[ND AGS.
N. MANCHESTER,
THE HECKMAN BINDERY, iNC.
AMERICAN
FERN
JOURNAL
Volume 70
1980
PUBLISHED BY THE AMERICAN FERN SOCIETY
EDITORS
David W. Bierhorst
Gerald J. Gastony
David B. Lellinger
John T. Mickel
MERCURY PRESS, ROCKVILLE, MARYLAND 20852
CONTENTS
Volume 70, Number 1, Pages 1-32, Issued March 28, 1980
Intersectional Hybrids in Isoétes BRIAN M. BOOM
Notes on Some Pleopeltis and Polypodium Species
of the Chihuahuan Desert Region TOM WENDT
The Deletion of Vittaria graminifolia
from the Flora of Florida GERALD J. GASTONY
New Taxa and Combinations of Pteridophytes
from Chiapas, Mexico ALAN R. SMITH
Shorter Notes; A New County Record for Pilularia americana
in Texas; Asplenium = gravesii Discovered in Arkansas;
Pilularia americana New to Tennessee; New Names for
Polypodium chnoodes and P. dissimile; A New Record for
Pellaea atropurpurea in Maryland; An Atypical Athyrium
from Eastern Tennessee
Reviews 4,
Volume 70, Number 2, Pages 33-80, Issued June 30, 1980
Equisetum * litorale in Illinois, lowa
Minnesota, and Wisconsin JAMES H. PECK
Gametophytes of Equisetum diffusum RICHARD L. HAUKE
A Double Spore Wall in Macroglossum LUIS D. GOMEZ P. and KERRY S. WALTER
Subdivision of the Genus Elaphoglossum
JOHN T. MICKEL and LUCIA ATEHORTUA G.
Notes on the Natural History
of Stylites gemmifera ERIC E. KARRFALT and DALE M. HUNTER
Reciprocal Allelopathy Between the Gametophytes
of Osmunda cinnamomea and Dryopteris intermedi
AYMOND L. PETERSEN and DAVID E. FAIRBROTHERS
Shorter Note: Thelypteris torresiana in Venezuela
Reviews 38, 68,
bho
39
Volume 70, Number 3, Pages 81-112, Issued September 29, 1980
The Distribution and Ecology of Phyllitis scolopendrium
in Michigan RICHARD P. FUTYMA 81
Supplemental Notes on Lesser Antillean Pteridophytes GEORGE R. PROCTOR 88
Additions to the Pteridophyte Flora of the Great Plains RALPH E. BROOKS 9]
Flavonoid Synthesis and Antheridium Initiation
in Dryopteris Gametophytes
RAYMOND L. PETERSEN and DAVID E. FAIRBROTHERS 93
Date of Publication of Sodiro’s
“Sertula Florae Ecuadorensis”’ DAVID B. LELLINGER 96
Reproductive Biology and Gametophyte Morphology
of New World Populations of Acrostichum aureum ROBERT M. LLOYD 99
Shorter Notes: Diplazium japonicum New to Alabama;
ths and Ferns; Three Additions to the
Pteridophyte Flora of Escambia County, Florida 11
Reviews 92, 98
Volume 70, Number 4, Pages 113-140, Issued December 30, 1980
A Range Extension for Dryopteris filix-mas ERWIN F. EVERT 113
Differential Germination of Fern and Moss Spores in Response
to Mercuric Chloride RAYMOND L. PETERSEN and PATRICK C. FRANCIS — 115
Differences in the Apparent Permeability of Spore Walls and ; .
Prothallial Cell Walls in Onoclea sensibilis JOHN H. MILLER 11
Allelopathy and Autotoxicity in Three Eastern 74
North American Ferns WILLIAM E. MUNTHER and DAVID E. FAIRBROTHERS = |2
Shorter Notes: Sandstone Rock Crevices, an Exceptional New Habitat
for Thelypteris simulata; A Second Alabama Locality for
the Hart’s-tongue
8
American Fern Journal :
Ee edt
a at :
Pied aieentos ee
AM ERICAN Volume 70
FERN ee
J 0 1 R NA L January-March, 1980
QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY
Intersectional Hybrids in Isoétes BRIAN M. BOOM I
— a Some Pleopeltis and ce ete Species
f the Chihuahuan Desert Regi TOM WENDT 4
The — of Vittaria graminifolia
e Flora of Florida GERALD J. GASTONY 12
New Taxa and Combinations of Pteridophytes
from Chiapas, Mexico ALAN R. SMITH _ 15
seucien: ree A New ag Record for Pilularia americana
as; Asplenium x gravesii Discovered in Arkansas;
cats americana New to Tennessee; New Names for
Polypodium chnoodes and P. dissimile; A New Record for
Pellaea atropurpurea in Maryland; An Atypical Athyrium
from Eastern Tennessee
Reviews
yaah
GARDEN LIBRARY
The American Fern Society
Council for 1980
ROBERT M. LLOYD, Dept. of Botany, Ohio University, Athens, Ohio 45701. President
DEAN P. WHITTIER, Dept. of Biology, Vanderbilt University, Nashville, TN 37235. Vice President
LESLIE G. HICKOK, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916.
Secretary
JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916.
Treasurer
JUDITH E. SKOG, Dept. of Biology, George Mason University, Fairfax, Va. 22030.
Records Treasurer
DAVID B. LELLINGER, Smithsonian Institution, Washington, D.C. 20560. Journal Editor
ALAN R. SMITH, Dept. of Botany, University of California, Berkeley, Calif. 94720
Memoir Editor
JOHN T. MICKEL, New York Botanical Garden, Bronx, N.Y. 10458. Newsletter Editor
American Fern Journal
EDIT
DAVID B. LELLINGER Smithsonian Institution, Washington, D. C. 20560
ASSOCIATE EDITORS
DAVID W. BIERHORST Rt. 3, Box 188, Picayune, MS 39466.
GERALD J. GASTONY Dept. of Biology, Indiana University, Bloomington, Ind. 47401
JOHN T. MICKEL New York Botanical Garden, Bronx, New York 10458
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AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 1 (1980) 1
Intersectional Hybrids in Isoetes!
BRIAN M. BOOM*
Engelmann (1886) and Campbell (1891) have described the simple procedure for
germinating /soétes spores in the laboratory to obtain micro- and megaga-
metophytes. Perhaps the absence of hybridization studies in this genus can be
explained partly by the fact that sexually mature living plants of more than one
species were rarely assembled at one time, or that when they were, the primary
aim was comparative embryology (La Motte, 1937). The present study was under-
taken to test for genetic compatibility among four selected /soétes species in three
sections of the genus (sensu Pfeiffer, 1922). All crossing combinations produced
progeny, and in at least one species apogamy may occur.
MATERIALS AND METHODS
Spores for the crosses were obtained from populations of plants as follows:
Isoétes (Reticulatae) macrospora Dur.—Monroe Co., TN, Boom 318, Shenan-
doah Co., VA, 8 Nov 1978, Evans; I. (Reticulatae) engelmannii A. Br.—Polk Co.,
TN, Boom 317, Putnam Co., TN, Boom 267; I. (Tuberculatae) flaccida
Shuttlew.—Dixie Co., FL, Boom 313, Wakulla Co., FL, Boom 314, 315; 1. (Cris-
tatae) riparia Engelm.—Tyrrell Co., NC, Boom 316. Voucher specimens have
been deposited in the Herbarium of the University of Tennessee (TENN). All
plants were collected during the summer or autumn of 1978, and were grown in the
greenhouse for a short time until the crosses were made in mid-January, 1979.
The crossing technique was quite simple, yet rigorously controlled. Forty-eight
glass vials were filled with about 1 ml of sterilized fine sand and 10 ml of sterilized
pond water. Each sporangium was dissected out of the sporophyll base, washed in
a sterile water bath, and then teased apart to release the spores into a vial, taking
great care to insure that the vials were not contaminated with unwanted spores.
Since microsporangia and megasporangia are usually found on the same plant,
spores were taken from completely intact sporangia to avoid the possibility of
using megagametophytes which already had been fertilized.
The crosses among the eight populations were set up in such a way that the
megaspores of each population were brought into contact with the microspores of
every other population. To test for spore viability and self compatibility, one plant
from each population was selfed. To provide controls for the crosses and to test
for apogamy, one vial was set aside for each population in which only megaspores
were placed. All vials were kept in the greenhouse at about 25° C, where they
were exposed to normal ambient light fluctuations, and were not disturbed except
for the occasional addition of sterile pond water.
*Department of Botany, University of Tennessee, Knoxville, TN 37916.
‘Contribution from the Botanical Laboratory, The University of Tennessee, N. Ser. no. 516.
Volume 69, number 4, of the JOURNAL was issued December 31, 1979.
AMERICAN FERN JOURNAL: VOLUME 70 (1980)
FIGS. 1-2. Sporophytes resulting from a cross between Isoétes flaccida and I. macrospora, two
species thought to have had very different evolutionary histories. FIG. 1. Sporophytes in vial at two
months after the cross was made. FIG. 2. Hybrid sporophytes * four months.
B. M. BOOM: INTERSECTIONAL HYBRIDS IN ISOETES 3
RESULTS AND DISCUSSION
About seven weeks after the crosses were made, the first green shoots of young
Isoétes sporophytes were observed in a number of the vials (Figs. ] and 2).
Although not every individual cross was successful, every possible crossing com-
bination was successful within the first two months of the experiment. None of the
control vials showed any signs of growth at that time, and the genetic compatibil-
ity of the four species presumedly has been demonstrated.
No new sporophytes were observed until 3.5 months after the experiment be-
gan, when a single sporophyte appeared in the /. macrospora control vial (Boom
318). Since no microspores had been introduced into this vial, either the
megagametophyte had somehow become fertilized before it was introduced into
the vial or the species is capable of reproducing apogamously. Considering the
precautions taken by using only fully intact sporangia as a source of spores,
apogamy seems more likely. Eight months after the crosses were made, no
sporophytes had developed in any of the control vials of the other three species.
Easily hybridized species of Jsoétes means that hybridization followed by
polyploidization may be a mode of evolution from time to time in the genus. The
occurrence of facultatively apogamous taxa is consistent with such a process. If,
as is suggested by the experimental observations, /. macrospora can be apogam-
ous, this could help explain the Virginia and Tennessee populations disjunct from
the typical northeastern range of this species (Dennis et. al., 1979). The reticulate
distal face and the cristate proximal face of the megaspores of J. macrospora
suggest a possible hybrid origin for this species.
Some Jsoétes populations on the Coastal Plain of the Carolinas have various
characters, primarily megaspore ornamentation, which clearly are intermediate
between typical /. engelmannii and/. riparia; supposedly these plants are hybrids
between the two. Specimens from such populations occasionally have been anno-
tated as /. engelmannii var. georgiana Engelm. or var. caroliniana Eaton.
The results of this study also support Matthews and Murdy’s (1969) interpreta-
tion of the often confusing /soétes populations on the granite outcrops of the
Piedmont of the southeastern United States. Introgression apparently is taking
place in pools which are ecologically intermediate between the habitats typical of
I. piedmontana (Pfeiffer) Reed and those of /. melanospora Engelm. For an alter-
nate explanation, see Rury (1978), who suggests that intermediates represent de-
velopmental stages of one polymorphic species.
The naturalness of Pfeiffer’s (1922) sections of the genus is suspect now more
than ever in light of the artificial intersectional hybridizations. The infrageneric
classification of Isoétes should be reexamined by means of an extensive genetic,
cytogenetic, and phytochemical survey, as well as by using traditional morpholog-
ical characters. :
This report of intersectional genetic compatibility need not necessarily affect
Isoétes taxonomy at the species level, however. In natural circumstances, the
taxa generally are isolated by geographic, ecological, or phenological barriers, and
they can be distinguished morphologically from one another. The amount of gene
va
: AMERICAN FERN JOURNAL: VOLUME 70 (1980)
flow between the typically isolated populations must be relatively small. If this is
not the case, it remains a challenge to explain why selection has not favored the
establishment of reproductive barriers between species.
The present study was initiated to test the potential for genetic experimentation
in /soétes. The preliminary results were very successful and indicate further and
wider genetic studies would be beneficial. Such future hybridization research
should take advantage of the artificial crossing technique recently described for
Selaginella (Webster, 1979). The method appears to be well suited for /soétes
crossing with little or no modification, and will permit more critical experimenta-
tion than ever could be possible with the non-sterile technique employed in the
present study.
Appreciation is extended to Dr. A. Murray Evans for critically reviewing this
paper. Field work was aided by a Grant-in-Aid of Research from Sigma Xi, The
Scientific Research Society.
LITERATURE CITED
CAMPBELL, D. H. 1891. Contributions to the life-history of Isoétes. Ann. Bot. 5:231-358.
DENNIS, W. M., A. M. EVANS, and B. E. WOFFORD. 1979. Disjunct populations of Isoétes
macrospora in southeastern Tennessee. Amer. Fern J. 69:97-99.
ENGELMANN, G. 1886. The genus Isoétes in North America. Trans. St. Louis Acad. Sci. 4:358—390.
LA MOTTE, C. 1937. Morphology and orientation of the embryo of Isoétes. Ann. Bot. 1:695—715.
MATTHEWS, J. F. and W. H. MURDY. 1969. A study of Isoétes common to the granite outcrops of
he southeastern piedmont, United States. Bot. Gaz. 130:53—61.
PFEIFFER, N. E. 1922. Monograph of the Isoétaceae. Ann. Mo. Bot. Gard. 9:79-232.
RURY, P. M. 1978. A new and unique, mat-forming Merlin’s-grass (Isoétes) from Georgia. Amer.
Fern J. 68:99-108.
WEBSTER, T. R. 1979. An artificial crossing technique for Selaginella. Amer. Fern J. 69:9-13.
REVIEW
“THE ECONOMIC USES AND ASSOCIATED FOLKLORE OF FERNS AND
FERN ALLIES,” by Lenore Wile May, Botanical Review 44:491-528. 1979-—As
stated by the author, this paper is not taxonomic in nature, but discusses fern
folklore and to a lesser extent their economic history. It provides an easily read
text for the generalist and a good bibliography for those persons interested in
pursuing this topic further. Some of the section titles include: Folklore, Fern
Dyes, Fern Fibers, Fern Foods, Medicinal Uses of Ferns, The Male Fern, and
The Bracken Fern. The section on medicinal uses occupies forty percent of this
article, with related medicinal notes in the folklore portion. The author mentions
the following about Ophioglossum vulgatum: ‘‘This plant is called adder’s tongue
because out of every leaf it sendith forth a kind of pedestal like an adder’s tongue,
it cureth the biting of serpents.’”-—J. Scott Peterson, Dept. of Botany & Plant
Pathology, Colorado State University, Ft. Collins, CO 80523.
AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 1 (1980) 5
Notes on Some Pleopeltis and Polypodium Species
of the Chihuahuan Desert Region
TOM WENDT”
In preparing a treatment of the Polypodiaceae s. str. for the forthcoming
Chihuahuan Desert Flora (M. C. Johnston, et al.), I found several taxonomic
changes to be necessary. A new variety of Polypodium thyssanolepis A. Br. ex
Kl. is described, and Pleopeltis erythrolepis (Weath.) Pic. Ser. is lowered to
varietal rank within Pleopeltis polylepis (Roem. ex Kunze) Moore. Material from
GH, LL, NY, TEX, and US was consulted in preparation of the treatment.
POLYPODIUM
Polypodium thyssanolepis A. Br. ex KI. is a species of lithophilic fern which
ranges from the southwestern United States to South America and the West
Indies. Originally described from Colombian material (Klotzsch, 1847), it was not
reported from the United States until 1913, when Maxon noted it among the
collections of L. N. Goodding from the Huachuca Mountains of southeastern
Arizona. Maxon (1913) stated that Goodding’s specimens were ‘“‘perfectly typical
of the species as it exists from Mexico to the Andes and in Jamaica.’’ In his
revision of several groups of squamate American polypodies (Maxon, 1916), the
most recent revision in which P. thyssanolepis has been treated, he recognized no
varieties within the species. However, in recording the species from Texas,
Maxon (1923) noted that specimens from both Texas and Arizona “‘have fronds
only scantily scaly beneath, in marked contrast to tropical material.”’
A number of characters are correlated with the sparser indument of the fronds
of these northern populations. Material from western Texas, southeastern
Arizona, northern Coahuila, and parts of Chihuahua appears to represent a
strongly marked new variety of P. thyssanolepis (Fig. 1). None of the synonyms of
P. thyssanolepis (see Maxon, 1916; Morton, 1973) refer to this new variety, which
may be distinguished from the typical variety by the following key:
Stipes sparsley scaly, the scales mostly ovate or lance-ovate, suborbicular scales few or none; scaly
indument of the lower lami fi tsod to ot th ; venation mostly free,
with fewer than 30%%(40%) of the sori within areoles; basal lobes of the lamina distinctly
alternate. P. thyssanolepis var. riograndense
Stipes densely scaly, the scales mostly suborbicular; scaly indument of the lower lamina surface dense,
typically entirely obscuring the surface; venation mostly areolate, with more than 70% (usually
nearly 100%) of the sori within areoles; basal lobes of the lamina opposite or subopposite.
P. thyssanolepis var. thyssanolepis
*Rama de Botanica, Colegio de Postgraduados, Chapingo, Edo. de México, México. ae
!Work accomplished at University of Texas at Austin and Gray Herbarium of Harvard University. I
thank Alan R. Smith of the University of California at Berkeley for unpublished data.
6 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
Polypodium thyssanolepis var. riograndense Wendt, var. nov.
A var. thyssanolepide stipitibus multo minus squamatis squamis suborbicularis
paucis vel absentibus, laminis minus squamatis, venatione libera pro parte max-
ima, et positione lobarum laminarum basilarium distincte alterna recedit.
Small lithophilic ferns. Rhizome slender, wide-creeping, 1.5-2.5 mm thick,
densely scaly; rhizome scales subulate to lanceolate-acuminate, (1.0) 1.5-3.0 mm
long, imbricate, light brown with a darker central stripe composed of dark-walled
cells with clear lumina, the margins slightly erose to irregularly ciliate. Fronds
distant or occasionally a few somewhat crowded, to 15(20) cm tall, usually much
smaller; stipes usually slightly shorter than to slightly longer than the laminae, but
varying from one-half to twice as long, sparsely scaly, the scales mostly subulate
to lanceolate-acuminate, peltate, to 3 mm long, erose to fimbriate, the larger ones
usually brown with a blackish central stripe, these scales often continuing into the
lower part of the rachis, with nearly orbicular, irregular, peltate scales scattered or
absent; laminae oblong or ovate to triangular-oblong to deltate, to 10 cm long, to
5.5 cm wide, acuminate or acute, deeply pinnatifid into up to 9(11) segments on
each side (usually fewer), glabrous above, sparsely to moderately densely scaly
below but the scales usually not completely obscuring the green of the surface, the
lamina scales peltate, ovate to lanceolate, usually attenuate-acuminate, 0.8-
2.0(3.0) mm long, light reddish-brown, darker at the point of attachment, weather-
ing gray, the margin remotely toothed to lacinate, orbicular scales few or none, the
lobes distant to fairly close, linear or spatulate to oblong, entire, obtuse to broadly
acute, regular or irregular in length on the same frond, perpendicular or slightly
ascending relative to rachis, the lowest pair distinctly alternate, venation mostly
free, fewer than one-half of the sori (usually many fewer) within areoles; sori
roundish, in a single row on each side of midvein of lobe, usually obscured by
scales; spores 64 per sporangium.
TYPE: Uncommon in crevices of cliffs and boulders in sheltered canyon with
Quercus grisea, Juniperus sp., Ungnadia speciosa, Garrya ovata, etc., lower
Indian Cave Canyon (side canyon of Dead Horse Canyon), north side of Chinati
Mountains, Presidio Co., Texas, 16 Oct 1977, M. L. Butterwick & E. J. Lott 3897
(TEX; isotypes GH, MEXU).
Polypodium thyssanolepis has generally been characterized as having areolate
venation and pinnatifid fronds (Maxon, 1916). A rare form of the species with
bipinnatifid fronds is known to show partial loss of areolation; this form occurs
with the normal form in Central America (Maxon, 1916), but is not known from
northern Mexico or the United States. It agrees in density of indument and all
other characters with typical var. thyssanolepis. On the other hand, the new
variety differs strongly and consistently from var. thyssanolepis in venation (Fig.
2). These venation characters are constant regardless of size; occasional speci-
mens of var. thyssanolepis from Chihuahua in which the fronds are much reduced
(laminae as small as 0.5 cm long), with many fronds nearly entire, nevertheless
display the areolate venation (and all other characters) typical of much larger
tropical plants of the variety. South American material, including all specimens
seen from Colombia (at GH), the type locality of the species, agrees with var.
thyssanolepis as here circumscribed.
Fronds of var. thyssanolepis may reach much larger sizes (to 60 cm or more)
than those of var. riograndense, but this probably is a direct environmental effect.
T. WENDT: NOTES ON PLEOPELTIS AND POLYPODIUM 7
Specimens of var. thyssanolepis from northern Mexico are generally much smaller
than tropical material; indeed, the reduction found in certain Chihuahuan speci-
mens, noted above, is unparalleled in var. riograndense.
Ss cea wre
ee
Pay
AS
ie ‘
!
FIG. 1. Distribution of Polypodium thyssanolepis varieties in the United States and Mexico excluding
Chiapas. Black squares = var. thyssanolepis; open squares = var. riograndense; half-squares =
intermediates.
The chromosome number of P. thyssanolepis var. thyssanolepis has been re-
ported as n=37 from South American material (Evans, 1963) and n=74 from a
Jamaican population (Wagner & Wagner, 1975). Polypodium thyssanolepis (vari-
8 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
ety unknown) has been reported as n=ca. 72 from material from ‘‘Mexico’’ (Sorsa
in Fabbri, 1965; Sorsa, 1966).
A limited number of varietal intermediates is found in Chihuahua (Fig. /), but
most specimens from this area are easily placed in one of the varieties. Several
collections (e.g., Pringle 443) include fairly representative plants of both varieties.
Further work, especially chromosomal, may reveal that var. riograndense would
be better treated as a separate species.
FIGS. 2 and 3. Venation of Polypodium thyssanolepis varieties. FIG. 2. Type of var. riograndense
(Butterwick & Lott 3897, TEX); arrows indicate scattered areoles. FIG. 3. Var. t/ lepis from San
Luis Potosi, Mexico (Johnston, Chiang & Wendt 12275, LL).
Figure I shows the geographical origin of the United States and Mexican speci-
mens examined. All Chihuahuan material for the species, along with representa-
tive specimens of the new variety, is cited below.
Polypodium thyssanolepis var. riograndense:
MEXICO: Chihuahua: Mountains between Guadalupe y Calvo and Nabogame, 7200 ft, on large
boulders in pine-oak forest, Correll & Gentry 23052, p. p. (GH, LL); 12 mi W of Cuauhtémoc, in
crevices of cliff, Correll & Johnston 21610 (LL); Between Yepomera and Babicora, in crevices of
boulders in pine-oak-juniper open mountain forest, Correll & Johnston 21624 (LL, US); Vicinity of
village of Majalca, in crevices of boulders, Correll & Johnston 21780 (LL); 12 mi W of Cuauhtémoc,
steep, rocky (granitic) slope, in pinyon pine-scrub oak association, Gould 8958 (LL); Rio Negro and
vicinity, LeSueur 1273 (TEX); Rocky hills near Chihuahua, cold cliffs, Pringle 443, p. p. (GH, NY,
US); 16 mi W of Cuauhtémoc, rolling terrain with scattered junipers, pinyons, and oaks, 7200 ft, in
rocky i teep slope, Reeder, Reeder, & Soderstrom 3477 (GH). Coahuila: Sierra del Jardin,
Canyon Hundido on N side of Pico Centinela, 8 km E of Rancho El Jardin by winding road, 1500-2250
m, steep canyon through igneous sierra, Johnston, Chiang, Wendt & Riskind 11803A (LL). Sonora:
Loop of the Rio de Bavispe, S of Aribabi, Sierra de Huépari, 1495 m, Harvey 1706 (US).
T. WENDT: NOTES ON PLEOPELTIS AND POLYPODIUM 9
UNITED STATES: Arizona: Cochise Co.: Conservatory Canyon, Huachuca Mts., vga Sept 1882,
Lemmon s. n. (GH, NY, US); Chiricahua Mts., Peebles & Loomis 5415 (US). Pima Co.: Baboquivari
Mts., Gilman 15 (US). Santa Cruz Co.: Sycamore Canyon, Patagonia Mts., 2800 ft, Ripley & Barneby
822 (NY). Texas: Brewster Co.: Chisos Mts., Boot Spring, 30 June 1932, Mueller s. n. (GH, NY,
TEX). Jeff Davis Co.: Near Fort Davis, in clefts and crevices of porphyritic rocks, E. J. Palmer 32196
(TEX, U
Polpodium thyssanolepis var. thyssanolepis:
XICO: Chihuahua: Minas Nuevas, ca. 8 mi NW of Parral, 6000 ft, Correll & Gentry 22764 (GH,
ee ai Ca. 5.5 mi NW of Parral, 5800 ft, Correll & Gentry 22723 (LL); Sierra de Santa Barbara, ca. 4
mi SW of Villa Matamoras, 6300 ft, Correll & Gentry 22802 (NY, LL); Along old railroad W toward
Rancho Ojito, Correll & Johnston 21488 (LL, NY); 25 mi SE of Cuauhtémoc, Correll & Johnston
21597 (LL); 11 mi S of Matamoras (Cuevas), 1950-2100 m, Gentry & Arguelles 18037 (LL, US);
Majalca (Pilares), 2075 m, Harvey 1463 (GH, US); La Bufa, on Rio Batopilas, Knobloch 578 (US);
Canyon E of Hidalgo de Parral, Knobloch 751 (US); Cerocahui-Cuiteco Road, Knobloch 882 (US);
Barranca Guerachic, between Agua Blanca and Guerachic, Knobloch 1849 (LL); Rocky hills near
Chihuahua, cold cliffs, Pringle 443, p. p. (GH); Potrero Peak, Pringle 977 (NY); Between San Fran-
cisco del Oro and Santa Barbara, near Arroyo de Granadefia, ca. 7000 ft, Soderstrom 894 (LL)
Intermediates between var. thyssanolepis and var. riograndense:
MEXICO: Chihuahua: Small mountain on NE edge of cal Correll 22688 (GH, LL); Mountains
just SE of Nabogame, 6000 ft, Correll & Gentry 23033 (LL); Mountains between Guadalupe y Calvo
and Nabogame, 7200 ft, Correll & Gentry 23052, p. p. (GH, LL, US); Majalea, Knobloch 329 (GH,
US), LeSueur 476 (US).
PLEOPELTIS
A study of material of Pleopeltis erythrolepis (Weath.) Pic. Ser. and P. polylepis
(Roem. ex Kunze) Moore throughout their ranges in Mexico and the United States
has led to the conclusion that they represent geographical varieties of the same
species. The following new combination is necessary:
hers cose polylepis vi var. erythrolepis (Weath.) Wendt, comb. & stat. nov.
Polypodium erythrolepis Weath. Contr. Gray Herb. 65:11. 1922. TYPE: Cold ape dea [Pot-
rero] Peak, Chihuahua, Pha 825 (GH!; isotypes, GH!, LL!, NY-2 sheets!, US-3 sheets!).
Phlebodium erythrolepis (Weath.) Conzatti, Fl. Tax. Mex. 1:95. 1946.
Pleopeltis erythrolepis (Weath.) Pic. Ser., Webbia 23:189. 1968.
Various characters have been used to distinguish the taxa. Weatherby (1922), in
his description of Polypodium erythrolepis, emphasized the long stipe and imbri-
cated, fimbriate-ciliate lamina scales of this ‘‘well-distinguished’’ species. How-
ever, in his treatment of the ferns of north-central Mexico (Weatherby, 1943) he
states:
[These new collections] go very far to break down the differences between P.
erythrolepis and P. peltatum [Pleopeltis polylepis]. In them, the abundant, ovate,
deeply lacerate-margined scales of the former, which seemed so distinctive when it
was proposed, nearly disappear and are replaced by suborbicular ones. The: surviv-
ing distinctions are: P. erythrolepis, stipe nearly as long as the blade, costa green on
the lower alee P. peltatum, stipe conspicuously shorter than the blade, costa
black on lower surface. In addition, P. erythrolepis tends to have narrower
rhizome-scales with narrower, more definitely erose-serrulate hyaline margins; but
this is os a tendency. Furthermore, the collection here cited under P. pel-
tatum . . . is also transitional . . . In all probability, P. erythrolepis would best be
treated as a vay of P. palates.
10 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
Knobloch and Correll (1962) ‘‘more or less avoided the taxonomic issue by recog-
nizing as P. peltatum those plants that have a distinctly blackish costa and placing
those plants that lack this characteristic into P. erythrolepis,’’ and in doing so
recognized both species from Chihuahua.
In the present study it was found that both costa color and stipe length are too
variable within both taxa to be taxonomically useful. The characters for distin-
guishing the varieties are given in the following key:
Scales of the lower surface of mature laminae entire or merely erose, mostly orbicular, usually not
densely imbricate; rhizome scales ovate to broadly lanceolate P. polylepis var. polylepis
Scales of the lower surface of mature laminae fimbriate or ciliate and/or mostly acuminate, usually
densely imbricate; rhizome scales lanceolate-acuminate to broadly lanceolate.
P. polylepis var. erythrolepis
Populations from Sonora, Chihuahua, northern Durango, and Texas fit easily
within var. erythrolepis as here circumscribed, and almost all central and southern
Mexican populations (southern Durango and Guanajuato south to Oaxaca, also
Baja California Sur) are ‘‘good”’ var. polylepis. However, a broad range of inter-
gradation between the varieties occurs in the northern Sierra Madre Oriental of
southern Coahuila, Nuevo Leon, and San Luis Potosi, where individuals referable
to both varieties as well as a preponderance of intermediates occur. Furthermore,
northern Coahuilan populations (Muzquiz to Sierra del Carmen) include many
intermediate types in additon to those referable to var. erythrolepis.
The problem is compounded not only by intrapopulational variation in lamina
scales, but also by the fact that scaliness of the fronds changes with age. Young
fronds of all populations tend to have many acuminate scales; these are then
apparently shed quite early in the case of var. polylepis, but are retained much
longer in var. erythrolepis. Mature, preferably fertile fronds therefore are neces-
sary for specimen identification.
Only one glaring exception to the above-mentioned geographical pattern was
found in the material studied. A specimen from the state of Mexico (Parque
Nacional de Laguna Zimpoala, Barkley, Webster & Rowell 7420, TEX) is refera-
ble by all characters to var. erythrolepis, although it is well south of the range of
that variety.
There also appears to be some problem in the differentiation of Pleopeltis
polylepis var. polylepis from P. macrocarpa var. trichophora (Weath.) Pic. Ser. in
central Mexico, particularly in the general area of Mexico City. The taxonomic
problems involving P. polylepis and P. macrocarpa are emphasized by the fact
that a variety originally described by Weatherby (1944) within P. polylepis (as
Polypodium peltatum var. interjectum) appears to belong closer to Pleopeltis
ee a (A. R. Smith, pers. comm.). Further studies are much needed in this
complex.
LITERATURE CITED
EVANS, A. M. 1963. New chromosome observations in the Polypodiaceae and Grammitidaceae.
Caryologia 16:671-677.
T. WENDT: NOTES ON PLEOPELTIS AND POLYPODIUM 11
FABBRI, F. 1965. Secondo supplemento alle tavole cromosomiche delle Pteridophyta di Alberto
iarugi. Caryologia 18:675—731.
KLOTZSCH, J. F. 1847. Beitrage zu einer Flora der Aequinoctial-Gegenden der neuen Welt. Filices
[Pt. 2]. Linnaea 20:337-445.
KNOBLOCH, I. W. and D. S. CORRELL. 1962. Ferns and Fern Allies of Chihuahua, Mexico. Texas
Research Foundation, Renner.
MAXON, W. R. 1913. Some recently described ferns from the Southwest. Amer. Fern J. 3:109-116.
—_—_——.. 1916. Studies of tropical American ferns-No. 6. Contr. U. S. Natl. Herb. 17:541-608.
—_—. 1923. Notes on American ferns—XIX. Amer. Fern J. 13:73-75.
MORTON, C. V. 1973. Studies of fern types, II. Contr. U. S. Natl. Herb. 38:215-281.
SORSA, V. 1966. Chromosome studies in the Polypodiaceae. Amer. Fern J. 56:113-119.
WAGNER, W. H., Jr. and F. S. WAGNER. 1975. A hybrid polypody from the New World tropics.
Fern Gaz. 11:125-135.
WEATHERBY, C. A. 1922. The group Polypodium lanceolatum in North America. Contr. Gray Herb.
65:3-14.
. 1943. Polypodiaceae. Jn I. M. Johnston. Plants of Coahuila, eastern Chihuahua and adjoin-
ing Zacatecas and Durango, I. J. Arnold Arbor. 24:306-339.
. 1944. A southern variety of Polypodium peltatum. Amer. Fern J. 34:17-19.
REVIEW
‘‘THE EXPERIMENTAL BIOLOGY OF FERNS,” by A. F. Dyer (ed.). Ex-
perimental Botany: An International Series of Monographs, vol. 14, 657 pp. 1979.
Academic Press, London and New York, ISBN 0-12-226350-2. $79.00—Over the
past forty years a significant amount of research has been devoted to the experi-
mental biology of pteridophytes. Although information has accumulated and de-
velopmental and genetic problems have been better circumscribed, there has been
no attempt to organize this into a fashion which would make the relevant ideas and
literature easily available to experimental biologists and botanists. This volume
attempts to review comprehensively nearly all of the significant studies in fern
experimental biology in a context which stresses the controversies currently ex-
tant and the problems and avenues of approach which promise to provide the most
productive and interesting rewards. In essence, it is a technical introduction to the
literature, with over 2000 reference citations. The textual contents reflect accu-
rately the current state of knowledge with heavy emphasis on morphogenetic
studies of the gametophyte generation. The 16 chapters, contributed by 16 au-
thors, detail our knowledge of meiosis; spore initiation, morphogenesis, and ger-
mination; structural, physiological, and biochemical aspects of the filamentous
gametophytic stage; differentiation from one-dimensional to two-dimensional
growth; antheridiogens; sporophyte development; apogamy, genetics, cytogenet-
ics, and hybridization; and experimental ecology. Although many of the contribu-
tions are excellent, some are superficial, reflecting in my estimation the lack of
experimental studies in those fields. However, because of the comprehensive
literature surveys and the emphasis on ideas and problems, this volume will be a
very valuable source book for experimental biologists and pteridologists for many
years to come.—Robert M. Lloyd, Department of Botany, Ohio University,
Athens, OH 45701.
12 AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 1 (1980)
The Deletion of Vittaria graminifolia
from the Flora of Florida
GERALD J. GASTONY*
Interest in new state and national records, range extensions, and the status of
rare or endangered species of ferns is probably nowhere more keen than in
Florida. Critical field and herbarium work to upgrade our knowledge of the dis-
tribution and habitat requirements of ferns, especially in the subtropical southern
region of the state, assumes political as well as scientific significance at a time
when state and federal socio-economic decisions are influenced by the ecological
status of species as humble as a lousewort or a snail darter. Efforts to increase the
accuracy of our floristic records for Florida ferns have been quite evident in recent
years, for example in the work of Messler (1974), Evans (1975), Ward and Hall
(1976), Nauman and Austin (1978), Nauman (1978), Austin et al. (1979), Adams
and Tomlinson (1979), and Nauman (1979a, 1979b). Such efforts, however, must
include the deletion of erroneous records as well as the addition of new records.
The deletion of one such erroneous record, the natural occurrence of Vittaria
graminifolia Kaulf. in Collier County, Florida, is the subject of this report.
The belief that V. graminifolia occurs in Collier County, Florida, its only re-
ported occurrence in the United States, is based on a statement appended to the
discussion of V. lineata (L.) J. E. Smith in Wherry’s (1964) Southern Fern Guide.
Wherry stated that V. filifolia Fée was found in 1960 in Collier County, and he
distinguished it from V. lineata by the weak iridescence and width of its scales. In
another context, Tryon (1964a) showed that V. filifolia is an incorrect name for
this species and that its correct name is V. graminifolia.
In an effort to bring cytological evidence to bear on the identity of the Appala-
chian gametophyte by counting its chromosomes and those of V. lineata and V.
graminifolia Gastony, 1977), I undertook a search for V. graminifolia in Collier
County. Dr. Wherry responded to my request for more information relating to his
1964 report by noting (in litt., 15 Aug 1976) that at the age of ninety and a half he
was no longer able to recall more specific locality data or whether an herbarium
voucher documented his report. He did recall, however, that he had visited the
living fern collection assembled by John Beckner in St. Petersburg, Florida and
that Beckner had there ‘‘two Vittarias,”’ one less winter hardy than the other. The
less hardy one from Collier County was what Dr. Wherry took to be V.
graminifolia (as V. filifolia).
Wherry’s information enabled me to contact John Beckner, who agreed to take
me to the site from which he had collected the Vittaria in question. In the com-
clearly remembered having made the original collection, and we eventually found
several specimens of what Beckner said was V. graminifolia if anything in that
*Department of Biology, Indiana University, Bloomington, IN 47401.
G. J. GASTONY: DELETION OF VITTARIA GRAMINIFOLIA 13
area was. These were immature, somewhat depauperate specimens epiphytic on
the trunk of Persea palustris, unlike the usual Sabal palmetto epiphytism of V.
lineata. They were not fertile and did not survive cultivation efforts in the Indiana
University greenhouses.
Beckner was certain, however, that Wherry had taken a specimen of his original
collection back to Pennsylvania and that a voucher specimen documenting Wher-
ry’s (1964) statement was to be found there. Subsequent inquiries led to a speci-
men (number 0925236) at the herbarium of the Academy of Natural Sciences in
Philadelphia (PH) bearing the stamp of the herbarium of the University of
Pennsylvania, which is on permanent loan to PH. The label identifies the speci-
men as Vittaria filifolia Fée and indicates that it had been cultivated from a plant
collected by John Beckner west of Deep Lake in Collier County, Florida. The
specimen was made on 3 January 1962, and has been annotated by Dr. Wherry as
V. filifolia. Beckner (pers. comm.) has since assured me that the locality from
which this specimen was taken is identical to the swampy locality we had visited
west of Copeland and Deep Lake. On 4 January 1980, I returned to this site and
established in a discussion there with Park Ranger Robert Goble that this locality
is in the center of what is now the Fakahatchee Strand State Preserve protected by
the Department of Natural Resources of the State of Florida.
I have analysed Wherry’s specimen from PH, utilizing the characters employed
by Tryon (1964b, pp. 212-215) in distinguishing V. lineata and V. graminifolia in
the Ferns of Peru. Comparable or identical characters are used by Lellinger (pers.
comm.) and by Stolze (pers. comm.) in distinguishing these species in their forth-
coming treatments of the ferns of Costa Rica, Panama, and the Chocé and the
ferns of Guatemala, respectively. Perhaps the most absolute criterion employed in
the discriminatory sets of characters used in these three major floristic treatments
is the incidence of tetrahedral-globose, trilete spores in V. graminifolia, as op-
posed to reniform, monolete spores in V. lineata. In this regard and in the other
characters examined, the specimen upon which the record of V. graminifolia in
Florida rests is surely V. lineata. I sent Beckner a photocopy of Wherry’s her-
barium specimen and he is certain (pers. comm.) that this specimen is the basis of
Wherry’s (1964) report.
It is interesting that Lakela and Craighead (1965), Long and Lakela (1971), and
Lakela and Long (1976) discussed V. /ineata in their treatments of the ferns of
south Florida but made no reference whatever to V. filifolia or V. graminifolia.
The reason for omitting V. graminifolia from these three works is unknown and is
particularly curious since the work by Long and Lakela (1971) does cite Wherry’s
book (1964) as a selected reference on the ferns of Florida. Long is deceased and
Craighead (pers. comm.) says that the decision as to what to include in their
checklist was entirely that of Lakela. Lakela, now in retirement, does not recall
the reason for this omission from any of these works (pers. comm.). There is no
indication that any of these authors ever consulted the specimen at PH. Based on
my experience with Beckner in revisiting the collection site of Wherry’s specimen
in the Fakahatchee Strand, I suspect that the difference in the cold-hardiness of
the ‘‘two Vittarias’’ was most likely due to the sub-optimal substrate of the
14 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
hardwood host tree and perhaps to a variant allelic constitution correlated with the
occurrence of these individuals on this unusual host.
It is always possible that V. graminifolia or any other species common in
tropical America may be carried into southern Florida by hurricane winds or other
means of dispersal and that such adventives may become temporarily or perma-
nently established in subtropical Florida. Because of Wherry’s report, Austin and
Nauman (pers. comm.) have searched extensively for V. graminifolia in the
Fakahatchee Strand but have never found it and have concluded that it is not
there. Critical examination of the morphology and ecology of the specimen dis-
cussed above indicates that there is no longer any reason to believe that V.
graminifolia ever did or does now occur in Florida. It should therefore be deleted
from the floristic record for Florida and thus from the flora of the United States.
I am grateful to Ross and Priscilla Stanley of Port Charlotte, Florida for their
hospitality during the field work and manuscript preparation for this paper and to
John Beckner for his aid in the field and in locating the specimen upon which Dr.
Wherry’s report was based. I thank Dr. Michael Madison for help in the field, Dr.
Wherry for help in interpreting his report, the officers of the herbarium of the
Academy of Natural Sciences (PH) for the loan of the specimen discussed, and
Ranger Robert Gobel for aid in interpreting the specimen locality data in terms of
the Fakahatchee Strand.
LITERATURE CITED
ADAMS, D. C. and P. B. TOMLINSON. 1979. Acrostichum in Florida. Amer. Fern J. 69:42—46.
AUSTIN, D. F., G. B. IVERSON, and C. E. NAUMAN. 1979. A tropical fern grotto in Broward
County, Florida. Amer. Fern J. 69:14—16.
EVANS, A. M. 1975. Cheilanthes in Florida. Amer. Fern J. 65:1-2.
GASTONY, G. J. 1977. Chromosomes of the independently reproducing Appalachian gametophyte: A
new source of taxonomic evidence. Syst. Bot. 2:43—48.
LAKELA, O. and F. C. CRAIGHEAD. 1965. Annotated Checklist of the Vascular Plants of Collier,
Dade and Monroe Counties, Florida. Fairchild Tropical Garden and the Univ. of Miami Press,
Coral Gables, Florida. ;
, and R. W. LONG. 1976. Ferns of Florida. An Illustrated Manual and Identification Guide.
Banyan Books, Miami.
LONG, R. W. and O. LAKELA. 1971. A Flora of Tropical Florida. A Manual of the Seed Plants and
Ferns of Southern Peninsular Florida. Univ. of Miami Press, Coral Gables, Florida.
Ce i 1974. The natural history of Ophioglossum palmatum in south Florida. Amer. Fern
33-39.
NAUMAN, C. E. 1978. A checklist of the ferns an
d primitive vascular plants of southeastern Florida.
Castanea 43:155-162
I
Amer. Fern J. 68:65-66.
TRYON : = M. 1964a. Taxonomic fern notes. IV. Some American vittarioid ferns. Rhodora 66:110—
—————. 1964b. The ferns of Pery Polypodi (D iti Ol 1 )
Meesiki ins ns ace Vieandreae). Contr. Gray Herb.
WARD, D. B. and D. W. HALL. 1976. Re-
introduction of Marsilea vestita into Florida. Amer. Fern J.
66:113-115.
WHERRY, E. T. 1964. The Southern Fern Guide. Doubleday, Garden City, N.Y.
AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 1 (1980) 15
New Taxa and Combinations of Pteridophytes
from Chiapas, Mexico
ALAN R. SMITH*
This is the second and, it is intended, final report on new taxa and new combina-
tions (beginning on p. 26) of pteridophytes in the state of Chiapas, Mexico. For the
first report, see Proc. Calif. Acad. Sci., Ser. IV, 40:209-230. 1975. All of the taxa
are to be included in the treatment of the pteridophytes of Chiapas, a part of the
‘*Flora of Chiapas’’ project headed by Dennis Breedlove, California Academy of
Sciences. English descriptions, fuller synonymies, and additional discussion will
be found in the floristic account. Certain of the new names are also needed by
Robert Stolze for his forthcoming treatment of the Polypodiaceae in the *‘Ferns
and Fern Allies of Guatemala.”’
I thank Colleen Sudekum for preparation of the illustrations. Scanning electron
micrographs of spores were made with a Coates and Welter 50 microscope, ob-
tained by the Electron Microscope Laboratory at the University of California,
Berkeley, under a grant from the National Science Foundation (GB-38359). Breed-
love collections were made with the help of National Science Foundation grants
GS-383, GS-1183, and GB-29483. I am grateful to A. M. Evans, who has collabo-
rated in the description of Polypodium chiapense.
yyy Asplenium insolitum A. R. Smith, sp. nov. Figs. 1-2.
Rhizomata suberecta, caudices ca. 1 cm diametro; frondes 35-45 cm longae,
stipitibus laminas fere aequantibus; stipites brunnei vel griseo-brunnei, non lus-
trati, ca. 1.5 mm diametro, adaxialiter viridi-alati, glabri, basi paleis paucis atro-
brunneis ovatis; paleae ca. 2 mm longae, obscure clathratae, parietibus crassis et
luminibus congestis parvulis; laminae ovato-lanceolatae, 18-25 cm longae, bipin-
natae, apice attenuatae sed nec flagelliformes nec proliferae; rhachides adaxialiter
viridi-alatae, abaxialiter brunneolae, epaleatae; pinnae 15—20-jugae, usque 6 cm
longae, 2.5 cm latae; pinnae infimae (1 vel 2 paria) aliquantum reductae, deflexae;
pinnulae usque 8-jugae per pinnam, saepe lobo acroscopico, aliter dentatae vel
bidentatae secus marginem, basi cuneatae, plerumque inaequilaterae (sub-
dimidiatae), latere basiscopico exciso; pagina laminae et axes subter glabri vel
pilis minutis (0.1 mm longis) adpressis capitatis; venae pinnatae, usque 4-jugae per
pinnulam; sori 1.0—2.5 mm longi, indusia tenui albido.
