“1000,” a d for fig. 3, for "150," read 300. 
Page 282, 1s e 6: For “n ” read ee = 40,7 


gend, figure 6: T 50, 
Page 286, legend, figure 18: For 35, 2 oe 100.” 
Page 296, line 7: For “cet,” read ‘ 


51. 
Page 5, line 5: For “or,” read “of.” 
age 9. 


Page ies Transfer footnote 7 sd page 13. 
Page 16. Plat bs is upside dow 
Page 17, line 6: For “tht,” ad “that 
Page 20. For “G ymnoearpiune,” read “Gymnocarpium.” 
Page 26, line 16: F which, as one would expec e quite variable, 
alt ough many,” Tead “tal entities are involved is ppc oper favored, 
many.” 


; 10 to. 22 of last eae peo should be lowered one line. 
Page 70, line 7 te or “Maxon,” read “Maxon.’ 
ge 71. 


Add ‘to pry of plate: eines arrows indicate farthest extent 
of darkening PM e or rachis 

Page 148. Item 2 in Tebliogs raphy: For telat » read “Christensen.” 
Page 150, line 10: For of, yO rea ¢ 


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