“TYPE: Terrestrial in montane rain forest, 11 km NW of junction of road to
Motozintla along road to El Porvenir and Siltepec, southwest side of Cerro
Mozotal, Munic. Motozintla de Mendoza, Chiapas, Mexico, 2100 m, 21 Nov 1976,
Breedlove 41653 (DS).
PARATYPE: Same locality, 27 Jun 1972, Breedlove 25760 (DS). 3,
Asplenium insolitum has no obvious close relatives. In dissection, it is similar to
some of the more divided members of the A. radicans complex (e.g., A. flabel-
lulatum Kunze var. partitum Klotzsch), but the blade apex is neither flagelliform
nor budding and the stipes and rachises are not shining. A closer relative is
perhaps A. cuneatum Lam., but that species differs in the flabellate venation of
the segments, longer sori, and obviously clathrate scales. Another possible rela-
Berkeley, CA 94720.
*University Herbarium, Department of Botany, University of California,
AMERICAN FERN JOURNAL: VOLUME 70 (1980)
| 5¢m
FIGS. 1-4. New Asplenium species. FIGS. 1-2. Type of A. insolitum, habit and lower pinna. Breed-
love 41653 (DS). FIGS. 3-4
. Paratype of A. sphaerosporum, habit and lower pinna, Breedlove 32512
(DS). Line scale for habit drawings.
Jo3¢
A. R. SMITH: NEW PTERIDOPHYTES FROM CHIAPAS 17
often a few minute clathrate scales in the pinna axils.
Asplenium sphaerosporum A. R. Smith, sp. nov. Figs. 3-6.
Rhizomata erecta; frondes plerumque 45-70 cm longae, usque 18 cm latae;
stipites atri, ca. 2 mm diametro, glabri, longitudine 0.5-0.75 partes laminarum
aequantes; rhachides atrae vel virides, distaliter viridi-alatae; laminae lanceolatae,
ad apicem acuminatae; pinnae patentes, vulgo 25 vel plus, ad apicem acuminatae,
paribus infimis plene bipinnatis, distaliter pinnis pinnatisectis, denique pinnis
serrato-incisis; segmenta obovata, usque 1.3 cm longa, 5 mm lata, basi cuneata,
apice dentata vel denticulata (dentibus 2-7), usque 12 paribus per pinnam, prope
apices pinnarum segmentis adnatis et decurrentibus; paginae laminarum at-
rovirides vel aeruginosae, crassae, glabrae; sori usque 2—3 per segmentum; sporae
grandes, globosae (interdum ellipsoideae), 32 per sporangium.
“TYPE: SE side of Cerro Baul (16 km NW of Rizo de Oro), Chiapas, Mexico,
Breedlove 21805 with Smith (DS).
tive is A. solmsii Baker ex Hemsl., but that species has tripinnatifid blades and
S
te
oe G-— < 6)
FIGS. 5-6. Scanning electron micrographs of spores of Asplenium sphaerosporum, Breedlove 28733
(DS), x 1500.
PARATYPES:
MEXICO: Chiapas: Lagos de Montebello, Breedlove 22303 with Smith (DS); Jct. of T:
anaté River
and river from Yochib, paraje Mahben Chauk, Munic. Tenejapa, Breedlove 6368 (DS, US); SE of
Cerro Baul (16 km NW of Rizo de Oro), Breedlove 21810 with Smith and 31338 with Smith (both DS); 7
km NE of Bochil along road to Simojovel, Munic. Bochil, Breedlove 28723, 28733 (DS), Breedlove
32311 with Smith (DS); Near summit of Chuchil Ton, NE of Bochil, Munic. San Andres Larrainzar,
Breedlove 29244 (DS); 7 km NE of Jitotol—-Pichucalco jct. on road from Bochil to Simojovel, Munic. El
Bosque, Breedlove 32512 with Smith (DS); 6.5 km N of Jitotol, Munic. Jitotol, Breedlove 32758 with
Smith (DS); Ocotepec, Rovirosa 1049 (PH); 20 km S of Ocozocoautla, Munic. Ocozocoautla -
Espinosa, Breedlove 29138 p. p. (DS), Miinch s.n. (DS); Ghiesbreght 404 (K, NY, PH). Lenaenden
Techolo, Sanchez 6 (UC, US); along Camino Real, near Jalapa, Weatherwax 176 (UC). —_ sarin
Schnee s.n. (K); Bourgeau 2365 (K). GUATEMALA: Fuego ?, Salvin & Godman 368 (K); Heyde
Lux [Donnell-Smith 3231] (K, US).
18 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
a y a Ly
BLOND
aN BS it ep PN \
en oe.
SAN Sars a
oem
FIGS. 7-10. New Cheilanthes taxa. FIGS. 7-8. Type of C. complanata, habit and base of lowermost
pinna, Breedlove 41747 (DS). FIGS. 9-10. Type of C. — var. fimbriata, habit and pinnule,
Breedlove 39018 (DS). Line scale for habit drawings
A. R. SMITH: NEW PTERIDOPHYTES FROM CHIAPAS 19
This species resembles somewhat A. achilleifolium, but I do not think that they
are closely related. The affinity seems closest to A. monodon Liebm. and A.
cuspidatum Lam. The former has large, globose spores, 32 per sporangium, like
those of A. sphaerosporum (Figs. 5-6); all specimens of A. cuspidatum that I have
looked at have small, reniform spores, 64 per sporangium. It is possible that A.
sphaerosporum arose through hybridization between some member of the A.
auritum group and A. cuspidatum. Alternatively, it could have speciated from A.
monodon. Additional studies are needed to understand the evolutionary relation-
ships within this complex group.
Asplenium auritum Swartz has often been applied in a broad sense, encompas-
sing plants that are simply pinnate (sometimes with a basal auricle) to fully bipin-
nate. I would restrict the application of the name to those plants of the complex
that are simply pinnate; such plants also have tan, reniform, relatively small
spores, 64 sporangium. In Chiapas (and apparently elsewhere in the range), A.
auritum s. s. occurs only at low elevations, 200-500 m.
Asplenium sphaerosporum occurs at higher elevations—(900)1250-2700
m—than any other member of the A. auritum complex in Chiapas, at elevations
where A. cuspidatum can occur. The latter is chiefly from Montane Rain Forests,
while A. sphaerosporum is most common in Pine-Oak-Liquidambar Forests.
? 084 Cheilanthes complanata A. R. Smith, sp. nov. Figs. 7-8.
701
.
Differt a C. hirsuta Link paleis rhizomatis distincte bicoloris, ad marginem
cinnamomeis, ad medium nigrescentibus; laminis pentagonis, latitudine lon-
gitudinem fere aequantibus, planis, segmentis ultimis non pendulis; segmentis
ultimis obovatis vel anguste ellipticis, 2—4-plo longioribus quam latioribus; laminis
atroviridibus, utrinque glabris; indusiis membranaceis, non valde dissimilibus
laminae, integris (sine glandibus vel pilis), non vel leviter ad axe decurrentibus.
TYPE: North and west slope of Cerro Mozotal below microwave tower along
road from Huixtla to El Porvenir and Siltepec, Munic. Motozintla de Mendoza,
Chiapas, Mexico, 3000 m, Breedlove 41747 (DS).
PARATYPE: Same locality, Breedlove 40335 (DS).
The best characters for separating C. complanata from its nearest relatives, i
marginata H.B.K., C. chaerophylla (Mart. & Gal.) Kunze, and C : hirsuta
(synonym of C. pyramidalis Fée), are the planar blades and the thin, entire indusia
that completely lack trichomes or marginal papillae. Another relative may be C.
cuneata Link, but that species has black or nearly black stipes and rachises, larger
blades, darker, concolorous rhizome scales, and more sharply differentiated in-
dusia.
Cheilanthes microphylla var. fimbriata A. R. Smith, var. nov. Figs. 2-18,
Differt a var. microphyllo indusiis fimbriatis pilis usque 5 mm longis;
trichomatibus numerosioribus albidis prope margines laminae supra; et laminis
generaliter parvioribus bipinnatis subdimorphis.
“TYPE: Along road to Ciudad Cuauhtemoc 6-8 km E of Frontera Comalapa,
Munic. Frontera Comalapa, Chiapas, Mexico, Breedlove 39018 (DS).
20 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
|| RE pepe ree EY ee eI cee Seti srs
S| eee pentnse a eee EEE EE I sea
5 Sa YP
| semeas Se
Baa se gas
ee se
FIGS. 11-12. Type of Polypodium alavae, habit and base of pinna, Alava 1287 (UC). Line scale for
habit drawings.
314!
Te
A. R. SMITH: NEW PTERIDOPHYTES FROM CHIAPAS 21
PARATYPES:
GUATEMALA: Petén: Lake Petén Itza, NW of San Andres, Contreras 3571 (US); Lake Petén Itza,
along shore W of San Andres, Lundell 17251 (US). MEXICO: Chiapas: E] Carmen, Miinch 184 (DS);
without locality, Miinch (DS); Same locality as type, Breedlove 26976 (DS); Munic. Ocozocoautla de
Espinosa, Rio de la Venta at the Chorreadero near Derna, Breedlove 36556 (DS); Munic. Ocozocoautla
de Espinosa, 13-18 km S of Ocozocoautla, Breedlove 37838 (DS); Villa de Yajalén, Rovirosa 971 p. p.
(PH). Tamaulipas: 2 mi S of Tres Palos and 1 mi down road to Loreto, Johnston 4884 (TEX). Yucatan:
San Anselmo, Gaumer 1238bis (US); Izamal, Gaumer 534 (UC, US), Gaumer 1409 (US); Chichan-
kanab, Gaumer 1473 (US), P. Valdez 65 (US), Gaumer 533 (US); Ruins of Nojpat, Schott 686 (US);
Mérida, Schott 135 (US).
Variety microphylla is known from the Antilles, southeastern United States,
and eastern and southern Mexico. I have not seen collections from the Yucatan
peninsula, where var. fimbriata is common. In Chiapas, var. fimbriata seems to
be more common than the type variety and does not grow with it.
Diplazium drepanolobium A. R. Smith, sp. nov.
Differt a D. lonchophyllo Kunze segmentis pinnarum magis obliquis et fal-
catioribus; frondibus plerumque grandioribus, pinnis vulgo 20-25 cm longis, 3.5-
6.0 cm latis.
-TYPE: 10 km above Rayon, Chiapas, Mexico, Breedlove 26122 (DS).
PARATYPES:
MEXICO: Chiapas: 26-28 km N of Ocozocoautla, Breedlove 22451 with Smith (DS); 45 km N of
Ocozocoautla, Breedlove 20760, 32852 (DS); 2-4 km below Ixhuatan, Breedlove 24163 (DS); 32 km N
of Ocozocoautla, Breedlove 38162 (DS); 46 km N of Ocozocoautla, Breedlove 38676 (DS); Without
precise locality, Ghiesbreght 361 (K); Arroyo de Ona, cerca Ixtacomitan, Rovirosa 59 (K, PH).
Veracruz: Schaffner 470 (P).
It is possible that this is only an extreme variant of D. lonchophyllum Kunze,
but D. drepanolobium has different blade dissection and seems at least as distinct
as some other segregates of D. lonchophyllum (e.g., D. prominulum Maxon and
D. subsilvaticum Christ). Relatively few specimens are intermediate between D.
drepanolobium and D. lonchophyllum; one such specimen is Breedlove 21624-A
(Chiapas, 13 km N of Berriozabal). This collection has abortive spores and may be
a hybrid. More numerous are the intermediates between D. drepanolobium and a
third species of the complex, D. franconis Liebm., which includes the following
synonyms: D. camptocarpon Fée, D. hahnii (Fourn.) C. Chr., and D. donnell-
smithii Christ. Several such intermediates from Chiapas are 22293, 22484, and
27480 (all DS). The best course in this difficult group seems to be to recognize all
three species. The alternative is to recognize a single species, a treatment that has
little merit.
On the basis of certain collections identified by Fée (e.g., Schaffner 470), for-
‘merly I applied the name D. acutale Fée to this species. However, the type of D.
acutale is referable to D. lonchophyllum.
Figs. 11-12.
: i : : ; : : est
Rhizomata repentia, 5-8 mm diametro, paleis aurantiaco-fulvis, aliquan ,
patentibus, concoloribus, glabris, ca. 5-10 mm longis; ——. ne ; ee - oe
gae; stipites straminei vel brunneoli, parce pubescentes, longitu JU.
partes laminae aequantes, (1)3—5 mm diametro;
22 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
5cm
‘e" 15
eat 13-17. New Polypodium species. FIGS. 13-15. Ty
and rhizome, Breedlove 37155 (DS). FIGS. 16-17.
Bebdins 27453 (DS). Line scale for habit drawi ings
pe of P. fuscopetiolatum, habit, portion of
Type of P. chiapense, habit and segment,
A. R. SMITH: NEW PTERIDOPHYTES FROM CHIAPAS 23
(7)14-30 cm latae, basi latissimae, pinnatae praeter apicem, apice Rep Te vel
subconformi, confluenti; rhachides moderate vel conspicue pubescentes, sine
16 cm lo 7-10 m
paleis; pinnae plerumque 7— ongae, 7-10 mm latae, integrae vel leviter
crenulatae, ad apicem acutae, a rotundatae, Grecian parallels, pinnis ses-
silibus dimidio proximali laminae, adnatis dimidio distali lam , Nunquam di-
latatis; venae ie pean liberae; paginae laminarum — ne : aarviees vel
olivaceae, subcoriaceae, utrinque glabrae vel sparse pubescentes; costae ven-
aeque infra pilis onan is vel numerosis, laxis, septatis, 0.4-0.8 mm longis; sori
rotundati, mediales vel ee vulgo ca. 2mm diametro; sporangia pilis
0 3 mm longis, laxis, ca. 6 pilis per capsulam
“TYPE: About 3 mi from race ibe fecal from Sols to Chanal, Chiapas, Mexico,
Alava 1287 (UC; isotype DS).
PARATYPES:
HONDURAS: Francisco Morazan: Km 24 on Tegucigalpa—Zamorano Rd., Kimnach 449 (UC).
MEXICO: Chiapas: Lagos de Montebello, Breedlove 22337 with Smith and 32126 with Smith (both
DS); same locality, Breedlove 37005 (DS).
Polypodium alavae is related to P. adelphum Maxon, from which it differs in
the narrower, more parallel-sided, mostly entire pinnae, long-hairy sporangia, and
more coriaceous, dark-green leaf tissue. It is also related to P. puberulum
Schlecht. & Cham., but it lacks the densely hairy leaf tissue on both surfaces and
the lower pinnae are constricted at the base.
B35 ghd as ane her igh on & Smith, sp. nov. Figs. 16-17.
entia, usque 10 cm longa, 3-5 mm diametro; paleae
rhizcmiatis’ cee e pasi fate rants dilatatae, renee acuminatae, incon-
spicue como atae, a inem interdum denticulatae; frondes
e sae, non r
12-40 cm longae, distantes (ca. 7-10 mm), paid 1-2(3) per plantam; stipites
longitudine 0.1-0.2 partes laminae aequantes, 0.5-1.0 mm diametro, stra minei vel
longae, 2.5— latae, anguste ovatae vel lanceolatae, basi subtruncatae vel
truncatae; ewan sine paleis, utrinque laxe villosae pilis 1-2 mm longis; seg-
menta 13—40 mm longa, 3—6 mm lata, patentia, lineari-lanceolata, apice obtusa vel
subacuta, basi Satie ‘dilatata, integra vel versus apicem obscure undulata; venae
l-furcatae, liberae; paginae laminarum dilute virides, pilis numerosis, 1-2 mm
longis, laxis, argenteis, secus costas ene costae decurrentes ad rhachim;
sori rotu ndati, inframediales; sporangia glabra
“TYPE: Selva del Ocote, 32 km NW of Ocozocoauitla, Munic. Ocozocoautla de
Espinosa, Chiapas, Mexico, Breedlove 27453 (DS).
PARATYPES:
MEXICO: Chiapas: Finca Mexiquito, Purpus 6754, p. p. (UC, US);
US); 13 km N of Berriozabal, Breedlove 20281, 31231 (both DS); Same locality,
Smith, 31537 with Smith (both DS).
Polypodium chiapense is similar to P. hygrometricum
stramineous to tan stipes, these villose with soft, silvery, lax,
mm long; rachises stramineous with similar hairs 1-2 mm long; :
perpendicular to the rachis; leaf tissue with numerous, lax, silvery, septate airs
1-2 mm long, these densest along the costa, and in the glabrous sporangia.
Finca Irlanda, Purpus 7225 (UC,
Breedlove 21671 with
Splitg., but differs in the
septate hairs ca. |
4 at
24 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
Polypodium fuscopetiolatum A. R. Smith, sp. nov. Figs. 13- 15.
Rhizomata repentia, 3-6 mm diametro, brunneola, paleis arte adpressis, lan-
ceolatis; paleae fulvae, in medi vittatae, margine m
plerumque castaneae, sine paleis; pinnae lanceolatae, apice acutae ve
acuminatae, plerumque 9-15 mm latae; venae 3- vel 4-furcatae, anastomosantes,
areolis 1-seriatis in quoque latere costae; pagina laminae utrinque pubescens vel
glabrescens vel fere glabra; venae costaeque pubescentes; sori rotundi, ca. 1.0—
1.5 mm diametro, mediales vel inframediales; sporangia glabra vel setosa, setis
minus quam 0.1 mm longis
VTYPE: 6-8 km WNW of Soyalo, Chiapas, Mexico, Breedlove 37155 (DS).
- PARATYPES:
EL SALVADOR: San Salvador: Santa Tecla, Jaurequi 67 (UC); Same locality, Garcia 22 (UC).
GUATEMALA: Santa Rosa: Jumaytepeque, Heyde & Lux [Donn. Smith 4090] (US). Solola: Near
Pueblo San Jorge, Hatch & Wilson 317 (US). Suchitepequez: Cuyotenango, Rojas 146 (US).
Guatemala: Sapoti barranca, Hayes (US); Pinala, Hayes (US); San Gerénimo, Salvin & Godman (K).
MEXICO: Chiapas: Salto de Agua, Escuintla, Matuda 18397 (MEXU, DS); Cacaluta, Escuintla,
Matuda 17005 (MEXU, DS); Finca Mexiquito, Purpus 6755, 6858 (UC, US); Huitla, Purpus 7223
(UC); El Sumidero, 22 km N of Tuxtla Guti Breedlove (all with Smith) 21584, 21593, 21595 (DS);
3 km N of Ocozocoautla, Breedlove 21923 with Smith (DS); 5-6 km W of Teopisca, Breedlove 22867
(DS); NW side of Cerro Vernal, 25-30 km SE of Tonala, Breedlove 25616 (DS); 6-8 km E of Frontera
Comalapa, Breedlove 26978, 39083 (DS); 27 km NE of Huixtla, Breedlove 28612, 28653 (DS); 65 km S
Hwy. 190 on rd. to Nuevo Concordia, Breedlove 37784 (DS); Cerro Vernal, 21 km S of Tonala,
Breedlove 38135 (DS). Guerrero: 25 mi S of Chilpancingo, Storer 11] (US); Dist. Mina, Manchon,
Hinton et al. 9451 (K); Dist..Montes de Oca, Vallecitos, Hinton et al. 11390 (K). Jalisco: Sierra del
Halo, 7 mi SSW of Tecalitlan, McVaugh 16189 (US). México: Dist. Temascaltepec, Rinc6én del Car-
Ps Hinton 1737 (K, UC, US). Michoacan: Dist. Coalcoman, Pto. Zarzamora, Hinton et al. 12249
(K).
This appears to be one of the more common species of Polypodium at lower
elevations (500-1350 m in Chiapas) on the Pacific slope of southern Mexico and
northern Central America. Polypodium fuscopetiolatum is closely related to P.
hispidulum Bartlett, but it can be distinguished by the narrower rhizome scales
that are denticulate or papillate on the margin and filiform at the tip and also by the
stipes and rachises usually castaneous and shining. There is also a resemblance to
P. plesiosorum Kunze, which has broader, ovate rhizome scales and grows at
higher elevations. Specimens of P. fuscopetiolatum have often been identified as
P. plesiosorum in herbaria.
Several Costa Rican collections are close to P. fuscopetiolatum but differ in
having smaller, thicker-textured fronds, broader, squared sinuses, and broader
and lighter-colored rhizome scales. I am not certain that they are conspecific with
P. fuscopetiolatum, but they appear to be closely related: Pittier 904, Tonduz
mine Standley 41904, Tonduz 8804, Mickel 2394, Stork 2987, and Allen 547 (all
A. R. SMITH: NEW PTERIDOPHYTES FROM CHIAPAS 25
Co mickelii A. R. Smith, sp. nov.
7st
Rhiz erecta, a ae 1.5-3.0 cm diametro; frondes 45-110 cm longae;
stipites aes 0.6—1. 0 partes laminarum aequantes, basi paleis ovatis mar-
gine erosis usque rae ae usque 15 mm longis et 6 mm latis, bicoloribus,
paleis fulvis in isa a guste brunneo-vittatis vel a ge oncoloribus et uni-
formiter fulvis; lam ovato-attenuatae, 12-35 cm latae; rhachides ae pro-
liferae, Slabieeseutes Gel paleis capillaceis praesertim basi pinnarum; pinnae pin-
natae, pinnulae lobo parvo deltoideo acroscopico, aliter integrae, vu in frondis
grinds pinnulis crenulatis vel grosse dentatis et lobo fere libero elliptico acro-
scopico; costulae infra glabrescentes vel basi cee fibrillosis; pagina laminarum
et venarum infra plus minusve oe Soria: supra nitida, atroviridis; sori
parvi, ut videtur exindusiati, non con
-TYPE: NE slope of Cerro Zhitibodbeepetl trail from La Candelaria to
Zacatepec, Dist. Mixes, Oaxaca, Mexico, Mickel 4836 with Leonard (NY).
PARATYPES:
GUATEMALA: San Marcos: Near Aldea Fraternidad, between San Rafael Pié de la Cuesta and Palo
Gordo, Williams et al. 26103 (F), 26299 (F, US). Above Finca El Porvenir, up Cerro de Mono, Volcan
ree deguens as Steyermark 37397 (F). HONDURAS: Intibuca: Quebrada del Pelon de Guise, Molina R.
6375 (F). MEXICO: Chiapas: SE side of Cerro Tres Picos, Breedlove 25379, 34385 (DS); Without
precise locality, Ghiesbreght 401 (YU). Oaxaca: Type locality, mn 4823 with Leonard (NY); Dist.
Mixes, vicinity of Zacatepec, Mickel 1565 (NY); Dist. Choapan, Lovani to river toward La Selva,
Hallberg 1577 (NY); Dist. Tuxtepec, 24km S of Valle = et “Mickel 5929 (NY); Dist. Ixtlan, 5km S
of Vista Hermosa, Mickel 7189 (NY, UC); Dist. Ixtlan, 29 km S of Valle Nacional, Mickel 6363 (NY,
UC). Veracruz: Munic. Yecuatla, El Haya, Ventura A. 3431 (NY); Munic. Yecuatla, El Cajon,
Ventura A. 4812 (NY :
In Oaxaca and Veracruz this species grows at rather low elevations, 450-1450
m; the Chiapas collections were made at 2100-2500 m. At the lower elevations,
the only other Polystichums encountered in Mexico are P. platyphyllum (Willd.)
Presl and occasionally P. muricatum (L.) Fée. Polystichum mickelii appears to be
without close relatives in Mexico and Central America. It is possibly of the group
of P. platyphyllum, as indicated by the exindusiate sori. It resembles somewhat
species from southern Brazil, e.g., P. montevidense (Spreng.) Rosenst.
Selaginella chiapensis A. R. Smith, sp. nov
Species heterophylla ex affinitate S. idiosporae Alston, sed foliis intermediis
Ovatis, acumnens (nec aristatis), foliis argenteis subtus, rhizophoris seacilnadiean
.2-0.4 mm dia iffert.
“TYPE: 18-20 ‘eon N of Ocozocoautla, Munic. Ocozocoautla de Espinosa,
Chiapas, Mexico, 800 m, Breedlove 28159 (DS).
PARATYPE: Above Rancho San Luis, ca. 2 mi N of Ocozocoautla, Chiapas,
Mexico, Carlson 2127 (BM, US).
In addition to the differences between S. chiapensis and S. idiospora mentioned
above, the new species seems to have the median leaves more obviously in two
ranks adjacent to. one another, the ranks scarcely or not at all overlapping. The
paratype cited differs from the type in having acute median leaves, eciliate lateral
leaves, and stouter rhizophores. It was originally annotated by Alston as “‘S.
idiospora ined.’’, and presumably later in pencil, “‘sp. nov.’ In the sum of its
characters, S. chiapensis seems more different from S. idiospora than the latter
does from S$. guatemalensis Baker.
26 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
uy
Diplopterigium bancroftii (Hook.) A. R. Smith, comb. nov.
Li4¢@ Gleichenia bancroftii Hook., Sp. Fil. 1:5, t. 4A. 1844’ LECTOTYPE: Jamaica, Bancroft (K!; ee
totype BM!), chosen by Proctor in apie ae Less. Antill. 60. 1977). Hooker may never have see
the other syntype, which is Jamaica, Sw
Nakai (Bull. Natl. Sci. Mus. 59: ‘50. 1950) regarded Gleichenia bancroftii as a
synonym of Diplopterigium farinosum (Kaulf.) Nakai, but the original description
and subsequent illustrations suggest that Mertensia farinosa Kaulf. is really a
species of Sticherus. Love, Léve, and Pichi Sermolli (Cytotax. Atlas Pterid.,
1977), apparently following Nakai, also adopt the name Diplopterigium farinosum
for Gleichenia bancroftii.
yb | Grammitis xiphopteroides (Liebm.) A. R. Smith, comb. nov.
25? = Polypodium xiphopteroides Liebm., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd., V.
1:196. 1849. MC ECTOTY PE sie chosen): Mexico, Veracruz, ‘‘Hac. de Mirador,’’ Liebmann [PI.
Mex. 2548, Fl. Mex. 189] (C
I regard Grammitis eee (Maxon) Proctor as a taxonomic synonym.
2%, Pellaea cordifolia (Sesse & Moc.) A. R. Smith, comb. nov.
Tie me cordifolium Sessé & Moc., Naturaleza Seeoy City), ser. II, 1(App.):182. 1890. “TYPE:
o, Cuyuacan and San Agustin near Mexico City
=" rceaid Pellaea cordata (Cav.) J. Smith ye Fée, 1852), P. cardiomorpha
Weath., and P. sagittata (Cav.) Link var. cordata (Cav.) A. Tryon as taxonomic
synonyms.
Pellaea cordifolia differs from P. sagittata in having segments rotundate-
cordate (vs. ovate-triangular to sagittate), rachis and segment stalks glabrous (vs.
usually puberulous), and spores tetrahedral-globose, 64 per sporangium (vs. ellip-
soidal, 32 per sporangium) (A. Tryon, Ann. Missouri Bot. Gard. 44:125-193.
1957). Pellaea cordifolia is a sexual diploid (2n=29 II), whereas P. sagittata s. s. is
an apogamous triploid (n =2n =87). The two species are partially sympatric, but P.
cordifolia has a much more restricted distribution than P. sagittata. The mor-
phological, chromosomal, and chorological differences between the two taxa
seem of a magnitude to justify recognition of two species.
que Pleopeltis macrocarpa var. interjecta (Weath.) A. R. Smith, comb. nov.
434 & Polypodium peltatum var. interjectum Weath. Amer. Fern J. 34:17. 1944. TYPE: Cerro de Tecpam
near Santa Elena, Chimaltenango, Guatemala, Standley 60957 (F).
As Weatherby stated, var. interjectum is closely related to var. trichophora
(Weath.) Pic. Ser. and var. macrocarpa (Weatherby’s var. lanceolata) on the one
hand and Pleopeltis polylepis (Roem. ex Kunze) Moore (= Polypodium peltatum
Cav., according to Christensen, Dansk. Bot. Ark. 9(3):11. 1937) on the other
hand. It is quite possible that var. interjectum is an evolutionary link to P.
polylepis from P. macrocarpa. My reason for placing it with the latter is that I find
that scale size and number (large and numerous in P. polylepis, small and sparse in
P. macrocarpa including var. interjectum) on the abaxial surface of the blade are
more consistent than characters of scale margin (entire in P. polylepis and entire
to erose to denticulate in P. macrocarpa). Weatherby used characters of the scale
margin to distinguish between P. macrocarpa and P. polylepis.
A. R. SMITH: NEW PTERIDOPHYTES FROM CHIAPAS 27
7%) Polystichum fournieri A. R. Smith, nom. nov.
1712 Polystichum muelleri Fourn., Mex. Pl. 1:91. 1872 (non Schum., 1803) "LECTOTYPE (here chosen):
‘‘In pinetis prov. Chiapas,’’ Mexico, Linden (P!). The other syntypes (all from Mexico) are: “*San Luis
de Potosi,’ Virlet d’Aost 46 (P-4 sheets!); ‘‘Orizaba,”’ F. Miiller 1496 (not found at P); and ‘‘Prope
ignovomum Rio Frio,’ Bourgeau (P!). Of the syntypes I have seen, only the Linden specimen is
ertile.
744 S§ticherus brevipubis (Christ) A. R. Smith, comb. nov.
THIS Gleichenia brevipubis Christ, Bull. Herb. Boissier, II. 6:280. 1906. LECTOTYPE: Valle del Rio
Navarro, Cartago, Costa Rica, Wercklé (P; isolectotype US), chosen by Lellinger, Proc. Biol. Soc.
Wash. 89:713. 1977.
742? Tectaria transiens (Morton) A. R. Smith, comb. & stat. nov.
7428 Tectaria incisa subsp. transiens Morton, Amer. Fern. J. 56:133. 1966. TYPE: Cordoba, Veracruz,
Mexico, Finck 57 (US).
In the sum of its characters, T. transiens is intermediate between T. hera-
cleifolia (Willd.) Underw. andT. incisa, and it is possible that it has arisen through
hybridization. However, in Chiapas it has been collected in areas where neither of
the suspected parents have been found. It differs from 7. heracleifolia in its
nonpeltate indusia, greater number of lateral pinnae, shorter-stalked lower pinnae,
and lack of strongly developed acroscopic lobes on lower pinnae. FromT. incisa,
it differs in the pinnae being serrately incised most of their length with two or three
large basal basiscopic lobes. From both species (Chiapas material only), 7. trans-
iens differs in the presence of short, glandular hairs on the costae, veins, and leaf
tissue below and also on the indusia. Spores of several collections appear well-
formed. Throughout its range, T. transiens seems to grow at higher elevations
than either 7. heracleifolia or T. incisa.
Additional collections seen are Sanchez 63 (UC) from Veracruz; Tuerckheim
839 (UC) from Guatemala; and Brade 47 (UC), Stork 2107 (UC), and Stork 1546
(UC) from Costa Rica. Similar collections have been seen from Ecuador and Peru.
28 AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 1 (1980)
SHORTER NOTES
A NEW COUNTY RECORD FOR PILULARIA AMERICANA IN TEXAS. —
While collecting on the interesting granite outcrops of Central Texas on July 20,
1979 I came across a local, but extensive population of Pilularia americana A. Br.
in a half-acre stock pond. The plants formed dense mats in mud at the margin of
the pond, and in several places the mat extended onto the bare granite under the
water surface. Livestock had uprooted many plants, and these were floating on
the pond surface. Each rhizome measured 10-14 cm long. The plants were all in
fruit, the sporocarps varying from olive-green to brown in color. Associated plants
included Eleocharis sp., Bacopa rotundifolia, and Lindernia anagallidea. The
pond in which the plants occurred had been enlarged from a previous vernal pool
by means of an earth dam, which seems to have helped the Pilularia poplation.
The population is the first known record for the species in Mason County. Other
populations are known in Texas only from near Marble Falls, Burnet County,
about 55 miles southeast, in similar habitats over granite. The Mason County
outcrop is known locally as ‘‘Spy Rock’’ and is located on a private ranch 8.8 mi E
on the north side of Ranch Road 1222 from the intersection with Hwy. 57, in Camp
Air, near Fredonia. Specimens are being distributed to the following herbaria: B,
F, GA, GH, K, MARY, MO, NCU, NY, SMU, TAES, TEX, US, VT.—Steven
R. Hill, Department of Botany, University of Maryland, College Park, MD 20742.
ASPLENIUM XGRAVESII DISCOVERED IN ARKANSAS.—In addition to
serving as an addendum to the Pteridophytes of Arkansas by Taylor (Rhodora
81:503—-548. 1979), this note is intended to document the value of the A.F.S.
annual fern forays as a stimulus to discovery. One of us (Werth) went to Arkansas
this past summer not only for the pure joy of a fern foray, but also with the ulterior
motive of obtaining Ozarkian and Ouachitan materials for his studies on genetic
variation in Asplenium. After a very successful foray, and at the suggestion of the
other of us (Taylor) and David Johnson of the University of Michigan, Werth
ea Hot Springs National Park in Garland County, Arkansas, on 13 August
It was known that A. pinnatifidum and A. bradleyi grew sympatrically in the
park on the novaculite outcroppings of the Gulpha Gorge. This population was
easily located, although fewer than thirty individuals of each species were found.
(Asplenium platyneuron was also present and in much greater abundance.)
Nonetheless a robust individual of A. x gravesii Maxon with leaves intermediate
between the two parent species was discovered. Fronds and a portion of the
rhizome were taken, and later examination of the spores showed them to be
abortive. Comparison of the leaves with the drawings appearing in Wagner and
Darling (Brittonia 9:57—63. 1957) confirmed that the plant was the hybrid.
This hybrid is quite rare, having been reported previously from only five states,
apparently as a consequence of the limited concurrence of the uncommon parent
species. It is likely that plants of A. gravesii have appeared sporadically in Gulpha
Gorge and died, never having been observed until the arrival one summer of an
unsated fern forayer. Voucher specimens of Werth 39K8 have been deposited at
MU and MIL.—Charles R. Werth, Department of Botany, Miami University,
Oxford, OH 45056 and W. Carl Taylor, Department of Botany, Milwaukee Public
Museum, 800 W. Wells St., Milwaukee, WI 53233.
AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 1 (1980) 29
PILULARIA AMERICANA NEW TO _ TENNESSEE.—The Pillwort, P.
americana A. Braun, a diminutive, aquatic pteriodophyte in the family Mar-
sileaceae, was first collected near Fort Smith, Arkansas by Thomas Nuttall in
1819. Since that time, its known range in the United States has been slowly
expanded.
Presently it is known in the United States from Crook (collected in 1894) and
Lake Counties, Oregon; in California from Siskiyou and Modoc Counties south-
ward to San Diego Co.; from Cherry Co., Nebraska; Reno and Harvey Counties,
Kansas; Comanche Co., Oklahoma; Burnet Co., Texas; Conway, Faulkner, Gar-
land, Logan, Sebastian, and Washington Counties, Arkansas; and Barrow, Wal-
ton, Washington, and Oglethorpe (9.3 mi NE of Lexington at Echols Mill, 11 Nov
1962, D. Blake & F. Montgomery s. n., MO) Counties, Georgia. The range of P.
americana has also been stated to include Louisiana by H. B. Correll (Amer.
Fern. J. 57:31-32. 1967), but D. S. Correll (pers. comm. with D. B. Lellinger) has
confirmed that this was an error
On 12 Aug 1979, the second author discovered P. americana washed up on the
shore of man-made Fall Creek Lake directly behind the Park Inn at Fall Creek
Falls State Resort Park, Van Buren Co., Tennessee (A. J. Petrik-Ott 1379 & F. D.
Ott, US). Following the initial discovery, we were amazed to find the shore line
behind and ca. 100 yards to the east and west of the Inn literally covered with
stranded plants of P. americana, and there were equally as many adrift in the
water along the shore. These plants ranged from near perfect specimens to those
in various stages of decay. In places along this shore, abundant stranded plants of
P. americana formed drifts up to six inches wide and one inch deep. There were
no rooted plants at this site.
The first author found rooted plants on the southeast side of the dam, growing in
sand and about six inches of water (4. J. Petrik-Ott 1380 & F. D. Ott, US). A
search of the northwest side of the dam yielded an unbelievably large plant of P.
americana which was floating and caught among the stem bases of a cattail popu-
lation (A. J. Petrik-Ott 1381 & F. D. Ott, US). This plant consisted of an ex-
tremely branched, continuous rhizome bearing numerous leaves (morphologically
rachises and stipes) and tufts of roots. There was sufficient material from this one
plant to make five rather crowded herbarium specimens. Other floating plants
were caught among the rocks of the dam on the lake side in great quantity.
Rooted plants of P. americana (only 1 to 2 cm tall) were found to bear several
sporocarps, but those found floating and stranded on shore (up to 6 cm tall) bore
only occasional sporocarps. Before our collections were made, there had been a
good deal of rain and the lake appeared to be up about a foot above normal. Many
of the stranded and floating plants may have been washed free from their rooted
places; indeed many still had sediment clinging to their roots. However, it 1s
difficult to believe that the large, extremely rhizomatous plant found on the
northwest side of the dam had been uprooted. Its rhizome was quite green, not
whitish like the rhizomes of rooted plants, and its roots free of sediment. This
plant certainly would have broken into many pieces had it previously been rooted
in the soil.
30 AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 1 (1980)
This site for P. americana extends its known range by approximately 400 miles
ENE from Faulkner County, Arkansas and 170 miles NW of Walton County,
Georgia.—Aleta Jo Petrik-Ott and Franklyn D. Ott, Department of Biology,
Memphis State University, Memphis, TN 38152.
NEW NAMES FOR POLYPODIUM CHNOODES AND P. DISSIMILE.
—Among the species of Polypodium subg. Goniophlebium, few are more distinct
than P. chnoodes Spreng. (Neue Entdeck. 3:6. 1822). Specimens of this species
have very large (5-8 mm long), strongly clathrate, blackish, spreading rhizome
scales and are weakly and evenly pilosulous on both lamina surfaces. The pinnae
are fully to partially adnate to the rachis and, in the latter case, often have a
conspicuous, basal auricle overlapping the rachis. Small, round sori are borne in 2
or 3 rows on each side of the pinna midrib. The veins anastomose in a typical
goniophlebioid pattern. The species is found in the Antilles, on Trinidad, and from
Guatemala to Venezuela and Colombia. In looking at type photographs and
specimens of New World Polypodium, I was surprised to see that the type of P.
dissimile L. (Syst. Nat. ed. 10, 2:1035. 1759), a specimen collected by Browne in
Jamaica (LINN 1251.24), is exactly the same as P. chnoodes. Apparently every-
body has adopted Sprengel’s name; nevertheless it must be put in synonymy
under P. dissimile.
The name P. dissimile has been applied consistently but incorrectly to a species
of Polypodium subg. Polypodium which has small (2—4) mm long), non-clathrate,
reddish-brown, appressed rhizome scales and which is glabrous on both lamina
surfaces, except for minute hairs on the costae. Most pinnae are partially adnate
to the rachis and are abruptly contracted at the base, often from a rather dilated
supra-basal portion. Small to medium, slighty elongate sori are borne in a single
row on each side of the pinna midrib. The veins are 2- or 3-forked and do not
anastomose. The species is found over roughly the same range as *‘P. chnoodes,”’
and in addition extends to Mexico, Peru, and Suriname. Now that P. dissimile has
to be used for what was called P. chnoodes, the next available name is P. sororium
Humb. & Bonpl. ex Willd. (Sp. Pl. ed. 4, 5:191. 1810), based on a specimen
collected by Humboldt and Bonpland near Caripe, Venezuela (B-Hb. Willd.
19684). The name P. sororium has been used occasionally in the past for some
Venezuelan specimens, but generally has been thought to be a synonym of P.
dissimile.—David B. Lellinger, U.S. Nat’l. Herbarium NHB-166, Smithsonian
Institution, Washington, DC 20560.
A NEW RECORD FOR PELLAEA ATROPURPUREA IN MARYLAND. —
While collecting specimens for the Herbarium at Towson State University, I
discovered a small colony consisting of six plants of the Purple-stemmed Cliff-
brake, P. atropurpurea (L.) Link, growing in east-facing crevices of an old rail-
road trestle at Rowlandsville, in Cecil County, Maryland. This is a new county
record for Maryland, as well as the first record of this species for the Delmarva
Peninsula. A voucher, Redman 3698, has been placed in the Towson State Uni-
versity Herbarium (BALT).—Donn E. Redman, Herbarium, Towson State Uni-
versity, Towson, MD 21204.
AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 1 (1980) 31
AN ATYPICAL ATHYRIUM FROM EASTERN TENNESSEE. — A recent bo-
tanical exploration of the Doe River gorge in Carter County, Tennessee has led to
the discovery on 8 August 1979 of an unusual growth form of Athyrium
asplenioides (Michx). A. A. Eaton. A single population consisting of several
frond-bearing rhizomes occurs along a moist, northeast-facing rock face at Pardee
Point, adjacent to the abandoned roadbed of the East Tennessee & Western North
Carolina Railroad, 1.5 miles southeast of Hampton (Wofford, Smith & Collins
79-222, TENN, US). The identity of the specimen was confirmed by A. M. Evans.
These plants resemble vigorously growing parsley and have crispate, fasciated
pinnae crowded toward the lamina apex (Fig. 1). Although this is a dramatic
departure from the usual frond morphology of Athyrium, the fragile, pale brown
scales of the stipe base, the sparsely glandular indusia, and the paired vascular
strands uniting above the stipe base into a U-shaped midvein are distinguishing
characters of the genus. Sori are infrequent, are borne intramarginally, and have
typical hammate-asplenioid indusia attached near the lateral vein. Examination of
the sori reveals aborted sporangia and a complete absence of spores. The apparent
adaptation to the rock face, atypical growth form, incomplete sporogenesis, and
FIG. 1. Specimens of Athyrium asplenioides mutant from Carter Co., Tennessee (Wofford, Smith &
Collins 79-222).
32 AMERICAN FERN JOUNRAL: VOLUME 70 NUMBER 1 (1980)
the fact that typical A. asplenioides occurs in more mesic sites along the base of
the same rock face favors the soe pe that these plants are morphological
mutants, rather than of hybrid ori
In addition to this unusual atleirtiin. Sanguisorba canadensis, Scirpus ces-
pitosus, Paronychia argyrocoma, and Drosera rotundifolia occur at the same
locality. Except for D. rotundifolia, these species are of restricted occurrence in
Tennessee and are included in the list of rare vascular plants of Tennessee (Com-
mittee for Tennessee Rare Plants. 1978. J. Tenn. Acad. Sci. 53(4):128-133).—
David K. Smith and B. Eugene Wofford, Department of Botany, University of
Tennessee, Knoxville, TN 37916, andJ. L. Collins, Division of Forestry, Fisheries
and Wildlife Development, TVA, Norris, TN 37828
Contributions from the Botanical Laboratory, University of Tennessee, n. s.
No. 520.
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AMERICAN
FERN
JOURNAL
Volume 70
Number 2
April-June, 1980
QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY
Equisetum x litorale in eae lowa
Minnesota, and Wisco
Gametophytes of Equisetum diffusum
A Double Spore Wall in Macroglossum
Subdivision of the Genus Elaphoglossum
Notes on the Natural History
of Stylites gemmifera
Ls)
a
JAMES H. PECK
RICHARD L. HAUKE 39
>
a
LUIS D. GOMEZ P. and KERRY S. WALTER
=
|
JOHN T. MICKEL and LUCIA ATEHORTUA G.
ERIC E. KARRFALT and DALE M. HUNTER 69
Reciprocal Allelopathy Between the Gametophytes
of Osmunda cinnamomea a sy Dryopteris inte
rmedia
YMOND L. PETERSEN and DAVID E. FAIRBROTHERS
Shorter Note: Thelypteris torresiana in Venezuela
Reviews
B
80
38, 68, 79
.
i
MITSOUM BOTAN RAT
JUL 15 1980
GARDEN LIBRARY
The American Fern Society
Council for 1980
ROBERT M. LLOYD, Dept. of Botany, Ohio University, Athens, Ohio 45701. President
DEAN P. WHITTIER, Dept. of Biology, Vanderbilt University, Nashville, TN 37235. Vice President
LESLIE G. HICKOK, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916
Secretary
JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916.
Treasurer
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Records Treasurer
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ALAN R. SMITH, Dept. of Botany, University of California, Berkeley, Calif. 94720
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American Fern Journal
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AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 2 (1980) 33
Equisetum X litorale in Illinois, lowa, Minnesota,
and Wisconsin
JAMES H. PECK*
The cross between Equisetum arvense L. (Field Horsetail) and E. fluviatile L.
(Water Horsetail) results in the hybrid E. x litorale Kuhl. (Shore Horsetail). It is
distinguished from its parents by its abortive spores and intermediate anatomical
and morphological traits (Hauke, 1978). The hybrid occurs less frequently in the
Midwest than hybrids between unbranched species of Equisetum. The hybrid was
reported from one county in Illinois (Mohlenbrock & Ladd, 1978) and four coun-
ties in Wisconsin (Hauke, 1965). It was not reported from lowa (Peck, 1976) nor
from Minnesota (Tryon, 1954), even though both parents occur together in those
states. Its recent discovery in Minnesota (Peck & Swanson, 1978) initiated a field
and herbarium search that uncovered a second locality in Illinois, first and second
localities in lowa, second, third, and fourth localities in Minnesota, and eight new
localities in Wisconsin. Most of these new localities are on or adjacent to the
flood-plain of the Mississippi River or its tributaries. The 2-3 ha stand located in
Allamakee County, Iowa, is probably the largest stand of the hybrid in North
America (pers. comm. with Drs. Hauke and Wagner). Consquently, efforts were
made to study the habitat, stand dynamics, and reproductive biology of the hybrid
at this locality to identify factors which clarify the origin, abundance, and persis-
tence of the hybrid in this locality. Observations also were made at 16 of the 20
localities of the hybrid in these four states, but in less detail.
A summary of locality data is provided by Fig. ] and by citations from her-
barium vouchers (new county records indicated by *). The original Illinois locality
(Lee County) is accepted from the report by Mohlenbrock & Ladd (1978); voucher
citation was not given and the specimen was not seen.
ILLINOIS: *Carroll Co.: Wet grounds near entrance to Mississippi Palisades State Park, N of
Savannah, Wunderlin 2668 (MWI).
IOWA: *Allamakee Co.: Lansing Twp.: Lansing Wildlife Refuge, 2-3 ha marsh, T99N R4W S12,
Peck 78-54 (ISTC, KIRI, MICH), Farrar 78-6-3-1 (ISC), Roosa 1759 (ISTC). Banks of Mississippi
River, June 1900, Orr s.n. (EMNM). *Des Moines Co.: Huron Twp.: Near pumping station No. 4, lowa
Slough, adjacent to Mississippi River, T72N R1W S4, Lammers 1542 and 2087 (1A, ISC, ISTC), Peck
78-300 (ISTC, KIRI, MICH).
SOTA: *Houston Co.: Mississippi River flood plain in wetlands of Crooked Creek at Reno,
Peck 79-734 (KIRI, MICH, MIN, UWL). Washington Co.: Confluence of Valley Branch Creek and St.
Croix River in thicket of Salix interior, Swanson 2878 (MIN, UWL). *Wabasha Co.: Weaver Bottoms,
floodplain of Mississippi River, 2 mi N of Weaver, Peck 79-709 (KIRI, MICH, MIN, UWL). *Winona
Co.: On floodplain of Mississippi River, W of Red Oak Island in Lake Onalaska, Peck 79-727 (KIRI,
MICH, MIN, UWL).
WISCONSIN: *Buffalo Co.: Nelson Twp.: Nelson-Trevino Bottoms of Mississippi River floodplain,
T22N R14W S36, Peck 79-824 (KIRI, MICH, WIS, UWL). *Crawford Co.: Emergent along shore near
bridge over Swamp Creek, 0.5 mi E of Lynxville on County Road B, TIN R6W S23, 8 Jul 1973,
Dawson s. n. (UWL). Grant Co.: Wilderness area of Wyalusing State Park, T6N R6W $20/21, 19 June
1959, Patman s. n. (WIS). Green Lake Co.: Marsh shore on S side of Lake Puckaway, Marquette,
Fassett 8799 (WIS). *La Crosse Co.: Barre Twp.: Seepage area at roadside ditch along Swamp Road,
*Department of Biology, University of Wisconsin-La Crosse, La Crosse, WI 54601.
34 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
TI6N R6W S820, Peck 79-803 (KIRI, MICH, UWL, WIS). *Pierce Co.: Clifton Twp.: Marshy thicket at
mouth of Kinnicinick River under Salix interior, T27N R20W, Peck 79-824 (KIRI, MICH, UWL,
WIS). Isabella Twp.: Marshy slough, backwater of Mississippi River. 0.5 mi W of Bay City, T24N
RI7W S7, Peck 79-838 (KIRI, MICH, UWL, WIS). Richland Co.: Shallow water springhole in slough
of Wisconsin River, 3 mi E of Gotham, T8N R2E S4, Hartley 5266 (IA, WIS). *Rock Co.: Marsh and
lowlands east of cooling canal on Rock River near Beloit, Rice 1649 (UWJ, WIS). *Trempealeau Twp.:
Marshy edge of backwater area of Mississippi River floodplain in Delta Fish and Fur Farm, T18N
R1OW SII, Peck 79-814 (KIRI, MICH, UWL, WIS). *Vernon Co.: Genoa Twp.: Shore to Bad Axe
River near Mississippi River, T12N R7W S12, Peck 79-813 (KIRI, MICH, UWL, WIS). Winnebago
Co.: Springy shore of Fox River near Eureka, Fassett 13243 (WIS).
The habitat of the hybrid is a shallow marsh or slough adjacent to a watercourse
which has a fluctuating water level, periodically flooding or stranding the site
where the hybrid occurs. The hybrid occurs in stands ranging from 1 m? to 2-3 ha,
with many stands 0.05-0.1 ha in extent. Species diversity within the stand is very
low compared to adjacent marshes. The most common associates are Sagittaria
latifolia, Salix interior, and Typha latifolia. The parent species were found at the
periphery of the hybrid’s stand, but rarely within the stand. The hybrid appears to
be quite aggressive in a habitat which is subjected to repeated disturbance by
flooding and sediment deposition.
TABLE 1. CHANGES IN EQUISETUM x LITORALE STAND HEIGHT AND DENSITY IN
ALLAMAKEE COUNTY, IOWA, DURING 1979.
Stand height
Stand density
Survivorship
Sample Date (¥ + sd) (stems/m?) (% of 21 April)
21 April 0.05 + 0.012 2809 + 61.8 100
12 May 0.49 + 0.049 2560 + 82.9 91
12 June 1.03 + 0.101 1792 2-133 64
30 July 1.52 + 0.082 1324 + 148 47
20 Aug! 0 0 0
15 Sep? 0.34 + 0.031 32. db 1
: Stand lodged; all aerial parts senescent.
Aerial stems newly arisen from subterranean stems.
Stand dynamics were monitored in 1979. A transect was established the length
of the stand. Twenty-five 0.25 m2 quadrats were selected at random points along
quadrat. Stand height was assessed by measuring the height of two corner plants
per quadrat. Subsequent observations were made from a series of transects 5 m
distant from and parallel to the previous transect. An average value for stand
density (stems/m?) and stem height was calculated for each transect. Survivorship
of aerial stems through the growing season was calculated based upon the average
greatest in spring and declined until summer, when only 47% of its initial stand
density was present. Considering the initial stand density (2,809 stems/m?), exten-
sive self-thinning was expected. Stand height increased through time resulting in
mean stand height of 1.52 m, with some exceptionally tall specimens over 2 m.
J. H. PECK: EQUISETUM x LITORALE 35
The stand was flattened by a severe storm in early August 1979. Consequently,
no measurements could be taken on August 20th. However, by September 15th,
the stand had sprouted new aerial stems from subterranean stems. These new
stems produced a comparatively sparse growth of limited stature. The new growth
Wi
FIG. 1. Distribution of Equisetum ~ litorale in the four states of the upper Mississippi River Valley.
protruded through a 10-18 cm thick mat of senescent and decaying aerial stems
that lodged following the storm. The stand also lodged in 1978 following a storm in
July. The new growth that year attained a mean height of 1.2 m. The extent of
recovery, therefore, is probably influenced by the time of lodging, with an earlier
date favoring a stronger recovery.
36 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
Observations on stand dynamics were also made on an adjacent pure stand of
the parent species. Those stands were less densely stocked and did not recover
following lodging to the extent the hybrid did. Whereas the hybrid occurs with
stands of low species diversity, the parent species exist with many more species
present within their stands. The lack of colonization or establishment within the
hybrid’s stands and the extensive colonization in parent stands in autumn suggests
that the thick mat of lodged stems in the hybrid’s stand physically prevents suc-
cessful invasion. Consequently, the aggressive growth of aerial stems and capabil-
ity to recover from mid-summer lodging contribute to the hybrid’s ability to main-
tain stands of low species diversity.
In considering the reproductive biology of the hybrid, it should be noted at the
outset that the hybrid’s spores are unable to produce gametophytes and, thus,
complete the sexual cycle. Furthermore, the general lack of strobilus production
by the hybrid in the upper Mississippi River valley essentially precludes the rare
event of production of an occasional viable spore. Consequently, sexual origin of
the hybrid is restricted to hybridization between its parents. Frequent flooding
along watercourses in the upper Mississippi River valley forms extensive sand and
mud flats that provide suitable conditions for Equisetum gametophytes and ample
opportunities for hybridization. The hybrid’s occurrence in small, discrete stands,
with uniform growth within a stand and readily demonstrated physical connec-
tions underground, suggests that each stand is the result of a single or a few
successful hybridizations followed by establishment of a clone by vegetative
growth. Therefore, the stand at each locality may represent an independent origin
of the hybrid.
Once formed, a hybrid plant can persist indefinitely as long as it avoids cata-
Strophic factors, such as desiccation during drought years. An indication of the
longevity of the hybird is given by the continued presence of the hybrid in Alla-
makee Co., Iowa, for at least the last 80 years. Since 1900, when Ellison Orr first
noted it on the flood plain of the Mississippi River, establishment of navigation
lock and dams has changed the hydrology of the shoreline environment. The
presence of the hybrid in that locality today suggests that its historical presence
and persistence has not been eliminated by alterations to the shoreline.
Vegetative proliferation of hybrid stands, on the other hand, was suggested by
the occurrence of small stands and isolated stems 50-350 m downstream from the
2-3 ha stand in Allamakee Co., Iowa. Fragmentation followed by water dispersal
of stems and their subsequent establishment downstream would result in new
stands from the same original plant. The propensity for vegetative proliferation of
Equisetum was discussed by Hauke (1963) and experimentally investigated by
Wagner & Hammitt (1970). In early August 1979, an experiment was conducted to
verify the hybrid’s ability to undergo vegetative proliferation and to contrast this
ability with that of its parents. Aerial stems and subterranean stems of the hybrid
in early September. The results (Table 2) indicate that burial increased the chance
that a fragment would form a new plant, that subterranean parts withstood the
J. H. PECK: EQUISETUM x LITORALE 37
stress of fragmentation, dispersal, and establishment better than aerial stems, and
that the hybrid’s ability is equal to or superior to that of its parents.
In summary, the origin, maintenance, and dispersal of the hybrid is favored by
the physical results of flooding. Flooding can form mud flats where sexual forma-
tion of the hybrid can occur, lodge aerial stems into a mat that prevents invasion of
the hybrid’s stands by potential colonizers, and facilitate vegetative proliferation
by physically breaking stems, dispersing them downstream and leaving them
stranded or buried on mud flats where new stands can become established. Al-
though vegetative proliferation probably has expanded the hybrid within its lo-
cality, evidence of long distance dispersal is lacking, in that both parents occur
with the hybrid in these states.
TABLE 2. COMPARATIVE ABILITY OF E. x LITORALE AND ITS PARENT SPECIES TO
REGENERATE PLANTS FROM AERIAL AND SUBTERRANEAN STEM FRAGMENTS
AFTER BEING LODGED ON OR BURIED IN A MUD FLAT
Species Cut and Lodged Cut and Buried
Aeria Subterranean Aerial Subterranean
E. arvense 0% 20% 60% 68%
E. x litorale 0% 16% 40% 96%
E. fluviatile 0% 8% 0% 20%
These observations suggest that analysis and integration of habitat, stand
dynamics, and reproductive ecology are feasible with Equisetum, and possibly
with other pteridophytes. Additional observations are needed on changes in nutri-
tional status (calories, macronutrients, and protein) of aerial and subterranean
stems through the year. Additional measures on stand dynamics (height, density,
biomass) are needed to contrast peripheral stands reported on here with stands to
the north and east. Competitive experiments between the hybrid, parent species,
and flowering plant associates would improve our understanding of Equisetum
ecology.
Dr. Richard L. Hauke, University of Rhode Island, and Dr. Warren H. Wagner,
Jr., University of Michigan, are thanked for verifying hybrid material and for
comments on the ecology of Equisetum. Curators of the following herbaria are
thanked for permission to examine specimens: Effigy Mounds National Monu-
ment, Marquette, [A (EMNM), Iowa State University (ISC), Mankato State Uni-
versity (MANK), University of Iowa (IA), University of Minnesota (MIN), Uni-
versity of Northern Iowa (ISTC), University of Western Illinois (MWI), Univer-
sity of Wisconsin-Janesville (UWJ), University of Wisconsin-La Crosse (UWL),
and University of Wisconsin-Madison (WIS).
LITERATURE CITED
HAUKE, R. L. 1963. A taxonomic monograph of the genus Equisetum subgenus Hippochaete. Nova
Hedw. Beih. 8:1-123.
. 1965. Preliminary reports on the flora of Wisconsin. No. 54. Equisetaceae—Horsetail Fam-
ily. Trans. Wis. Acad. Sci. 54:331-346.
. 1978. A taxonomic monograph of Equisetum subgenus Equisetum. Nova Hedw. 30:385—-
455.
38 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
MOHLENBROCK, R. H., and D. M. LADD. 1978. Distribution of Illinois Vascular Plants. Southern
Illinois University Press, Carbondale, IL.
PECK, J. H. 1976. The pteridophyte flora of Iowa. Proc. Iowa Acad. Sci. 83:143-160.
———.,, and S. D. SWANSON. 1978. Equisetum x litorale recorded for Minnesota. Amer. Fern J.
TRYON, R. M. 1954. The Ferns and Fern Allies of Minnesota. University of Minnesota Press,
Minneapolis, MN.
WAGNER, W. H., Jr., and W. E. HAMMITT. 1970. Natural proliferation of floating stems of
scouring-rush, Equisetum hyemale. Mich. Bot. 9:166-174.
REVIEW
“FLORA DE LA PROVINCIA DE JUJUY REPUBLICA ARGENTINA, PARTE
Il. PTERIDOPHYTA,” by E. R. de la Sota. Angel L. Cabrera, general editor.
Instituto Nacional de Tecnologia Agropecudria, Buenos Aires, Argentina, 1977.
Ps. 22,000 (ca. $28.00).—INTA is publishing a series of very attractive Flora
volumes for the provinces of Entre Rios and Jujuy (both in northern Argentina)
and for the Patagonian region. This volume is the first on ferns for the three series.
The endpapers have useful physiographic, phytogeographic, and political maps of
the province. The book begins with a few pages concerning the morphology,
cytology, reproduction, and systematics of the Pteridophyta. General keys lead to
23 families, which are used in a modern sense similar to that in Jermy and Mickel’s
classification. Each species treatment includes a synonymy, description, notes,
and list of specimens. Each is illustrated with a nicely drawn habit sketch plus
sketches of morphological details. Some 244 taxa are treated, judging by the index
that concludes the volume. The printing is mostly of high quality, although a few
plates had some light areas across them and a few typographical errors can be
found. The weakest point of the book is the binding. This book will be highly
useful to all who need to know the pteridophytes of northwestern Argentina and
vicinity. Orders should be sent to INTA Publicaciones, Chile 460, 1098 Buenos
Aires, Argentina.—D.B.L.
AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 2 (1980) 39
Gametophytes of Equisetum diffusum
RICHARD L. HAUKE*
Equisetum gametophytes have been studied for many years (Buchtien, 1887;
Hauke, 1967, 1968, 1969, 1971, 1977; Duckett, 1970, 1972, 1973, 1977; and litera-
ture cited in these references). The gametophytes of all Equisetum species except
E. diffusum Don have been described. In a recent monograph of Equisetum subg.
Equisetum (Hauke, 1979), I presented an evolutionary sequence of species ar-
ranged according to gametophyte specialization which did not parallel the se-
quence based on sporophyte specialization. Subsequently I was able to obtain
viable spores of E. diffusum and to culture them. The purpose of this paper is to
describe gametophytes of Equisetum diffusum and to discuss their evolutionary
implications.
MATERIALS AND METHODS
On 24 February 1974, I collected living rhizomes of E. diffusum along the road
between Chail and Kandaghat, near Simla, Himachal Pradesh state, India. These
flourished in pots in the greenhouse at the University of Rhode Island. In August
1978, I first noticed cones developing on the ends of unbranched, new stems in
two pots. The plants in these pots apparently had died back and regrown. The first
cone was removed before it expanded, surface sterilized with 50% commercial
sodium hypochlorite bleach, rinsed with sterile distilled water, and dissected in
sterile distilled water. Ten drops of the spore suspension were inoculated onto
petri dishes containing Bold’s Basal Medium (BBM) in 1.5% agar. Subsequent
cones were allowed to open naturally, and the spores were shaken onto the sur-
face of the solidified nutrient medium. The culture dishes were placed in a growth
chamber on a 12 hr light/dark cycle at a temperature of 20/15°C under 40 watt cool
white fluorescent tubes yielding 8000 ergs/cm?/sec radiant energy at the surface of
the cultures, as measured with a YSI radiometer.
Gametophytes were grown in isolation to determine crossability, self-
compatibility, and the possible occurrence of apogamy. Spores that had begun to
germinate in petri dishes were transferred aseptically into 15 mm diameter test
tube slants of BBM agar capped with metal caps and placed in racks for incubation
in the growth chamber. Initially the isolated gametophytes grew more slowly than
those left in petri dishes, possibly because the test tube caps shaded the
gametophytes; when the tubes were positioned to allow full light intensity on the
agar surface, the growth rate accelerated.
RESULTS
The spores of E. diffusum, like those of all other species of Equisetum, are
spherical, chlorophyllous, thin-walled, alete, and have two hygroscopic elaters
attached at their middles, which form four strap-like arms with spoon-shaped tips.
Under suitable conditions of moisture, temperature, and light, they germinate
readily in 1-2 days by dividing eccentrically to produce a small rhizoid cell that
*Department of Botany, University of Rhode Island, Kingston, RI 02881.
40 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
FIGS. 1+. Gametophytes of Equisetum a. FRG. i; Gametophyte a4 days old showing rhizoids,
eal (m), and plates, x 70. FIG. 2 64davsold. x 1.8.
G. 3. Archegonial gametophyte 50 Pee old, showing broad plates with. irregular margins, X nes
FIG. Gametophytes 158 days old with sporophytes. Left =selfed bisexual gametophyte, sporophyte
visible 20 days earlier. Right=crossed iia gametophyte, sporophyte visible <5, days earlier. Note
smaller size of gametophyte on right, x
R. L. HAUKE: GAMETOPHYTES OF EQUISETUM DIFFUSUM 4}
loses its chloroplasts and a large, green somatic cell. Further division of the latter
produces a flattened, linear gametophyte which branches to form several plates of
cells. Eventually a parenchymatous cushion bearing plates dorsally and rhizoids
ventrally is established by a marginal meristem. (Fig. 1).
Sex organs begin to appear 35 days after inoculation on the basal cushion
meristem of initially unisexual gametophytes. Young male and female gameto-
phytes look alike, but with continued growth they become dimorphic (Fig. 2).
The female gametophytes become larger than the male, develop numerous
plates, and assume a grass-green color (Fig. 3). The plates of E. diffusum are up to
2 mm broad and have irregular margins and thickened bases. Archegonia develop
at the base of the plates and consist of three tiers of four neck cells each. The egg
is embedded in the cushion. The terminal neck cells elongate to four times longer
than broad and spread apart in an arching manner. Seen from above, the spread
terminal necks cells resemble a pinwheel.
The male gametophytes remain smaller than the female, with only sparse plate
development, and are yellowish to pinkish. Antheridia develop from the cushion
and protrude somewhat at maturity, becoming twice as long as wide. They dis-
charge sperm by two or four cap cells. The cap cells in this species are not so
distinctive as in other species and apparently may divide anticlinally, so that at
times there are three or five cap cells.
Female gametophytes that are not fertilized within a certain time begin to pro-
duce antheridia. The marginal meristem that has been producing archegonia grows
out into an antheridial lobe. In petri dish cultures, where interaction between
gametophytes is possible, the bisexual condition may become apparent within 50
days. In isolation tubes, where interaction is not possible, bisexuality is delayed to
90 days or later. Some isolated gametophytes 150 days old still appear only
female. Gametophytes apparently cease growth when they begin bearing
sporophytes (Fig. 4). If they are unisexual at that time, they never become bisex-
ual. Unlike the situation usually seen in other pteridophytes, Equisetum
gametophytes normally bear several sporophytes per gametophyte (Fig. 4).
Whereas more than 50% of the gametophytes in plate cultures are antheridial, in
isolation tubes, in the absence of interaction between gametophytes (but presum-
ably with better nutrition), only 5 out of 42 (12%) were male. The other 37 were
female. One tube inadvertently received two gametophytes initially, one of which
became male and the other female. Three of the male gametophytes were trans-
ferred to tubes with female gametophytes and flooded. The tube with two
gametophytes was flooded (Fig. 4), as were 17 tubes with only female
gametophytes. The four tubes with both male and female gametophytes all showed
sporophytes a month later. None of those with only female gametophytes did. The
tubes originally containing only female gametophytes were reflooded, and eventu-
ally they became bisexual and selfed (Fig. 4). At 158 days post inoculation the
experiment was terminated and the tubes refrigerated to stop growth. Later they
were examined and discarded, at which time 14 of the 17 selfed gametophytes had
visible sporophytes. In two cases, the sporophytes all looked achlorophyllous.
42 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
Of the 17 tubes never flooded, one had a single visible sporophyte and another
had four. In both cases, antheridial lobes had arched over to press antheridia
against the archegonial meristem, and presumably sperm had been able to enter
the archegonia in the absence of free water. The other 15 tubes showed no visible
sporophytes. In general, flooded gametophytes grew slightly more than did un-
flooded ones.
DISCUSSION
The gametophytes of E. diffusum are similar to those of other species of
Equisetum in basic form. They consist of a parenchymatous cushion bearing
rhizoids ventrally and photosynthetic plates dorsally. In sexual condition, they
produce separate antheridial and archegonial gametophytes, with the latter sub-
sequently becoming bisexual if unfertilized. Sexual dimorphism, cessation of
growth in sporophyte-bearing gametophytes, and production of numerous
sporophytes per gametophyte are all seen in other species of Equisetum. The last
character is rather uncommon in archegoniates, where gametophytes usually have
several archegonia, but normally produce only one sporphyte per gametophyte.
Bruce and Beitel (1979) tabulated 476 Lycopodium gametophytes, of which 17%
had more than one sporophyte and one had 13 sporphytes. Mesler (1977) reported
up to 27 sporophytes on a wild gametophyte of Equisetum hyemale. The increased
percentage of archegonial gametophytes in isolation culture and the delay in their
becoming bisexual also occurs in other species of Equisetum (Hauke, 1977).
Certain features of E. diffusum gametophytes indicate that they are relatively
primitive and more closely allied to those of E. telmateia than to any other
species. Their plates are broad, with irregular margins and thick bases. Their
archegonia have terminal neck cells about four times longer than wide. Their
antheridia have 2—4 cap cells and are about twice as long as wide. Plate develop-
ment on antheridial gametophytes is more extensive than in other Equisetum
species except E. telmateia.
Duckett (1973) carefully described the morphology of several species of subg.
Equisetum; using that information plus my own observations (Hauke, 1968), I
aligned the gametophytes of subg. Equisetum in a presumed evolutionary se-
quence (Hauke, 1979), with the most primitive species being E. telmateia:
arvense
telmateia __—_- pratense me = PALUStTe me =f Viatile ep = bOZOteNnse
sylvaticum
From my current observations of E. diffusum gametophytes, it is now obvious
that in an evolutionary scheme based on gametophytes, E. diffusum is inter-
mediate between E. telmateia and the three species of sect. Heterophyadica sensu
Hauke: E. arvense, E. pratense, and E. sylvaticum.
One problem raised by this sequence is that it does not correlate well with the
apparent evolutionary sequence based on sporophyte character:
fluviatile > palust > bogot > diff i sich
p > gotense —P CITUSUM —eempe telmateia ————e pratense
sylvaticum
R. L. HAUKE: GAMETOPHYTES OF EQUISETUM DIFFUSUM 43
That sequence, it is true, placed E. diffusum and E. telmateia side by side, but it
also considered E. bogotense to be most like E. diffusum. This is the greatest
discrepancy between the schemes based on sporophyte and gametophyte evolu-
tion. Perhaps there is no reason why the two generations should be correlated
evolutionarily, since they presumably evolve for different environmental fitness,
but it seems appropriate to consider the whole plant in a single evolutionary
scheme. In that case, one might expect the gametophyte to reflect more conserva-
tive traits, as the reproductive structures of other plants are assumed to do. For an
example of independent selection for floral and vegetative fitness in flowering
plants, however, see Wilken (1978). If taxonomy is intended to be phylogenetic,
should it emphasize gametophyte or sporophyte? One factor which diminishes the
usefulness of the gametophytic stage in pteridophyte taxonomy is the paucity of
characters it possesses. A compromise scheme, utilizing characters of both
gametophytes and sporophytes might be:
arvense
pratense
palustre sylvaticum
bogotense diffusum
telmateia
It injures my sense of the fitness of things to look at two species with sporophytes
as similar as those of E. bogotense and E. diffusum and to separate them widely in
a classification. Yet the gametophyte as well as the sporophyte must be consid-
ered in arriving at any taxonomy which claims to be phylogenetic. In fact, if it is
more conservative in evolution, then it should be given greater emphasis in
taxonomy.
Isolation experiments showed that the gametophytes of E. diffusum outcross
readily. They also self readily in most cases, but the absence of any detectable
sporophytes on three of 17 selfed gametophytes, and the chlorotic sporophytes on
two others, indicates some lethal load (see Lloyd, 1974). The absence of
sporophytes on individuals that were not flooded indicates the absence of
apogamy in E. diffusum. Ease of selfing and absence of apogamy are also found in
the other species of Equisetum.
I wish to thank Dr. Stoddard Malarky for helping me collect E. diffusum and Dr.
Roger Goos for reading the manuscript.
ADDENDUM
In a paper which appeared while this article was awaiting publication, Duckett
(1979) made several observations which are pertinent here. He reported that pro-
longed culture and numerous attempted fertilizations were required to obtain
maximal sporophyte frequencies, and suspected ‘‘leaky lethals,”’ but on the basis
of crossing and selfing tests discounted that possibility. He noted that the initia-
tion of sporophytes is accompanied by cessation of gametophyte growth, and
attributed this to allelopathic substances from the sporophyte. He reported that
polyembryony is present in all species, but is uncommon and occurs mostly in
ot AMERICAN FERN JOURNAL: VOLUME 70 (1980)
Species with rapid sex change from female to bisexual, or with numerous archego-
nial lobes. | have observed polyembryony to be common in isolated, selfed
gametophytes not only of E. diffusum, but also of E. fluviatile, E. hyemale var.
affine, and E. arvense, and suspect that they were all larger and therefore with
more receptive archegonia when flooded than were those of Duckett’s e xperi-
ments
LITERATURE CITED
BRUCE, J. G., and J. M. BEITEL. 1979. A community of Lycopodium gametophytes in Michigan.
Amer. Fern J. 69:33-41.
BUCHTIEN, O. 1887. Entwicklungsgeschichte des Prothallium von Equisetum. Biblio. Bot 2(8):1-49.
DUCKETT, J. G. 1970. Sexual behaviour of the genus Equisetum, subgenus Equisetum. Bot. J. Linn.
Soc. 63:327-352.
. 1972. Sexual behaviour of the genus Equisetum, subgenus Hippochaete. Bot. J. Linn. Soc.
65: 87-108.
. 1973. Comparative morphology of ai gametophytes of the genus Equisetum, subgenus
auisetian, Bot. J. Linn. Soc. 66:1-2
. 1977. Towards an understanding of sex | detienination j in Equisetum: an analysis of regenera-
tion in gametophytes of the subgenus Equisetum. Bot. J. Linn. Soc. 74:215-242.
. An experimental study of the reproductive biology and hybridization in the European
and North American species of Equisetum. Bot. J. Linn. Soc. 79:205-
HAUKE, R. L. 1967. Sexuality in a wild population of Equisetum arvense gametophytes. Amer. Fern
57:59-66.
. 1968. Gametangia of Equisetum bogotense. Bull. Torrey Bot. Club 95:341-345.
. 1969. Gametophyte development in Latin American horsetails. Bull. Torrey Bot. Club
96: 568-577.
- 1971. The effect of light quality and intensity on sexual expression in Equisetum
gametophytes. Amer. J. Bot 73-377.
1977. Experimental studies on growth and sexual determination in Equisetum
Co Amer. Fern J. 67:18-31.
979. A taxonomic monograph of Equisetum subgenus Equisetum. Nova Hedwigia 30:385—
od
LLOYD, R. M. 1974. Reproductive biology and evolution in the Pteridophyta. Ann. Missouri Bot.
Gard. 61:318-331.
MESLER, M. R., and K. L. LU. 1977. Large
California. Amer. Fern J. 67: 97-98.
1978. Vegetative and floral relationships among western North American popula-
tions of Collomia linearis Nuttall (Polemoniaceae). Amer. J. Bot. 65:896-901.
gametophytes of Equisetum hyemale in northern
AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 2 (1980) 45
A Double Spore Wall in Macroglossum
LUIS D. GOMEZ P. and KERRY S. WALTER*
Since the original description of Macroglossum alidae by Copeland (1909a, p.
343; 1909b, p. 9), little has been learned about this strange and primitive member
of the Marattiales. Campbell (1911, 1914a, 1914b) has dealt with the anatomical
aspects of the gametophytes and sporophytes but, surprisingly, he gives no de-
scription of the spores. In Erdtman (1957, p. 79), the proximal and distal faces of
M. alidae are illustrated both in surface view and in optical section; the spore is
globose but distally flattened, with an irregular and sparsely tuberculate exine.
Kremp & Kawasaki (1972, p. 4) described the spores from seven specimens as
rounded-triangular (32.4 x 28.7 wm), trilete, and scabrate.
The observation of spores of M. alidae (Molesworth-Allen 3197, US) under the
SEM shows a regularly and densely tuberculate-subbacillate perispore (Fig. 1),
the shape and dimensions of which are in agreement with the foregoing authors.
An unusual case of double-walled spores is shown in Fig. 2, a preparation from the
same specimen. In it, the outer exine layer is cracked to reveal a smaller but
morphologically perfect spore inside, one per *‘parent spore.’’ This phenomenon
is hitherto unreported in the fern literature but may not be rare, for we have
observed a ‘“‘parent spore’ of Botrychium sp. containing four ‘‘daughter spores,”
and it is quite possible that such ‘‘angiospores’’ occur in other pteridophytes.
The biosystematic implications of angiospore production are, as yet, unknown.
Research is needed to elucidate various questions that come to mind, including:
(a) What, if any, percentage of angiospores is viable? (b) What is their genotype
and resulting phenotype? (c) What is the ploidy level of angiospores in relation to
‘‘parent spores’’? (d) Do angiospores represent a reduction mechanism for poly-
ploidal pteridophytes? (e) Within a sporangium, what percentage of ‘parent
spores’’ contain angiospores? (f) What effect would this ratio have on the popula-
tion structures of the resulting gametophytic and sporophytic generations? (g)
Does the smaller size the angiospores have any effect on the range and pattern of
their dispersibility? (h) Is angiospory a primitive trait only to be found in eu-
sporangiate pteridophytes?
pore size and shape, of themselves, are not indicative of viability. Recent
literature abounds with examples of the germination of supposedly non-viable
abortive spores in hybrids and of larger than normal diplospores formed through
ameiotic apogamy. At present, questions (b) and (d) are unanswerable due to the
lack of appropriate materials. The fact that the cytology of Macroglossum has
never been investigated prevents speculation on whether angiospores represent
any change in ploidy level, be it reduction or augmentation. The Marattiales have
high chromosome numbers as do the Ophioglossales, the only other instance in
which we have as yet observed angiospory. Until a large enough quantity of both
‘‘parent spores’ and angiospores are cultured, questions (e) and (f) also remain
unanswerable.
*Museo Nacional de Costa Rica, Apartado 749, San José, Costa Rica.
46 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
The Marattiales show a high degree of endemism; Macroglossum, for instance,
is confined to Borneo. It is logical to assume that smaller spores, such as angio-
spores, might be more easily dispersed. On the other hand, smaller spores could
be short-lived, reducing their dispersibility and enhancing endemism. The genus
Botrychium is cosmopolitan, but throughout its range its species show complex
patterns of geographically separated cytological races. This genetic variation may
be partly responsible for the taxonomic chaos within the genus. It may be that
angiospory is related to these races.
-
(1)
FIG. 1. Normal spore of Macroglossum alidae, x 3075. FIG. 2.
1535.
2]
Double-walled spore of M. alidae, x
LITERATURE CITED
CAMPBELL, D. H. 1911. The Eusporangiatae. Carnegie Inst. Wash. Publ. 140.
. 1914a. The genus Macroglossum Copeland. Phil. J. Sci. 9:219-226, t. I.
: — The structure and affinities of Macroglossum alidae, Copeland. Ann. Bot. 28:651-
See ae B. 1909a. New genera and species of Bornean ferns. Phil. J. Sci. 3:343-352, t.
1909b. The ferns of the Mala
; y-Asiatic region. Part I. Phil. J. Sci. 4:1— 7
ERDTMAN, G. 1957. Pollen and Spor. hil. J. Sci. 4:1-66, 1. I-XXI
€ Morphology/Plant Taxonomy. Almavist & Wiksell, Stock-
holm.
KREMP, G. O. W. and T. KAWASAKI. 1972. The Spores of the Pteridophytes. Hirokawa, Tokyo.
AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 2 (1980) 47
Subdivision of the Genus Elaphoglossum
JOHN T. MICKEL and LUCIA ATEHORTUA G.*
The genus Elaphoglossum is one of the largest and most complex of fern genera.
It is composed of perhaps 600 species, which frequently are difficult to distin-
guish. The genus has not been afforded careful study in the past. As a result, some
names have been misapplied and others not used at all, and sometimes new names
have been made for old species. Consequently, much herbarium material is either
misidentified or unidentified. ‘‘Elaphoglossum is more in need of a taxonomical
revision than other fern genera, all the more so as many species are imperfectly
known or badly delimited in comparison with their allies.’’ (Pichi Sermolli, 1968).
Elaphoglossum is a remarkably uniform genus of mostly simple-bladed ferns
with acrostichoid sori. The veins are free (with two exceptions), the rhizomes
scaly, and the blades densely scaly to nearly glabrous. The taxonomy of the genus
is based to some extent on frond form and rhizome habit, but more importantly on
the scales of the rhizome and blade.
Until now there has not been a useful treatment that conveniently breaks the
large number of species into smaller, more coherent units. Modern keys have been
made for a few areas, such as tropical Africa (Schelpe, 1969), Brazil (Alston,
1956a), Guatemala (Mickel, 1980b), Malaysia (Holttum, 1978), and India (Sledge,
1967), but these do not provide insight into treatment of the genus as a whole. The
purpose of this paper is to take the first step in revising the genus by breaking it
into infrageneric units that subsequently can be monographed. Even so, this
treatment is provisional because a complete understanding of the genus can be had
only after the species are better known.
HISTORY OF THE GENUS
Linnaeus (1753) described Elaphoglossum crinitum under Acrostichum, which
included all ferns with sporangia covering the dorsal blade surface. Schott (1834)
first proposed the name Elaphoglossum, but it was not formally described until
later by John Smith (1841, p. 148), and was not widely accepted until the end of the
century.
The first broad treatment of the genus was prepared by Fée (1845) under Acro-
stichum. He divided the elaphoglossoids into two groups, Oligolepideae and
Polylepideae. These in turn were subdivided on the basis of frond size and scale
characters. Later, Fée (1852) used four primary groups without further subdivi-
sion: Oligolepideae, Polylepideae, Pilosellae, and Chromatolepideae.
Moore (1857-1862) was the first to use the genus name Elaphoglossum exten-
sively; he made many new combinations under it. He utilized a generic breakdown
of Oligolepidum (‘‘fronds naked, or with but few scales’’) and Polylepidum
(‘fronds clothed with numerous scales’’). Sodiro (1897, under Acrostichum) used
more group names (Glabra, Setosa, Oligotrichia, Polytrichia, Squamosa,
Oligolepidia, Laciniata, Polylepidia), but without designating nomenclatural rank.
*New York Botanical Garden, Bronx, NY 10458.
48 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
Diels (1899) made two sections of the genus: Eu-Elaphoglossum for the bulk of
the species and Hymenodium for the distinct, broad, net-veined E. crinitum. His
informal subsectional groups largely followed those of Sodiro.
In the same year, Christ (1899) published his classic Monographie des Genus
Elaphoglossum, which stands as the most detailed treatment of the genus, cover-
ing 142 species (and numerous synonyms) in 32 groups. The primary division was
based on venation: ordo Stenoneura with veins running all the way to the margin
without thickened vein ends and ordo Condyloneura with veins ending just short
of the margin with swollen vein ends (hydathodes). Christ further subdivided
these into sections, subsections, and divisions, using characters such as blade
scales, frond size, stipe articulation, and rhizome habit and thickness. Basically,
his species groups were natural and recognizable (nearly all of our ultimate divi-
sions are based on them). Unfortunately he placed what we believe to be unrelated
divisions and subsections together and did not provide a usable key to the groups.
For example, Christ placed some of the species with black, marginal, subulate
scales in each of the two ordos, when in fact they are very closely related and none
shows the hydathodes of ordo Condyloneura. By the same token, his subsections
Dimorpha, Petiolosa, Pilosa, and Ovata in Condyloneura show no signs of
hydathodes and belong with close relatives in ordo Stenoneura. This is not to say
that hydathodes are a poor character; we would have adopted Christ’s two ordos
as subgenera were it not for misgivings about possible convergent evolution of
subulate scales and hydathodes or lack thereof.
As generally accepted, the species of Elaphoglossum have narrowly elliptic,
undivided fronds, but several species of elaphoglossoid affinity with divided
fronds are known. Several small, flabellately divided species were first generically
segregated by Link (1841) as Peltapteris and pinnately divided plants by Presl
(1851) as Microstaphyla. Although some of the American species had the pinnate
form of Microstaphyla and had been placed in that genus, Gomez (1975) saw that
their true relationship was with Peltapteris. He left only the type species (from St.
Helena) in Microstaphyla. One large, pedately divided species, E. cardenasii, was
described from Bolivia by Wagner (1954); strangely, no one has proposed a sepa-
rate genus for it. Christ placed the species of Peltapteris (as Rhipidopteris) and
Microstaphyla under Elaphoglossum and, after showing that frond architecture
was the sole character for distinguishing these groups from species of Elaphoglos-
sum, Mickel (1980a) concurred in keeping them all in Elaphoglossum.
RELATIONSHIPS OF THE GENUS
The relationships of the elaphoglossoid ferns to other ferns are unclear.
Elaphoglossum is placed close to the lomariopsid genera by some authors because
of the acrostichoid sori, largely epiphytic habitat, monolete spores, and chromo-
some number of x=41. Holttum (1947) placed it in the Lomariopsidoideae of his
Dennstaedtiaceae, and Alston (1956b) placed it in the Lomariopsidaceae. Crabbe,
Jermy, and Mickel (1975) placed it in the Elaphoglossoideae of the Aspleniaceae
and close to the Lomariopsidoideae. Pichi Sermolli (1968) pointed out the dis-
tinctness of the group from the lomariopsids and erected a new family for it, the
MICKEL & ATEHORTUA: SUBDIVISION OF ELAPHOGLOSSUM 49
Elaphoglossaceae, but placed this next to the Lomariopsidaceae. Christensen
(1938) gave it its own subfamily Elaphoglossoideae in the Polypodiaceae, but
could not determine its relationship with certainty.
Sporne (1975) and Holttum (1947) have suggested a davallioid relationship be-
cause of similarities of stele and chromosome number. We have found some
spores which closely resemble those of Bolbitis appendiculata of the lomariopsids
(Hennipman, 1977, pl. 4). Many other spores are similar to those of various
species of Oleandra and Arthropteris (illustrated by Liew, 1977). This is especially
interesting because Oleandra, like Elaphoglossum, has simple blades, straight,
free veins, phyllopodia, hydathodes, and chromosome number of x=41.
MORPHOLOGICAL CHARACTERS
Rhizome habit.—Most species have short-creeping rhizomes, but they range
from moderately to long-creeping and from ascending to erect, and are 1-12 mmin
diameter. Members of a single subsection usually have a similar rhizome habit,
but the entire spectrum can be found within subsection Pachyglossa.
Aerophores.—Elaphoglossum aerophores are pale, aerenchymatous out-
growths from the lenticel line on the stipe base or adjacent rhizome, one on each
side of the stipe. They may appear as fleshy wings about 1 mm broad along the
phyllopodia or as tongue-shaped emergences from the stipe base or adjacent
rhizome. They are not conspicuous on dried specimens, especially in those
species with fasciculate fronds, which makes a survey based on herbarium mate-
rial difficult. Lloyd (1970) made a study of living specimens in Costa Rica and
found aerophores in 92% of the species studied. More extensive systematic and
morphological study of this structure is needed.
Phyllopodia.—These dark, sclerified stipe bases are not found in all species and
are concentrated in a few subsections, particularly those with coriaceous, sub-
glabrous fronds (esp. subsects. Pachyglossa and Huacsaro and sect. Amyg-
dalifolia). Pichi Sermolli (1968) distinguished Peltapteris from Elaphoglossum on
the basis of the former’s lacking phyllopodia, but Peltapteris’ closest relatives,
which are in Elaphoglossum sect. Squamipedia, also lack them. Most sections or
subsections either lack phyllopodia or have them only poorly developed.
Frond size.—Christ (1899) distinguished some of his glabrous fasion by frond
size. To some extent this is valid. Section or subsection members are consistently
approximate in size. Subsections may have small and medium fronds or medium
and large fronds, but generally not both small and large, although there are di-
minutive specimens of nearly all species. There are also very long individual
specimens in groups with normally medium-sized fronds (e.g., E. vestitum to 1.7
m long and E. herminieri to 2 m long).
Blade shape.—Although the blades are basically simple and unlobed, there is
some variation in form. They are fairly uniformly ovate-elliptic in the E. lindenii
complex (subsect. Setosa), lanceolate with an obtuse apex in subsect. Muscosa,
narrowly elliptic with an obtuse apex in subsect. Huacsaro, and linear-elliptic in
subsect. Eximia. Blade shape seems to be more consistent in species groups or
series than in sections or subsections.
50 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
Venation.—Elaphoglossum fronds are basically free-veined. Elaphoglossum
crinitum has net venation and has been set aside as the genus Hymenodium by
some authors, but E. crassifolium, which also has net venation, does not seem to
be closely related. Elaphoglossum decoratum has an occasional anastomosis, and
in several species the vein endings are laterally extended and even fused occa-
sionally (e.g., E. acutissimum) to form a commissural vein. In most species the
veins diverge from the midvein and go to the margin at an angle of 70—80°. In
several sections (esp. Undulata, Setosa, Eximia, and Amygdalifolia), they diverge
at a narrower angle (40-60°), are farther apart (ca. 2 mm), and terminate about 1
mm short of the margin, often ending in a round thickening or hydathode. Appar-
ently in all species the veins end slightly short of the margin, but in many species
the blade is coriaceous with slightly recurved margins, making the actual vein
endings difficult to distinguish. Therefore, some authors (including Christ) have
claimed that the veins go to the margin.
Hydathodes.—The veins in the blade generally end near the margin and are at
least slightly swollen. In certain groups, the veins end 1-2 mm short of the margin
and are greatly enlarged to form hydathodes. Whether these function differently
from those not enlarged is not known. Hydathodes are found in all species of
sects. Setosa, Eximia, Undulata, and Amygdalifolia, and are not known from any
species of the other sections. These sections comprise what Christ designated
ordo Condyloneura. We do not recognize these as subgenera here because it is
unclear whether there are close relationships between members of the two ordos,
as perhaps between sects. Setosa (with hydathodes) and Polytrichia (lacking
them)
Blade texture.—Among members of a subsection and closely related groups,
texture seems to be nearly uniform. Coriaceous fronds generally are nearly gla-
brous, but whether all species with such fronds are related is questionable. The
absence of blade scales makes it difficult to assess relationships. Most species
have herbaceous to firm laminae, but those with hydathodes are usually thin.
Rhizome scales.—The rhizomes are generally densely scaly, with linear, lan-
ceolate, ovate, or rarely even round scales, which are attached at a cordate to
peltate base. Scale color ranges from bright orange to maroon, brown, or black.
Texture varies from thin to sclerotic. Scale margins in most species are entire or
have weak teeth, but in sect. Lepidoglossa they often are fringed with slender,
hair-like teeth. Differences in cell patterns have not yet been examined systemat-
ically, but probably will be helpful taxonomically. In some groups within sect.
Elaphoglossum, the rhizome scales may be deciduous, leaving in some cases
completely glabrous creeping rhizomes. In a few species the naked rhizome is
black and glutinous.
Blade scales.—There is great diversity in scale morphology, but the blade scales
are basically lanceolate. They may be erose-margined (E. muscosum), entire (E.
decoratum), or deeply ciliolate-toothed (subsect. Polylepidea). Some are so
deeply ciliolate-toothed there is barely any scale body (E. vestitum) or are reduced
to stellate hairs (E. pilosum). In E. tectum, the stipe and upper blade scales are
MICKEL & ATEHORTUA: SUBDIVISION OF ELAPHOGLOSSUM 51
round-peltate with erose margins, whereas those of the lower blade surface are
stellate hairs. In E. petiolatum, E. huacsaro, and their relatives, the blade scales
are further reduced to resinous dots. Coriaceous, subglabrous groups often have
reduced, minute, black, linear or stellate blade scales.
There are no multicellular hairs in Elaphoglossum; those thought to have hairs
(sects. Setosa and Polytrichia) have in reality hair-like subulate scales which are
linear-lanceolate with inrolled margins. It is not clear whether all groups with
subulate scales are related or not. Although they have distinctive, minute, glandu-
lar hairs in common, they differ in their spores.
Stipe scales.—They are like those of the rhizome at the base and like those of the
blade near the apex. Sometimes these are transitions from one to the other type,
but in other cases, two distinct types are intermixe
Glandular hairs.—Minute, erect, unicellular, sland. tipped hairs are located on
the stipe in sects. Polytrichia, Setosa, and Eximia. Their presence seems to be
correlated with the presence of subulate scales. In subsect. Apoda, these hairs
also are found on the blade surface.
Fertile fronds.—In most species, the fertile fronds have narrower and shorter
blades and proportionally longer stipes than the sterile ones. The fertile fronds
may be longer, essentially equal, or noticeably shorter than the sterile ones. The
relationship does not seem to be consistent in whole sections or subsections, but is
so in species groups or occasionally in subsections (fertile longer in subsects.
Huacsaro and Pilosa).
In some species, the fertile frond is folded in half lengthwise along the midvein
until maturity. This conduplicate fertile blade is found in E. lindenii and E. pilosel-
loides of sect. Setosa, but not in all species of the section; in subsect. Petiolosa,
both species show this condition.
The indument of the fertile frond generally is the same on the upper surface as
on that of the sterile blade. However, there are a few species that have scales on
the lower surface among the sporangia, and their presence is especially notewor-
thy. It is not consistent with subsections, occurring in E. villosum, E. muscosum,
E. siliquoides, all of different subsections, but seems to be common to all mem-
bers of subsect. Plumierana.
In certain species, there seems to be a conspicuous sterile margin, that forms a
pale border 1-2 mm wide outside the sporangial mass. It is especially noticeable in
some thin-textured species such as E. albomarginatum. Although it is not readily
apparent in most other species, close examination shows that the sterile margin is
underrolled and thus concealed. Probably some sort of sterile margin exists in
most or all species, which is to be expected since the veins do not reach the
margin.
Spores.—Elaphoglossum spores usually have been described as having narrow
crests with minute spicules on the surface (Erdtman, 1957, based on E. vieillardii
from New Caledonia), but our survey of spores using the scanning electron micro-
scope has shown considerable variation within the genus. We have examined 163
species, including representatives of all sections and subsections. Figures 1-18
are based on specimens in the New York Botanical Garden herbarium.
52 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
Spores of most species have slender crests or low folds or ridges. Section
Squamipedia (Fig. 7) also has low ridges, but the entire surface is densely or-
namented with distinctive spicules found also in the flabellately or pinnately di-
vided subsect. Peltapteris. High slender crests are found in sects. Amygdalifolia,
Decorata, and in some members of sects. Setosa and Elaphoglossum. Subsection
Pachyglossa, probably the largest in the genus, displays several different spore
types: some species have tall, nearly spine-like, smooth or perforated crests, to
highly fenestrated, lace-like crests; others have long, slender crests lacking holes;
and others have low, broad ridges.
Spores with low, broad ridges are found in sects. Lepidoglossa (Fig. 9), Poly-
trichia (Fig. 12), and part of sect. Elaphoglossum subsect. Pachyglossa (Fig. 3).
The ridges tend to be short (Figs. 9 and /2) rather than elongate, and often have
small verrucae in the valleys between the ridges, which suggests a possible rela-
tionship among these groups. On the other hand, the lack of other ornamentation
could indicate either a common or a primitive spore type or a condition arrived at
independently. Correlations of additional characters are needed to resolve this.
Spores of subsect. Muscosa are unique in having verrucate surfaces (Fig. //).
The spores of sects. Eximia, Undulata, and part of sect. Setosa subsect. Setosa
lack ridges or crests of any sort. Instead, they have a dense covering of short
spines which branch at the base to form a reticulum (Figs. 14 and /6-/8). In sect.
Undulata, the spines occasionally fuse to form small crests (Fig. 18). In E,
beaurepairii (sect. Eximia, Fig. 17) the reticulum is very Open, whereas in sect.
Setosa subsect. Setosa (E. crinipes et aff.) the branches often fuse laterally to
make a more dense covering on the spore surface (Fig. 14). Spores of some
members of subsect. Setosa (E. lindenii et aff.) and Alpestria have low, slender
crests which often have holes of various sizes in them (Fig. 13), somewhat like
those mentioned for subsect. Pachyglossa. Holes also occur in the spore surface,
which in some cases resembles somewhat the spore surface reticulum of sects.
Undulata and Eximia. Perforations, at least in the body of the spore, also are
found in some species of subsect. Pachyglossa, but whether this indicates a rela-
tionship has not yet been determined.
Not every species fits into the spore pattern for its group. Some species of
subsect. Pilosella have holes in the surface and others do not. In subsect.
Plumierana, E. buchii (Fig. 15) and E. plumieri have solid crests, whereas the
morphologically similar E. lanceum has highly fenestrated crests. In general,
however, spore morphology has been quite useful in confirming relationships
presumed on other grounds and in Suggesting new phyletic interpretations.
CONSPECTUS OF THE GENUS ELAPHOGLOSSUM
(INCLUDING CHRIST’S INFRAGENERIC GROUPS IN PARENTHESES)
sect. Elaphoglossum (sect. Craspedoglossa)
MICKEL & ATEHORTUA: SUBDIVISION OF ELAPHOGLOSSUM 93
sect. hi emia Mickel & Atehortuta
sect. Squamipedia Mickel & Atehortua (subsect. Pachyglossa div. Squamipedia)
pa oa aghiapten (Link) Mickel & Atehorttia (subsect. Pachyglossa div. Rhipidopteris)
subsect. Ova
subsect. Bt Christ
sect. Decorata Mickel & Atehortta (subsect. Platyglossa div. Decorata)
- sect. Lepidoglossa Christ (sect. Gymnoglossa)
subsect. Polylepidea Christ (subsect. Polylepidea div. Auricoma)
a Microlepidea Christ (subsect. Microlepidea div
subsect. Pilosa Christ (subsect. Pilosa div. Grata; subsect: Microlepidea div. Viscosa; subsect.
cans lepidea div. Stipitata; subsect. Dimorpha)
subsect. Petiolosa Chris
subsect. Huacsaro Ee & Atehortua
aba Muscosa Mickel & Atehortia (subsect. Polylepidea divs. Muscosa and Bellerman-
niana)
sect. eter Chris
subse emeilouiok (Fée) Christ
eee Hybrida Christ (subsect. Platyglossa div. Melanolepidea; subsect. Hybrida by type and
description but not by other included species)
subsect. Apoda Mickel & Atehorttia
sect. Setosa (Christ) Mickel & Atehortua
subsect. Alpestria Mickel & Atehortta
ec Plumierana Mickel & Atehortta
sect. Eximia Mickel & Atehortua
subsect. Eximia Mickel & Atehorttia (subsect. Hybrida by included species but not by type or
escription)
subsect. aca Mickel & Atehortua
sect. Wali
sect. Amygdalifolia (cist Mickel & Atehortta
The following of Christ’s groups are of unknown relationship: subsect. Glos-
soides, subsect. Coespitosa, subsect. Pachyglossa div. Micradenia, subsect.
Polylepidea div. Argyrophylla, subsect. Polylepidea div. Fimbriata, subsect.
Pilosa div. Boraginea, and subsect. Pilosa div. Gardneriana.
KEY TO THE SECTIONS OF ELAPHOGLOSSUM
1. Veins ending short of the margin, enlarged at the tip to form generally conspicuous pneeene
2. Rhizome long-creeping; blade glabrous; phyllopodia present, but short. ...... sect. mall
2. Rhizome short- to long-creeping; blade scaly, only rarely glabrous; gral aaa
3. Blade linear-elliptic or pedately divided: blade scales very small, dark, lanceolate, not ig ane
sect. Eximia
3. Blade narrowly iced or ovate-lanceolate; blade scales subulate or cordate- sr once pale.
4. Blade scales subulat t. Setosa
4. Blade scales tdi ene. erose or toothed sect TF lakes
1. Veins ending at or very close to si margin, not ending in hydathodes.
5. Blade glabrous or subglabro
6. Rhizome long-creeping, font very small (2-20 cm long), usually lacking lg anes blade
simple to finely dissected. Squamipedia
6. Rhizome erect or short- to clei ade fronds small to large (12-200 cm m lone) phyllopodia
distinct or indistinct but always present t. Elaphoglossum
94 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
ong 1-6. Spores of Elaphoglossum sect. oe FIG, 1. E. pteropus ipo 19180), x
ac it leg a e598 ai Pa Ae IG. 3. E. dussii rie ay $.n.), FIG. 4.E
ra ;
Wacket 7849), pg ), * 1200. FIG. 5. E. bicolor (Wacket 121), x 1560. FIG. i E. wacketii
MICKEL & ATEHORTUA: SUBDIVISION OF ELAPHOGLOSSUM 95
be Pee aly.
7. Blade scales subulate. sect. Polytrichia
7. Blade scales lanceolate, reniform, round-peltate, or stellate, not subulate.
8. Blade scales only at frond margin and along midvein. sect. Decorata
8. Blade scales not restricted to frond margin and midvein sect. Lepidoglossa
215" Elaphoglossum sect. Elaphoglossum
258}° Sect. ee ossa Christ, Monogr. Elaph. 20. 1899.
e long- to short-creeping or suberect; rhizome scales linear to lanceo-
late; shvllosodie distinct or indistinct; blade glabrous or subglabrous, coriaceous
to very thin; hydathodes lacking; blade scales minute, laciniate-pectinate to stel-
late; spores With various forms of crests or low ri
YPE SPECIES: Acrostichum conforme Swartz [= ean conforme
(Swartz) Schott ex J. Smith]
KEY TO THE SUBSECTIONS OF SECT. ELAPHOGLOSSUM
1. Blade coriaceous; veins obscure; phyllopodia conspicuous subsect. Pachyglossa
1. Blade very thin, translucent; veins evident; phyllopodia inconspicuous........... subsect. Tenuifolia
_ 21585 Elaphoglossum subsect. Pachyglossa Christ, Monogr. Elaph. 20. 1899. ued
28% \Subsect. cineeie! Christ, Monogr. Elaph. 20. 1899. CECTOTYPE (chosen here): Acrostichum
latifolium Swartz [=Elaphoglossum latifolium (Swartz) J. Smith]. Christ did not designate a t
species for pie subsection, but chose E. latifolium as the type of his first and most typical division,
atotula.
2597 — Flaccida Christ, Monogr. Elaph. 20. 1899.
Subsect. Herminieriana Christ, Monogr. Elaph. 21. 1899.
eee of the section, with the blade coriaceous and the phyllopodia espe-
cially distinct.
AYPE SPECIES: Acrostichum conforme Swartz [=Elaphoglossum conforme
(Swartz) Schott ex J. Smith]7~ !°97%°
SELECTED SPECIES EXAMINED:
Elaphoglossum acrostichoides (Hook.) Schelpe, E. acutifolium Rosenst., E. affine (Mart. & Gal.)
E. angulatum (Blume) Moore, E. angustatum (Schrad.) Hieron, *E. bicolor Rosenst., E.
callifolium (Blume) Moore, E. chartaceum (Baker) C. Chr., E. crassifolium (Gaud.) Anderson &
Crosby, E. crassinerve (Kunze) Moore, *E. funckii (Fée) Moore, *E. gayanum (Fée) Moore, E.
glabellum J. Smith, E. glaucescens Rosenst., E. glaucum (Fée) Moore, *E. glossophyllum Hieron., E.
guatemalense Klotzsch, E. herminieri (Bory & Fée) Moore, E. hoffmannii (Mett.) Hieron, E.
hymenodiastrum (Fée) Brade, E. inaequalifolium (Jenm.) C. Chr., E. latifolium (Swartz) J. Smith, E.
a (Fée) Moore, *E. lingua (Presl) Brack., E. longifolium (Presl) J. Smith, E. maxonii
x Morton, E. pteropus C. Chr., E. rigidum (Aubl.) Urban, E. schiffneri Christ, E. schom-
sehr (Fes) Moore, E. simplex Nagi J. Smith, E. sporadolepis (Kunze) Moore, E. oo
(Liebm.) Moore, E. tovarense (Moritz ex D. C. Eat.) Moore, E. tuckerheimii Brause, *E. vagans
(Mett.) Hieron., and E. wawrae (Luerss yc Chr.
This is the largest and taxonomically most complex subsection of the
genus. There are relatively few available diagnostic characters, and the varia-
tion within and between species is difficult to interpret. Several groups are
vaguely discernible, but whether they are all closely allied or have evolved to the
coriaceous, glabrous condition independently is still in question. We have re-
frained from distinguishing taxonomic groups until more information is gathered.
One group has very short to suberect, stout (4-12 mm diam.) rhizomes and
spores with many short crests that resemble broad spines (Fig. 2). In some of
56 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
oage ane oe of Elaphoglossum. Sect. Squamipedia. FIG. 7. E. squamipes (Wurdack 692),
; roi eet ow 540), x 1200. Sect. Lepidoglossa. FIG. 9. E. paleaceum (MacBride
1uacsaro ey nct Annie FOF) 1200. FIG. 11. E. bellermannianum
(Camp E 3938B), x 960. FIG. 12. E. auripilum (Mickel 2657), x 1800. ate sae
MICKEL & ATEHORTUA: SUBDIVISION OF ELAPHOGLOSSUM 37
these there are perforations in the crests and in the spore surface as well (Figs. 4
and 5). On the other hand, some species (e.g., E. pteropus) have spores with tall,
slender, solid crests (Fig. /).
Many species have more slender rhizomes (2-4 mm diam.) that have dark,
sclerotic rhizome scales which may become deciduous, as in E. guatemalense and
E. glabellum. These have spores with low, broad ridges (Fig. 3
There is a species complex, including E. affine, E. tenuifolium, E. leptophyllum,
and E. schiffneri, which has ovate-lanceolate, orange to tan rhizome scales with
varying degrees of dark, sclerotic streaks in them. Their rhizomes range from
short-creeping to rather long-creeping. Quite possibly this complex is related to
what looks like a distinct group with very long-creeping, slender, cord-like
rhizomes with black, ovate-lanceolate rhizome scales (species marked with an
asterisk in the above list). Elaphoglossum hoffmannii has the most slender, nearly
naked rhizomes and smallest fronds of the group and may not belong here.
Elaphoglossum subsect. Tenuifolia Mickel & Atehortua, subsect. nov.
Rhizomata gracilia, saepe nuda; phyllopodia inconspicua; lamina angusta
tenuissima; nervi visibiles; sporae dense spiculatae cristatae
TYPE SPECIES: Elaphoglossum acutissimum Christ. - Fibs
SELECTED SPECIES EXAMINED:
Elaphoglossum burchellii (Baker) C. Chr., E. slay dase (Jenm.) Urban, E. praelongum (Fée) C.
Chr., E. sherringii (Baker) C. Chr., and E. wacketii Rose
This group is distinct in its very Gis eshived blades and often naked or even
black and glutinous rhizome. The glabrousness of the blade induces us to place
these species in sect. Elaphoglossum, but they may well belong elsewhere,
perhaps nearer to subsect. Pilosa. At least one member of this group has resinous
dots on the dorsal surface of the blade and spiculate spores (Fig. 6), such as are
found in E. huacsaro (Fig. 10) in subsect. Huacsaro and in some members of
subsect. Pilosa, but the naked rhizome and glabrous blade resemble subsect.
Pachyglossa.
Elaphoglossum gramineum has a glabrescent rhizome and resin-dotted blade,
but the blade is coriaceous, unlike that of other members of this subsection.
Subsection Flaccida was intended by Christ to include the thin-bladed species
we treat as subsect. Tenuifolia, but unfortunately he selected as type E. flac-
cidum, which generally is regarded today as a synonym of E. rigidum, a member
of the coriaceous subsect. Pachyglossa.
ge eam sect. Squamipedia Mickel & Atehortua, sect. nov.”
Rhiz a gracillima longe repentia; rhizomatis stipitisque paleae ovato-
iliceolntes. eaten vel lacerato-pectinatae; phyllopodia rara; lamina parva;
laminae paleae parvae fuscae reductae, saepe hastatae; nervi inconspicul;
hydathodi nulli; sporae plerumque multispiculatae cristis demissis latis anas-
tomosantibus ornatae.
? Although the sectional names Squamipe edia, Setosa, Ber Eximia repeat subsectional names, Latin
— oses are given for both levels since at the subsect
type subsections, and some botanists might not seam he subsectional names as properly described
without separate Latin diagnoses.
58 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
FIGS. 13-18. Spores of Elaphoglossum.
Sect. Setosa. FIG, |
1402. crinipes (Standley 85982), x 1200. FIG. 15. E. scan (Jenman. s.n.), x 1200. Sect. Eximia.
- E. eximium (Mickel 3417), x 1080. FIG. Sect
17. E. beaurepairei (Brade 20937), x 1200.
FIG. 18. E. hirtum (Mickel 2978), x 1200.
E. moritzianum (Fendler 362), x 1440.
FIG.
U, adulata
arty
MICKEL & ATEHORTUA: SUBDIVISION OF ELAPHOGLOSSUM 59
TH
TY PE SPECIES: Acrostichum squamipes Hook. |=Elaphoglossum squamipes
(Hook.) Moore}.
KEY TO THE SUBSECTIONS OF SECTION SQUAMIPEDIA
1. Stipe scales deeply lacerate-pectinate subsect. Ovata
1. Stipe scales entire, ovate- aes lat
2. Sterile blade pinnately or Aabelieesl? divided, rarely undivided but then es: flabellat
bsect. Aciedies
2. Sterile blade entire, linear-elliptic or oblanceolate.
3. Blade margin crenulate; blade thin; spores with narrow crests subsect. Feeana
3. Blade margin entire; blade coriaceous; spores with low folds or oy ot e — icules.
sect. fe area
Elaphoglossum subsect. Squamipedia Mickel & Atehortua, bie no
Rhizomatis stipitisque paleae ovato-lanceolatae peltatae cakes lamina
simplex; sporae spiculatae (Fig. 7).
ATYPE SPECIES: Acrostichum squamipes Hook. |=Elaphoglossum squamipes
(Hook.) Moore}.
OTHER SPECIES EXAMINED:
Elaphoglossum cardiophyllum (Hook.) Moore, E. craspedariiforme (Fée) Brade ex Alston, E.
deltoideum (Sod.) Christ, E. lloense (Hook.) Moore, and E. revolutum (Liebm.) Moore
This is one of the most distinctive groups in the genus, with its small fronds,
broadly ovate rhizome and stipe scales, and lack of phyllopodia. In blade form E.
cardiophyllum seems to belong here, but phyllopodia are present and spore
spicules are lacking. This subsection is extremely closely related to subsect. Pel-
tapteris and agrees in all characters except frond architecture.
5%) [Elaphoglossum subsect. ane oo Mickel & ee stat. nov.
~st}4
Peltapteris Link, Fil. Sp. Hort. Reg. eet Berol. em 147. 1841
Rhipidopteris Schott ex Fée, ag gf ta . 1845, nom. illeg.
Similar to subsect. RE iiecatia: but at Serie blade pinnately or flabellately
divided, rarely undivided but flabellat
TYPE SPECIES: Osmunda ante Swartz [=Elaphoglossum peltatum
(Swartz) Urban].
OTHER SPECIES EXAMINED:
Elaphoglossum ie seer rr este dager - moorei (E. Britt.) Christ, E. peruvianum (Gomez)
Mickel, and E. tripartitum (Hook. & Grev.) M
This group is usually treated as a ect genus (Gomez, 1975; Morton, 1955),
but can be distinguished from species of subsect. Squamipedia only by the frond
dissection.
Elaphoglossum subsect. Ovata Christ, Monogr. Elaph. 23. 1899.
Similar to subsect. Squamipedia, but the scales of the blade, stipe, and rhizome
deeply lacerate-pectinate; phuilapedia distinct; spores with broad folds or ridges
but lacking spicules (Fig. 8).
TYPE AND SOLE SPECIES: Acrostichum ovatum_ Hook. & Grev.
[=Elaphoglossum ovatum (Hook. & Grev.) Moor rs, ~ 12974
nd asing sect. D d sub dia and Muscosa on Christ's divisions
egies sts some Sot tanists would argue that “ cited divisions pa have been described in
Latin to be valid. Since Christ’s descriptions were in German, we here supply Latin po to
avoid any confusion.
60 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
In frond form and slender rhizome this looks strikingly like E. sguamipes, but
the deeply lacerated scales are quite distinct.
oP Elaphoglossum subsect. Feeana Christ, Monogr. Elaph. 22. 1899,
ronds linear-lanceolate or oblanceolate, thin, with crenulate margin; phyl-
lopodia short but distinct; spores with crests and sparse to abundant spicules.
TY PE SPECIES: Acrostichum feei Bory [=Elaphoglossum feei (Bory) Moore].
OTHER SPECIES EXAMINED: q ro88t
Elaphoglossum procurrens (Mett.) Moore and E. wrightii (Mett.) Moore.
These species are close to subsect. Squamipedia in the very slender, long-
creeping rhizome and entire scales, but the thin blade with crenulate margin and
spores with narrow ridges or crests distinguish them as a separate group.
Rg sect. Decorata Mickel & Atehortua, sect. noy.?
hizomatis erecti crassi paleae lineares; phyllopodia brevia; stipitis paleae
magnae patulae; laminae magnae ellipticae praeter costam Mmarginesque paleis
imbricatis aureis vestitas glaberrimae; nervi conspicui hinc inde anastomosantes;
hydathodi nulli; sporae breviter cristatae.
TYPE AND SOLE SPECIES: Acrostichum decoratum Kunze | =Elaphoglos-
sum decoratum (Kunze) Moore]. Lays
This species is remarkably distinct and without Close relatives.
\
ee Elaphoglossum sect. Lepidoglossa Christ, Monogr. Elaph. 21. 1899,
ger Sect. Gymnoglossa ea ey sash Elaph. 22. 1899. LECTOTYPE (chosen here): Osmunda bifur-
cata Jacq. [=Elaphoglossum ifurcatum (Jacq.) Mickel]. Christ did not designate a type for sect.
Gymnoglossa, but he did choose E. furcatum ( syn. E. bifurcatum) as type of his subsect. Dimorpha,
which is the first named and most typical subsection of sect. Gymnoglossa.
7 ad . s
toothed, ciliolate or rarely entire, or round-peltate, or modified to stellate hairs;
veins evident to obscure, free; hydathodes lacking; spores with low ridges, rarely
ridges.
4148 ee paleaceum (Hook. & Grev.) Sledge] (syn. Acrostichum sSquamosum
KEY TO THE SUBSECTIONS OF SECTION LEPIDOGLOSSA
1. Blade densely scaly; scales lanceolate, toothed, not stellate or reduced to resinous dots.
2. Blade Ovate-lanceolate, coriaceous, often obtuse at apex; blade scales erose; spores verruculate.
2. Blade linear-lanceolate or narrowly elliptic, acuminate a
Spores with low, non-verruculate Leip. ek noe: subsect. Polylepidea
modified to stellate hairs and/or resinous dots
Itate.
ha oeatheslecdeveseesaty rts oi subsect. Microlepidea
MICKEL & ATEHORTUA: SUBDIVISION OF ELAPHOGLOSSUM 61
4. Blade glabrous, elliptic with a caudate apex. subsect. Petioloa
4. Blade scaly to subglabrous, linear to lanceolate, acuminate to obtuse at apex
5. Rhizome short-creeping; blade acute to acuminate, rarely crenulate to pinnate divided.
sect. Pilosa
5. Rhizome long, ascending; blade obtuse. See Huacsaro
5k? Elaphoglossum subsect. Polylepidea Christ, Monogr. Elaph. 21. 1899.
Rhizo short-creeping to ascending; rhizome scales dark, ciliolate; blade
scales a ciliolate; blade narrowly elliptic, densely ay spores with low
ridges, usually with small verrucae between the ridges (Fig. 9).
“LECTOTYPE SPECIES (chosen here): Acrostichum paleaceum, Hook. &
Grev. [=Elaphoglossum paleaceum (Hook. & Grev.) Sledge]. — #!9
SELECTED SPECIES EXAMINED:
Elaphoglossum acuminans C. Chr. ex Urban, E. auricomum Seapey Moore, E. casanense
Rosenst., E. deckenii (Kuhn) C. Chr., E. dombeyanum Moore & Houlst., E. eatonianum (E. Britt.) C.
eggersii (Baker) Christ, E. fuertesii Brause, E. vain ype E. kuhnii Hieron., E.
laminarioides (Bory) Moore, E. langsdorfii (Hook. & Grev.) Moore, E. meridense ies Moore,
E. orbignyanum (Fée) Moore, E. plumosum (Fée) Moore, E. eens (Kuhn) Christ, E. rufescens
(Liebm.) Moore, E. vestitum (Schlecht. & Cham.) Schott ex J. Smith, and E. wagneri (Kunze) Moore
This is one of the two largest, most complex, and unmanageable groups in the
genus. The toothed or ciliolate scales on the blade and usually also on the rhizome
are the principal distinguishing features.
2589" Elaphoglossum subsect. Microlepidea Christ, Monogr. Elaph. 22. 1899.
Rhizome scales Bneay auceoine. entire; vac’ scales stellate below, round and
peltate above; spores with low, oth r idge
“LECTOTYPE SPECIES Taina here): yee tectum Humb. & Bonpl.
7445 ex Willd. [=Elaphoglossum tectum (Humb. & Bonpl. ex Willd.) Moore]. Christ
selected this species as type of his div. Tecta, which is the first-named and most
typical of the divisions under subsect. Microlepidea.
OTHER SPECIES EXAMINED: Elaphoglossum furfuraceum (Mett.) Christ.
This subsection is very close to subsect. Pilosa, but the round, peltate scales are
not found in any other group.
258aC- Teper subsect. Pilosa Christ, Monogr. Elaph. 23. 1899.
staphyla Presl, Epim. Bot. 160. 1851
LSev -Elapodlssum subsect. Dimorpha Christ, Monogr. Elaph. 22.
me short-creeping; rhizome scales usually on. entire to pinnately di-
vided; bhade salen resembling stellate hairs or lanceolate-toothed; blade often
with resinous dots; spores with low ridges or narrow crests.
-LECTOTYPE SPECIES (chosen here): Acrostichum pilosum Humb. & Bonpl
ex Willd. [=Elaphoglossum pilosum (Humb. & Bonpl. ex Willd.) Moore].
Elaphoglossum pilosum is the type species of Christ’s div. Grata, which was the
first and most typical division of subsect. Pilosa.
SELECTED SPECIES EXAMINED:
Elaphoglossum bifurcatum (Jacq.) Mickel, E. dimorphum (Hook. & Grev.) Moore, E. gratum (Fée)
Moore, E. lagesianum Rosent., E. lepidotum J. Smith, E. mathewsii (Fée) Moore, E. nervosum (Bory)
Christ, E. petiolatum (Swartz) Urban, E. rosenstockii Christ, E. salicifolium (Willd. ex Kaulf.) Alston,
and E. viscosum (Swartz) J. Smith.
"920 |
62 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
The degree of scaliness and variation from scales to resinous dots within one
species is not well understood. This subsection is very closely allied to subsects.
Huacsaro and Microlepidea, and the demarcation between them is not clear.
Elaphoglossum dimorphum displays an intermediate frond morphology be-
tween simple fronds, such as those of E. nervosum, and the pinnately dissected
fronds of E. bifurcatum. All three species occur on St. Helena, and Mickel (1980a)
has shown that they differ essentially only in dissection. Whether these are in fact
three distinct species or all forms of one species is still a question.
aoe Elaphoglossum subsect. Petiolosa Christ, Monogr. Elaph. 23. 1899.
Rhizome short-creeping; rhizome scales dark, linear, bristle-like; blade elliptic
with caudate tip; fertile blade folded; blade scales lanceolate or lacking, occasion-
ally with resinous dots; spores with low ridges. ¥e iver
VTYPE SPECIES: Acrostichum petiolosum Desv. [=Elaphoglossum petiolosum
(Desv.) Moore}.
OTHER SPECIES EXAMINED: Elaphoglossum trianae Christ.
The relationships of this subsection are not clear. The resinous dots are re-
miniscent of those of subsects. Pilosa and Huacsaro, the bristle-like rhizome
scales are similar to those of some members of subsect. Hybrida, and the condup-
licate fertile blades are similar to those found in some members of subsect. Setosa.
The blade shape and spore details are unique.
9G)
ae Elaphoglossum subsect. Huacsaro Mickel & Atehortua, subsect. nov.
zomata adscendentia longa; rhizomatis paleae fuscae integrae; lamina ellip-
tica, apice obtusa; laminae saepe resinoso-punctatae; paleae lanceolatae den-
ticulatae vel pilis stellatis conspersae; sporae dense spiculatae late cristatae.
“ae PE SPECIES: Acrostichum huacsaro Ruiz [=Elaphoglossum huacsaro c
(Ruiz) Christ]. e 7"
Pica oe SPECIES EXAMINED:
aphoglossum alfredii ili
dence Ce ten Roses and acne eke en ml Mose
This group probably is closely related to subsect. Pilosa, as shown by the scales
and resinous dots, but the long ascending rhizome, obtuse blade apex, and the
highly spiculate spores are distinctive (Fig. 0).
ase Elaphoglossum subsect. Muscosa Mickel & Atehortua, subsect. noy.?
paar ar a Magee repentia; phyllopodia nulla; stipites laminaeque dense
ovate lance ane Is paleae latae patulae; lamina anguste elliptica ad lanceolata vel
a uy COrlacea, apice plerumque obtusa; paleae saepe cum sporangiis
atae. a1
“TYPE SPECIES: Acrosti
: chum muscosum Swa
(Swartz) Moore]. as
SELECTED SPECIES EXAMINED:
Elaphoglossum aschersonii Hieron., E. bellermannianu
e. corderoanum (Sod.) Christ, E. decipiens Hieron., E.
Christ, and E. plicatum (Cav.) C. Chr.
Although the scales of E. engelii approach those of the subsect. Polylepidea, the
spores are unique in the genus in having verruculae covering the surface (Fig. 1/).
bi he
=Elaphoglossum muscosum
m (Klotzsch) Moore, E. blandum Rosenst.,
engelii (Karst.) Christ, E. leamannianum
|
si
25345,
MICKEL & ATEHORTUA: SUBDIVISION OF ELAPHOGLOSSUM 63
Elaphoglossum sect. Polytrichia Christ, Monogr. Elaph. 22. 1899.
Rhizome short- -creeping to erect; rhizome scales linear to linear-lanceolate;
phyllopodia inconspicuous or lacking: stipe and blade, especially blade midvein
and margin, with subulate scales and also bearing minute glandular hairs; hy-
dathedee lacking; spores with low ridges and often small verrucae between them.
“LECTOTYPE SPECIES (chosen here): Acrostichum crinitum L. [=Elapho- >
glossum crinitum (L.) Christ]. This was the type of Christ’s first-named and most
typical subsection, Hymenodium.
KEY TO THE SUBSECTIONS OF SECTION POLYTRICHIA
1. Blade broadly elliptic and fleshy; veins netted, obscur subsect. Hy see ee
1. Blade pee et narrowly elliptic, or ovate- ete subcoriaceous or papyraceous; vein
free, usually eviden
2. Stipe short to si blade scales dark brown to black, located mostly on the margin and midvein.
subsect. Hybrida
2. Stipe nearly lacking; blade scales orange, distributed subuniformly over the blade surfaces.
subsect. Apoda
Elaphoglossum subsect. Hymenodium — Christ, Monogr. Elaph. 23. 1899.
Hymenodium Fée, Mém. Fam. Foug. 2: 20. 1845.
eae ne fleshy, oe ‘slip veins obscure, netted; spores with
owr
TYPE eae SOLE SPECIES: Acrostichum crinitum L. [=Elaphoglossum
crinitum (L.) Christ].
Although this is occasionally distinguished as a separate genus, it agrees
very Closely with subsect. Hybrida.
25246 Elaphoglossum subsect. Hybrida Christ, Monogr. Elaph. 23. 1899.
izome short-creeping or ascending; stipe long; blade usually 1 stool
ous; blade scales especially on the margin and midvein, black or da rk bro
spores with low ridges (Fig. /2). — 1S ey
“TY PE SPECIES: Acrostichum hybridum Bory | =Elaphoglossum hybridum
(Bory) Moore].
SELECTED SPECIES EXAMINED:
Elaphoglossum albomarginatum A. Reid Smith, E. auripilum Christ, E. cordifolium Rosenst.,
E. denudatum (Jenm.) Maxon ex Morton, E. erinaceum (Fée) Moore, E. lindbergii (Mett.)
Rosenst., E. melanopus (Kunze) Moore, E. prestonii J. Smith, E. scolopendrifolium (Raddi) J.
Smith, E. spannagelii Rosenst., and E. tambillense (Hook.) Moore.
This subsection is very complex, and the species limits are not well under-
stood. The variation in rhizome scales is especially perplexing. Most species
have linear, orange rhizome scales, and others have bristle-like, maroon
scales, but the differences are not always clear-cut. At least two species have
a glabrous or subglabrous blade.
This subsection is composed mostly of Christ’s ‘‘divisio Melanolepidea™ of
ordo Stenoneura, since they lack hydathodes. Christ referred subsect. Hy-
brida to ordo Condyloneura, although E. hybridum, the type species, lacks
hydathodes and belongs to div. Melanolepidea. All other species Christ in-
cluded in subsect. Hybrida have hydathodes and make up our subsect.
Eximia.
_ 2244
64 AMERICAN FERN JOUNRAL: VOLUME 70 (1980)
nid
o> Elaphoglossum subsect. Apoda Mickel & Atehortua, subsect. nov.
Stipites fere nulli; laminae paleae aurantiacae subulatae, per laminae ssa
ficiem regulariter conspersae; sporae breviter cristatae. ras
AY PE SPECIES: Acrostichum apodum Kaulf. [=Elaphoglossum apodum
(Kaulf.) Schott ex J. Smith].
OTHER SPECIES EXAMINED:
Elaphoglossum cubense (Mett. ex Kuhn) C. Chr. and E. siliquoides (Jenm.) C. Chr.
Members of this subsection closely resemble those of subsect. Setosa in
their orange to brown subulate blade scales, but seem to belong to sect.
Polytrichia on the basis of no hydathodes and spores with low ridges and
.. perforated crests. They also differ from sect. Setosa in their very short stipes.
iW Elaphoglossum sect. Setosa (Christ) Mickel & Atehortua, stat. nov.”
1b">) Elaphoglossum subsect. Setosa Christ, Monogr. Elaph. 23. 1899.
izome short- to long-creeping or erect; rhizome scales linear; phyllopodia
lacking; stipes with minute, erect, glandular hairs; plants mostly small; veins
evident, spaced well apart, ending well short of the margin in distinct
hydathodes; scales subulate, orange to brown; spores with many low crests
and usually with a perforate surface, or not crested and the surface echinate-»
reticulate. 25
TYPE SPECIES: Acrostichum villosum Swartz | =Elaphoglossum villosum
(Swartz) J. Smith].
KEY TO THE SUBSECTIONS OF SECTION SETOSA
1. Rhizome long-creeping; rhizome scales dark brown to black subsect. Alpestria
1. Rhizome ascending to erect: rhizome scales pale to dark brown.
2. Blade margin usually crenulate.
2. Blade margin entire.
3. Plants very small (2-15 cm tall): blade spatulate, rarely sublinear; hydathodes inconspicuous.
subsect. Pilosella
); blade narrowly elliptic to linear-lanceolate: veins
Pisin tees seine tev akan andenstescaey wik iG ur) itso subsect. Setosa
2¢\ 4) Elaphoglossum subsect. Setosa Christ, Monogr. Elaph. 23. 1899.
hizome ascending to erect; blade narrowly elliptic to linear-lanceolate; veins
and hydathodes evident: Spores with low crests (Fig. 13) or spines (Fig. /4).
“TYPE SPECIES: Acrostichum villosum Swartz [=Elaphoglossum villosum
(Swartz) J. Smith]. L_ 3sst\
SELECTED SPECIES EXAMINED:
Elaphoglossum costaricense Christ, E. lindenii (Bory ex Fée) Moore, E. moritzianum (Klotzsch)
oore, E. ocoense C. Chr., E. omphalodes (Fée) Brade, E. Palorense Rosenst., and E. setosum
Moore.
ree thicss tiseb eles see Oro subsect. Plumierana
3. Plants small to medium-sized (540 cm tall
and hydathodes evident.
Some other species, such as E. crinipes C. Chr., E. oblanceolatum C. CAG, Ee.
papillosum (Baker) Christ, and E. setigerum (Sod.) Diels, are included here be-
cause they look like species of subsect. Setosa in their external morphological
characters, but they differ significantly in Spore architecture. Rather than having
crests, their spores are densely covered with Short spines whose bases branch to
form a reticulum (Fig. 14). Elaphoglossum fluminense Brade may belong here
also. It looks like a slender member of subsect. Pilosella, but has perforate spores
with very low crests like those of most members of subsect. Setosa.
ast
2890
MICKEL & ATEHORTUA: SUBDIVISION OF ELAPHOGLOSSUM 65
0
Elaphoglossum subsect. Pilosella Christ, Monogr. Elaph. 23. 1899.
hizome erect; scales subulate; hydathodes inconspicuous; plants especially
small (2-15 cm tall); spores non-perforate with the ridges low and broad, usual
lacking spicules. ~
“TYPE SPECIES: Acrostichum piloselloides Pres| |=Elaphoglossum pilosel-
loides (Presl) Moore].
OTHER SPECIES EXAMINED:
Elaphoglossum hayesii (Mett.) Maxon, E. horridulum J. Smith, E. jamesonii (Hook. & Grev.)
Moore, E. pusillum (Mett.) C. Chr., and E. spatulatum (Bory) Moore.
Elaphoglossum horridulum and E. jamesonii look like they belong here, but are
different in their spores having spicules. This subsection is close to subsect.
Setosa, although its spores have ridges rather than crests and hardly any are
perforate, while the hydathodes are less conspicuous than in subsect. Setosa.
Elaphoglossum subsect. Alpestria Mickel & Atehortua, subsect. nov.
Rhizomata longe repentia; rhizomatis paleae fuscae; phyllopodia inconspicua,
nervi raro furcati; hydathodi conspicui; stipitis laminaeque paleae subulatae, raro
lanceolatae; paleae inter sporangia nullae; sporae perforatae spiculatae breviter
cristatae. 3%
L-TYPE SPECIES: Acrostichum alpestre Gardn. [=Elaphoglossum alpestre
(Gardn.) Moore}.
OTHER SPECIES EXAMINED:
Elaphoglossum barbae Rosenst., E. chiapense A. Reid Smith, E. hirtipes (Fée) Brade, and E.
leptophlebium (Baker) C. Chr.
This subsection is close to subsect. Setosa in the subulate scales and perforate
spores, but differs in the rhizome scales and habit. Elaphoglossum yatesii (Sod.)
Christ seems to fit here, except that its blade is densely clothed with lanceolate
scales.
759°” Elaphoglossum subsect. Plumierana Mickel & Atehortua, subsect. nov
<5
rn.
e qo i
hizomata breviter repentia usque erecta; lamina margine crenulata; laminae
paleae subulatae aurantiacae; sporae cristatae (Fig. 15).
ATY PE SPECIES: Elaphoglossum plumieri Moore.~ !
OTHER SPECIES EXAMINED:
Elaphoglossum buchii C. Chr., E. lanceum Mickel, and E. smithii (Baker) Christ.
In their thin, crenulate blades these species resemble somewhat the species of
subsect. Feeana, but are distinct in rhizome habit, rhizome and blade scales,
phyllopodia, and hydathodes. There is considerable spore variation within the few
species of this group. Elaphoglossum lanceum has highly perforate, lace-like
crests, whereas the other species have solid, slender, nonperforate crests.
a37F
Elaphoglossum sect. Eximia Mickel & Atehortua, sect. nov.” site
Rhizomata breviter repentia vel adscendentia; phyllopodia nulla; stipitis paleae
lanceolatae minimae vel saepe subulatae; nervi distantes, angulo 40—60° abe untes;
hydathodi conspicui; laminae paleae sparsae minimae, non subulatae; sporae
reticulato-echinatae ecristatae. a5
YPE SPECIES: Acrostichum eximium Mett. [=Elaphoglossum eximium
(Mett.) Christ].
Cy
ly
P20
0
97°F
66 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
KEY TO THE SUBSECTIONS OF SECTION EXIMIA
1. Blade entire, linear to linear-elliptic; stipe scales subulate. subsect. Eximia
1. Blade pedately divided; stipe and rhizome scales small, lanceolate. .......... subsect. Cardenasiana
is Elaphoglossum subsect. Eximia Mickel & Atehortua, subsect. nov.
Lamina integra, linearis vel lineari-elliptica; stipitis paleae subulatae; laminae
paleae sparsae minusculae. !
YPE SPECIES: Acrostichum eximium Mett. [=Elaphoglossum eximium
(Mett.) Christ].
OTHER SPECIES EXAMINED:
Elaphoglossum aubertii (Desv.) Moore, E. beaurepairii (Fée) Brade, E. brachyneuron (Fée) J.
Smith, E. gracile (Fée) Christ, E. lineare (Fée) Moore, and E. stenopteris (Klotzsch) Moore.
In the subulate scales, hydathodes, and reticulate-echinate spores (Figs. 16 and
17), this group is similar to some members of subsect. Setosa and to some extent
to sect. Undulata.
scat Elaphoglossum subsect. Cardenasiana Mickel & Atehortua, subsect. nov.
A subsect. Eximia paleis stipitis rhizomatisque minoribus, rhizomate magis
carnoso, laminaque pedatim divisa diversa. 4G
/TYPE AND SOLE SPECIES: Elaphoglossum cardenasii Wagner.
This species is unique in the genus in its pedately divided fronds, but in other
characters shows close relationship to subsect. Eximia.
ot Elaphoglossum sect. Undulata Christ, Monogr. Elaph. 24, 1899.
hizome short-creeping to erect; phyllopodia lacking; blade ovate-lanceolate;
blade scales subulate to deltate-lanceolate, erose or toothed; hydathodes con-
Spicuous; spores without ridges, openly reticulate-echinate, the spine bases di-
verging and forming a reticulum occasionally with irregular verrucae or perforated
crests (Fig. 18).
TYPE SPECIES: Acrostichum hirtum Swartz [=Elaphoglossum_ hirtum
(Swartz) C. Chr.] (syn. E. undulatum (Willd.) Moore).
OTHER SPECIES EXAMINED:
Elaphoglossum bakeri (Sod.) Christ, E. boryanum (Fée) Moore, and E. proliferans Maxon & Mor-
ton ex Morton.
This subsection is closely related to subsect. Setosa and possibly to subsect.
Eximia, with which it has subulate scales and echinate spores in common.
Elaphoglossum castaneum (Baker) Diels is similar to E. hirtum in size and
shape and In spores with very small, crest-like projections perforate at the base
(Fig. 18). This could represent a condition intermediate between sects. Undulata
and Setosa, but the rhizome is longer-creeping, the rhizome scales are small,
_ Sclerotic, and resinous, and blade scales are lacking.
oa tO
rs Bes secs te Amygdalifolia (Christ) Mickel & Atehortua, stat. nov.
AGA a saree mas oe obs Monogr. Elaph. 22. 1899. :
‘a eate cae, 2 A ia dy ender; phyllopodia short; rhizome scales round
Ee : €s linear-lanceolate; veins evident: hydathodes conspicu-
ous; blade subglabrous with irs: Bie meee tt
: g ith minute stellate hairs; spores with narrow crests with
1\F%
TYPE AND SOLE SPECIES: Acrostichum amy gdalifolium Mett. [=Elapho-
glossum amygdalifolium (Mett.) C hrist]. — “\4y
This species is quite distinct and has no close relatives.
MICKEL & ATEHORTUA: SUBDIVISION OF ELAPHOGLOSSUM 67
ACKNOWLEDGMENTS
This study was supported by National Science Foundation grant DEB 77-25582
to the senior author. We gratefully acknowledge the kind help of Dr. Rupert
Barneby for his assistance with the latin and his criticism of the manuscript, Mr.
Joel Huang for his technical work with the scanning electron microscope and for
preparing the photographs, and Drs. Arthur Cronquist and David Lellinger for
nomenclatural advice.
LITERATURE CITED
ALSTON, A. H. G. 1956a. The Brazilian species of Elaphoglossum. Bol. Soc. Brot. 32:1-32.
_______, 1956b. The subdivision of the Polypodiaceae. Taxon 5:23-25.
CHRIST, H. 1899. Monographie des Genus Elaphoglossum. Neue Denkschr. Allg. Schweiz. Naturf.
. Gesammten Naturwiss. 36:1-159, t.1-4.
CHRISTENSEN, C. 1938. Filicinae. Jn F. Verdoorn. Manual of Pteridology. Nijhoff, The Hague.
CRABBE, J. A., A. C. JERMY, and J. T. MICKEL. 1975. A new generic sequence for the
pteridophyte herbarium. Fern Gaz. 11:141-162.
DIELS, L. 1899. Polypodiaceae. In A. Engler & K. Prantl. Die Natiirlichen Pflanzenfamilien
1(4): 139-339.
ERDTMAN, G. 1957. Pollen and Spore Morphology/Plant Taxonomy. Almgvist & Wiksells, Uppsala,
Sweden.
FEE, A. L. A. 1845. Histoire des Acrostichées. (Mem. Fam. Foug. 2.) Veuve Berger-Levrault,
Strasbourg.
—_—___—.. 1852. Genera Filicum. (Mém. Fam. Foug. 5.) Veuve Berger-Levrault, Paris & Strasbourg.
GOMEZ, L. D. 1975. Contribuciones a la pteridologia costarricense. VI. El género Peltapteris Link en
osta Rica. Brenesia 6:25-31.
HENNIPMAN, E. 1977. A monograph of the fern genus Bolbitis (Lomariopsidaceae). Leiden Bot.
Ser. 2:1-331.
HOLTTUM, R. E. 1947. A revised classification of leptosporangiate ferns. J. Linn. Soc., Bot.
$1:123-158
————. 1978. Elaphoglossum. Flora Malesiana II, 1(4):289-314.
LIEW, F. S. 1977. Scanning electron microscopical studies on the spores of pteridophytes. XI. The
family Oleandraceae (Oleandra, Nephrolepis, and Arthropteris). Gard. Bull. Singapore
30:101-110, ¢.J-VI.
LINK, H. F. 1841. Filicum Species in Horto Regio Botanico Berolinensis Cultae. Veit, Berlin.
LINNAEUS, C. 1753. Species Plantarum, vol. 2. Salvi, Stockholm.
LLOYD, R. M. 1970. A survey of some morphological features of the genus Elaphoglossum in Costa
Rica. Amer. Fern J. 60:73-83.
MICKEL, J. T. 1980a. Relationships of the dissected elaphoglossoid ferns. Brittonia 32:109-1 17.
—_— . 1980b. Elaphoglossum. /n R. G. Stolze. Ferns and Fern Allies of Guatemala. Fieldiana, Bot.
39: in :
MOORE, T. 1857-1862. Index Filicum. Pamplin, London.
MORTON, C. V. 1955. Notes on Elaphoglossum, III. The publication of Elaphoglossum and
Rhipidopteris. Amer. Fern. J. 45:11-14.
PICHI SERMOLLI, R. E. G. 1968. Adumbratio Florae Aethiopicae. 15. Elaphoglossaceae. Webbia
PRESL, C. B. 1851. Epimiliae Botanicae. Abh. Konigl. Bohm. Ges. Wiss. V, 6:361-624.
SCHELPE, E. A. C. L. E. 1969. Reviews of tropical African Pteridophyta, 1. Contr. Bolus Herb.
1:1-132.
SCHOTT, H. 1834. Genera Filicum. Wallishausser, Vienna.
SLEDGE, W. A. 1967. The genus Elaphoglossum in the Indian peninsula and Ceylon. Bull. Brit. Mus.
(Nat. Hist.), Bot. 4:79-96.
68 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
SMITH, J. 1841. An arrangement and definition of the genera of ferns, with observations on the
affinities of each genus. J. Bot. (Hook.) 4:38-70, 147-198.
SODIRO, A. 1897. Cryptogamae Vasculares Quitenses. Typis Universitatis, Quito.
SPORNE, K. R. 1975. The Morphology of Pteridophytes, ed. 4. Hutchinson, London.
WAGNER, W. H., Jr. 1954. A Bolivian Elaphoglossum of unique leaf structure. Bull. Torrey Bot.
Club 81:61-67.
REVIEW
“HOW TO KNOW THE FERNS AND FERN ALLIES,” by John T. Mickel,
1979. Wm. C. Brown Company Publishers, Dubuque, Iowa. $7.95 hardcover,
$5.95 wire coil bound.—In the standard format of the Pictured Key Nature Series,
this book provides the information needed to identify North American ferns and
fern allies. Introductory chapters on structure, life history, hybridization, cultiva-
tion, collection, and nomenclature provide a basic understanding of ferns and fern
allies and the terminology needed to identify these plants successfully. This also
makes the work useful as a textbook or handbook for amateur pteridologists.
The greatest part of the volume consists of bracketed keys which lead the user
to the appropriate genus and species. Diagnostic characters for each species,
provided in the annotated keys, are supplemented by a brief description, habitat
preference, frequency of occurrence, and distribution map, as well as an illustra-
tion for nearly every species. Limited synonymy is also included. Hybrids and
infraspecific taxa are mentioned with related species. Problems in taxonomy are
explained so that the basis for confusion can be understood. The genera are listed
alphabetically, enabling the experienced pteridologist to turn to the appropriate
genus and begin keying at that point. The uninitiated can begin with the generic
key that is found near the beginning of the book. Edgar Paulton’s line drawings are
quite good and in general capture the distinguishing characteristics of each
species. The book concludes with a listing of state and regional identification
pteridologists alike have long awaited.—W. C
: : -—W. Carl Taylor, D ;
Milwaukee Public Museum, Milwaukee, WI 53233. ne lado ls
AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 2 (1980) 69
Notes on the Natural History of Stylites gemmifera
ERIC E. KARRFALT and DALE M. HUNTER*
Several unexpected observations were made recently while collecting plants of
Stylites E. Amstutz for anatomical study. The plants were collected near the end
of the growing season (18-19 April 1979) so as to obtain young plants which had
just completed various numbers of growing seasons. In the type locality, Srylites
is described as forming pure colonies which stand just above the water level in the
lacustrine bog in which they are found (Rauh & Falk, 1959); but the plants we
collected at 4100 m altitude (Karrfalt & Hunter 22, NY) near Lago Junin, 14 km
north of Junin, Peru, were invariably growing in association with various flower-
ing plants (Figs. 1 and 2) and were frequently submerged. The colonies at the
Junin locality were generally in the form of radially symmetrical, dome-shaped
hummocks (Fig. 1), but various other rounded shapes occurred as well. The
hummocks ranged in diameter from 20 to about 200 cm. The larger hummocks
generally were found to contain a larger proportion of other plants in addition to
Stylites than did the smaller ones. The plants in the hummocks were extremely
densely packed and usually stood above the water level, but some hummocks
were partially or completely submerged. The submerged portions of these hum-
mocks were populated nearly exclusively by Stylites, but the emergent portions
included other plants as well (Fig. 5). These plants usually were rather small (with
stems a few millimeters in length), but some were quite good-sized (stems 2-4 cm
long) and bore about 40 leaves up to 7 cm long, as well as numerous gemmae.
Their leaves did not have the typical flattened form with deflexed tips, but rather
were subtriangular to terete in cross section and ascending. All intermediate forms
between these atypical leaves and those described by Rauh and Falk (1959) were
also seen; the variation in leaf morphology will be described in detail in a sub-
sequent report. The plants with the atypical leaf form always were submerged and
not densely crowded. On the other hand, plants bearing typical leaves occurred
both above and below the water level in the bog, but these plants always were
densely crowded. Leaf form correlates with population density rather than with
emergence or submergence. The nature of this correlation is not certain, but
experiments in progress suggest that it is largely or entirely environmental.
In contrast to its very limited geographical range, Stylites is extremely vigorous
and abundant where appropriate conditions for its growth exist. The J unin locality
is a bog which has been used as a pasture for many years. It is heavily grazed by
sheep and llamas, as indicated by the cropped herbage (Figs. 1 and 4) and abun-
dant llama dung. The Stylites plants, however, very rarely show any evidence of
even accidental damage by the animals. The Junin locality occupies at least sev-
eral acres; we were unable to determine its full extent due to our anoxemia and
consequent lack of energy.
The leaves of Stylites are coated with considerable quantities of mucilage, as are
the basal parts of the leaves of all /soétes species of which we have seen living
*Department of Biology, Brooklyn College, Brooklyn, NY 11210.
70 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
material. Also as in /soétes, the leaves are replaced annually (Rauh & Falk, 1959).
As the new leaves grow and expand within the hummock, the dead, mucilage-
coated leaves of the largest plants are extruded en masse onto the surface of the
fie a at i FIG. 1. A typical hummock. FIG. 2. Close up of part of the hummock
Gouden e ig ? os Submerged plants with atypical leaves. FIG. 4. A hummock and surrounding
g . . 5. A partially submerged hummock. Unlabelled arrows = Stylites plants; E = pat-
rice ea (Fig. 2, E). Once out on the surface of the hummocks, the individual
ea sporophylls become separated from one another (Figs. 1, 5, S). The extru-
sion of the old sporophylls would seem to be advantageous for spore dispersal.
Indeed, an analogous process has been shown to be involved in spore dispersal in
KARRFALT & HUNTER: NATURAL HISTORY OF STYLITES 71
Isoétes drummondii (Osborn, 1922). However, examination of large numbers of
extruded sporophylls of Stylites never revealed any discernible evidence of the
establishment of gametophytes by the spores carried with the extruded
sporophylls. Field observations of gametophytes were necessarily limited to those
which could be made with a hand lens; that is, only megaspores were examined
and these only for the opening of the trilete scar. Any megagametophytes which
were contained within unopened spore walls were not distinguished from un-
germinated spores.
Rauh and Falk (1959) found very few megagametophytes and no mi-
crogametophytes of Stylites. In our material, gametophytes were likewise very
infrequently encountered. Only megagametophytes were identified in the field and
these were found only in association with adult sporophytes which showed some
evidence of recent damage or injury, such as relatively few leaves or a reduced
stem diameter near the apex. Six gametophytes were found, all of which bore
sporophytes with one or two leaves and roots. The occurrence of the rare
megametophytes only in association with the rare, injured sporophytes suggests
that the absence of gametophytes from other locations is the result of unequal
competition between the gametophytes and the much larger, densely crowded
adult sporophytes and gemmae.
It was not possible to determine the specific source of the spores which gave
rise to the gametophytes we collected. Our gametophytes were probably derived
from the massive quantities of spores produced by the immediately adjacent
sporophytes, but the possibility cannot be excluded that the successful spores
may be have been transported with old sporophylls which had been extruded onto
the surface of the same or some other hummock.
Many of the plants which we collected bore abundant gemmae and therefore
must be assigned to S. gemmifera W. Rauh, inasmuch as S. andicola E. Amstutz
has no vegetative reproduction. The other criteria by which Rauh and Falk (1959)
distinguished the sporophytes of S. andicola from those of S. gemmifera are
merely quantitative and are of questionable value. For example, the leaves of S.
andicola are said to be 5—7 cm long, whereas those of S. gemmifera are said to be
3.5-5 cm long, but as noted above some gemma-bearing plants in our collection
had leaves as long as 7 cm. Although these longest leaves did not have the mor-
phology typical of Stylites, the leaves of plants collected from hummocks invari-
ably had the typical form, and some of these were as long as 5.5 cm. According to
Rauh and Falk, the stem of S. andicola is mostly unbranched and up to 20 cm
long, whereas that of S. gemmifera is frequently branched and not more than 8 cm
long; obviously these characters would be of no use in identifying an unbranched
plant whose stem was not more than 8 cm long. Also, S. andicola is supposed to
form hummocks in which all individuals are the same age, whereas colonies of >.
gemmifera contain both old and young plants. Although they did not explicitly
state their method, Rauh and Falk seem to have used size as an indicator of
relative age. In any case, the hummocks which contain unbranched plants of a
uniform large size and no gemmae (i.e., hummocks of *’S. andicola’’) may simply
72 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
be relatively old colonies in which the intense competition for space has been
resolved in favor of the largest and most vigorous individuals which neither
branch nor produce gemmae. The only qualitative distinction between the
sporophytes of the two species of Stylites is the presence of gemmae in S. gem-
mifera and their absence in S. andicola. However, the number of gemmae on a
plant is highly variable. In our material, from one to eight were seen, and many
plants had no gemmae at all. According to the criteria given by Rauh and Falk,
sporophytic specimens without gemmae and with stems shorter than 8 cm may be
distinguished as to species only by the length of their leaves. Unfortunately, as
noted above, we have gemma-bearing plants, obviously assignable to S. gemmi-
fera, which have leaves longer than 5 cm. Thus it appears to us that the distinct-
ness of the two species of Stylites is in sufficient doubt that a critical reexamina-
tion of these two taxa is in order. Moreover, inasmuch as the separation of Stylites
from /soétes already has been questioned (Kubitzki & Borchert, 1964; Bierhorst,
1971), this reexamination also should review the generic assignment of these
species.
LITERATURE CITED
BIERHORST, D. W. 1971. Morphology of Vascular Plants. Macmillan, New York.
KUBITZKI, K. and R. BORCHERT. 1964. Morphologische Studien an Isoétes triquetra A. Braun und
Bemerkungen uber das Verhaltnis der Gattung Stylites E. Amstutz zur Gattung Isoétes L.
Ber. Deutsch. Bot. Ges. 77:227-233.
OSBORN, T. G. B. 1922. Some observations on Isoetes Drummondii, A. Br. Ann. Bot. 36:41-54.
RAUH, W. and H. FALK. 1959. Stylites E. Amstutz, eine neue Isoetacee aus den Hochanden Perus.
I. Teil: Morphologie, Anatomie, und Entwicklungsgeschichte der Vegetationsorgane. Sitz-
ungber. Heidelberger Akad. Wiss. 1959:1-83.
AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 2 (1980) 73
Reciprocal Allelopathy Between the Gametophytes
of Osmunda cinnamomea and Dryopteris intermedia
RAYMOND L. PETERSEN* and DAVID E. FAIRBROTHERS**
Allelopathy is the chemical inhibition of growth and/or development of one
organism by another. The literature on allelopathy is very extensive and has been
summarized in several reviews (Muller, 1970; Pickett & Baskin, 1973; Rice, 1974).
See also Swain (1977) for a synoptic review of secondary compounds as al-
lelopathic agents.
In the life cycle of any species, one portion, designated by Petersen & Fair-
brothers (1973) as the weakest link, is likely to be the most vulnerable to al-
lelopathic interactions. As an evolutionary strategem, allelopathy would be de-
veloped most effectively against the weakest link in an organism’s life cycle, such
as germinating spores and developing prothalli of ferns or germinating seeds and
seedlings of higher plants. Furthermore, it is at these critical points that one ought
to be able to best detect allelopathy.
Intra- and interspecific allelopathic interactions occur in four ways: (1)
sporophytes acting on sporophytes; (2) sporophytes acting on gametophytes; (3)
gametophytes acting on sporophytes; and (4) gametophytes acting on
gametophytes. Seed plants reduce gametophytic vulnerability structurally by en-
closing the gametophyte and ovule and functionally by pollination and fertiliza-
tion. Therefore, the weakest link in seed plants is shifted from their gametophytes
to their germinating seeds and seedlings. In pteridophytes, presumably the
gametophyte generation—spore, prothallus and gametes—is the most vulnerable
portion of the fern life cycle with regard to interspecific interactions for habitat
maintenance (allelopathy and competition), for a smaller amount of chemicals
would be required to eliminate a gametophyte from effective competition than a
sporophyte.
Of the four interactions, gametophytes acting on gametophytes of different
species permits study of the weakest link hypothesis in an experimental design
where interspecific competition is minimal. Fern gametophytes were selected as
experimental organisms because they are easily cultured and amenable to the
experiments’ requirements of control, replication, manipulation, and minimiza-
tion of competition. Six fern species usually found growing in similar natural
habitats were selected for the initial survey experiment which was conducted to
determine if allelopathic effects were detectable between any of the species. By
similar natural habitats we mean that, among the species selected, there is some
overlap between their respective ecological amplitudes so that it is possible that
they would be in competition for the same space.
The literature on fern allelopathy is limited but increasing. Froeschel (1953)
reported that water extracts of Polypodium aureum and Lycopodium clavatum
*Department of Botany, Howard University, Washington, D.C. 20059.
**Department of Botany, Rutgers University, New Brunswick, NJ 08903.
AMERICAN FERN JOURNAL: VOLUME 70 (1980)
74
FIGS. 1-3, Gametophytes of Dry.
growth. FIG. 1. Gametophytes o
opteris intermedia and Osmunda cinnamomea after 30 days of
f D. intermedia (2 prothallial cell Stage) and O. cinnamomea (multi-
al D. intermedia gametophyte from a control monoculture. FIG. 3.
phyte from a control monoculture.
PETERSEN & FAIRBROTHERS: RECIPROCAL ALLELOPATHY 75
decreased the growth rates of gametophytes of four fern species. Bell (1958) found
that an aqueous extract of Dryopteris filix-mas gametophytes stimulated spore
germination and prothallium growth in D. borreri, but that the gametophytic de-
bris of D. filix-mas, when incorporated into agar medium, prevented D. borreri
spore germination, an example of gametophyte—gametophyte allelopathy.
Fukuzumi (1971) reported allelopathic effects from Preris japonica frond extracts
on Impatiens balsamina root growth. Gliessman and Muller (1972) investigated
the phytotoxic effects of compounds from Preridium aquilinum on the surround-
ing vegetation. (See Miller, 1968 for an excellent, though now dated, comprehen-
sive review of fern gametophyte literature.)
Davidonis and Ruddat (1973, 1974) reported sporophyte—gametophyte al-
lelopathy in ferns. The roots and fronds of Thelypteris normalis produce two
allelopathic chemicals, thelypterin A and B, which inhibit cell division in the
gametophytes of TJ. normalis, Pteris longifolia, and Phlebodium aureum.
Thelypterin A has been tentatively identified as an indole derivative. Davidonis
(1976) found that 7. normalis gametophytes produce thelypterin A; the
sporophytes of other ferns also contain these compounds: thelypterin A and B in
T. dentata roots, thelypterin A in T. noveboracensis leaves, and thelypterin B in
the roots of two Pteris species. Davidonis (1976) also reported the presence of an
unidentified growth inhibitor in the leaves of Osmunda cinnamomea. Star and
Weber (1978) reported that sporophyte exudates from Pityrogramma species in-
hibit spore germination and gametophyte development in P. calomelanos. In-
hibitors were identified as a dihydrochalcone and a flavonol. Munther and Fair-
brothers (1980), testing leaf leachates and extracts of Dennstaedtia punctilobula,
Osmunda cinnamomea, and Osmunda claytoniana obtained from New Jersey and
Vermont populations, demonstrated geographic differences in allelopathic and
autotoxic responses among these species based on the amount of spore germina-
tion.
MATERIALS AND METHODS
Mature spores were collected from the following species: Osmunda cin-
namomea, O. claytoniana, O. regalis, Matteuccia struthiopteris, Onoclea sen-
sibilis, and Dryopteris intermedia. White's minimal nutrient medium at one-half
strength at pH 5.5 was used to culture the gametophytes. Both liquid and 1% agar
cultures were used. Spore density was approximately 1,000 spores per 90 mm
diam. plate. All cultures were grown under axenic conditions (Steeves, 1955), ina
growth chamber with fluorescent light (300 ft-c) at 20° C under a diurnal cycle of
12/12 hr.
In the initial survey experiment, spores of the six species were sown on agar
plates in paired strips adjacent to one another in all possible combinations. Con-
trol plates containing spores from one species also were sown. Plates were
examined daily at the interfaces of adjacent species for symptoms of allelopathy or
competition such as decreases from the control plates in percent germination or
growth.
76 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
PETERSEN & FAIRBROTHERS: RECIPROCAL ALLELOPATHY 77
On the basis of this experiment, D. intermedia and O. cinnamomea were
selected as the most promising taxa for further experimentation because the
gametophytes of these species appeared to inhibit each other’s growth. A
minimum of 10 replicates was run for each experiment. Initially, control plates and
experimental plates containing a mixture of Dryopteris and Osmunda spores were
prepared. The next phase was designed to eliminate the possibility of competition.
Separate liquid cultures of D. intermedia and O. cinnamomea gametophytes (0.5g
spores/liter of half-strength White’s Medium at pH 5.5) were initiated and grown
for two weeks. The gametophytes were then filtered off and the supernatants were
conserved. Agar plates were prepared as above. Control plates consisted of
spores of one species sprayed with one ml of the supernatant of the same species;
experimental plates contained spores of one species sprayed with one ml of the
supernatant of the other. This experimental design eliminates the possibility of
interspecific competition (e.g., differential nutrient assimilation by one species
over the other).
RESULTS AND DISCUSSION
In the survey of six species for allelopathic symptoms, a clear area was detected
on the plates at the interface between O. cinnamomea and D. intermedia
gametophytes. This was the result of progressive inhibition of D. intermedia
gametophyte growth, which was proportional to the proximity of Osmunda
gametophytes.
In the first phase of the Dryopteris—Osmunda growth rate analyses, Dryopteris
spore germination was initially lower on the experimental plates (60% germination
after 7 days) compared to the control plates (90% germination after 7 days). But
after 10 days, 90% germination was reached on the experimental plates. Post-
germination rate data was discontinued because it soon became apparent that the
gametophytes of both species on the experimental plates were growing very
slowly (Fig. 1) and no longer had significantly different growth rates. In contrast,
the control plate gametophytes developed normally (Figs. 2 and 3).
The gametophytes of D. intermedia and O. cinnamomea inhibit the growth and
development of one another, but one cannot distinguish from these results
whether the inhibition is the result of allelopathy or competition, and so a set of
supernatant experiments was designed to do so. These experiments essentially
reproduced the results of the preceeding set of experiments. Control plates
sprayed with supernatant from the same species produced normally de veloped
gametophytes (Figs. 4 and 5). But experimental plates sprayed with supernatant
from the other species yielded gametophytes having a severely retarded growth
rate; after germination and a few cell divisions, development essentially ceased
(Figs. 6 and 7).
This reciprocal supernatant experiment proves that the gametophytes of each
species were suppressing cell division of the gametophytes of the other species
and that this was done through the release of inhibitory compounds, rather than by
competition.
78 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
This is the first recorded example in ferns of reciprocal allelopathy, in which
two antagonistic species act on one another, that has been demonstrated in vitro
between gametophytes.
Previous investigators working with ferns have demonstrated unidirectional
allelopathy in the following systems: sporophyte acting on gametophyte (Froe-
schel, 1953; Davidonis & Ruddat, 1973, 1974; Star & Weber, 1978; Munther &
Fairbrothers, 1980) and gametophyte acting on gametophyte (Bell, 1958).
We wish to acknowledge financial aid from NSF Grant GB-13202 and a Rutgers
Research Council Grant awarded to D. E. Fairbrothers.
LITERATURE CITED
BELL, P. R. 1958. Variations in the germination-rate and development of fern spores in culture. Ann.
Bot -5S11.
DAVIDONIS, G. H. 1976. The occurrence of thelypterin in ferns. Amer. Fern J. 66:107-108.
—_—_——., and M. RUDDAT. 1973. Allelopathic compounds, thelypterin A and B in the fern Thely-
pteris normalis. Planta (Berlin) 111:23-32.
UDDAT. 1974. Growth inhibitor in gametophytes and oat coleoptiles by the
thely plerin A and B released from roots of the fern Thelypteris normalis. Amer. J. Bot.
1:925-930.
FROESCHEL, P. 1953. Remstoffen by lagere plantaardige Organismen. Natuwuv. Tydschr. 35:70-75.
FUKUZUMI, K. 1971. cr a effect of Pteris japonica on Impatiens balsamina root growth
from leaf extracts. Biol. J. Naro Women’s Univ. 21:8-9.
GLIESSMAN, S. R., and C. H. MULLER, 1972. The phytotoxic potential of bracken, Pteridium
aquilinum (L.) Kuhn. Madrono 21:299-304.
MILLER, J. H. 1968. Fern gametophytes as experimental material. Bot. Rev. 34:361-440.
MULLER, C. H. 1970. The role of allelopathy in the evolution of wea In K. L. Chambers, ed.
Biochemical Coevolution. Oregon State Univ. Press, Corv
MUNTHER, 4s E., and D. E. A 1980. Allelopathy iad autotoxicity in three species
of ferns. Amer. Fern J. 70: i S:
PETERSEN, . L. and D. E. FAIRBROTHERS. 1973. Allelopathy: Gametophytic chemoantagonism
between Dryopteris and Osmunda or supporting the ‘‘weakest link’’ hypothesis. Amer. J.
Bot. 60:32. ( Abstr.)
PICKETT, S. T., and T. M. BASKIN. he Allelopathy and its role in the ecology of higher plants.
Biologist (Phi Sigma Soc.) 55:49-73.
RICE, E. L. 1974. Allelopathy. pn a Press, New York.
STAR, A. E., and P. WEBER. 1978. Sporophytic exudate inhibition of pe development in
Pityrogramma calomelanos. Plant Sci. Conf. VPI, Blacksburg, Va. (Abs
STEE VES, T. A., I. M. SUSSEX, and C. R. PARTANEN. 1955. In vitro sides on | abnormal growth
of prothalli of the bracken fern. Amer. J. Bot. 42:232-245.
SWAIN, T. 1977. Secondary compounds as protective agents. Ann. Rev. Plant Physiol. 28:479-501.
AMERICAN FERN JOURNAL: VOLUME 70 (1980) 79
REVIEW
“FLORA DEL AVILA” by Julian A. Steyermark and Otto Huber. Publication
Especial de la Sociedad Venezolana de Ciencias Naturales, Caracas. 1978. 971 pp.
+ 308 plates. Bs. 150 (ca. $35.00).—This is a flora of the Parque Nacional ‘El
Avila,’ part of a small mountain range lying between the city of Caracas and the
Caribbean Sea. The area is about 130 square kilometers and the major peaks reach
an altitude of 2000 to 2700 meters.
Introductory chapters describe the history of botanical exploration of the re-
gion, the soils, geology, climate, principal vegetation types, and the geographic
relations of the flora. There is also a series of color photographs of flowers, the
vegetation, and three pteridophytes.
The flora consists of 127 families of flowering plants, 809 genera, and 1741
species. Each family treatment has an illustrated key to the species and a list of
them with ecological notes. There are also line drawings of a selection of species.
The ‘Flora of Avila’ is a major addition to neotropical floras, and its utility
extends well beyond the region covered.
The ferns and fern-allies are all treated under the Pteridophyta, rather than by
families, with 52 genera and 151 species included. The key to these is rather long,
but it is well organized and accurate. Marginal illustrations of characters aid in the
use of the headings. Following the list of species, there are excellent drawings of
99 species. The largest genus is Polypodium (sens. lat.) with 19 species, followed
by Elaphoglossum with 15 and Asplenium with 13. About a third of the genera are
represented by a single species.
The fern flora consists mostly of rather common and widely distributed species
of the montane forest and cloud forest of northern South America. There are no
ednemics, but about 25 species are of restricted distribution or are otherwise of
geographic interest. For example, the rare Lycopodium caracasicum, restricted to
the coastal Cordillera, reaches its easternmost station here, and both
Phanerophlebia juglandifolia and Pellaea ovata are represented by disjunct sta-
tions at the eastern limit of their range. About two-thirds of the species grow in the
cloud forest zone and some are restricted to it, for example the five species of
Cyatheaceae and several species of Elaphoglossum and Polypodium. There is a
subparamo zone on the highest parts of the mountains, and it is here that the
strongest relations with the flora of the high Andes are found. Asplenium
monanthes and Lycopodium vestitum are examples among the pteriodphytes.
The ‘‘Flora de Avila’’ may be obtained from: Sociedad Venezolana de Ciencias
Naturales, Calle Arichuna, Apartado de Correos 76771, Urb. El Marques,
Caracas 107, Venezuela. The price if 150 bolivares (ca. $35.00).—Rolla Tryon,
Gray Herbarium, Harvard University, C ambridge, MA 02138.
80 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
SHORTER NOTE
THELYPTERIS TORRESIANA IN VENEZULEA. — For several years, the
senior author has been intrigued by the sudden appearance and persistence in his
Caracas garden of a terrestrial, acaulescent fern, most attractive with its pale
green, large, gracefully thrice-cut fronds. Specimens sent to Dr. John Mickel were
identified as Thelypteris torresiana (Gaud.) Alston, a species originally described
from Guam and found native elsewhere in the Asian tropics. In the New World,
this species has become introduced and naturalized in the southeastern United
States, Cuba, Jamaica, the Lesser Antilles, Trinidad, Tobago, Honduras, Ven-
ezuela, Brazil, and Argentina.
Vareschi in Lasser (Fl. Venez. 1:439. 1969) treats this species under the in-
validly published name ‘‘Lastrea setigera.’’ Morton (Amer. Fern J. 52:27-29.
1962) gives a correct synonymy and has shown that the species should not be
confused with the rare Old World T. setigera (Blume) Ching.
Leonard (Amer. Fern J. 62:97-99. 1972) observed the preference of 7. tor-
resiana for moist ravines and stream banks in the southeastern United States. It
occurs in similar habitats in Venezuela, in such places as moist forests along roads
and trails at 400-1400 m altitude. It is common in cool cloud forests, but also
grows in warmer zones, both in deciduous and evergreen tropical forests. In the
senior author’s garden in Caracas, it is aggressive, weedy, and often invasive,
characteristics which have facilitated its spread in natural habitats. In the lo-
calities where it has become naturalized, it appears to be part of the native vegeta-
tion.
According to specimens in the Herbario Nacional de Venezuela (VEN), T.
torresiana was first found in Venezuela in 1943 in the Parque Nacional Pittier,
Estado Aragua (Killip & Lasser 37797, US, VEN). Since then it has spread in the
Coastal Cordillera throughout northeastern Venezuela to the states of Portuguesa,
Yaracuy, Carabobo, Guarico, Miranda, Sucre, and Monagas, and to the Distrito
Federal.—Julian A. Steyermark, Instituto Botanico, Apartado 2156, Caracas,
Venezuela and Francisco Ortega, Estacion Biolégica ‘‘Pozo Blanco,’’ Apartado
116, Acarigua, Edo. Portuguesa, Venezuela.
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AMERICAN volume 7
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JOURNAL
cee en
QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY
The Distribution and Ecology of Phyllitis scolopendrium
in Michi RICHARD P. FUTYMA
Supplemental Notes on Lesser Antillean Pteridophytes
GEORGE R. PROCTOR
Additions to the Pteridophyte Flora of the Great Plains
RALPH E. BROOKS
Flavonoid Synthesis and Antheridium Initiation
in Dryopteris Gametophytes
RAYMOND L. PETERSEN and DAVID E. FAIRBROTHERS
Date of Publication of Sodiro’s
**Sertula Florae Ecuadorensis”’
DAVID B. LELLINGER %
Reproductive Biology and Gametophyte Mor
of New World Populations of Acrostichum aureum ROBERT M. LLOYD 99
Shorter Notes: Diplazium japonicum New to Alabama;
Moths and Ferns; Three Additions to the
Pteridophyte Flora of Escambia County, Florida MISSOUR) BWR 111
OCT 16
GARDEN LIBRARY
The American Fern Society
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ROBERT M. LLOYD, Dept. of Botany, Ohio L University, Athens, Ohio 45701. President
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AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 3 (1980) 81
The Distribution and Ecology of Phyllitis scolopendrium
in Michigan
RICHARD P. FUTYMA*
The Hart’s-tongue, Phyllitis scolopendrium, has been noted for its circumboreal
and North American disjunct distributions (Fernald, 1935; Wagner, 1972). On this
continent P. scolopendrium var. americana Fern. is known to occur in Ontario
(Soper, 1954), Michigan (Hagenah, 1954, 1956), New York, Tennessee, and
Alabama (Short, 1979). By far the majority of the Hart’s-tongue sites are associ-
ated with the limestones and dolomites of the Niagara escarpment. This geological
formation can be traced from central New York westward into Ontario, where it
turns northwestward near the head of Lake Ontario, and through the Bruce Penin-
sula and Manitoulin Island in Lake Huron, into the upper peninsula of Michigan.
From there it arcs southwestward through Wisconsin’s Door Peninsula to the east
of Green Bay and disappears to the south.
This paper will deal principally with the northernmost American Hart’s-tongue
colonies, those in upper Michigan, and with some ideas concerning the factors
determining its distribution in that region.
A NEW LOCALITY IN MICHIGAN
On 3 August 1978, I discovered a previously unreported locality for P.
scolopendrium in Mackinac County, Michigan, while I was botanizing along the
slopes of a bedrock knob on the Niagara escarpment. The site is strewn with low,
moss-covered dolomite boulders under a tree canopy almost completely domi-
nated by Acer saccharum, with only minor numbers of other hardwood species
(Fig. Ty
At the time of the discovery of the site, two fronds of P. scolopendrium were
collected as a voucher specimen and deposited in the herbarium of the University
of Michigan Biological Station (UMBS). Also found were Polystichum lonchitis
and Geranium robertianum (Fig. 2), two plants frequently associated with Phyl-
litis scolopendrium (Hagenah, 1956). Walking Fern, Camptosorus rhizophyllus,
occurs locally at the site, densely covering large boulders at least 1.5 m above the
ground surface. Dr. W. H. Wagner, Jr. was among those visiting the site shortly
after its discovery. He compiled the following list of pteridophytes on 29 August
1978: Asplenium trichomanes, Botrychium virginianum, Camptosorus rhizophyl-
lus, Cystopteris bulbifera, C. fragilis, Dryopteris filix-mas, D. intermedia, D.
spinulosa, Equisetum arvense, E. scirpoides, E. sylvaticum, Matteuccia
struthiopteris, Onoclea sensibilis, Polypodium virginianum, Polystichum braunii,
and P. lonchitis. All were found within 50 m of the Phyllitis colony. Asplenium
viride also has been reported at the site (D. Henson, pers. comm.
This new Hart’s-tongue site is situated within the Hiawatha National Forest,
and will be referred to here as the ‘East Lake station.”’ In the Fall of 1978, the
U.S. Forest Service decided to survey and map the extent of the Phyllitis colony.
*Department of Botany, University of Michigan, Ann Arbor, MI 48109.
Volume 70, number 2, of the JOURNAL was issued June 30, 1980.
2 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
8
tube (horizontal) in the right foreground is 65 cm long. FIG
low boulder, along with Polystichum lonchitis and Geraniu
. 2. Phyllitis scolopendrium growing on a
m robertianum on top of boulder.
R. P. FUTYMA: PHYLLITIS SCOLOPENDRIUM IN MICHIGAN 83
In all, 64 individuals were counted, their locations mapped, and the tree nearest to
each group of individuals was marked. In combination with long-term observa-
tion, this information has potential for use in studying the population ecology of
this rare fern.
During the summer of 1979, the U.S. Forest Service undertook an inventory of
all prospective localities within the borders of the Hiawatha National Forest in
hopes of uncovering other unreported Hart’s-tongue sites. Unfortunately, no suc-
cess was reported.
DISTRIBUTION IN UPPER MICHIGAN
Hagenah (1954, 1956) reported on the first two upper Michigan sites for P.
scolopendrium. The East Lake station is situated between these two, which are 30
km. apart (Fig. 3). The population at the Trout Lake station in Chippewa County
apparently is extinct. The easternmost station, known as the Hagenah site, has
recently been acquired by the Michigan Nature Association as a plant preserve.
The location of these sites and the major North American concentration of P.
scolopendrium in Ontario, south of Lake Huron, are shown in Figure 3.
All three Michigan localities are similarly located on prominent hills that are
part of the Niagara escarpment. Along much of its length in Mackinac and Chip-
pewa counties, the escarpment is obscured by thick deposits of glacial drift. The
position of the escarpment is manifested mainly by a series of bedrock knobs
scattered from east to west across the region. These hills rise 30-100 m above the
surrounding plain and range in area from 150 to over 3000 hectares. It is highly
unlikely that they were ice-free nunataks during the Wisconsinan glaciation, as
suggested by Fernald (1935) in explaining the occurrence of Hart’s-tongue on the
highest outcrops of the same escarpment in Ontario.
Another similarity shared by the three localities is that the Hart’s-tongue col-
onies are situated at elevations near or above that of the ancient shoreline of Lake
Algonquin. In fact, the East Lake station was discovered in the course of floristic
reconnaissance along one such shoreline. The plants are growing on boulders
uncovered by wave action. Lake Algonquin, a precursor of lakes Huron and
Michigan, covered much of upper Michigan immediately upon retreat of the con-
tinental ice sheet about 11,000 years ago. At that time the bedrock hills that define
the Niagara escarpment formed an archipelago in the lake. By about 10,400 years
ago Lake Algonquin ended when the waters fell to lower levels and more of the
present land area was uncovered.
The fact that P. scolopendrium has been found in upper Michigan only in places
that were islands in Lake Algonquin may have some special significance.
Throughout Mackinac and parts of Chippewa counties, there is a large area which
was inundated by Lake Algonquin but now is covered by deciduous forests of the
sort preferred by the fern, in which there are limestone and dolomite outcrops
(e.g., Drummond Island) or concentrations of glacially-transported boulders. Al-
though actively sought, Hart’s-tongue has not been seen in those places. This
distribution pattern may be explained in several ways; here we will consider three
hypotheses.
84 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
Hypothesis 1.—Phyllitis scolopendrium was dispersed to upper Michigan, pre-
sumably from the south, at the time of the existence of Lake Algonquin, ca. 10,500
years ago. These rocky islands with their depauperate flora may have offered
suitable substrates and conditions of low competition favorable to the establish-
f=
TL ee :
eu rignlent ide — vet ‘um in the upper Great Lakes region. The three stations of
sine re indicated by dots. The general : pe
shown by stippling. (modified from Soper 1904: g area where the fern occurs in Ontario is
ment of this fern. When more land area was uncovered by the recession of lake
levels and closed forests covered the region, P. scolopendrium may not have been
a es aggressive Colonizer to spread from its territory acquired earlier.
Se deed tat dp pcroer of P. scolopendrium to the former islands may be a
of environmental differences between hilltop rock outcrops and those at
R. P. FUTYMA: PHYLLITIS SCOLOPENDRIUM IN MICHIGAN 85
lower elevations. The hills along the escarpment have thin soils with more out-
crops, and the fact that they were high enough to escape submergence under Lake
Algonquin may be only a coincidence. Fewer favorable sites for the fern exist at
lower elevations because thicker deposits of glacial till and lake sediments cover
the bedrock. At those lowland sites where rock surfaces are available, other
environmental factors may be unfavorable.
Hypothesis 3.—P. scolopendrium had (or has) a wider distribution in upper
Michigan than is known at present. Logging of forests may have opened the
vegetation at many former localities of the fern, making the sites unsuitable and
leading to its extinction. Therefore, the original distribution of the Hart’s-tongue
in upper Michigan prior to European settlement had little to do with the geography
of Lake Algonquin.
DISCUSSION
The suggestion that P. scolopendrium first reached upper Michigan during the
existence of Lake Algonquin has some appeal. Such an hypothesis could explain
why extensive areas of limestone outcrops which are situated between the Michi-
gan stations and the main North American concentration of Hart’s-tongue in
Ontario, and which were inundated by Lake Algonquin, such as Manitoulin Island
and Drummond Island, are devoid of the fern. The Bruce Peninsula, where many
of the Ontario stations are located, also was completely submerged at that time,
but one may propose that its connection to the mainland at a point where many
non-submerged Hart’s-tongue localities exist facilitated its colonization at a later
date.
However, fossil pollen studies by the present author and others (Brubaker,
1975; Saarnisto, 1974) indicate that the late-glacial forests of the region, during and
after the existence of Lake Algonquin, comprised mainly spruce (Picea spp.) and
Jack pine (Pinus banksiana). A salient feature of the ecology of Phyllitis scolopen-
drium var. americana is that it is never found in coniferous forests, even when
adjacent tracts of deciduous forests contain the fern. In Ontario it is seen under
deciduous canopies ranging from successional poplar stands to climax maple-
beech forest, but never under conifers (A. Reznicek, pers. comm.). If the Hart’s-
tongue is a strict associate of the northern hardwoods forest, then it might have
reached northern Michigan only within the past 5000 years, which is when this
vegetation type became most widespread in the region. Thus, unless the ecology
of this species was different 10,500 years ago from what it is today, we should be
safe in rejecting the first hypothesis.
The effect of forest clearance on populations of P. sclopendrium is poorly
known. Most of the localities in Ontario and Michigan have been logged at one
time or another. In the case of the East Lake station, logging did take place, but
perhaps not to the extent of clearcutting. The example of the Ontario Hart’s-
tongue colonies found in early successional Populus woods shows that it can be an
aggressive colonizer little affected by logging, provided that spore sources exist
nearby. There is still the possibility that small, isolated colonies could become
extinct and not be recolonized after logging and forest regrowth.
86 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
Most North American P. scolopendrium sites appear to be associated with
moist slopes or hillsides, such as bouldery talus slopes and crests of escarpments
(Soper, 1954) and sinkholes (Short, 1979). Along the Michigan outcrops of the
Niagara escarpment, there are very few places where there is a high, steep cliff
below which a rocky talus has accumulated. The Michigan Hart’s-tongue colonies
are found where bedrock just breaks through the surface on a moderately steep
hillside or on slopes with a high concentration of low boulders that are separate
from the bedrock. At the East Lake station, and possibly the second, easternmost
locality described by Hagenah (1956), the boulders on which the ferns are growing
represent a lag deposit formed by the removal of the surrounding sandy till by the
action of the waters of Lake Algonquin. The boulders themselves had been quar-
ried from nearby outcrops by the glacier and carried only a short distance before
being deposited.
Extensive flat areas with limestone bedrock near or outcropping at the surface,
such as Manitoulin and Drummond islands, do not appear to be suitable. Possibly
these present too dry a habitat (Hagenah, 1956) or the forests are too open for the
Hart’s-tongue.
Another indicator of this fern’s requirement for moist conditions is its prefer-
ence for growing on low boulders no more than 30 or 40 cm above the forest leaf
litter (Fig. 2) or in crevices of limestone pavement. The plants would be more
exposed and subject to desiccation on sheer cliff faces or higher on boulders. Low
position may also be a consequence of insulation and protection from desiccation
provided by winter snow at northern localities.
Large, glacially transported dolomite and limestone boulders exposed above the
soil are scattered throughout much of Mackinac County south of the Niagara
escarpment. Most are close to | m in diameter, but individuals 2-3 m in diameter
are not unusual. Seldom do these boulders occur in concentrations similar to those
seen in typical Hart’s-tongue habitat. Although the overall forest setting may seem
Suitable, these boulders may present desiccation problems and may be too few at
any given location to provide a sufficient number of microsites for a viable colony
of P. scolopendrium to become established.
CONCLUSIONS
Phyllitis scolopendrium has now been reported from three localities on the
Niagara €sCarpment in upper Michigan. All three stations are similar in that they
are situated on hills that were islands in Lake Algonquin, which existed ca. 10,500
years ago. Despite assiduous searching by botanists, P. scolopendrium has never
been found in this regi
This interesting distribution pattern probably does not indicate that P.
scolopendrium first reached these sites while Lake Algonquin was in existence,
€getation was coniferous forest, a vegetation type at present
this fern. A more likely explanation is that these hillside sites
vironmental requirements in terms of topography, moisture,
and microsite abundance more adequately than other sites where limestone out-
crops and boulders are available.
not associated with
R. P. FUTYMA: PHYLLITIS SCOLOPENDRIUM IN MICHIGAN 87
We can never be certain that P. scolopendrium did not occur in a greater
number of localities in upper Michigan at some time in the last 10,000 years. More
intensive botanical exploration in the region may eventually confirm or refute the
apparent correspondence between Hart’s-tongue fern localities and island areas in
Lake Algonquin. In this regard it may be productive to pay particular attention to
boulder concentrations along the ancient shorelines of this glacial lake.
With respect to determining the time of immigration of P. scolopendrium to
upper Michigan, we can only say that it arrived less than 10,000 years ago and
possibly only within the last 5000 years. There is little hope of being able to
pinpoint this date more exactly, for it is unlikely that the spores or other parts of
this rare plant will be found in the fossil condition.
Therefore, the first hypothesis is the least likely and the second is the most
plausible explanation for the distribution of Hart’s-tongue in Michigan. We do not
have sufficient information to reject the third hypothesis.
In order to understand the factors determining the geographic distribution of
rare plant species such as P. scolopendrium, we must take into account the vege-
tational history of the region, as well as the ecological relationships between rare
species and the biotic and abiotic components of their immediate environment.
Further contributions in this regard can be made by studying the population
dynamics of known colonies of P. scolopendrium. Such a study will be possible at
the East Lake station, where an entire Hart’s-tongue colony has been counted and
mapped.
I would like to thank the following people for their help in providing information
at various times during my research and for their comments and criticisms of the
manuscript: Joseph Beitel, William S. Benninghoff, Don C. Henson, Anton
Reznicek, Charlotte Taylor, Edward G. Voss, and Warren H. Wagner, Jr. The
work was supported in part by a National Science Foundation Doctoral Disserta-
tion Improvement Grant.
LITERATURE CITED
i coc atc LB 1975, sidecases forest patterns associated with till and outwash in northcentral
L Michigan. Quaternary Res. 5:499-527.
FERNALD. M. L. 1935. Critical plants of the Upper Great Lakes region of Ontario and Michigan.
Rhodora 37:197—222, 238-262, 272-301, 324-341.
a cae D. J. 1954. The Hart’s-tongue in Michigan. Amer. Fern J. 44:2-7.
————. 1956. More Hart’s-tongue in Michigan. Amer. Fern J. 46:70-74.
SAARNISTO, M. 1974. The deglaciation history of the Lake Superior region and its climatic implica-
tions. Quarternary Res. 4:316-339.
SHORT, J. W. 1979. Phyllitis scolopendrium newly discovered in Alabama. sy Fern J. 69:47-48.
SOPER, J. H. 1954. The Hart’s-tongue fern in Ontario. Amer. Fern J. 44: 129-14
WAGNER, W. H, Jr. 1972. Disjunctions in homosporous vascular plants. Ann. panos Bot. Gard.
59:203-217.
88 AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 2 (1980)
Supplemental Notes on Lesser Antillean
Pteridophytes
GEORGE R. PROCTOR*
During the several years since publication of the author’s pteridophyte volume in
R. A. Howard’s “Flora of the Lesser Antilles” (vol. 2, 1977), a number of errors
and a few omissions have come to light. For the convenience of persons interested in
the ferns of this geographic area, it seems desirable to bring together and make
available the more important corrections and augmented facts. I am indebted to Dr.
David Lellinger of the Smithsonian Institution for placing many of these data at my
disposal. The information is presented according to the page-numbers of the volume.
Emendations of geographic range, minor corrections of spelling, etc., are omitted.
. 7.—In citing collectors for the various islands, the name of Stehlé, 1953-1954,
was unfortunately omitted from the lists for Guadeloupe, Les Saintes, and Marti-
nique. My apologies to Dr. Stehlé!
. 16.—The correct citation for this plant is Psilotum nudum (L.) Beauv., Prodr.
Fam. Aethéog. 112. 1805. Also, the correct date of J. Bot. Schrader 1800(2) should
be 1802, not 1801. This correction should be made at numerous places where it
appears throughout the volume.
p. 18.—H. & M. Stehlé (Mém. Soc. Bot. France 1953-54, p. 45) reported
Psilotum complanatum Swartz from Martinique on the basis of unpublished reports
by Fée and Urban. No specimens were cited. The Urban record (Symb. Ant. 9:391.
1925) merely cites the Fée reference. Fée’s record (Mém. Foug. 11:133. 1866) cites
a collection by Mlle. Rivoire, “sur un carapa, prés de Saint-Pierre.” This report
should be considered doubtful unless it can be substantiated by an authentic
specimen.
pp. 75-78.—In subgenus Mecodium, two possible additional species should be
noted. One of these, known only from Guadeloupe, was given the illegitimate name
Hymenophyllum caespitosum by Fée (1866, non Gaud., 1825) and an unpublished
one by Maxon and Morton. This entity, based on a L’Herminier specimen at Paris,
ra pee be considered a diminutive form of H. undulatum, but requires further
study.
The other species is Hymenophyllum I’ herminieri Mett. (Linnaea 35:392. 1868).
This plant has the frond shape of H. fucoides (subg. Hymenophyllum), but its entire
margins place it in subg. Mecodium. Although the original L’Herminier specimens
were found in Guadeloupe, more recently similar material was collected in Domin-
ica (G. Proctor Cooper 106, US, collected Jan. 31, 1933, “on rocks”). This may
well be a valid species.
pp. 132-133.—The correct generic name of this plant is Lonchitis L., Sp. Pl.
2:1078. 1753, and the correct species citation is Lonchitis hirsuta L., loc. cit. The
name Anisosorus falls into synonymy, but the relevant typification remains the same.
136.—the correct citation for the Pteridium is P. aquilinum var. arachnoideum
(Kaulf.) Brade (Zeitschr. Deutsch. Ver. Wiss. Kunst. S. Paulo 1:56. 1920).
*Arnold Arboretum Herbarium, 22 Divinity Avenue, Cambridge, MA 02138.
G. R. PROCTOR: LESSER ANTILLEAN PTERIDOPHYTES 89
pp. 136—138.—The plant described and illustrated here should bear the generic
name Blotiella R. Tyron (Contr. Gray Herb. 191:96. 1962), and the species should
be known as Blotiella lindeniana (Hook.) R. Tyron, based on a type from
Venezuela. The name Lonchitis should not be associated with this taxon. As the
only record of this species from the Lesser Antilles has been shown by Lellinger
(Taxon 26:578-580. 1977) to be based on a misidentification, the genus Blotiella
cannot be listed as occurring in the Lesser Antilles, and this entire entry should be
deleted from the book.
p. 176.—The type species of Cheilanthes, by conservation, is C. micropteris
Swartz, of South America.
p. 183.—The correct authority of Adiantum lucidum is “(Cav.) Sw.”, with the
basionym Pteris lucida Cav., based on a specimen collected by Née in Ecuador; this
was cited by R. Tryon (Contr. Gray Herb. 194:148. 1964). The phrase “not Preris
lucida Cav.” on line 1 should therefore be deleted.
p. 194.—The correct name of this plant is Oleandra articulata (Swartz) C. Presl,
Tent. Pterid. 78. 1836, based on the same type cited for O. nodosa. The latter name
should be considered illegitimate. This question was thoroughly discussed by G. J.
de Joncheere (Taxon 18:538—-541. 1969).
p. 219.—Bolbitis aliena (Swartz) Alston (Kew Bull. 1932:310. 1932) was record-
ed from the Lesser Antilles by Hennipman (Leiden Bot. Ser. 2:135. 1977). He cited
the following specimens: St. Eustatius: Boldingh 44B (U); Guadeloupe: L’ Hermin-
ier 21 (CAL); and “Leeward Is.,” Holme s. n. (K).
This species can be distinguished from B. nicotianifolia by the lobed pinnae and
the absence of a separate terminal pinna. From B. portoricensis it is distinguished
by the vein-areoles lacking (or nearly lacking) included free veinlets and by the
Sterile blades being neither elongate nor proliferous.
p. 288.—The correct citation for the species on this page should be Nephrolepis
rivularis (Vahl.) Mett. ex Krug in Urban (Engl. Bot. Jahrb. 24:122. 1897).
p. 265. Species 8, listed as Diplazium limbatum, should be removed from
Diplazium and restored to the monotypic genus Hemidictyum. The correct name of
this species is therefore Hemidictyum marginatum (L.) C. Presl, based on
Asplenium marginatum L.
p. 292.—The correct name of species no. 20 is Thelypteris opulenta (Kaulf.)
Fosberg (Smiths. Contr. Bot. 8:3. 1972), based on Aspidium opulentum Kaulf. The
type is Chamisso s. n. (LE), from Guam.
p. 295.—The correct name of species no. 22 is Thelypteris kunthii (Desv.) Morton
(Contr. U. S. Natl. Herb. 38:53. 1967), based on Nephrodium kunthii Desv. The
type is a Venezuelan specimen without stated collector, ex. herb. Desvaux (P.)
p. 296.—The correct name of species no. 24 is Thelypteris hispidula (Dene. )
Reed (Phytologia 17:283. 1968), based on Aspidium hispidulum Dene. (Nouv. Ann.
Mus. Hist. Nat. 3:346. 1835). The type is said to be a Riedlé or Guichenot
specimen from Timor (P. .
p. 297.—Note 24a, Thelypteris hispidula var. hispidula and 24b, Thelypteris
hispidula var. inconstans (C. Chr.) Proctor, comb. nov., based on Dryopteris dentata
var. inconstans C. Chr. (Kungl. Sv. Vet. Akad. Handl. 16(2):27. 1936). The
90 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
lectotype of the latter is Ekman H 10524 (S, isolectotype US), selected by A. R.
Smith (Univ. Calif. Publ. Bot. 59:67. 1971).
p. 329.—The correct name of species no. 3. is Polypodium sororium Humb. &
Bonpl. ex Willd. in L. (Sp. Pl. 5:191. 1810). The type is Humboldt 424 (B—Herb.
Willd. 19684-1) from near Caripe, Venezuela.
p. 331.—The correct name of species no. 6 is Polypodium dissimile L., the type
being P. Browne (LINN 1252.24) from Jamaica.
p. 338 or 339. Polypodium palmeri Maxon (Contr. U. S. Natl. Herb. 17(7):600.
1916) should be added to the Lesser Antilles list. The type is Palmer 308 (US
572544 from Mexico. This member of subg. Microgramma was collected long ago
in Barbados by Jenman (NY, US), unfortunately without further data. This species
is largely Central American in distribution, but also has been found once in Jamaica.
This species is somewhat similar to P. lycopodioides, but differs in its larger size
(sterile fronds 5—20 cm long, 2-4 cm broad) and thicker texture, and in the thicker,
rope-like, whitish-scaly rhizomes.
p. 344.—Polypodium decurrens Raddi (Opusc. Sci. Bol. 3:287. 1819; Pl. Bras.
1:23, t. 33. 1825), based on material from Brazil, has been confirmed as occurring
in the Lesser Antilles. There is a good Martinique specimen of Duss 1568 at US,
whose label says, “Terrestre, rare. Piton Marcel, entre la montagne Pelée et le
Precheur; tres rare dans les pitons de Fort de France,” collected July 1885. The
Plumier plate cited on p. 344 (Tr. Foug. 99, t. 1/4) unquestionably belongs to the
same species.
Polypodium decurrens is referred to subg. Campyloneurum, and is unique among
the lesser Antillean species of this subgenus in having pinnate instead of simple
fronds. The individual pinnae are not unlike the entire blade of P. repens in shape,
texture, and venation.
p. 344.—Polypodium recurvatum Kaulf. can definitely be excluded from the
Lesser Antilles list; the record was based on Duss 4093, which was referred to P.
y Weenes & Bonpl. ex Willd. by A. M. Evans (Ann. Mo. Bot. Gard. 55(3):
p. 361.—In the caption for Figure 60b, the name for “e” should be G. taxifolia
(not G. taenifolia).
_ p. 366. A recently described species of Cochlidium from the Lesser Antilles is C.
jungens L. E. Bishop (Amer. Fern J. 68:84. 1978), based on Nicolson 1975 (US)
from Morne Micotrin, St. George Parish, Dominica. Among the Lesser Antillean
species of Cochlidium, C. jungens is distinguished from C. seminudum by the
smaller size of the fronds (2-8 cm long vs. 8-20 cm), but also by the non-contracted
and non-acuminate fertile fronds; the sterile blades are also narrower, being mostly
less than 3 mm wide. From C. rostratum it is distinguished by the superficial sori,
not immersed in a deep central groove.
p. 368.—The name of species no. 2. should be Cochlidium rostratum (Hook. )
Maxon ex C. Chr. (Dansk. Bot. Ark. 6(3):23. 1929), based on Wright s. n. (K.,
weed US) from Omotepe Island, Nicaragua. Additional Lesser Antillean records
of this species include Stehlé 341 and 1096 (both US) from Guadeloupe.
Pp. 374. The correct citation for Vittaria lineata is (L.) J. E. Smith (Mém. Acad.
Turin 10:421, ¢. 9, f. 5. 1793).
AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 3 (1980) 91
Additions to the Pteridophyte Flora
of the Great Plains
RALPH E. BROOKS*
Recent herbarium studies made while preparing manuscript for the forthcoming
manual of the Great Plains flora have led to the discovery of several specimens
representing new state records or significant range extensions apparently overlooked
by Petrik-Ott in “The Pteridophytes of Kansas, Nebraska, South Dakota and North
Dakota” (Beih. Nova Hedwigia 61:1—332. 1979).
Botrychium lunaria (L.) Swartz var. lunaria.—This taxon has been reported
previously for the Great Plains from North Dakota by Petrik-Ott (1979, p. 37);
however, that specimen is B. minganense Vict. The South Dakota collection
represents a southern range extension for this circumboreal species.
SOUTH DAKOTA: Lawrence County: Northern Black Hills: Old Balmoral Mine, NW from Crown
Hill, shrubby glade on plateau at old mine, 6100 ft alt., June 1930, Mrs. F. L. Bennett s.n. (BHSC).
Botrychium lunaria var. onondagense (Underw.) House.—This plant previous-
ly was known from scattered localities in the northwestern and northeastern United
States.
NORTH DAKOTA: Burke County: 12 mi SE of Lignite, N-exposed wooded ravine, 11 June 1971, G.
D. Hegstad 7855 (NDA).
Botrychium matricariifolium A. Braun.—Petrik-Ott (1979, p. 294) stated that
she had seen no collections of this species from South Dakota, and so she excluded
it from the Great Plains flora. In 1978 I visited the U.S. National Herbarium and
found the specimen cited by Clausen (Mem. Torrey Bot. Club 19:87. 1938) to be
determined correctly. This was verified by Dr. David Lellinger (pers. comm., 1980),
and so B. matricariifolium must remain a part of the Great Plains flora.
SOUTH DAKOTA: Custer County: Black Hills: Custer, 5500 ft alt., 15 Aug 1892, P. A. Rydberg
1186 (US).
Botrychium minganense Vict.—Petrik-Ott (1979, pp. 34-36) annotated, de-
scribed, and illustrated this specimen as typical -B. /unaria. Of the five plants on the
cited sheet, two are immature. The remaining three are typical B. minganense, they
have distinctly pinnatifid or pinnate lower pinnae, with only the uppermost pinnae
flabellate. This determination was kindly verified by Dr. Warren Wagner, Jr. (pers.
comm., 1979). The collection represents a slight southern range extension since the
species previously was known from Labrador west to Alaska and south to Michigan,
Minnesota, Colorado, Nevada, and California.
NORTH DAKOTA: McHenry County: Towner, sandy prairie, 11 June 1955, O. A. Stevens 1530
(NDA)
Ophioglossum vulgatum var. pseudopodum (Blake) Farw.—This specimen
was first reported as O. vulgatum L. by Clausen (1938, p. 126), who did not
recognize any infraspecific taxa within this species. Petrik-Ott (1979, p. 295) listed
“Herbarium, University of Kansas, Lawrence, KS 66044.
92 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
the record as unverified since she had not examined the specimen. I examined it in
1978, and found it to be the northern var. pseudopodum. The collection was made at
the southernmost limit of the range in our region. The variety previously was
reported from southern Canada south to Virginia, Indiana, Illinois, Nebraska, and
California. I have visited the Kansas locality in recent years and, although the
habitat is suitable for this plant, the chances are that it is now extirpated. Road
construction and housing developments have drastically altered the area since 1929.
NSAS: Crawford County: Pittsburg, | mi W of Broadway, in woods on low, rich slopes and in
draws, rare clusters, 15 June 1929, F. A. Riedel s.n. (NY).
Polystichum lonchitis (L.) Roth.—This collection is a slight southern and
eastern range extension from northeastern Wyoming. Both the South Dakota and
northeastern Wyoming sites are disjunct from the primary range of this circumboreal
species, which lies more than 150 miles to the west and many more miles to the
ortn.
SOUTH DAKOTA: Lawrence County: Black Hills, SW1/4, Sec. 36, T5N, RIE, S of Roughlock Falls,
mossy loam underwoods, over talus below limestone bluff, N-facing slope, 22 July 1971, C. A. Taylor,
W. Casper & A. Glynn 10918 (SDC).
I wish to thank the following curators for the loan of specimens and for aid in
various other ways: Dr. William T. Barker, North Dakota State University (NDA);
Dr. Gary Larson, South Dakota State University (SDC); Dr. John Mickel, New York
Botanical Garden (NY); and Dr. Joseph Thomasson, Black Hills State College
(BHSC).
REVIEW
“DAS BUCH DER FREILANDFARNE,” by R. Maatsch. 196pp. illustr. Paul
Parey, Berlin and Hamburg. 1980. ISBN 3-489-61422-4. DM. 68. (ca. $40.00).—
This book is intended for serious hardy fern growers. An introductory portion
contains notes on nomenclature, taxonomy, morphology, and fern habitats illustrated
with black-and-white photographs and line drawings. About half the book is a useful
alphabetical list of fern species and cultivars, concentrating on those grown in
Europe, and giving Latin and common names, a brief description of the plant, and
other useful notes. The last quarter of the book concerns fern culture. Unusual and
helpful information on flowering plants suitable for growing with ferns is included. |
hope the publisher will prepare an English edition so that Prof. Maatsch’s book
receives the wide circulation it deserves in the English-speaking world.—D.B.L.
AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 3 (1980) 93
Flavonoid Synthesis and Antheridium Initiation
in Dryopteris Gametophytes
RAYMOND L. PETERSEN* and DAVID E. FAIRBROTHERS**
There is now a fairly extensive vascular crytogam flavonoid literature and there
are a number of researchers actively engaged in this research field (Swain &
Cooper-Driver, 1973). All vascular crytogam flavonoid work has been done on the
sporophyte generation, with one exception: Laurent (1966) determined that
Blechnum brasiliense gametophytes produced the flavonoid kaempferol, which is
one of the flavonoids produced by the B. brasiliense sporophyte. Reasons for the
paucity of information on fern gametophyte flavonoids include the easy accessibil-
ity of sporophytes and the now disreputed opinion that flavonoids, being associ-
ated with lignin synthesis, are exclusive to vascularized plant bodies. This
exclusivity has been lost because flavonoids have been isolated and identified in
various non-vascular plant groups: certain algal divisions, bryophytes (Swain,
1974), and fern gametophytes (Laurent, 1966).
Initially our investigation was undertaken to determine if Dryopteris intermedia
A. Gray and D. marginalis A. Gray gametophytes produce flavonoids and, if so,
were these the same flavonoids produced by their sporophytic counterparts
(Petersen, 1976). Because of the unusual results of this first portion of the re-
search, the inquiry was amplified to include an analysis of flavonoid content along
a developmental profile of the gametophytes.
Half-strength White’s minimum nutrient medium adjusted to pH 6.0 was used to
culture the gametophytes. Liquid cultures were prepared by placing 0.25 g of
spores in a 41 flask and adding 2 | of nutrient solution. Separate cultures of D.
intermedia and D. marginalis were grown at 22°C under 300 ft-c of illumination
from cool-white fluorescent lights in a 12/12 hr diurnal cycle.
Gametophytes were harvested and assayed for flavonoids at three de velopmen-
tal stages: (1) pre-antheridial initiation (0 antheridia/gametophyte), (2) antheridial
initiation (0 or 1 antheridia/gametophyte), and (3) post-antheridial initiation (4-6
antheridia/gametophyte).
Ten-gram samples of harvested gametophytes were immediately extracted in
methanol and re-extracted repeatedly until a colorless supernatant was obtained.
Concentrated extracts were spotted onto Whatman 3MM chromatography paper
(42 x 55 cm). Chromatograms were developed in two dimensions employing the
two standard solvent systems for the separation of flavonoids: t-butanol, acetic
acid, water (3:1:1) for the first dimension and 15% acetic acid, water for the
second dimension. Completed chromatograms were inspected under UV light,
both in the presence and absence of NH. Spot color changes under both condi-
tions were noted and R; values determined. Spots were excised, eluted in spectral
grade methanol, and UV spectral data obtained using standard procedures (Mabry
*Department of Botany, Howard University, Washington, D.C. 20059.
**Department of Botany, Rutgers University, New Brunswick, New Jersey 08903.
94 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
et al., 1970). Positive determinations of isolated flavonoid aglycones were done by
co-chromatographing them against authentic compounds. Quantitative scoring for
flavonoid content was done by comparative visual inspection of spot intensity and
size.
In the initial experiment, D. marginalis gametophytes were cultured for 30 days
and then harvested. Because of extreme crowding, most of the gametophytes
formed filaments rather than plates, and most bore a number of antheridia lat-
erally. Flavonoids were detected in these gametophytes, and they were the same
ones that occur in D. marginalis sporophytes (Table 1).
TABLE 1. IDENTIFICATION DATA FOR FLAVONOID GLYCOSIDES OF QUERCETIN AND
KAEMPFEROL FOUND IN DRYOPTERIS INTERMEDIA AND D. MARGINALIS GAMETO-
PHYTES AND SPOROPHYTES.
Quercetin (A) Kaempferol (B) Kaempferol (C)
CHROMATOGRAM SPOT APPEARANCE
UV Violet Violet Violet
UV/NH3 Yellow Yellow Yellow
CHROMATOGRAM SPOT Rr VALUES
TAB 0.50 0.60 0.63
HOAc 0.44 0.56 0.45
UV SPECTRAL DATA (A max., nm)
MeOH 256, 282, 306, 357 266, 292, 347 265, 302, 349
NaOMe Zils 325, 412 274, 324, 401 275, 325, 401
AICIs sige sp 362sh, 274, 302, 349, 395 273, 302, 348, 394
AIChs/HCI 268, 300sh, 358, 273, 302, 343, 392 273, 298, 343, 392
NaOAc 273, 415 273,302, 381 273, 301, 375
maoeres of inadequate material for spectrometric analysis, new cultures of D.
marginalis were Started, and cultures of D. intermedia were begun to determine if
they likewise produced the same flavonoids as D. intermedia sporophytes. After
three weeks, the cultures were harvested and assayed for flavonoids. No
flavonoids were detected on the chromatograms, and examination showed that the
ns Jas had not produced antheridia. Therefore, more gametophytes were
ema wea flavonoid content could be analyzed at three developmental
The last experiments showed that the same flavonoids are produced by the
gametophytes of these two species as are produced by their respective
sporophytes (Table 1). They are quercetin and kaempferol glycosides. (See Mabry
et al., 1970, for data comparisons and structural details.) Dryopteris intermedia
(Co rophytes and gametophytes produced two flavonoids: a quercetin glycoside
ompound A) and a kaempferol glycoside (Compound B). Dryopteris marginalis
produced these two compounds, a wa: é
(Compound C), , as well as an additional kaempferol glycoside
PETERSEN & FAIRBROTHERS: ANTHERIDUM INITIATION IN DRYOPTERIS 95
The experiments (Table 2) also show flavonoids to be absent (—) during the
pre-antheridial stage. They were first detected as faint spots (+) at the onset of
antheridium formation. Flavonoid concentration is much higher (+ +) in the post-
antheridium initiation stage than at the onset of antheridium formation, as re-
vealed by greater intensities of spot fluorescence. Antheridium formation and
TABLE 2. FLAVONOID CONTENT OF DRYOPTERIS GAMETOPHYTES AT THREE DE-
VELOPMENTAL STAGES.
Pre-antheridial Antheridial Post-antheridial
initiation initiation initiation
COMPOUNDS (A) 4B) 6 = (Cf @EAP 2ONBP AB) ay ay €C)
D. intermedia = = as ae ee i at oh a
D. marginalis - _ ue - iy =e aie a7. vet
— = compound not detected; + = compound detected but at a low concentration relative to ++.
flavonoid synthesis clearly are associated, but whether the two events are interre-
lated (cause and effect) or merely a coincidence remains to be determined. Al-
though nothing comparable has been discovered in flowering plants, Barber (1956)
identified a glucose-rhamnose glycoside of quercetin in staminate squash flowers
that was absent from the pistillate flowers.
We wish to acknowledge financial aid from NSF Grant GB-13202 and a Rutgers
Research Council Grant awarded to D. E. Fairbrothers.
LITERATURE CITED
BARBER, G. A. 1956. Flavonoids of staminate and pistillate squash flowers. Arch. Biochem.
Biophys. 64:401-411.
LAURENT, S. 1966. Contribution a I’étude des tanins et des autres substances phénoliques hy-
drosolubles, élaborées par les prothalles de filicinées. Thése Docteur I’ Université de Paris.
MABRY, T. J., K. R. MARKHAM, and M. B. THOMAS. 1970. The Systematic Identification of
Flav onoids. Springer-Verlag, ek York, Heidelberg, and Berlin.
PETERSEN, R. L. 1976. Chemical research in the genus Dryopteris Adanson: systematics, mor-
aoe and allelopathy. Ph.D. Thesis. Rutgers, The State University, New Brunswick,
Jer
New
SWAIN, T. 1974. | Biochestica) evolution in plants (Chapter II). In nai Florkin and E. H. Stotz, eds.
Comprehensive Biochemistry, vol. 29A. Elsevier, Amster
» and G. COOPER-DRIVER. 1973. Biochemical anes in the Filicopsida. Jn A. C.
Jermy, J. A. Crabbe, and B. A. Thomas, eds. The Phylogeny and Classification of the Ferns.
Bot. J. Linn. Soc. 67, Suppl. 1. Academic Press, New York and London.
96 AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 2 (1980)
Date of Publication of Sodiro’s
‘‘Sertula Florae Ecuadorensis”’
DAVID B. LELLINGER*
Several years ago, Morton (Amer. Fern J. 62:57-64. 1972) published on the
dates of original publication of the parts of Father Luis Sodiro’s ‘*Cry ptogamae
Vasculares Quitenses.”’ In that case, original publication in periodicals antedated
the reprint ‘‘Cryptogamae,”’ sometimes by several years. Sodiro’s **Sertula
Florae Ecuadorensis’’ exhibits similar, but shorter differences between original
and reprint publication dates. The portions concerning ferns first were published
in four parts in the ‘‘Anales de la Universidad Central’ in Quito, Ecuador. These
parts were reprinted a few months later with changes of pagination in two parts
under the general title ‘‘Sertula Florae Ecuadorensis.’’ The details are presented
in Table 1. Some pages apparently were not reset to produce the reprint, but
others obviously were. For instance, the bottom line of page 191 of the first part of
the ‘‘Anales’’ ends ‘‘linea-’’, but in the repaged reprint this is six lines from the
bottom and has been misspelled ‘‘ilnea-’’! This might lead one to think that the
reprint actually is a preprint. However, the ‘‘Anales’’ issue is dated January 1905
and the reprint 1905 with no indication of month, and so it is unlikely that the
reprint was issued before the ‘‘ Anales.”’
TABLE 1. DATES OF PUBLICATION OF “SERTULA FLORAE ECUADORENSIS”’ PARTS
CONTAINING FERNS.
Original publication in Repaged and reprint publication as
Anal. Univ. Central Sert
19(135)191-200. Jan 1905. I. Acrosticha. pp. 1-12, t. J. 1905.
22(158-159):21-30. Jan/Feb 1908. II. Pteridophyta. pp. 1-12(part). June 1908.
Ps Raa Mar 1908. II. Pteridophyta. pp. 12(part)-27(part). June 1908.
(161):161-176. Apr 1908. II. Pteridophyta. pp. 27(part)-42. June 1908.
Fortunately, not much time elapsed between the original and reprint publication
dates; probably no botanical names will be affected by the adoption of the earlier
dates. Many authors, including Christensen in his ‘‘Index Filicum,”’ have cited
pages and dates from the reprints, rather than from the original publication, as
should be done. Because the original publication is rare, the names and biblio-
graphic data for correct citation of new fern names and taxa are given in Table 2.
I am indebted to Dr. Dan Nicolson and Mr. James Zarrucci for calling this
easy to my attention and for obtaining xerocopies of the original ‘‘Anales”’
* , . ° .
U. S. Nat’L. Herbarium, Smithsonian Institution, Washington, DC 20560.
D. B. LELLINGER: SODIRO’S “SERTULA FLORAE ECUADORENSIS” 97
TABLE 2. NEW FERN SPECIES AND VARIETIES PUBLISHED BY SODIRO IN THE
‘**ANALES DE LA UNIVERSIDAD CENTRAL.”
Acrostichum actinolepis Sodiro, op. cit. 19(135):199. Jan 1905.
angamarcanum Sodiro, op. cit. 19(135):193. Jan 1905.
antisanae Sodiro, op. cit. 22(161):164. Apr 1908.
chodatii Sodiro, op. cit. 22(161):174. Apr 1908.
christii Sodiro, op. cit. 19 135):192. Jan 1905.
cinereum Sodiro, op. cit. 22(161):172. Apr 1908.
cladotrichum Sodiro, op. cit. 19(135):197. Jan 1905.
diversifolium Sodiro, op. cit. 22(161):166. Apr 1908.
ellipsoideum Sodiro, op. cit. 22(161):164. Apr 1908.
engleri Sodiro, op. cit. 22(161):167. Apr 1908.
fulvum Sodiro, op. cit. 22(161):168. Apr 1908.
gossypinum Sodiro, op. cit. 22(161):172. Apr 1908.
guamanianum Sodiro, op. cit. 22(161):169. Apr 1908.
hieronymi Sodiro, op. cit. 19(135):199. Jan 1905.
hikenii Sodiro, op. cit. 22(161):169. Apr 1908.
litanum Sodiro, op. cit. 19(135):198. Jan 1905.
longissimum Sodiro, op. cit. 19(135):191. Jan 1905.
molle Sodiro, op. cit. 22(161):171. Apr 1908.
muriculatum Sodiro, op. cit. 22(161):174. Apr 1908.
oleandropsis Sodiro, op. cit. 19(135):195. Jan 1905.
pangoanum Sodiro, op. cit. 19(135):194. Jan 1905.
pellucidum Sodiro, op cit. 19(135):195. Jan 1905.
pichinchae Sodiro, op. cit. 22(161):173. Apr 1908.
pruinosum Sodiro, op. cit. 22(161):166. Apr 1908.
pteropodum Sodiro, op. cit. 19(135):196. Jan 1905.
rupicola Sodiro, op. cit. 22(161):175. Apr 1908, as ‘*rupicolum.”’
subsessile Sodiro, op. cit.. 19(135):194. Jan 1905.
trichophorum Sodiro, op. cit. 19(135):198. Jan 1905.
urbani Sodiro, op. cit. 22(161):170. Apr 1908.
viscidulum Sodiro, op. cit. 22(161):165. Apr 1908.
Alsophila bilineata Sodiro, op. cit. 22(160):90. Mar 1908.
A. christii Sodiro, op. cit. 22(160):89. Mar 1908.
Asplenium anomalum Sodiro, op. cit. 22(160):95. Mar 1908.
chimboanum Sodiro, op. cit. 22(160):102. Mar 1908.
costale Sodiro, op. cit. 22(160):95. Mar 1908.
crassifolium Sodiro, op. cit. 22(160):97. Mar 1908.
heterolobum Sodiro, op. cit. 22(160):98. Mar 1908.
hieronymi Sodiro, op. cit. 22(160):99. Mar 1908.
humile Sodiro, op. cit. 22(160):99. Mar 1908.
melanosorum Sodiro, op. cit. 22(160):101. Mar 1908.
oxylobum Sodiro, op. cit. 22(160):96. Mar 1908.
procerum Sodiro, op. cit. 22(160):96. Mar 1908.
tungurahuae Sodiro, op. cit. 22(160):97. Mar 1908.
Mt a aM eg gl gd) okt at al all cl le aaa ot
Preer rr? ee
98 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
A. vesiculosum Sodiro, op. cit. 22(160):100. Mar 1908.
Cyathea asperata Sodiro, op. cit. 22(158-159):27. Jan/Feb 1908.
asperata var. minor Sodiro, op. cit. 22(158-159):28. Jan/Feb 1908.
brachypoda Sodiro, op. cit. 22(158-159):26. Jan/Feb 1908.
canescens Sodiro, op. cit. 22(158-159):22. Jan/Feb 1908.
furfuracea Sodiro, op. cit. 22(158-159):25. Jan/Feb 1908.
muriculata Sodiro, op. cit. 22(158-159):28. Jan/Feb 1908.
nitens Sodiro, op. cit. 22(158-159):21. Jan/Feb 1908.
ochroleuca Sodiro, op. cit. 22(158-159):29. Jan/Feb 1908.
oxyacantha Sodiro, op. cit. 22(158-159):24. Jan/Feb 1908.
parvifolia Sodiro, op. cit. 22(158-159):25. Jan/Feb 1908.
subinermis Sodiro, op. cit. 22(158-159):28. Jan/Feb 1908.
. tungurahuae Sodiro, op. cit. 22(158-159):30. Jan/Feb 1908.
Nephrodium cinereum Sodiro, op. cit. 22(160):103. Mar 1908.
N. cinereum var. intermedium, Sodiro, op. cit. 22(160):104. Mar 1908.
N. longipilosum Sodiro, op. cit. 22(160):103. Mar 1908.
Polypodium scutulatum Sodiro, op. cit. 22(161):163. Apr 1908.
Pteris aspidioides Sodiro, op. cit. 22(160):91. Mar 1908.
biternata Sodiro, op. cit. 22(160):94. Mar 1908.
esmeraldensis Sodiro, op. cit. 22(160):92. Mar 1908.
falcata Sodiro, op. cit. 22(160):93. Mar 1908.
procera Sodiro, op. cit. 22(160):94. Mar 1908.
rigida Sodiro, op. cit. 22(160):92. Mar 1908.
rimbachii Sodiro, op. cit. 22(160):91. Mar 1908.
robusta Sodiro, op. cit. 22(160):93. Mar 1908.
aanaanaaaaa
MO OO TO
pe REVIEW
‘TAXONOMY OF THELYPTERIS SUBGENUS STEIROPTERIS, INCLUD-
ING GLAPHYROPTERIS (PTERIDOPHYTA), by Alan R. Smith, Univ.
Calif. Publ. Bot. 76:1-38, t. 1-4. 1980.—Among the large neotropical fern genera,
none 1s more complex than Thelypteris. Even though subg. Steiropteris is but a
small portion of the genus (22 species, some divided into varieties), this monograph
is very welcome. Over half the taxa are new species or required a new name or
combination, which is a measure of the confusion which the author has resolved.
Spores and some details of morphology are illustrated with photographs, and
distributions are shown by means of maps. The author has seen many type
specimens, and so the synonymies are doubtless accurate. A list of exsiccatae is
included but, unfortunately, an index to scientific namies is not. The work is
available for $5.00 from University of California Press, 2223 Fulton St., Berkeley,
CA 94720.—D.B.L.
AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 3 (1980) 99
Reproductive Biology and Gametophyte
Morphology of New World
Populations of Acrostichum aureum
ROBERT M. LLOYD*
The majority of homosporous ferns are characterized by a life-cycle which
permits the production of a genetically homozygous zygote following self-
fertilization of a single gametophyte (intragametophytic selfing). This homozygos-
ity leads to the expression of recessive deleterious or lethal genes (genetic load, as
defined here) present in the genotype, unless this expression is buffered by the
polyploid system. Sporophytes expressing such genes will be eliminated rapidly
and the spore genotypes produced individually by the remaining viable
sporophytes will be genetically uniform, barring mutation and meiotic ir-
regularities (e.g., homeologous pairing; Klekowski, 1979). In species which regu-
larly undergo selfing, genetic load will be absent or will be expressed at low levels
(Klekowski, 1979). Thus, analysis of genotypes for genetic load allows for an
estimate of the genetic variability in a population.
The fern life-cycle also permits reproduction which is genetically analogous to
inbreeding and outbreeding in angiosperms, the latter facilitating the storage of
recessive deleterious and lethal genes (Wallace, 1970). Although none of the
above patterns of reproduction are mutually exclusive, work of the past decade
has led to the hypothesis that specific morphological and developmental features
of the gametophyte generation will increase the probability of selfing or crossing
(intergametophytic mating) and that these probabilities can be correlated with
estimates of heterozygosity in the form of genetic load (Lloyd, 1974). However,
more recent work with Ceratopteris (Lloyd & Warne, 1978) and Acrostichum
(Lloyd & Gregg, 1975) suggests that the past hypotheses are insufficient to explain
the genetic diversity expressed in these species and that other factors are in-
volved. This paper summarizes our most recent work on the gametophyte mor-
phology, reproductive biology, and genetic diversity in a number of populations of
Acrostichum aureum distributed from Florida to the northern coast of South
America and attempts to circumscribe the current problems in this field.
The genus Acrostichum consists of at least three species: A. danaeifolium
Langsd. & Fisch., a New World endemic which is widely distributed in fresh
water and slightly saline swamps (Adams & Tomlinson, 1979); A. aureum Ais
circumtropical in distribution and usually most abundant in mangrove habitats
where it can withstand partial tidal immersion (Holttum, 1954; Small, 1938); and
A. speciosum Willd., a species of tropical Asia and Australia which is abundant in
mangroves through Malaya in areas frequently inundated by tides (Holttum,
1954). These types of habitats are extreme; few species of plants have evolved the
necessary physiological and morphological features to successfully colonize them.
Previous work on the gametophyte generation in Acrostichum includes mor-
*Department of Botany, Ohio University, Athens, OH 45701.
100 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
FIGS. 1-3. Spores of A. aureum. FIG. 1. Papua, Brass 518 (UC), x 1000. FIG. 2. Florida, Curtiss 5463
(UC), x 10000. FIG. 3 3. Papua, Brass 518 (UC), x 8000.
R. M. LLOYD: REPRODUCTIVE BIOLOGY OF ACROSTICHUM 101
phological studies of A. speciosum by Stokey & Atkinson (1952) and a study of the
morphology and reproductive biology of Mexican populations of A. danaeifolium
by Lloyd & Gregg (1975).
MATERIALS AND METHODS
Spores of A. aureum were collected from 39 plants from eight populations as
follows:
Culture No. 146: 1.1 mi W of U. S. Highway 41 on State Highway 92, Collier Co., Florida. 148: 2.1
mi W of Westlake on State Highway 27, Everglades National Park, Dade Co., Florida. 150: 30.5 mi SW
of entrance station on State Highway 27 at road to Westlake, Everglades National Park, Monroe Co.,
Florida. 190: 0.25 mi E of Negril on road to Savana la Mar, Westmoreland Parish, Jamaica. /9/: Mile
post 57, 57 mi E of Georgetown on Public Road East, Guyana. 192; 8 km N of Governor’s Palace,
Parimaribo, near end of road to Leonsburg, Suriname. /93: 0.2 mi from road to Colon on road to Coco
Solo, Canal Zone, Panama. 194: Lowland area near Pacific Ocean at N end of the Bridge of the
Americas, Canal Zone, Panama.
Spores were sown and gametophytes grown aseptically on sterile inorganic
nutrient medium solidified with 1% agar (for composition see Klekowski, 1969) in
100 x 15 mm petri dishes. Gametophytes were grown under continuous illumina-
tion by fluorescent and incandescent lamps at an intensity of 210 to 290 ft-c at
temperatures of 19-24° C. Prothallial morphology was studied using living mate-
rial as well as that mounted in Hoyer’s medium mixed with acetocarmine. Spore
sizes were determined by mounting spores in diaphane and calculating their
equatorial diameters with a calibrated ocular micrometer. Other methods utilized
in specific experiments are described below.
Spores observed by the scanning electron microscope were dry-mounted on
double-stick tape, coated with gold ca. 10 nm thick, and observed at 20 kv ac-
celerating voltage with a Hitachi HHS-2R scanning electron microscope.
RESULTS
Spores of A. aureum are tetrahedral and are (37)45—-72 (mean + s.d. = $6.7 =
4.58) um in diameter. The spore surface is minutely tuberculate (Fig. 1). The
tubercle-like structures on the surface bear varying numbers of projecting papil-
lae. Spores from plants from Papua (Figs. 1, 3) and Fiji exhibit numerous but
somewhat irregularly shaped and oriented papillae. Spores from Florida plants
(Fig. 2) and other Fijian plants appear to have more numerous papillae as well as
other types of superficial deposits. In contrast, spores examined from plants from
Australia and Trinidad are tuberculate but appear either to lack papillae or to have
thickened superficial deposits which more or less obscure their presence. Spores
examined of A. speciosum from Papua exhibit surface features highly similar to
those from Florida plants of A. aureum. eS
Spore germination is usually initiated by the emergence of a rhizoid five days
following sowing. Gametophytes produce a one-dimensional filament up to 10
cells in length before initiation of two-dimensional growth (Fig. 4). In some in-
Stances, cells near the base of the filament will divide, producing a second one-
dimensional filament (Fig. 5).
102 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
ait
wd
Tt)
ee
*
;
eee
r)
ee
a ee?
QS
ta
ec
c
|
cc
ce <
/
FIGS. 4-11. Stages in gametophyte development and sexually mature gametophytes ot A. aureum.
FIG. 4. One-dimensional filament with initiation of two-dimensional growth in basal cell, 185 wm long,
days after sowing. FIG. 10. Y ancia. 4.0 mm long,
= days after sowing. FIG. 11. Mature gametophyte showing sequential pattern of archegonial-
antheridial-archegonial production and branched meristem, 7.5 mm long, 96 days after sowing.
R. M. LLOYD: REPRODUCTIVE BIOLOGY OF ACROSTICHUM 103
Two-dimensional growth is initiated by a longitudinal division in a central or
more basal cell of the one-dimensional filament. Further longitudinal divisions in
the filament may follow one or more patterns: cells throughout the filament,
except for the terminal and basal cells, may divide longitudinally and produce a
two-dimensional filament two cells wide (Fig. 6); less frequently, central cells of
the 1-dimensional filament may divide sequentially, producing an area up to four
or more cells in width before further divisions occur in the more basal and terminal
cells of the filament. In both pathways, 2-dimensional growth ultimately results in
a broadly linear or spatulate gametophyte four to six cells wide. Further divisions
of cells along one of the lateral margins of these prothalli will produce a lateral
meristematic region located near the basal region of the gametophyte. Subsequent
growth produces an asymmetrical ovate prothallus with different sized wings (Fig.
8). In older gametophytes, growth frequently produces more or less symmetrical
wing tissue on both sides of the lateral meristem, resulting in a mature prothallus
which appears to have an apical meristematic notch (Figs. 9 and /0)). This notch
area remains shallow in most prothalli observed; in some, however, the meristem
exceeds the wing tissue and no notch is evident. In some older gametophytes, the
meristematic region becomes quite broad, and, rarely, may divide into two sepa-
rate regions (Fig. 11) with non-meristematic tissue between.
Gametangia initiation in culture was rapid. All cultures, except 193-K!, exhib-
ited a female to hermaphroditic gametangial sequence of development (Fig. 8)
with the exception of occasional gametophytes which precociously initiated an-
theridia (Fig. 7). Of the 20 cultures studied in detail, seven produced some male
gametophytes, but the percentage of such gametophytes in culture (except 193-K)
was less than 6.0 (Tables ] and 2). In all cases these prothalli rapidly became
hermaphroditic.
The length of the unisexual female gametophytic stage varied from culture to
culture. In one culture (190-D), hermaphroditic prothalli were produced simulta-
neously with female prothalli. In other cultures (150-I, 193-E, 193-L),
gametophytes remained unisexual and female throughout the culture period. In
the remaining cultures, hermaphroditic prothalli were produced (2)6—28 (mean =
s.d. = 15.6 + 5.8) days following appearance of female prothalli (Table 1).
Gametangial sequences of individual gametophytes are diverse, but the vast
majority of gametophytes exhibited a female to hermaphroditic sequence. Arche-
gonia were initiated on the cushion immediately behind the young lateral meristem
and were produced continuously until the gametophytes were fully cordate witha
pronounced elongate cushion with numerous senescent gametangia (Fig. 8). An-
theridia were initiated on wing tissue near the apical notch, initially along the
margin of the cushion and later outward toward the wing margins (Figs. 9 and / 0).
Fully mature hermaphroditic gametophytes exhibited antheridia covering both
not been observed in the distal portions of the prothallus. In culture 190-A, sam-
phyte
‘Here and elsewhere in this paper the letter following the culture number designates a gameto
population originating from a specific sporophyte.
104 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
ples of gametophytes 46 days after sowing indicated that some of them had pro-
duced up to 90 antheridial initials. These prothalli exhibited up to 28 or more
senescent, 8 mature, and 50 immature archegonia. Six days later (52 days follow-
ing sowing), fully mature hermaphroditic prothalli were present, exhibiting up to
60 senescent, 10 mature, and 35 immature archegonia, and over 325 antheridia per
wing of which about 12% contained mature spermatozoids.
Gametophytes which were initially male produced only one or two antheridia
prior to the initiation of archegonia. In most cases, following maturation and
dehiscence of these antheridia, gametophytes became functionally unisexual and
female and their subsequent ontogeny paralleled that described above.
TABLE 1. DAYS FROM SOWING TO APPEARANCE OF GAMETOPHYTE-TYPES IN
CULTURES OF A. AUREUM.
Culture Days to appearance of
number Density/cm? Male Female Hermaphroditic
146-C 27.7 28 2 38
146-G 6.5 35 29 (35)50
146-L 6.9 : 50
146-M 21:3 28 46
148-B 21 28 44
150-A 18.0 51 3 (35)56
150-B 29.7 44
150-C 28.2 56
150-E 3.9 } 50
150-I 8.1 3 -!
150-K 14.1 3 44
190-A 6.1 ] 49
se i | :
2. 37 : 6
190-F 31.5 59 oe
193-A 17.6 28 56
193-B 98.3 44 ) _*
re 45.7 29 3
5 191
193-L | id : “er
‘sampled up to 76 days
?sampled up to 85 days
3sampled up to 90 days
Some of the gametophytes expressed sequential patterns of functional unisexu-
ality. In 146-C, about 15% of the sampled gametophytes expressed a sequence of
archegonial initiation, maturation, and senescence, followed by initiation and
maturation of antheridia. Other gametophytes appear to have gone through func-
tional stages in sexual ontogeny from archegoniate to hermaphroditic to an-
theridiate to archegoniate. This sequence was noted in several older prothalli in
193-A (Fig. 11) and in one gametophyte of 150-K. In gametophytes producing
proliferations near the base, such as those with two 1-dimensional filaments aris-
ing from the single basal cell, some of the proliferations were covered with an-
theridia, whereas others were only archegoniate.
Culture 193-K was unique among those studied in its expression of large num-
bers of male prothalli (Table 2). The initial ratio of male to female prothalli,
excluding asexual prothalli, was 1:1. As the culture developed, male gameto-
R. M. LLOYD: REPRODUCTIVE BIOLOGY OF ACROSTICHUM 105
phytes increased in frequency to a 3:1 ratio; this; in turn, was followed by an
increase in both female and hermaphroditic gametophytes. It is of interest to note
that at the end of the observation period, there was a 1.1:1 ratio of male and
hermaphroditic prothalli to female prothalli. Male gametophytes in this culture
frequently were highly elongate and irregularly formed. Many of them initiated
antheridia in the early ontogenetic stages following the attainment of a
2-dimensional morphology. In contrast, female prothalli were larger and appeared
to be similar in all respects to the female prothalli of the other cultures.
TABLE 2. SEXUAL ONTOGENY IN AGAR CULTURES OF A. AUREUM.
‘Sexual expression (%)-
Days from sowing Neuter Male Female Hermaphroditic
Culture No. 146-C
21 97.5 25
28 48.6 5.4 45.9
35 26.4 74.6
38 8.7 78.3 13.0
44 10.5 2.6 68.4 18.0
47 : J 40.0 34.2
Culture No. 190-A
33 00.0
AY 64.7 bo
43 22.2 77.8
49 29.4 47.1 235
52 28.6 31.4 40.0
55 11.1 40.7 48.1
58 6.7 33.3 60.1
76 14.3 85.7
87 100.0
Culture No. 193-K
30-33 8.3 29. 1.9
39-45 217 56.7 19.1 25
48-54 15.9 32.9 50.0 1.2
63 24.4 46.3 29.3
Culture No. 193-L
30 28.6
40 58.3 41.7
42 38.5 61.5
100.0
90 100.0
Parameters of gametophyte morphology and ontogeny discussed above suggest
that intergametophytic mating should be prevalent in gametophyte populations of
Acrostichum aureum. Specific factors exhibited by the gametophyte generation
which increase the probability for intergametophyte mating include the female to
hermaphroditic gametangial sequence in most gametophytes studied, the dioeci-
ous condition expressed in some populations, and the sequential functional uni-
sexuality expressed in some gametophytes of some populations. Additional sup-
106 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
port for this assessment comes from observations on sporophyte production in
composite cultures and in timing of the appearance of sporophytes in isolate vs.
composite cultures. For example, in culture 146-M, after 83 days 90.9% of the
prothalli were unisexual female and 9.1% were hermaphroditic. Examination of
sampled prothalli indicated that all of the unisexual female gametophytes had
produced one or two young embryos, which would only be possible through
intergametophytic mating. In addition, in nearly all cultures sampled sporophyte
production in composite culture occurred between 17 and 31 days earlier than in
isolate culture. As the gametangial sequence is from female to hermaphroditic in
these prothalli, in composite cultures spermatozoids produced by the early her-
maphroditic gametophytes will fertilize many of the unisexual female prothalli. In
contrast, in isolate culture, each of the gametophytes must become hermaphrodi-
tic prior to sporophyte production. This sequence of events has been well
documented in studies on Ceratopteris by Klekowski (1970a).
TABLE 3. FREQUENCY OF DELETERIOUS SPOROPHYTIC GENOTYPES IN IN-
TRAGAMETOPHYTICALLY SELFED, ISOLATED HERMAPHRODITIC GAMETOPHYTES
OFA. AUREUM.
Number No. (%) No. (%) No. (%) late No. (%)
Culture tested No. (%) zygotic embryonic sporophytic lea
number prothalli normal lethals lethals lethals lethals
46-N 19 18(94.7) 1(5
148-B 20 19(95.0) 1(5.0)
150-A 20 19(95.0) 1(5.0)
150-B 20 17(85 .0) 1(5.0) 2(10.0)
150-I 18 17(94.4) 1(5.6)
190-F 19 15(78.9) 3(15.8) 1(5.3)
191-B 19 17(89.5) 1(5.2) 1(5.2)
192-A 40 39(97.5) 1(2.5)
193-C 20 17(85.0) 1(5.0) 2(10.0)
193-E 20 11(55.0) 9(45.0)
193-F 8 7(87.5) 1(12.5)
193-M 20 19(95.0) 1(5.0)
All others:! 511 $11(100.0)
Totals: 754 726(96.3) 20(2.65) 2(0.26) 1(0.13) 5(0.66)
‘Includes 27 cultures: 146-C, 146-G, 146-I, 146-J, 146-L, 146-M, 146-O, 150-C, 150-E, 150-H, 150-K,
Ee ee oe OS, 193-D, 193-G, 193-H, 193-1, 193-K, 193-L,
Additional information relative to reproductive biology can be obtained by
analyzing frequency of deleterious or lethal genes (genetic load) expressed in
sporophytes (Klekowski, 1979). To analyze genetic load in A. aureum, from 20 to
40 gametophytes per sporophyte (= a gametophyte family), prior to the attainment
of sexual maturity, were selected at random from stock cultures and individually
isolated in 60 x 20 mm petri dishes containing nutrient agar. Following growth,
cultures were watered twice weekly to facilitate fertilization, and the resultant
sporophytes were allowed to develop to the third frond stage. Results of these
studies are given in Table 3.
R. M. LLOYD: REPRODUCTIVE BIOLOGY OF ACROSTICHUM 107
Genetic load was determined as the percentage: of the hermaphroditic
gametophytes per gametophyte family which failed to yield normal sporophytes.
Families exhibiting genetic load in a portion of the gametophytes tested are con-
sidered to be expressing heterozygosity in their gametophytic genotypes. Expres-
sions of genetic load were in the form of zygotic lethals (in 2.65% of the 754
gametophytes tested), embryonic lethals (in 0.26%), late sporophytic lethals (in
0.13%) and leaky lethals (in 0.66%) (see Klekowski, 1970b, 1979, for complete
discussion of these genetic expressions). It is significant to note that 96.3% of the
tested prothalli did not exhibit any deleterious or lethal genotypes. Of the 39
sporophytes tested, 27 (69.2%) were devoid of genetic load (Table 4). In the 12
sporophytes expressing load, it varied from 5.0% (cultures 148-B, 150-A, 193-M)
to 45.0% (culture 193-E). The mean genetic load for all plants tested was 3.7%.
TABLE 4. GENETIC LOAD IN A. AUREUM RELATIVE TO SIZE AND LOCATION OF THE
POPULATION.
Population oe
Size (est. Range (x) % No. plants No. (%) plants
number Location no. plants) genetic load tested with genetic load
193 Panama 3000 0-45.0 (5.96) 13 4(30.8)
190 Jamaica 1000 021.1 (4.22) > 1(20.0)
191 Guyana 400 0-10.4 (5.2) 2 1(50.0)
150 Florida 75-100 0-15.0 (3.65) 7 3(42.8)
146 Florida 25-50 0-5.3 (0.66) 8 1(12.5)
148 Florida 20 5.0 1 1(100.0)
194 Panama 15 0 1 000.0)
192 Surinam 8 0-2.5 (1.25) 2 1(50.0)
Total: 0-45.0 (3.24) 39 12(30.8)
Although the number of plants tested from each population is insufficient for
Statistical comparison, it is of interest to note that those sporophytes exhibiting
the higher genetic load values are found in the larger populations and that the
small populations (with 50 or fewer individuals) have very low levels of recessive
deleterious or lethal genes (Table 4).
Leaky lethal expression (Klekowski, 1970b) was noted in three gametophyte
families. In 150-B, normal sporophytes appeared on the two prothalli 165 days
after sowing and 38 days following normal sporophyte production of the remaining
prothalli tested. Each of these two prothalli exhibited several abortive embryos,
indicating that previous selfing had occurred involving lethal genetic combina-
tions. In 193-M, the first sporophyte which appeared was abnormal and exhibited
a long, cylindrical, tubular growth with ruffled margins. Subsequent sporophytes
from other fertilizations produced normal fronds. ;
Apomictic proliferations were noted on only one gametophyte in 147-L. Ninety
days after sowing, this prothallus proliferated a blade of tissue bearing rhizoids on
one margin and small epidermal cells similar to those found on young
sporophytes. Irregularly organized vascular tissue was present near the base of
this blade, but no roots or stomata were noted.
108 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
DISCUSSION
Gametophyte Morphology.—The gametophyte morphology and ontogeny of A.
aureum is remarkably similar to that of A. danaeifolium (Lloyd & Gregg, 1975)
and agrees in most respects with that of A. speciosum Willd. (Stokey & Atkinson,
1952). Spores of A. aureum are almost identical in size and shape to those of A.
danaeifolium; however, there are minute differences in spore surface markings,
especially the more pronounced tuberculate pattern exhibited by A. aureum.
Other gametophyte features which are qualitatively similar between the two
species are formation of the 1-dimensional filament, the lateral meristematic re-
gion of the 2-dimensional prothallus, the relatively shallow apical notch region
(however, protruding beyond the wing tissue in some prothalli of A. aureum), the
female to hermaphroditic gametangial sequence, and the sexual expression in
gametophyte families grown in composite culture.
The major difference between gametophytes of the two species is the distribu-
tion of antheridia, which are mostly restricted to the apical wing and meristem
region of A. aureum, but also are found in more basal regions along the cushion
margins and among the rhizoids in A. danaeifolium. In addition, the sequential
production of archegonia-antheridia-archegonia in some prothalli of A. aureum is
unknown in the other species.
It is apparent from both sporophyte and gametophyte studies that these two
species are closely related. Further evidence in support of this is their ability to
freely hybridize in culture and to produce normal viable F: sporophytes, although
these sporophytes have not yet been grown to maturity to measure chromosome
homology (Lloyd, unpubl.).
Reproductive Biology.—Sex ontogeny in most cultures of Acrostichum aureum
sampled in this study is female to hermaphroditic or initially dioecious. The length
of the unisexual stage prior to the attainment of bisexuality is sufficient to facili-
tate intergametophytic mating. The facility for such mating is also evidenced in
culture by the rapidity of embryo formation in unisexual prothalli following the
initiation of antheridia on just one gametophyte in a composite culture. Thus, the
gametophytic developmental pathway must be considered as one which has a
higher probability of intergametophytic mating than of intragametophytic selfing.
However, correlative heterozygosity in the form of genetic load is insufficient in
naturally occurring sporophytes to suggest that outbreeding is a normal occul-
rence. For example, of the tested plants 69% exhibited no heterozygosity for
recessive deleterious genes and 15% exhibited such genes in less than 6% of the
genotypes sampled. As intergametophytic mating is strongly suggested by the
culture experiments, if the assumption is made that these plants are genetically
homozygous due to the lack of genetic load expression, other factors must be
superimposed upon the hypothesized mating system which are more significant in
determining the genetic composition of the populations as a whole.
First and foremost, the culture methodology as used in these experiments may
be insufficient to document with accuracy the gametangial sequences as they are
realized in nature. In parallel experiments on A. danaeifolium, gametophytes
R. M. LLOYD: REPRODUCTIVE BIOLOGY OF ACROSTICHUM 109
grown on soil exhibit greater antheridial production (Lloyd & Gregg, 1975). Al-
though some of these gametophytes undergo a male to hermaphroditic gametan-
gial ontogeny, dioecism in cultures was still highly prevalent, suggesting that soil
grown gametophyte populations in nature would have higher probabilities of in-
tergametophytic mating. As gametophytic ontogenies on agar cultures of A. au-
reum and A. danaeifolium are highly similar, it is reasonable to assume that the
gametophytes of A. aureum would present similar responses to soil culture. How-
ever, the habitat of A. aureum is at least partially inundated by tides, suggesting
that the soil component for gametophyte populations will contain higher levels of
salts. Brief experiments by Stokey & Atkinson (1952) using dilute sea water as
part of the culture medium induced restricted growth of gametophytes of A.
speciosum. This type of reduced growth under less than optimal conditions fre-
quently leads to the initial production of antheridia and can prevent formation of
viable archegonia (Page, 1979). Thus, it is possible that the gametophytic on-
togenies in the culture experiments reported here do not represent gametophytic
ontogenies as realized in nature.
Other factors which undoubtedly have a significant influence are population
size, spore output per plant, the influence of the specific aquatic habitat, and the
genetic system. It is of interest to note that the highest levels of genetic load were
found in the larger populations, suggesting that the frequency and success of
recombinants increases with number of individuals as well as age of the popula-
tion. As spore production by each individual of A. aureum is massive, it is proba-
ble that inbreeding (in this case, intergametophytic selfing) will occur until such
time as there is sufficient spore intermixing to increase the likelihood of outbreed-
ing.
The influence of the aquatic habitat may play an important role in the selection
of specific genotypes, perhaps perpetuated by intragametophytic selfing. It is
significant to note that work to date on other aquatic species, including Acro-
stichum danaeifolium, Ceratopteris thalictroides and C. pteridoides, has provided
highly similar results. These species are all characterized by a gametophyte on-
togeny which favors intergametophytic mating (including an antheridogen in
Ceratopteris spp.), but the vast majority of individuals tested express little or no
heterozygosity in the form of genetic load. In this regard, Baker (1965) cites
seashores and the margins of salt marshes as open habitats where species which
are inbreeding with ‘‘general purpose genotypes’? may be advantageous. Angio-
sperms which occupy these open and disturbed types of habitats are generally
found to be autogamous or apomictic and so are unable to build up recombinants
in the population rapidly. :
Lastly, the genetic system of pteridophytes must be considered. We still have
little understanding of the polyploid system and the maintenance and expression
of heterozygosity in these organisms. It is possible that most of them are highly
heterozygous and that genetic load is effectively screened from expression. If so,
our current methodology for analysis for heterozygosity is insufficient.
It is obvious from these studies that we have little understanding of fern mating
systems as they operate in nature and much further work, especially that oriented
110 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
toward the genetic system and natural populations of gametophytes and
sporophytes, is required before we will be able to circumscribe adequately these
phenomena as they operate in nature.
This work has been supported by National Science Foundation Grants Nos.
GB-36923, BMS 75-07191, and DEB 79-05079. I would like to thank T. R. Warne,
D. Buckley, and S. Buckley for their assistance in the laboratory.
LITERATURE CITED
ADAMS, D. C. and P. H. TOMLINSON. 1979. Acrostichum in Florida. Amer. Fern J. 69:42-46.
BAKER, H. G. 1965. Characteristics and modes of origins of weeds. Jn H. Baker and G. L. Stebbins,
eds. The Genetics of Colonizing Species. Academic Press, New York.
HOLTTUM, R. E. 1954. Flora of Malaya, Vol. II. Ferns of Malaya. Gov't. Printing Office, Singapore.
KLEKOWSKI, E. J., JR. 1969. Reproductive biology of the Pteridophyta. III. A study of the
Blechnaceae. Bot. J. Linn. Soc. 62:361-377.
. 1970a. Reproductive biology of the Pteridophyta. IV. An experimental study of mating
selene in Ceratopteris thalictroides (L.) Brongn. Bot. J. Linn. Soc. 63:153-169.
. 1970b. Populational and genetic studies of a homosporous fern—Osmunda regalis. Amer. a
Bot. 56:1122-1138.
. 1979. The genetics and reproductive biology of ferns. In A. F. Dyer, ed. The Experimental
Biology of Ferns. Academic Press, London
LLOYD, R. M. 1974. Reproductive biokiay and évohiion in the Pteridophyta. Ann. Missouri Bot.
Gard. 61:318-331.
, and T. L. GREGG. 1975. Reproductive =a and gametophyte morphology of Acro-
stichum danaeifolium from Mexico. Amer. Fern J. 65:105—120.
, and T. R. WARNE. 1978. The absence of aid load in a morphologically variable sexual
cee a cigar sae aia ce ponpanen “ie Bot. 3:20-36.
PAGE, C. N. 1979. Experimental as of _ ecology. In A. F. Dyer, ed. The Experimental
okey of Ferns. Academic eg Lon
SMALL, J. K. 1938. Ferns of the Southeastern ee Reprint ed., 1964. Hafner, New York.
STOKEY, A. G. and L. R. ATKINSON. 1952. The gametophyte of Acrostichum speciosum Willd.
Phytomorphology 2:105-11
WALLACE, B. 1970. Genetic Load, Its Biological and Conceptual Aspects. Prentice-Hall, En-
glewood Cliffs, NJ.
AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 3 (1980) 111
SHORTER NOTES
DIPLAZIUM JAPONICUM NEW TO ALABAMA.—In the summer of 1977
while I was a student at Auburn University, J. I. Glick, another student, asked me to
confirm his identification of a fern frond he had collected from a plant growing in a
steep-sided, wooded ditch on the Auburn campus in an area that had been allowed to
remain wild by the university groundskeepers. Glick had identified the frond as
Athyrium thelypteroides. However, Auburn is in Lee County of east-central Ala-
bama, far south of the range of that fern, and the fern did not look exactly like A.
thelypteroides. 1 identified the frond as Diplazium japonicum (Thunb.) Bedd., a
native of eastern Asia (Glick s.n., Short 979, both AUA). According to Wherry
(Southern Fern Guide, 1964), this fern had been found previously in one Florida
locality. Wherry was doubtful whether the fern was cultivated locally and did not
know the source of the spores which produced the plants. The species is not known
to be in cultivation in the Auburn area, and the source of these plants is equally
unknown. To my knowledge, D. japonicum has not been reported from any other
localities in the southeastern United States. The Auburn population consisted of two
mature, spore-bearing plants and several juveniles. The plants seem to be estab-
lished well enough to be considered naturalized. The winters previous to and
subsequent to the plants’ discovery were among the most severe on record, but they
had no adverse effect on the plants. Other ferns found in the ditch include Asplenium
platyneuron and the naturalized Lygodium japonicum, Pteris multifida, and Thely-
pteris torresiana.—John W. Short, 905 McKinley Ave., Auburn, AL 36830.
MOTHS AND FERNS.—In a previous paper (Amer. Fern J. 61:166-170. 1971), 1
reported on a nymphalid-like moth that oviposits on C. 'yathea holdridgeana in such
a fashion that the eggs mimic the immature sori of the fern. Recently, I have found
the larvae of a microlepidopteron predating the laminar tissue of Cnemidaria
mutica, C. choricarpa, and Sphaeropteris brunei, of the family Cyatheaceae. In
their last stages, the caterpillars weave cocoons with silk, spores, and sporangia.
The adults obtained from the three ferns are the same species of moth, a species
yet to be determined. Another moth lays eggs on the fronds of Botrychium dissec-
tum. The larval stages feed on it and later spin a cocoon by sewing together two
segments or pinnules.
The most interesting relationship between moths and ferns I have encountered
So far is that of Hymenophyllum myriocarpum and a microlepidopteron whose
larvae feed on the filmy fern and pupate in a case made up of the folded segments,
which then resemble mature involucres in their size, color, and position. It is
surprising to find that the insects only use the basal, lower, and middle pinnae of
the fern fronds to build their cocoons, perhaps to guarantee themselves an appro-
priate relative humidity among the mosses or a more efficient camouflage. Cer-
tainly, a careful survey of tropical ferns will reveal that they are not so impervious
to insect attack as they commonly are thought to be.—Luis D. Gomez P., Her-
bario Nacional, Museo Nacional de Costa Rica, Apartado 749, San Jose, Costa
ica.
112 AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 2 (1980)
THREE ADDITIONS TO THE PTERIDOPHYTE FLORA OF ESCAMBIA
COUNTY, FLORIDA. — In 1978 a number of students from the University of
West Florida conducted a field survey of the pteridophytes of Escambia County,
Florida, concentrating on the botanically largely unexplored northern part. Just
north of where Florida Highway 4 crosses Canoe Creek (R31W, TSN, Sec. 8),
eight fern species were found along the west bank, including two which we had not
observed previously in the county: Thelypteris torresiana (Gaud.) Alston (Burk-
halter & Booker 5844, 5912) and Athyrium asplenioides (Michx.) A. A. Eaton
(Burkhalter & Booker 5845, 5913). Vouchers have been deposited at the Univer-
sity of Florida, Gainesville (FLAS), and the University of West Florida, Pen-
sacola (UWFP). Dr. Daniel B. Ward (pers. comm., 26 Feb. 1979) verified that
these species had not been collected previously in Escambia County.
The collection site is an open area with mostly clay and sand soil. In addition to
a number of weedy flowering plants like Carex spp. and Boehmeria cylindrica, the
following ferns were found: Asplenium platyneuron, Lygodium japonicum, Os-
munda regalis, O. cinnamomea, Thelypteris normalis, and Woodwardia areolata.
Thelypteris torresiana and A. asplenioides are not found in the shaded swamp
forest understory somewhat to the north of the collection site; they were, how-
ever, observed at a few other scattered locations in northern Escambia County.
Later, a small, spontaneous colony of Nephrolepis cordifolia (L.) Presl was
discovered in downtown Pensacola on an old, brick building at the northeast
corner of Palafox and Main Streets. The plants were securely anchored in the
brickwork about seven feet above the sidewalk, and were situated below the
broken end of a raingutter downspout, from which they received plentiful water.
This is not an unusual habitat for certain ferns in urban areas like New Orleans and
Mobile, and the colony appeared to be quite old and healthy. Clifton Nauman
(pers. comm., 31 Aug 1978) commented that the specimen (Burkhalter 5919,
UWFP) constituted a new record for Escambia County.—James R. Burkhalter,
3703A W. Brainerd St., Pensacola, FL 32504.
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QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY
A Range Extension for Dryopteris filix-mas ERWIN F. EVERT 113
Differential Germination of Fern and Moss Spores in Response
to Mercuric Chloride RAYMOND L. PETERSEN and PATRICK C. FRANCIS |!
a
Differences in the Apparent Permeability of Spore Walls and
Prothallial Cell Walls in Onoclea sensibilis JOHN H. MILLER 119
Allelopathy and Autotoxicity in depen Easter d
North American Ferns LLIAM a MUNTHER and DAVID E. FAIRBROTHERS 124
Shorter Notes: Sandstone Rock Crevices, an Exceptional New Habitat
for Thelypteris simulata; A Second Alabama Locality for 136
the Hart’s-tongue is
‘ 138
American Fern Journal
138
Index to Volume 70
“
Errata es
_ A34
The American Fern Society
Council for 1980
ROBERT M. LLOYD, Dept. of Botany, Ohio University, Athens, Ohio 4570 President
DEAN P. WHITTIER, Dept. of Biology, Vanderbilt University, Nashville, TN ee Vice President
LESLIE G. HICKOK, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916.
Secretary
JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916.
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American Fern Journal
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DAVID B. LELLINGER Smithsonian Institution, Washington, D. C. 20560
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AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 4 (1980) 113
A Range Extension for Dryopteris filix-mas
ERWIN F. EVERT*
On 14 December 1979, I collected Dryopteris filix-mas (L.) Schott in a mesic
ravine in Glencoe, Cook County, Illinois. Two large plants and a smaller one were
growing within a few feet of each other on a steep, relatively undisturbed northwest-
facing slope of the ravine, about 400 feet west of Lake Michigan in T42N, R13E,
SE1/4, Sec. 6. Associates were Acer saccharum, Aralia nudicaulis, Hamamelis
virginiana, Quercus rubra, and Trillium grandiflorum. A search of other ravines in
the vicinity failed to reveal any other plants of D. filix-mas.
This fern has not been reported previously for Illinois (Mohlenbrock & Ladd,
1978). It is newly reported for Wisconsin (Brown County) by Peck and Taylor
(1980). The nearest stations appear to be in Marquette County, Michigan (Billing-
ton, 1952, p. 177) and in Brown County, Wisconsin, about 270 and 160 miles
distant.
Are these plants spontaneous on this site or is their occurrence due to the actions
of man? Although this question cannot be answered with absolute certainty, the
following points support natural occurrence:
(1) The plants grow in an environment typical for the species, in a relatively
undisturbed area with autochthonous associates.
(2) Pepoon (1927), Moran (1978), Swink and Wilhelm (1979), and personal
observation document the presence of many other uncommon plants for this area in
the ravines along Lake Michigan. Some of these are: Dryopteris intermedia, D.
marginalis, Equisetum scirpoides, Fagus grandifolia, Lycopodium lucidulum, Mit-
chella repens, Pinus resinosa, Polystichum acrostichoides, Shepherdia canadensis,
and Thelypteris hexagonoptera.
(3) Personal observation indicates that D. filix-mas is not commonly cultivated in
this area at present. Therefore, it is not likely that these plants have escaped from
cultivation. ‘Although the Male Fern formerly was grown for its medicinal properties,
the colony does not appear to have been established for a long time, and so it is
unlikely to have originated from plants that were cultivated in the past. It is also
unlikely that plants on such a steep and inaccessible site were deliberately planted.
(4) Single plants or small colonies of other ferns with northern distributions are
known to occur widely disjunct in southern Michigan (Wagner, 1972, p. 205 and
pers. comm.). Examples include Botrychium minganense, Gymnocarpium dryopter-
is, and Polystichum braunii.
(5) Dryopteris filix-mas, a homosporous pteridophyte, apparently is capable of
intra-gametophytic selfing (W. H. Wagner, pers. comm.), and so one wind-borne
Spore could produce a new, disjunct colony. ee
(6) The distribution pattern of D. filix-mas (Hultén, 1962, p. 119), with its widely
disjunct stations in California, Mexico, South America, Hawaii, Greenland, Iceland,
and Africa, indicates that this species is capable of wide dispersal.
*1476 Tyrell St., Park Ridge, TL 60068.
Volume 70, number 3, of the JOURNAL was issued September 29, 1980.
114 AMERICAN FERN JOURNAL: VOLUME 70
Specimens of Evert 1651 have been deposited at the Morton Arboretum Herbari-
um (MOR) and the University of Michigan Herbarium (MICH).
LITERATURE CITED
BILLINGTON, C. 1952. Ferns of Michigan. Cranbrook Institute of Science, Bloomfield Hills, MI.
HULTEN, E. 1962. The Circumpolar Plants 1, Vascular Cryptogams, Conifers, and Monocotyledons.
Kungl. ‘Ape Vetenskapsakad. Handl. 8(5):1-275.
MOHLENBROCK, R. and D. M. LADD. 1978. ae of Illinois Vascular Plants. Southern
Illinois pula cals Carbondale, IL.
MORAN, R. C. 1978. Vascular flora of the ravines along Lake Michigan in Lake County, Illinois.
0
M :
PECK, J. H. and W. C. TAYLOR. 1980. Check list and distributions of Wisconsin ferns and fern
allies. Michigan Bot. 19:252-268.
PEPOON, H. S. 1927. An Annotated Flora of the Chicago Area. Academy of Sciences, Chicago, IL.
SWINK, F. and G. WILHELM. 1979. Plants of the Chicago Region, rev. & expanded ed. Morton
Arboretum, Lisle, IL.
WAGNER, W. H., Jr. 1972. Disjunctions in homosporous vascular plants. Ann. Missouri Bot. Gard.
59:203-217.
AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 4 (1980) 115
Differential Germination of Fern and Moss Spores
in Response to Mercuric Chloride
RAYMOND L. PETERSEN and PATRICK C. FRANCIS*
The pervasive use of toxic substances and the synthesis of myriad new such
compounds requires that toxic substance limits be set and that environmental
monitoring be maintained in order to insure the health and integrity of the
biosphere. Bioassays are the primary mechanism for meeting these two require-
ments. A bioassay designed to determine the environmental impact of a particular
toxic substance, for example divalent mercury, should have the following attributes:
ease of performance, low cost, brevity, sensitivity, and high overall reflectivity of
environmental stress induced by the presence of the toxic substance. The appropri-
ateness of a bioassay is based on the selection of its two primary components—the
living system employed and the parameter measured.
Petersen et al. (1980) have demonstrated the feasibility of using the germination
of Onoclea sensibilis spores to gauge the toxicity of various heavy metal ions. It was
found that for the three metal ions tested, toxicity was directly proportional to
atomic weight. Hg** is twice as toxic as Cd** and four times as toxic as
Co**. The present study is a comparison of the germination responses of different
fern and moss spores to divalent mercury.
There are few investigations on the effects of metal ions and other potential
pollutants on fern spores and gametophytes that yield data pertinent to pollution
research and control. Nakazawa and Tsusaki (1959) determined that the fern spore
cytoplasm associated with rhizoid differentiation has a marked affinity for, metal
ions. Nakazawa and Otaki (1962) demonstrated the affinity of developed rhizoids for
metal ions. Fern rhizoids apparently function like the root hairs of vascular plants in
absorbing water and minerals from the soil. Therefore, their affinity for metal ions is
not surprising. Co** and Ni** were shown to prolong filamentous (one-
dimensignal) growth in Lygodium smithianum gametophytes (Parés, 1958) resulting
in a retardation of their development sequence. LiCl caused a precocious differentia-
tion of terminal papillae (gland-like hair cells produced by some fern gametophytes)
in Dryopteris varia (Nakazawa, 1960a, b). Several metal chlorides at concentrations
of 0.005—-0.08M decreased the period of fern sperm motility (Igura, 1958).
A few papers on ferns with direct application to pollution monitoring have been
published. Klekowski (1976), Klekowski and Berger (1976), and Klekowski and
Poppel (1976) found that meiotic chromosome behavior during fern sporogenesis
was correlated with the presence of toxic substances in the environment. Howar
and Haigh (1972) studied the effects of increasing doses of X-radiation on the first
mitotic division of Osmunda regalis spores. Edwards and Miller (1970, 1972a, b)
Studied the quantitative effects of ethylene on Onoclea sensibilis spore germination
and gametophyte growth. Fern gametophytes also have been successfully employed
In bioassay procedures for the plant hormones kinetin, gibberelic acid, and
antheridogen (Bopp, 1968; Brandes, 1973).
*Department of Botany, Howard University, Washington, DC 20059.
116 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
Mosses are especially adapted to absorb and concentrate substances present in the
atmosphere, and so moss gametophytes have proven to be sensitive indicators of
airborne pollutants (Huckabee, 1973: Skaar et al., 1973; Little & Martin, 1974).
Francis and Petersen (1980) reported on the synergistic effects that metal ion
combinations had on the germination of Polytrichum commune spores.
pores of Onoclea sensibilis, Osmunda cinnamomea, and Osmunda claytoniana
and the moss Polytrichum commune were selected for investigation because they are
of wide distribution and frequent occurrence and all produce copious quantities of
easily collected spores. In addition, O. sensibilis had been employed in the
development of the fern spore heavy metal bioassay (Petersen, et al., 1980). The two
Osmunda species represented a fern family distinct from that of Onoclea and would
yield data on intrageneric differences in response to heavy metal ions. Polytrichum
commune was selected to compare its heavy metal response with those of the ferns.
TABLE 1. SPORE GERMINATION IN VARIOUS Hg* * CONCENTRATIONS EXPRESSED AS A
PERCENTAGE OF CONTROL GERMINATION. '
Hg* +
ppm O. sensibilis O. cinnamomea O. claytoniana P. commune
0.0 100 100 100 100
0.2 101 98 96 77
0.4 98 91 89 30
0.6 98 Ti 33 8
0.8 88 60 20 0
1.0 85 13 14 0
LS 75 8 8 0
2.0 68 1 1 5
3.0 55 0 0 r
4.0 30 zi z
6.0 9 a a
8.0 l w *
10.0 0 i i
20.0 0
1 *
Tests were not run, or the experiment was lost, where no numerical value is given.
MATERIAL AND METHODS
_Approximately 10,000 spores of each species were cultured in a petri dish 60mm
diam. in 8 ml of full strength liquid Knudson’s medium at a pH 5.5 with 0-20 ppm
of divalent mercury ion (Hg* *) added as HgCl). The dishes were sealed in clear
Plastic sandwich bags and cultured in a Sherer Growth Chamber at 20°C in 300 ft-c
of continuous light from cool-white fluorescent lamps. Spore germination was
considered to have occurred when the spores produced a rhizoid or the first
prothallial cell divided. After eight days, 500 spores were examined per plate. Three
replicates were run for each species tested at each Hg* * concentration.
PETERSEN & FRANCIS: DIFFERENTIAL SPORE GERMINATION 117
RESULTS AND DISCUSSION
The germination of the controls (0 ppm HG**) was uniformly high; O.
sensibilis had 88%, O. cinnamomea 92%, O. claytoniana 96%, and P. commune
94%. Spore germination at each Hg* * concentration except one was lower than in
the controls. Germination was expressed as a percentage of each species’ control
germination (Table 1). These results are summarized in terms of the standard
bioassay toxicity values of LCs9 and LCj99, which are presented both in terms of
ppm and wM (Table 2). LCs is the concentration of a toxic substance necessary to
kill 50% of the organisms from a control value, and LCjo9 is the minimum
concentration necessary to kill 100%.
TABLE 2. Hg* * LCs) AND LCjo99 VALUES FOR FOUR TAXA.
Toxicity
i O. sensibilis O. cinnamomea O. claytoniana P. commune
LCso
ppm 3.2 0.82 0.53 0.30
uM 16 4.1 2.6 1.5
LCi
ppm 10 3.0 3.0 0.8
uM 50 15 15 4.0
A comparison of spore germination responses shows that P. commune is the taxon
most susceptible to HG** and is ten times more sensitive than O. sensibilis,
which is the least susceptible. The two Osmunda species have intermediate and
similar HG* * toxicity values. They are approximately four times more sensitive to
HG** than is O. sensibilis (Table 2). Therefore, based on the sensitivity of
response, P. commune would be the species of choice in a mercury ion bioassay.
We wish to acknowledge financial aid from NSF Grant SM177-05566.
LITERATURE CITED
BOPP, M. 1968. Control of differentiation in fern-allies and bryophytes. Ann. Rev. Pl. Physiol.
1-380
19:3 ;
BRANDES, H. 1973. Gametophyte development in ferns and bryophytes. Ann. Rev. Pl. Physiol.
24:115-128.
EDWARDS, M. E. and J. H. MILLER. 1970. Inhibition of cell division by ethylene in fern
gametophytes under various light conditions. Pl. Physiol. (Lancaster) 46(Suppl.):32.
, and J. H. MILLER. 1972a. Growth regulation by ethylene in fern gametophytes. Il.
Inhibition of cell division. Amer. J. Bot. 59:450-457.
. and J. H. MILLER. 1972b. Growth regulation by ethylene in fern gametophytes. III.
Inhibition of spore germination. Amer. J. Bot. 59:458-467. oe
FRANCIS, P. C. and R. L. PETERSEN. 1980. Synergistic behavior of heavy metal combinations by the
fern and moss spore germination bioassay. Assoc, Southeastern Biol. Bull. 27(2):46 (abstract).
HOWARD, A. and M. V. HAIGH. 1972. Radiation responses of fern spores during their first cell-cycle.
Int. J. Radiat. Biol. 18:147-158. ;
HUCKABEE, J. W. 1973. Mosses: sensitive indicators of airborne mercury pollution. Atmosph.
Environ. 7:749-754.
118 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
IGURA, I. ASS. Cytological and morphological apa on ie gametophytes of ferns. XI. The
ife of the fern spermatozoid. Bot Mag.
Tokyo 71:37-42.
pecan E. J., Jr. 1976. Mutational load-in a fern population growing in a polluted environment.
r. J. Bot. 63:1024-1030.
‘ — B..B. BERGER. 1976. reenine mutations: ina =m ar growing in a
polluted envi t r. J. Bot. 63:239-246.
; _ D. M. POPPEL. 1976. Ferns: potential in-situ. bioassay ile for aquatic-borne
tagens. ene Fern. J. 66:75-79.
LITTLE, . and oe . MARTIN. 1974. Biological monitoring of heavy metal pollution. Environ.
Pollut oe
NAKAZAWA, s. ne Cytodifferentiation patterns of Dryopteris protonema modified by some
chemical agents. Cytologia 25:352-361
1960b. Aceleracion y retardo en la morfogénesis de la fase protonema del protalo de los
helelchos. Anal. Inst. Biol. Univ. México 32:191-1
, and A. TSUSAKI. 1959. Appearance of * ‘metallphilic cytoplasm” as a prepattern to the
differentiation of rhizoid in fern protonema. Cytologia 24:378-388.
I. 1962. A prepattern to the papilla differentiation in Dryopteris gametophytes.
3 Phyton (Buenos Aires) 18:113-120.
PARES, Y. 1958. Etude experimentale de la morphogenése du gamétophyte de quelques Filicinées.
Ann. Sci. Nat. Bot., Il, 19:1-120.
PETERSEN, R. L., D. ARNOLD, D. E. LYNCH, and S. A. PRICE. 1980. A heavy metal bioassay
on percent spore germination of the sensitive fern, Onoclea sensibilis. Bull. Environ. Con-
tam. Toxicol. 24:489-4
SKAAR, H., E. OPHUS, and B. M. GULLVAG. 1973. Lead accumulation within nuclei of moss leaf
cells. Nature 241:215-216
AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 4 (1980) 119
Differences in the Apparent Permeability
of Spore Walls and Prothallial Cell Walls
in Onoclea sensibilis
JOHN H. MILLER
The spores of Onoclea sensibilis L. undergo marked changes in their apparent
permeability during germination. For example, spores do not stain with an
acetocarmine-chloral hydrate mixture until germination has proceeded for about 20
hr (Towill & Ikuma, 1975; Fisher & Miller, 1978), whereas after that time stain is
absorbed readily. During the early stages of germination, the spores of Onoclea,
Matteuccia struthiopteris (L.) Tod., and other species are difficult to prepare for
electron microscopy because embedding resins fail to penetrate adequately (Mar-
engo, 1973; Gantt & Arnott, 1976; Raghavan, 1976). Spores in later stages can be
processed with no difficulty. This impermeability seems to be associated with the
inner spore coat (intine). Fisher and Miller (1978) noted that when the intine of
Onoclea spores was artificially ruptured during the early stages of germination and
the protoplast was directly exposed to acetocarmine-chloral hydrate, the protoplast
stained rapidly, whereas no intact spores could be stained. The time at which
Matteuccia and Onoclea spores normally become penetrable by embedding resins
coincides with the time the intine ruptures naturally during germination (Gantt &
Arnott, 1965; Marengo, 1973). If the intine of Onoclea is caused to open by
treatment with sodium hypochlorite, even dormant spores and those in early stages
of germination may be infiltrated readily with embedding resin (Bassel, Kuehnert &
Miller, 1981).
Dormant spores of both Onoclea and Matteuccia have a loose outer spore coat
and a thick intine, along which there is a longitudinal seam, the raphe (laesura), on
the flattened, proximal face of the spore; the spore protoplast is naked within the
intine (Gantt & Arnott, 1965; Bassel, Kuehnert & Miller, 1981). Germinating
spores synthesize a new wall around the protoplast inside the intine between 8 and
16 hr. At the time the intine ruptures and is cast off, this new wall becomes the
bounding wall of the young protonema. Clearly there is a difference in which
materials will penetrate the spore intine and which will cross the normal prothallial
cell wall. Carpita et al. (1979) published a method for obtaining quantitative
information about the apparent capillary pore size of plant cell walls, which limits
the passage of solutes. The cells are placed in a solution of a non-ionic solute having
a water potential lower than that of the cells, which causes water to leave the cells. If
the solute can pass through the wall, and thus the solution can be in contact with the
plasma membrane, one observes plasmolysis (retraction of the protoplast from the
cell wall). If, however, the solute particles are too large to penetrate the wall, exit of
water from the cell causes cytorrhysis (collapse of the cell wall around the protoplast
as it shrinks). I applied this technique to spores and young gametophytes of Onoclea
and observed differences in the apparent capillary pore sizes of the spore intine,
Prothallial cell walls, and rhizoid walls.
“Department of Biology, Syracuse University, Syracuse, NY 13210.
120 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
MATERIALS AND METHODS
Sporophylls of Onoclea sensibilis were collected in the vicinity of Syracuse, NY
in the fall of 1978. They were stored in plastic bags and refrigerated until needed.
The methods by which spores were isolated from the fertile fronds and general
cultural methods are described in Miller and Greany (1974). An additional step was
taken in which the outer spore coat was removed from the dry spores by brief
treatment with sodium hypochlorite, following which the spores were redried and
stored (Vogelmann & Miller, unpubl.). Removal of the brown outer spore coat made it
easier to observe plasmolysis or cytorrhysis. Dormant spores were hydrated before
they were used by floating them on the surface of distilled water for three hours.
Young gametophytes were grown by floating spores on the surface of Knop’s
solution for three days in an air-conditioned growth chamber where the temperature
was 26 + 1°C and the light intensity was about 800 ft-c of continuous cool-white
fluorescent illumination. After three days, the gametophytes had 3—5 vegetative cells
and one primary rhizoid.
The solutes tested included NaCl, ethylene gylcol, glycerol, glucose, sucrose,
polyethylene glycol-600 (PEG-600) and PEG-1000 (all chemicals from Fisher
Scientific Co.). All compounds were dissolved in distilled water. The concentrations
of the solutions, which are given in Table ], were selected in preliminary experi-
ments so that responses were observed within 1-5 min. Spores or prothallia were
mounted directly in a drop of solution on a microscope slide and were covered with
a cover slip. Observations were made with a microscope at a magnification of
approximately 300 x. Photographs were made of representative cells.
TABLE 1. INDUCTION OF PLASMOLYSIS (P) OR CYTORRHYSIS (C) BY DIFFERENT SOLUTES.
Substance and
and olecular | Prothallial
Concentration diameter (nm)? Spores cells Rhizoids
Ethylene glycol (50, 15) 0.45 P P
Glycerol (50, 15) 0.55 P P 4
Glucose (50, 20) 0.88 . P
Sucrose (50, 20) 1.03 € P i
PEG-600 (50, 20) 2.9 c P and C r
PEG-1000 (—, 40) 35 — Cc C
% concentration (w/v). First concentration in parenthesis is for spores; second concentration 1s for
prothallia and rhizoids.
*Values for ethylene glycol and glycerol from Goldstein and Solomon , fe ei
(1960); for glucose and sucrose
rom Durbin (1960); for PEG-600 and PEG-1000 from Carpita, et al. (1979). '
RESULTS AND DISCUSSION
The phenomena of plasmolysis and cytorrhysis occurred in Onoclea spores,
prothallial cells, and rhizoids (Table 1). Collapse of the spore wall through
cytorrhysis was demonstrated in 50% sucrose (Figs. ] and 2 show two views of a
spore at different focal levels). Typically the spores became indented and bowl-
shaped; the indentation always occurred on the proximal face (Fig. 1). No plas-
molytic retraction of the protoplast took place at any point; even at the rim of the
J. H. MILLER: DIFFERENCES IN APPARENT WALL PERMEABILITY 121
- 1-10. Examples of plasmolysis and cytorrhysis in spores, rhizoids, and prothallial cells of O.
sensbils 50 um scale is the same for all photographs. FIGS. | and 2. Cytorrhysis in a spore
shown at different focal levels. FIGS. 3 and 4. Plasmolysis in spores shown at different focal levels.
FIG. 5. Control rhizoid. FIG. 6. Plasmolysed rhizoid. FIG. 7. Ribbon-like rhizoid, collapsed as a result
of oo FIG. 8. Control prothallus. FIG. 9. Plasmolysed prothallus. FIG. 10. Cytorrhysis in
Prothallus
122 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
bowl the granular cytoplasm was in contact with the intine (Fig. 2). A 15% NaCl
solution clearly induced plasmolysis (Fig. 4). Spore plasmolysis also was accompa-
nied by an indentation of the proximal spore face (Fig. 3), since there appeared to be
a firm adhesion of the protoplast to the intine in the area of the raphe, and when the
protoplast became plasmolysed, the intine was drawn in at that point. A control
rhizoid is shown in Fig. 5, and plasmolysis in 20% glucose is illustrated in Fig. 6.
Cytorrhysis in a rhizoid in 40% PEG-1000 resulted in the collapse of the cell into a
ribbon form (Fig. 7). The vegetative portion of a normal prothallus is pictured in
Fig. 8. Plasmolysis gave the appearance shown in Fig. 9, whereas the collapse and
crumpling of the cells through cytorrhysis is shown in Fig. 10. The figures give the
pictorial definition of the terms plasmolysis and cytorrhysis as they are used in this
paper.
The main results are summarized in Table /. Prothallial cells showed only
plasmolysis with compounds up to the size of sucrose. PEG-600 caused both
cytorrhysis and plasmolysis, whereas PEG-1000 produced pure cytorrhysis. Follow-
ing the reasoning of Carpita, et al. (1979), the limiting pore size of prothallial cell
walls appears to be between 2.9 and 3.5 nm. This is somewhat smaller than the
values found by Carpita, et al. (1979) for the cells of several species of angiosperms.
Rhizoids appear to have a slightly larger wall pore size than prothallial cells, since
PEG-600, which caused both cytorrhysis and plasmolysis in prothallial cells, caused
only plasmolysis without cell collapse in rhizoids. The overall permeability of
rhizoids was shown by Smith (1972) to be greater than the permeability of prothallial
cells of Polypodium vulgare. His measurements were made by following the uptake
of a vital dye into the protoplast, and thus reflect the permeability of the plasma
membrane. The results of the present study indicate that some of the difference
between the permeability of rhizoids and prothallial cells may be caused by
differences in the permeabilities of their walls. The spore intine clearly was much
more impermeable than the cell walls of the gametophyte. Glycerol was the largest
molecule which caused plasmolysis of the spore protoplast; glucose induced pure
cytorrhysis. The capillary pore size of the intine appeared to be less than 0.8 nm.
Spores were visibly affected only by higher concentrations of each of the
substances than were required to produce effects in rhizoids or prothallial cells. This
may reflect the fact that the spore cytoplasm is very dense and non-vacuolate, as
‘seen in electron micrographs (Bassel, Kuehnert & Miller, 1981). More of the water
of the spore may be bound in the hydration of proteins, for example, and relatively
little available for free osmotic exchange. The same concentrations of osmotic
solutions which plasmolysed or collapsed prothallial cells within 1-5 min acted
much more rapidly on rhizoids. In each test, rhizoids were affected in less than 30
sec, the tume necessary to prepare the sample and make the first observation. This
rapidity is probably another reflection of the greater permeability of rhizoids. When
young gametophytes were placed in plasmolysing solutions, the basal cell of the
plant was affected first, followed by the intermediate and more apical cells.
Deplasmolysis occurred in prothallial cells and rhizoids which were plasmolysed in
certain of the solutions. Both cell types deplasmolysed completely within 15 min
after immersion in ethylene glycol. Prothallial cells were deplasmolysed completely
in two hr in glycerol and were partially deplasmolysed in the same time in glucose.
J. H. MILLER: DIFFERENCES IN APPARENT WALL PERMEABILITY 123
Rhizoids: showed only a partial recovery in either of these two compounds. Plasmoly-
sis appeared to be permanent in solutions of any of the large substances. The
instances of deplasmolysis indicate that the plasma membranes of the cells were
permeable to the plasmolysing solute, and enough was taken up eventually to reverse
the flow and cause an influx of water into the protoplasts.
The results which are presented in this paper support the idea that the low
permeability of Onoclea spores results from the permeability properties of the
intine. The estimated capillary pore size of the intine is only about one quarter that
of prothallial cells and rhizoids. The intine should play a major role in determining
the entry and exit of materials into and from the spore during the first stages of
germination before the intine is ruptured. Some aspects of permeability seem not to
be explicable on this basis. Vogelmann (1980), for example, showed that colchi-
cine, griseofulvin, and isopropyl N-chlorophenyl carbamate produce striking effects
on spore germination, and each appears to enter the spore before the time of intine
rupture, although one would expect them to be excluded on the basis of their size.
One possible explanation for this type of anomaly is the suggestion by Carpita, et al.
(1979) that a small number of larger pores might provide access to the protoplast by
larger molecules, whereas osmotic effects may be governed by the more abundant
smaller pores.
This research was aided by grant PCM-7904593 from the National Science
Foundation.
LITERATURE CITED
BASSEL, A. R., C. C. KUEHNERT and J. H. MILLER. 1981. Nuclear migration and asymmetric cell
ees | in eaiaes sensibilis spores: an ultrastructural and cytochemical study. Amer. J. Bot
68:(in press).
CARPITA, N.. D. SABULARSE. D. MONTEZINOS and D. P. DELMER. 1979. Determination of the
pore size of cell walls of living plant cells. Science 205:1144 —114
DURBIN, R. P. 1960. Osmotic flow of water across permeable cellulose membranes. J. Gen. Physiol.
44:315-326.
FISHER, R. W. and J. H. MILLER. 1978. Growth regulation by ethylene in fern gametophytes. V.
Ethylene we the early ree of spore germination. Amer. J. Bot. 65:334 —339.
GANTT, E. and H. J. ARNOTT. 1965. Spore germination and development of the young gametophyte
of the ane fern (Matteuccia struthiopteris). Amer. J. Bot. 52:82-94.
GOLDSTEIN, D. A. and A. K. SOLOMON. 1960. Determination of oe pore radius for human
red cells by osmotic pressure measurement. J. Gen. Physiol. 44:1
MARENGO, N. P. 1973. The fine structure of the dormant spore of crac struthiopteris. Bull.
orrey Bot. Club 100:147-150.
MILLER, J H. and R. H. GREANY. 1974. Determination of rhizoid orientation by light and darkness
N germinating spores of Onoclea sensibilis. Amer. J. Bot. 61:296—302.
RAGHAVAN, V. 1976. Gibberellic acid-induced germination of spores of Anemia phyllitidis: Nucleic
acid and protein synthesis during germination. Amer. J. Bot. 63:96 60-972.
SMITH, D. L. 1972. Staining and osmotic properties of young gametophytes of Polypodium vulgare L.
and their bearing on rhizoid formation. Protoplasma 74:465—497.
TOWILL, L. R. and H. IKUMA. 1975. Photocontrol of the germination of Onoclea — II. Analysis
of the germination process by means of anaerobiosis. Plant Physiol. 55:150—15
VOGELMANN, T. C. 1980. Nuclear migration, asymmetric cell division and rhizoid aaa: in
germinating spores of the sensitive fern, Onoclea sensibilis L. Ph.D. dissertation, Syracuse
University.
124 AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 4 (1980)
Allelopathy and Autotoxicity
in Three Eastern North American Ferns
WILLIAM E. MUNTHER and DAVID E. FAIRBROTHERS*
Undoubtedly gametophytes are the most vulnerable stage in the fern life cycle.
Although individual sporophytes produce from 50 to 100 million haploid spores
(Shaver, 1954), few of the small, delicate gametophytes, which in most ferns are
less than 1.25 cm wide, survive and produce sporophytes. Conditions governing
spore germination are critical to gametophyte establishment. Conway (1953)
found that, in Scotland, few gametophytes or young sporophytes of Pteridium
aquilinum occurred under field conditions, despite heavy spore production by
mature plants. This suggests that the gametophyte and young sporophyte stages
may be limiting in the establishment of this species. Many studies (cited in Hill,
1971) suggest that fern gametophytes and sporophytes can possess quite different
habitat requirements and that one stage of the life cycle can be more sensitive than
the other to physical and biological conditions, and so limit the success of the
species.
Although environmental (abiotic) conditions are extremely important, biotic
conditions, particularly those created by the sporophytes or gametophytes, may
also be of critical importance in the establishment of new individuals of the same
or another species. Therefore, these factors may regulate both population density
and community composition. Chemical inhibition of one plant by another, or
allelopathy, has been known for over a century (Muller, 1966) and has been much
studied in flowering plants and conifers. The phenomenon of antibiosis is also well
known to microbiologists. However, relatively little research has been devoted to
rite: inhibitors produced by non-seed plants, other than microorganisms (Rice,
Bohm and Tryon (1967) reported that many species of ferns produce phenolic
compounds. They examined 46 species for the presence of hydroxylated cinnamic
and benzoic acids and found a basic complement of cinnamic acids (p-coumaric,
caffeic, and ferulic) in the ferns they tested. Also generally present were
p-hydroxybenzoic, protocatechuic, and vanillic acids. Sinapic, syringic, and
o-coumaric acids were reported to be less common. In a follow-up study, Glass
and Bohm (1969) found similar phenolic compounds in 46 additional species. The
presence of a basic complement suggests that the well established pathways of
phenolic metabolism in the seed plants also function in ferns.
Many of the phenolic compounds found in ferns are known to be allelopathic in
many species of higher plants, either directly or indirectly, such as after microbial
decomposition (Rice, 1974). Most phenolic acids are at least slightly soluble in
water. With the increasingly acidic rainfall in the northeastern United States (Li-
kens et al., 1970; Bormann & Likens, 1977), weak organic acids such as the
phenolics may be leached quite readily either from the leaves during the growing
season or from senescent plants. A wide variety of organic and inorganic sub-
*Department of Botany, Rutgers University, New Brunswick, NJ 08903.
MUNTHER & FAIRBROTHERS: ALLELOPATHY AND AUTOTOXICITY 125
stances known to be allelopathic are capable of being leached from the leaves of a
number of species of higher plants (Tukey, 1966, 1969; Rice, 1974).
Of the few allelopathic studies concerned with ferns, most have involved
Pteridium aquilinum. Del Moral and Cates (1971) reported that aqueous leaf ex-
tracts and litter extracts of P. aquilinum inhibit seeds of Hordeum vulgare,
Bromus tectorum, and Pseudotsuga menziesii. Gliessman and Muller (1972) found
that P. aquilinum inhibited germination and subsequent radicle growth of seeds of
Bromus rigida and Avena fatua. The toxic principle was detected in the fern leaf
leachates. Because the toxic principle was transported by some form of precipita-
tion, water-soluble phenolic acids were suspect, and cinnamic acid was tentatively
identified. In another study, Stewart (1975) found that water-soluble extracts from
Western Bracken (P. aquilinum var. pubescens) delayed germination of Rubus
spectabilis seeds and inhibited germination of R. parviflorus seeds, but had no
effect on the seeds of Pseudotsuga menziesii.
Glass (1976) prepared a solution of phenolic acids having the same composition
as was detected in the soil associated with the roots of P. aquilinum and tested its
effect on the growth of barley, wheat, oats, rye, rye-grass, barley grass (Hordeum
murinum), Clover, and Agropyron repens. Growth was inhibited in all species
investigated except A. repens. Whether P. aquilinum sporophytes in some way
inhibit the germination of their own spores or those of another species of fern was
not investigated in Glass’s or any other study reported in the literature.
Davidonis and Ruddat (1973) found that Thelypteris normalis sporophytes in-
hibit the growth of T. normalis gametophytes, as well as those of Preris and
Phlebodium. The inhibitors, which they termed thelypterins a and b, were similar
in many ways to indoleacetic acid (IAA) and were found to be exuded from
sporophyte roots. Davidonis and Ruddat (1974) also reported that gametophytes
grown in the immediate vicinity of a mature sporophyte of 7. normalis had a
reduced number of cells and an altered gross morphology. They observed the
greatest inhibition of T. normalis gametophytes when the thelypterins were added
before spore germination had occurred. :
Davidonis (1976) reported that T. noveboracensis, Pteris multifida , and P. vit-
tata also contained thelypterins. Since the thelypterins were detected in leaf diffu-
Sates of T. normalis and T. noveboracensis, she postulated that foliar leaching
may be one mechanism by which these inhibitors are released into the environ-
ment
Petersen (1976) found that Dryopteris intermedia and Osmunda cinnamomea
gametophytes inhibited each other’s development when grown together in culture.
€ also reported that gametophytes of these two species “lock” each other into
Perpetual juvenility. Petersen was unable to isolate and indentify the inhibitory
Substances since they occurred in very small amounts or were highly volatile.
Horsley (1977) reported that the presence of Dennstaedtia punctilobula and T.
noveboracensis sporophytes were correlated with reduced numbers of Prunus
Serotina, Acer rubrum, and A. saccharum seedlings under field conditions. —
Very limited research has been conducted either to detect possible allelopathic
interactions between species of ferns or to demonstrate the inhibition of one
126 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
generation by the other within a species (autotoxicity). It is not known whether
allelopathy and autotoxicity are widespread in natural fern communities.
Davidonis and Ruddat (1973, 1974) and Davidonis (1976) examined a small number
of species for inhibitors, but in many instances the species tested for allelopathic
interactions were those that are maintained in greenhouses, and not those which
occur together in natural situations. Petersen (1976) has shown that fern
gametophytes of one species produce substances that inhibit gametophytes of
others, the classic allelopathic inhibition of one species by another. Under field
conditions, however, the dominant and normally perennial sporophytes probably
produce much greater quantities of allelopathic compounds capable of affecting
spore germination or gametophyte growth than would the much_ smaller
gametophytes. Petersen’s research suggests that sporophytes might also inhibit
gametophytes, as they were found to have produced the same flavonoids, but no
experimental evidence to support such a hypothesis was presented.
There are reports suggesting allelopathy between fern generations in both
natural populations and the greenhouse. In greenhouses, potted ferns often pro-
duce large numbers of spores, but gametophytes are seldom found growing on the
soil surface directly underneath the sporophytes. It is also unusual to find
gametophytes growing underneath or even near mature sporophytes under field
conditions, even when spore production is heavy.
In this research, aqueous leaf extracts, leaf leachates, and leaf litter infusions
from Osmunda cinnamomea, O. claytoniana, and Dennstaedtia punctilobula
were examined for their allelopathic and autotoxic potential. Leaves of all three
species were also tested for the production of volatile inhibitors.
All three species occupy much the same habitat, although O. cinnamomea is
often found in moister areas than the other two. The geographical ranges of all
three species also are approximately the same, and all are native and common in
the northeastern United States. Although these three species occupy almost iden-
tical habitats, they seldom occur in close proximity to one another within a given
area.
MATERIALS AND METHODS
Mature spores of O. cinnamomea, O. claytoniana, and D. punctilobula were
collected in 1976 and 1977 from wild populations which were located in Sandy-
stone Township, Sussex County, New Jersey, and in the Town of Goshen, Addi-
son County, Vermont.
Spore-bearing leaves of both species of Osmunda were collected during the first
two weeks of May from New Jersey populations and during the last week of May
and first week of June from Vermont populations. Sporangia were allowed to
dehisce at/room temperature. Spores were refrigerated at 4.4” C as soon as possi-
ble after they were shed from the sporangia. Although Osmunda spores contain
chlorophyll and remain viable for only a few days at room temperature (Cobb,
1963), they will retain their viability for over a year if refrigerated (Stokey, 1951)-
Spore-bearing leaves of D. punctilobula were collected during the last two
weeks of July from the New Jersey populations and during the first week of
MUNTHER & FAIRBROTHERS: ALLELOPATHY AND AUTOTOXICITY 127
August from Vermont populations. After dehiscence of the sporangia, spores
were stored dry and at room temperature. The spores of D. punctilobula lack
chlorophyll and have a thicker, more resistant spore wall than Osmunda spores,
and so require no refrigeration to remain viable.
Fresh fronds from all three species were used to prepare aqueous extracts and
leachates. They were collected from both Vermont and New Jersey populations
throughout the growing season. Fronds were refrigerated immediately after col-
lection and were then stored at 4.4° C. The extracts and leachates were prepared
from the fronds within three or four days after collection.
All aqueous leaf extracts were prepared by first cutting into pieces 5 g of leaves
(fresh weight) and then placing the pieces in 80 ml of distilled water. The leaves
were boiled briefly (3 min) to stop enzyme activity and, it was hoped, to destroy
some of the microorganisms which might later contaminate cultures. After boiling,
the leaves were ground in a Virtis blender for 5 min. The ground mixture was then
vacuum filtered and the resulting aqueous extract was brought to volume with
distilled water (100 ml distilled water for each 5 g of ground leaves). Extracts were
stored at 4.4° C until they were used to prepare cultures, usually within 3-5 days.
Leaf leachates for each species were prepared by placing fronds (2 thick) on 1.5
mm mesh screen which was placed on top of a rectangular plastic box. Fronds
having an area of ca. 630 cm? were then misted with 300 ml of distilled water. The
droplets of water falling from the leaves were collected in the large plastic box.
Leachates were stored at 4.4° C until used.
Litter infusions were prepared by placing 10 g of chopped dry frond litter in a
500 ml beaker. Weekly additions of 100 ml of distilled water were made for a
period of 3 weeks. Beakers were placed in direct light in the greenhouse. Filter
paper was placed on top of each beaker to prevent external contamination. After 3
weeks, the aqueous portion of the infusion was decanted and vacuum filtered. The
filtered liquid was used to prepare the experimental cultures.
The methods used to prepare cultures were based on those reported by Munther
(1975). All cultures using extracts, leachates, and infusions were prepared as
follows: plastic seed germinators (clear plastic boxes measuring I1 x ikx 3 cm)
were filled with 0.138 1 (dry) of autoclaved horticultural grade (medium) vermicu-
lite, and various treatment solutions were added to the vermiculite. The treatment
solutions contained two parts of extract, leachate, or infusion and one part of 2X
Hoagland’s no. 1 solution plus trace elements (Hoshizaki, 1975). Fern spores
generally require only moisture to germinate (Miller, 1968), but a nutrient solution
was added because the cultures were examined over an extended time, and nutri-
ents are necessary after the first few cell divisions of germinating spores. A total of
75 ml of solution was added to each experimental box. In controls, distilled water
was mixed with Hoagland’s solution instead of ‘the extracts, leachates, or infu-
sions. Fern spores were sown on the upper surface of pieces of unglazed clay pots
placed on top of the vermiculite, 3 per germinator. New pots were used, and all
were from the same manufacturer’s lot. The pots, broken into pieces of approxi-
mately equal size, were boiled vigorously in water for 15 min three separate times,
using fresh water for each boiling to remove the impurities.
128 AMERICAN FERN JOURNAL: VOLUME 70 1980)
Cultures were placed under 285-315 ft-c of illumination provided by alternately
placed 40-watt plant grow (G.E.) and cool white fluorescent tubes with a photo-
period of 14 hrs. The light intensity used was based on field observations obtained
with a light meter and on a recommendation reported by Miller and Miller (1961).
Temperature was maintained at 22-24° C.
Each culture was checked daily after sowing until the first spores began to
germinate. At that time, a count was made to determine percent germination and
then repeated every other day for thirteen days. Since experimental data indicated
that the last two counts were not appreciably different, the time was later reduced
to 11 days. The criterion used to determine when spore germination had occurred
was the appearance of the rhizoid following the first division of the spore, not just
the uptake of water as shown by swelling. All counts were made under low power
(100) of a light microscope, and a mechanical counter was used to record the
number of germinated and non-germinated spores within a randomly selected
microscopic field. One count was taken from each chip in the box. All experi-
ments were conducted in triplicate, using three germinator boxes for each repli-
cate.
To test for the production of volatile compounds by the fronds of each of the
three species, 30 g of fresh fronds were placed in a 16 x 31 cm plastic box. Also
within this plastic box were 9-cm petri dish bottoms containing culture medium
similar to that used in the other experiments. Each plate contained 0.091 | (dry) of
vermiculite plus 50 ml of control solution on top of which were the pieces of clay
pot with spores. The box was sealed with clear plastic tape. For controls, 30 g of
cotton moistened with water was used in place of the fresh fronds. This method
was similar in principle to that described by Muller (1966), and allowed any vol-
atile compounds produced by the fronds to concentrate in the closed atmosphere
of the box. The only contact between the fronds and the spores was through the
air. The sealed boxes were placed under 300 ft-c of light for 14 days, after which
counts were taken to calculate percent germination. All experiments were con-
ducted in tripicate.
The replicated means of percent germination resulting from treatment with leaf
leachates and extracts were compared statistically using Duncan’s multiple range
test (Duncan, 1955; Steele & Torrie, 1960), with replicated control means, for
every possible autotoxic and allelopathic interaction (Munther, 1978). Separate
comparisons to controls were made using the litter infusion treatment means.
Only the means from the first (day 1) and last (day 11 or 13) counts were used for
statistical analysis. These statistical comparisons were therefore based on a
minimum (the presence of any germinated spores in any replicate of a species and
treatment) and a maximum (all spores capable of germination from a variable
population) level of germination for every possible interaction. Replicated volatile
treatment means were compared to the controls in a similar manner; however,
only one series of means representing the maximum level of germination was used
in these comparisons, since it was not possible to obtain a minimum level due to
the design of the experiments.
MUNTHER & FAIRBROTHERS: ALLELOPATHY AND AUTOTOXICITY 129
RESULTS AND DISCUSSION
In the following discussion, compounds produced in the leaves of a species
which inhibit the germination of spores of the same species will be termed ‘‘au-
totoxic.’’ The inhibitory effects on spores of another species will be termed “‘al-
lelopathic.”’
Spores of both Osmunda species began to germinate in five or six days in
Vermont and New Jersey populations. Dennstaedtia spores generally took longer
to begin germination. Those from Vermont required at least 11 days, and those
from New Jersey required at least 8 days.
TABLE 1. EFFECTS OF AQUEOUS LEAF EXTRACTS AND LEACHATES ON SPORE GER-
MINATION AND EARLY GAMETOPHYTE GROWTH IN DENNSTAEDTIA AND OSMUNDA.
Species Vermont New Jersey
Spores Leaves Leachate Extract Leachate Extract
FC EC FC | FC Gy FC i &
Autotoxic Interactions
O. cinn. O. cinn a - ~ - -
O. clay O. clay. _ “ a + ~ - - ~
D. punc D. punc. 2 = * aa = + 4
Allelopathic Interactions
O. cinn O. clay. S - + os = as +
O. cinn D. punc - ~ + + * - + sl
O. clay. O. cinn. & = bid - ~~ re _ -
O. clay D. punc. ~_ - - — ces
D. pune O. cinn. _ = + ~ - - - -
D. punc O. clay. - — + + = = = =
FC = first count; LC = last count a ies
+ = Statistically significant inhibition (0.05 level); — = no significant inhibition
In the Vermont populations, all species exhibited some degree of autotoxicity
(Table 1). In each species, the leaf extract was found to inhibit germination signifi-
cantly, particularly in the last count. InO. cinnamomea, leaf leachates also signif-
icantly inhibited spore germination, and the inhibition was nearly as great as that
caused by the extracts (Table 3). This differed considerably from the results
obtained from the New Jersey populations, where only one species, D
punctilobula, was found to be significantly autotoxic (Table 1). In this case, the
inhibition was caused by the extract.
Significant allelopathic interactions also were found in the Vermont popula-
tions. Leaf extracts of bothO. claytoniana and D. punctilobula inhibited spores of
O. cinnamomea, significantly lowering percent germination (Table /). Inhibition
of germination was quite severe, particularly as revealed in the final counts (Ta-
bles 4 and 5). Spores of D. punctilobula were inhibited by leaf extracts of O.
cinnamomea and O. claytoniana (Table 1). Inhibition in this case also was quite
marked, especially by extracts of O. claytoniana (Table 4). Aqueous leaf extracts
of O. claytoniana often severely inhibited spores of the other two species, but
€xtracts of both O. cinnamomea and D. punctilobula had little effect on O.
claytoniana spore germination (Table 1). Leaf leachates of the three species tested
Produced no significant allelopathic inhibition of spore germination.
130 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
The number of significant allelopathic interactions was lower in the New Jersey
populations than in the Vermont populations (Table 1). Spores of O. cinnamomea
were inhibited by leaf leachates of D. punctilobula, which was the only significant
allelopathic inhibition by leaf leachates in either the New Jersey or Vermont
populations (Tables ] and 5). The inhibition produced by the leachates was not so
severe as that produced by the extracts (Table 5). Also in New Jersey, germina-
tion of O. cinnamomea spores was inhibited by leaf extracts of both O.
claytoniana and D. punctilobula able 1). Osmunda cinnamomea and O.
claytoniana leaf extracts had little effect on germination of D. punctilobula spores
(Table 1). This result is quite different from that obtained for the Vermont popula-
tions (Table 1). However, as in the Vermont populations, spores of O. claytoniana
were unaffected by leaf extracts (or leachates) of both O. cinnamomea and D.
punctilobula (Table 1). Nevertheless, in the majority of cases, the Vermont and
New Jersey populations reacted quite differently in terms of the number and type
of allelopathic and autotoxic interactions.
TABLE 2. EFFECT OF LEAF LITTER INFUSIONS ON SPORE GERMINATION AND EARLY
GAMETOPHYTE GROWTH IN NEW JERSEY POPULATIONS OF DENNSTAEDTIA AND
OSMUNDA.
Species litter infusion
Spores Leaves FC LC
Autotoxic Interactions
cinn. O. cinn. - =
O. clay. O. clay. - =
D. punc D. punc. = =
Allelopathic Interactions
O. cinn O. clay. = =
O. cinn D. punc Sa x
O. clay O. cinn =
O. clay. D. punc = oa
D. punc O. cinn. “ —
D. punc O. clay. _ -
FC = first count; LC = last count
+ = statistically significant inhibition (0.05 level); — = no significant effect
Leaf litter infusions of New Jersey material did not inhibit germination greatly
(Table 2). Dennstaedtia punctilobulaspores were inhibited by O. cinnamomea leaf
litter infusions, the only allelopathic inhibition caused by treatment with litter
infusions. The only species exhibiting autotoxicity was O. cinnamomea (Table 2),
in which the litter infusion merely delayed germination, since there was no inhibi-
tion in the last count.
Two types of inhibition were observed in spores treated with extracts and
leachates. Spores of all three species imbibed water readily when treated with
leachates or extracts, but spores treated with extracts often did not divide. This
indicates that the inhibitor was able to enter the spore through the wall and to
prevent the first division. This type of inhibition causes a low percentage of spores
to germinate, is evident in the first count when compared to the controls, and
MUNTHER & FAIRBROTHERS: ALLELOPATHY AND AUTOTOXICITY 131
continues through the last count (Table 1). This was the most common type of
inhibition. In the second type, several divisions occurred and percent germination
was not affected in the first count (see O. claytoniana, autotoxicity, Table 1). The
inhibitor in this instance did not affect germination, but acted on the several-celled
stage (young gametophyte). In the later counts, if a spore had germinated but had
died (chlorophyll lost) at the several-celled stage, it was counted as not germi-
nated. Therefore, the last count measures the inhibitor’s effect on early
gametophyte growth and development. The presence of this type of inhibition can
be recognized only at the last count.
When leaf extracts caused significant inhibition of germination, gametophyte
development from spores which germinated usually was affected, with the
gametophyte usually arrested in the filamentous stage and little or no further
growth occurring. Leaf leachates usually did not cause this type of response.
Gametophytes resulting from spores which did germinate underwent normal de-
velopment, but often exhibited somewhat slower growth than did the controls.
TABLE 3. AUTOTOXIC EFFECTS OF LEAF LEACHATES AND EXTRACTS EXPRESSED AS
A PERCENTAGE OF SPORE GERMINATION AND EARLY GAMETOPHYTE GROWTH IN
VERMONT POPULATIONS OF O. CINNAMOMEA. STATISTICAL ANALYSIS USING DUN-
CAN’S MULTIPLE RANGE TEST WITH A 0.05 SIGNIFICANCE LEVEL. (ALL EXTRACT
AND LEACHATE MEANS SIGNIFICANTLY DIFFERENT FROM ALL CONTROL MEANS.)
First count (sx = 5.421)
is E2
Means El LI E3 ee eT eee ee
s= ‘a Oe Be SE 8
X= ny 6 UMA OOS ee ee ee
Last count (sx = 5.034)
Means B3 El E2 L3 a S66 6S
s= 7.1 ‘i a. ae 3 OU CU
x= ot OW Wh - Us. MA Oe ee ee LM.
SX = standard error (of the mean); s = standard deviation; X = mean percent germination
C1-3 = control means; L1-3 = leachate treatment means; E1-3 = extract treatment means
Phenolic compounds are known to inhibit ion uptake in flowering plants through
reversible alterations in membrane permeability (Glass, 1973, 1974) and may af-
fect IAA metabolism. The spore wall may prevent the phenolic inhibitor from
entering the spore, with the first divisions of the spore occurring on stored re-
serves. By the time the gametophyte reaches the several-celled stage, the phenolic
inhibitor may prevent ion uptake sufficiently to inhibit growth or further cell
division.
The effects of the extracts on spore germination and on early gametophyte
Stages were much greater than the effects of the leachates in almost every experi-
ment. While experimental effects of the leachates may be easily extrapolated to
field conditions, the extracts present an enigma. While nothing directly paralleling
an extraction procedure exists in nature, experimental results obtained using ex-
132 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
tracts do possess some validity in determining the phytotoxic potential of plants
under laboratory conditions. Extracts may concentrate compounds which are
leachable and concentrated in the soil under field conditions. However, some
compounds present in an extract may not be leached from healthy, growing tissue,
and many substances can be altered by the extraction procedure itself. On the
other hand, substances that are not normally leached from such tissue can be
released by senescent tissue quite readily (Gliessman & Muller, 1972; Stewart,
1975). Various forms of stress, such as temperature extremes, drought, attack by
pathogens, or mechanical injury also increase the leachability of metabolites from
foliage (Tukey, 1966, 1969; Rice, 1974). Soils also vary in their composition,
structure, moisture content, pH, and the kinds of microorganisms present. Thus,
they would have different affinities for various inhibitors, or may even render
them inactive or increase their activity due to microbial decomposition (del Moral
& Cates, 1971; Rice, 1967, 1969; Wang et al., 1967).
TABLE 4. ALLELOPATHIC EFFECTS OF O. CLAYTONIANA LEAF EXTRACTS AND
LEACHATES EXPRESSED AS A PERCENTAGE OF SPORE GERMINATION AND EARLY
GAMETOPHYTE GROWTH IN VERMONT POPULATIONS OF O. CINNAMOMEA AND D.
PUNCTILOBULA. STATISTICAL ANALYSIS USING DUNCAN’S MULTIPLE RANGE TEST
WITH A 0.05 SIGNIFICANCE LEVEL. (ALL EXTRACT (NOT LEACHATE) MEANS SIGNIFI-
CANTLY DIFFERENT FROM ALL CONTROL MEANS.)
. cinnamomea
Last count (sx = 8.450)
Means E2 E3 El L2 Li Cl C3 C2
s= 14.1 18.0 27.2 19.0 8.7 3.5 8.1 10.2 7.0
<= 12.2 24 39.4 43.6 54.8 58.4 76.8 80.5 81.3
D. punctilobula
First count (sx = 4.618)
Means El E3 E2 1 C3 L3 C2
S= 3.8 5.8 3.3 13.7 3.8 9.1 14.5 4.5 12
X= 15.1 18.5 28.0 36.0 41.9 47.5 49.7 S27 54.5
Last count (sx = 4.965)
Means El E3 E2 LI L2 L3 C3 Cl C2
s= =a 12.5 15.6 2.3 10.9 3 6.4 2.9 6.2
x= 11.0 26.0 32.8 64.0 65.1 67.3 74.9 78:7 76.2
sx = standard error (of the mean); s = standard deviation; X inati
cas ' 3 eviation; X = mean percent germination
C1-3 = control means; L1-3 = leachate treatment means; E1-3 = extract treatment means
In all samples except one of D. punctilobula from New Jersey, the pH of the
extracts was lower than that of the leachates (Table 5 ). The pH of the extracts and
leachates from New Jersey populations did not differ greatly from that obtained
from the Vermont material (Table 6). Although in all samples the pH of the
leachates from Vermont populations was higher than in leachates from New Jer-
sey, the difference probably is not enough to be significant. It can be deduced
from Table 6 that allelopathy is not strictly a low pH phenomenon. For example,
MUNTHER & FAIRBROTHERS: ALLELOPATHY AND AUTOTOXICITY 133
the leaf leachates and extracts from O. cinnamomea (New Jersey) exhibited very
little phytotoxicity, yet they possessed the lowest pH.
As already mentioned, litter infusions of O. cinnamomea were found to be
autotoxic (Table 2) only in the first count, indicating delayed germination. Such a
delay could be important under field conditions, however, increasing the chances
of attack by microbial pathogens which could cause damping off of the young
prothalli, as is often the case with seeds.
Within the species studied in New Jersey, the pH values obtained for leaf-litter
infusions were higher than expected when compared with those obtained for leaf
leachates and extracts (Table 6).
TABLE 5. ALLELOPATHIC EFFECTS OF D. PUNCTILOBULA LEAF EXTRACTS AND
LEACHATES EXPRESSED AS A PERCENTAGE OF SPORE GERMINATION AND EARLY
GAMETOPHYTE GROWTH IN VERMONT AND NEW JERSEY POPULATIONS OF 0. CIN-
NAMOMEA. STATISTICAL ANALYSIS USING DUNCAN’S MULTIPLE RANGE TEST WITH
A 0.05 SIGNIFICANCE LEVEL. (ALL EXTRACT (NOT LEACHATE) MEANS SIGNIFI-
CANTLY DIFFERENT FROM ALL CONTROL MEANS IN VERMONT; ALL MEANS SIGNIF-
ICANTLY DIFFERENT FROM ALL CONTROL MEANS IN NEW JERSEY.)
Vermont
Last count (sx = 5.488)
LI 2
Means E2 E3 El LS Cl C3 C2
S = 27 3.5 9.5 15.8 14.7 74 8.1 10.2 7.0
X= 27.8 33.2 38.8 oe RE OOS ES. ES
New Jersey
First count (sx = 4.569)
Means El E3 E2 L2 L3 Ll Cl C2 C3
a= 10.4 8.1 10.6 7.4 4.2 8.8 7.9 75 3.0
x= 29.9 36.7 37.1 ay. oe OF er
Last count (sx = 4.608)
Means El E2 E3 L3 Ll L2 C2 C3 Cl
= 12.2 12.3 5.3 3.9 6.1 4.4 8.3 6.1 8.2
X = MS Ob eo aS se
ee ee
mean per cent germination
SX = standard err -s= dard deviation; X =
rt Tere 8 oe 3 = extract treatment means.
C1-3 = control means; L1-3 = leachate treatment means; El-
llowing the first
he time of the
parent
Frond litter was collected while the fronds were still standing, fo
killing frost in 1976. It is unknown whether or not it rained between t
first frost and the time of collection. If it did, this may account for the ap
lack of phytotoxicity in the infusions, since any water soluble phytotoxins could
have been removed by the rainfall. The importance of the first rainfall after frond
senescence already has been cited by Gliessman and Muller (1972), who studied
the effects of P. aquilinum leaf litter extracts on Avena fatua and Bromus rigida
radicle growth, and by Stewart (1975), who examined the effects of P. aquilinum
litter extracts on seeds of Rubus sp. and Douglas-fir.
134 AMERICAN FERN JOURNAL: VOLUME 70 (1980)
No inhibitory volatile compounds were detected in biologically significant
levels from leaves removed from New Jersey or Vermont populations. Denn-
staedtia punctilobula leaves are quite fragrant, particularly when crushed, but the
substance producing this fragrance, presumably coumarin, had no apparent effect
on spore germination.
Water-soluble inhibitors predominate in more humid or wet environments, ac-
cording to the hypothesis developed by Whittaker (1970) relating toxin production
to climate. Since the Vermont populations exist in a wetter environment than do
the New Jersey ones (Climatological Data, U. S. Dept. of Commerce, NOAA,
1976-1977), this may partially explain the increased number of allelopathic and
autotoxic interactions caused by water-soluble leachates and extracts. Volatile
inhibitors, according to Whittaker (1970) and Muller (1970), would be most com-
mon in a hot, arid environment. The fact that none were found in either Vermont
or New Jersey populations would also lend support to their theory relating toxin
production to climate, as both areas are relatively moist. Summer temperatures in
the study areas in Vermont and New Jersey were nearly identical (Climatological
Data, NOAA, 1976-1977).
TABLE 6. pH VALUES OBTAINED FROM AQUEOUS LEAF EXTRACTS AND LEACHATES
FOR NEW JERSEY AND VERMONT POPULATIONS, AND LITTER INFUSIONS FOR NEW
JERSEY POPULATIONS.
pH
Species NJ VT
Osmunda cinnamomea
leaf extract 5.40 ia
leaf leachate 6.00 6.60
litter infusion 6.50 —
soil (5/77) aoa. & os
Osmunda claytoniana
leaf extract 5.60 5.80
leaf leachate 6.20 6.65
litter infusion 6.75 —_
Soi 6.40 —
Dennstaedtia punctilobula
leaf extract 5.70 5.80
leaf leachate 5.50 6.00
litter infusion 5.95 —
soil (5/77) 5.90 —
We express our appreciation to R. W. Willemsen for his help during a portion of
this research, and to B. F. Palser and J. A. Quinn for their helpful suggestions and
comments.
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136 AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 4 (1980)
SHORTER NOTES
SANDSTONE ROCK CREVICES, AN EXCEPTIONAL NEW HABITAT
FOR THELYPTERIS SIMULATA.—The Massachusetts or Bog Fern, Thelypteris
simulata (Davenp.) Niewl., is a common wetland fern in the New England states.
Most manuals give its typical habitat as very acid, shaded bogs and swamps,
frequently in association with sphagnum moss. In some New England cranberry
bogs, the fern is so abundant as to be a weed. Hartley (Rhodora 67:399— 404. 1965)
reported for the first time 7. simulata from Wisconsin, a disjunction of approximate-
ly 600 miles from the nearest station in West Virginia. Since the original discovery,
twelve other stations have been found for 7. simulata in west-central Wisconsin. The
Wisconsin habitats are all flat, low-lying woods with a moist layer of peat about one
foot thick overlying sand. The dominant trees usually are Pinus strobus and Acer
rubrum, and the most common understory species Alnus rugosa and Ilex verticillata.
The most abundant herbaceous plant at most sites is Osmunda cinnamomea. Other
characteristic ground layer associates include Carex brunnescens, C. folliculata, C.
trisperma, Dryopteris intermedia, D. spinulosa, D. X triploidea, Maianthemum
canadense, Mitchella repens, Osmunda regalis, Rubus hispidus, Sphagnum spp.,
and Viola incognita. It seems likely that in presettlement times, before the natural
character of central Wisconsin was vastly changed by drainage ditches, peat fires,
farming, etc., 7. simulata was a common member of the low, wet-acid woods flora
that once covered much of the sandy plain of extinct glacial Lake Wisconsin.
In mid-August 1979, I found a single plant of 7. simulata growing one foot above
the ground in the crevice of a sandstone cliff at Castle Mound, Castle Mound State
Park, Jackson County, Wisconsin (T2IN, R4W, SWYsSWYs sec. 23). To my
knowledge, there are no reports in the literature of 7. simulata occurring on rock
cliffs. The plant consisted of only eight sterile fronds, one of which was taken as a
voucher (Moran 995, MIL). Two more fronds were taken on 13 Aug 1980 and
checked by Dr. W. H. Wagner, Jr. for correct identity (Moran 1267, MICH). One of
the fronds was soriferous, although all the sporangia were still white and immature;
maturation was late compared with several other Wisconsin populations that were
already actively sporulating.
Castle Mound is composed of Cambrian sandstone that rises 180 feet above the
surrounding lake plain of extinct glacial Lake Wisconsin. It is typical of many
weathered, castellated mounds found in the central Wisconsin sand plains. The T.
simulata cliff habitat on Castle Mound faces north and is shaded by Pinus resinosa,
Fr strobus, and Quercus rubra. The soil reaction in the crevice where 7. simulata
was growing was pH 5.0. No other vascular plants were growing in the crevice with
T. simulata. The cliff face immediately surrounding the plant was barren, except for
one plant of Dryopteris spinulosa growing in a crevice about one foot above Z
simulata. About ten feet above the plant was a ledge with numerous individuals of
Polypodium virginianum, along with Aquilegia canadensis, Athyrium filix-femina,
and Betula papyrifera. :
AMERICAN FERN JOURNAL: VOLUME 70 NUMBER 4 (1980) 137
Although it seems odd that 7. simulata should occupy a dry sandstone cliff when
its typical habitat is wet, acid bogs, usually in association with sphagnum moss, the
two habitats are similar in certain respects. The wet, cold acid conditions of a bog
make it physiologically difficult for roots to absorb water and mineral nutrients.
Such a state of “physiological drought” simulates the dry, nutrient-poor crevices of
a sandstone cliff. This type of habitat switch is known to experienced field botanists
from other examples of swamp or bog plants growing on rocks, and vice-versa. A
few such examples are: Cystopteris bulbifera, Dryopteris marginalis, D. spinulosa,
Ledum groenlandicum, Lorinseria areolata, Matteuccia struthiopteris, Osmunda
cinnamomea, Phegopteris connectilis, Sphagnum spp., and Thelypteris palustris.
Although T. simulata may have been common in the low wooded acid swamps
that surrounded Castle Mound in presettlement times, the nearest presently known
locality is two miles away. The Castle Mound individual certainly is the result of
relatively wide-range spore dispersal. It is important to point out that, in view of
Klewkowski’s (Science 153:305—307. 1966) ideas on the adaptive value of polyploidy
in homosporous pteridophytes, the Castle Mound individual is most probably the
result of single spore establishment and intragametophytic selfing.—Robbin C.
Moran, Wisconsin Scientific Areas Preservation Council, Department of Natural
Resources, P.O. Box 7921, Madison, WI 53707.
A SECOND ALABAMA LOCALITY FOR THE HART’S TONGUE.—The
discovery of the Hart’s-tongue, Phyllitis scolopendrium (L.) Newm., in a sinkhole
in Jackson County, Alabama (Amer. Fern J. 69:47-48. 1979) generated interest in
further searches for this fern among members of the Huntsville Grotto of the National
Speleological Society who had participated in the find. According to Eric Bachel-
der, my guide to the Jackson County locality, these spelunkers found a second,
larger population in a sinkhole in Morgan County soon after the original discovery
(Huntsville Grotto Newsletter 20:49-50. 1979). On 31 May 1980, I visited the new
locality with Mr. Bachelder again as my guide. The population is in a deep sinkhole
in the area known as Newsome Sinks, a large sink-valley in northeastern Morgan
County about 25 miles southwest of the Jackson County locality and 65 miles
southwest of the one in Marion County, Tennessee. The sinkhole is about 70 feet
deep and has sheer walls. A small stream falls into the sink, making the air very
misty and humid, unlike the dry Jackson County sink. Also unlike the Jackson
County sinkhole, it is necessary to rappel down to a wide ledge about half-way
own, where most of the Phyllitis plants are. Fifty-three plants occur on the ledge,
along with luxuriant C 'ystopteris bulbifera and Wood Nettle (Laportea canadensis),
which may have obscured more Phyllitis plants. At least 20 Hart’s-tongues were
mature adults; the juveniles ranged from sporelings to almost adults. The ledge is
Partially overhung by the cliffs above, and the Phyllitis plants grow in a narrow strip
beneath the overhang. The left end of the strip contains mostly adults and large
Juveniles; the plants toward the right are gradually reduced in size, age, and density.
Apparently the population is spreading towards the right. Four fairly large Hart’s-
tongues also were seen at the bottom of the sinkhole. A number of fronds were
collected as a voucher (Short 1/95, AUA and duplicates to be distributed).—John
W. Short, 905 McKinley Ave., Auburn, AL 36830.
138
AMERICAN FERN JOURNAL: VOLUME 70 (1980)
AMERICAN FERN JOURNAL
Manuscripts submitted to the JOURNAL are reviewed for scientific content by
one or more of the editors, and,
ften, b
During the past year we have received the kind assistance of J. Beitel,
L. G. Hickok,
I. Cousens, A. M. Evans,
y one or more outside reviewers as well.
B. M. Boom,
A. C. Jermy, J. H. Miller, J. D.
Montgomery, D. H. Nicolson, H. E. Robinson, J. Skog, A. R. Smith, A. Star, R. G.
Stolze, W. C. Taylor, R. A. White, D. P. Whittier, and J. J. Wurdack, to whom we
are deeply indebted. We welcome suggestions of other reviewers.—D.
INDEX TO VOLUME 70
Acrostichum, 51 Se seat tad 97: agen 65: ep apni
102.
u istii Pe pone: um
cladotrichum, 97; conforme. 55: crinitum, 63 denaciioltarn,
99, 108, 109: decoratum, 60; diversifolium. 97; ellipsoideum
97: engleri, 97: eximium ei, 60: fulvum, 97; gossyp
97. guamanianum, 97; hieronymi. 97: hickenii, 97; hirtum, 66
huacsaro. 62; hybridum, 63: latifolium. 55: litanum, 97
longissimum. 97: molle, 97; muriculatum, 97; muscosum. 62:
eandro 7. ovat: paleaceum, 60, 61: pangoanum,
: pellucidum. 97; petiolosum. 62: sg ge 97: pilosel-
loides, 65: pilosum, 61: pruinos 97; pte cio 97.
i Hloen
m: ii, 61: maxonii, 55; melanopus, 63: meride
ubsect. Microlepidea 62: moorei, 59: moritzianum
ity 62: obl | 64: ocoense x
. 61: hace ie 48, 53. 59:
ssa, 49, igo Gots Be S
—— Polylepidia, 50, 53. 60-62:
rent . 63: seti
. 66: elect: Setosa, 49-53.
bie teil. an siliquoides, 51, 64: sim-
63: spatulatum, 65: spor-
D. Sh. 53¢ 3B, 57, 59, OO
Way caneri. 61: wawrae, 55: wrightii. ~~ vatesih.
9. 41-43; arvense. 33. 42-44. 81: bogoten
oe Sapiens 3
33-35,
81
Equisetum, 3. 36. 3
2
oe
lustre, 42, 43: pratense, 42. 43: ecnorvie ri
, 43, 81;
42.
gang x ictale in Illinois, ial. Minnesota, and Wiscon-
E
— ELF range extension for aaa filix-mas, 113
= aan psa of Se I
‘airbrot
worothers, D, E. Mur aig R. L. Petersen)
: columbianu 59;
orderoanum, 62: cordifolium, eae costari-
9; sect. Craspedoglossa. 52. 55;
jo apinip ii 62: deci
oratu . 60:
etophytes, 93
Flora del Avila (rev.). 79
Flora de la provincia de Jujuy Republica Argentina. parte II.
Pteridophyta (rev. ),
Francis, P. C. (see R. L. Petersen)
Futyma, R. P. bie banrenee and ecology of Phyllitis scolo-
pendrium in Mic
Gametophytes of sem diffusum, 39
Gastony, G. J. The deletion of Vietria graminifolia from the flora
of Florida, on
Gleichenia anions 26: — 27
Pb:
Omez P. oths and ferns, 111
Gomez P.. L a & K. S. Walter. A double spore wall in
Macroglossum, 45 ;
Grammitis rigens, 26; taxifolia, 90: taenifolia, 90; xipho-
teroides, 26
renee tg dryopteris.
ei. Ga
mnie Equisetum diffusum, 39
Henin margina’ 9
Hill, R.. A new pi record for Pilularia americana in
exas, 28
How to know the ferns and ferns allies (rev.), 68
Huber, O. (see J. A. Steyermark)
Hunter, D. M. (see E. E. Karrfalt)
Hymenodium, 50, 63
Hymenophyllum caespitosum, 88: fucoides, 88: subg. Hymeno-
phyllum, 88: — rminieri, 88: subg. Mecodium, 88: myriocar-
um ndulatum, 88
o
an
Intersectional ap in owe 1
soé 2: drummondii. 7h: Sonera i‘
. 3) Var. iana, 3: . 1,2: mac wipers;
1-3: melanospora, 3: siesieadind, a eae 3
alt, ELE. . M. Hunter. Notes on the natural history of
69
Lellinger, D. B. Date of publication of Sodiro’s “Sertula Florae
Ecuadorensis.” 96: New names for Polypodium chnoodes and
P. nantes 30
loyd, R. M. Reproductive biology and tea apt
ogy New World populations of Acrostichum au
riopsidaceae, 48,
Lonchitis, 88. 89; hirsuta, 88
rine areolata, 137
Lyco m, 42: caracasicum, 79: clavatum, 73: lucidulum. 113:
cana
astern japonicum, 111, 112; smithianum, 115
Maatsch, R.. Das Buch der Fendi (rev.), 92
Macroglossum, 45, 46; alidae
Marattiales, sf 46
arsileac
hodedcia aes. FOy8 15 a SF
ay W. The economic uses = saci folklore of ferns
and fern allies (rev.)
Mertensia farinosa, 2
Mickel How to know the ferns and fern allies (rev.). 68
Mickel. J. T. & - spheate G. Subdivision of the
Elaphoglossum, 4
Microstaphyla eed
Miller, J. H. peed rences in the apparent permeability of spore
walls my Serer cell walls in Onoclea sensibilis, 119
Sandstone rock crevices, an exceptional new
habitat . Te simulata. 136
Moths wet fern
Munt "ke pa EB; sap nici arsnhiggnre and auotoxicity
in ser eastern Northern A n ferns, 124
Nephrodium cinereum. 98, var. in haicaiaeds: 98: kunthii, 89;
140
ry: 98
Jifolia ke — 89
A new count y reco n Te
New ache Pol shar hnoodes and P. dsm. =
A new record for Pellaea Detach in Mar . 30
New taxa and combinations of cndiatione: frou See
Mexico,
aierst on some ——- and Polypodium species of the
Notes on the natural history of wa gemmifera, 59
Oleandra, 49: articulata
a. 116, rap 20. 12 ay sersibiis. 5-81, rps 119
podum
s Ree fa i : awe eee 60:
cinnamomea, 73-77. 112. 116. 17, 125, 126. 129-134. 136.
137; claytoniana, 75, 116. 117. 126, 129-132: peltata, 59:
E25
: . J, Petrik-Ott)
Peck. J. H. Equisteum litorale in Illinois. lowa. Minnesota,
and Wisconsin, 33
Pellaea oo 30: cardiomorpha, 26: cordifolia, 26: ovata.
26
79; sa sie var. cordata,
Peart -_ 9.39
Petersen. R. L. & D. E. Fairbrothers. onan synthesis and
ntheridium initiation in Dryopteris gametophytes, 93; Recipro-
cal all ween the gametophytes of back da cin-
namomea and Dryopteris intermedia. 73
Petersen. R. L & P. C. — Differential oe of fern
and moss spores in response reuric chloride. 113
oe ee 2 & D. Or. ” Pilularia americana new to
Pancopbi ee 79
P
Phlebod . 125; aureum. en erythrolepis. 9
Psi "i. 137: scolopendrium, 81-87, 137, var. americana,
si
hace 28: americana. 28-30
Pil caine americana new to Tennessee. 29
de rogramma = m calomelanos. ae
P y pis. 5. 9: mac . 10. 26. var. ——
26, var. macrocarpa. 26. var. chop 10, 26: polylepis. 5,
pis, 9. 10
u =
hispidum 24: hygro ‘ometricum, 23:
um. 26: lycopodioi su ae
scutulatum, “98: sororium, 30, is. 5-8: var.
+ 5-8. var. thysanolepis 5-9: virginianum. 81.
136: vulgare, 122: x roides.
seen [ R. Supplemental notes on Lesser Antillean pterido-
Polyps. 5. os psn Take 49
Polysti : bra . 113; fournieri, 27:
lonchitis. 81. 82. 92 asta A 25: muelleri.
27: m sie 25; plat tea 25
Psilotum complanatum, 88: nudum. 88
AMERICAN FERN JOURNAL: VOLUME 70 (1980)
Soy 88; acts 75. 124, 125, 133. var. arachonideum,
88, var. pubescens,
Pteris, med 125: sent ee 98: biternata. 98: esmeraldensis, 98:
falcata, 98: japonica, 75: lucida, 89; longifolia, 75; —
111, 125: procera, 98: rigida, 98: rimbachii, 98: robusta, 98:
vittata, 125
ete is filix-m
Reciprocal ‘tpt pent the adelante of Osmunda
and Dryo as intermedia,
eae D. ‘E. A new record for Pellaea etropusputes in Mary-
hol of New World
land, 30
scone biology and g
populat “nee paris 2 urew
h der ee
=
VIEWS: 2: The economic uses and
sea Siti of ferns a
bio
de Jujuy Republica Argentina, parte II. Pteridophyta, 38: How
o know the ferns and vee - es. ae Taxon nomy of Thelypteris
subg. rei 2 (Pteridophyta), 98
hipidopteris.
Sandstone rock crevices, an exceptional new habitat for Thely-
Fciren simulata, 136
ond Alabama locality for the Hart’s-ton 137
sant 3; chiapensis, 25; See 25: panel: 25
Sho W. Di
lazium japonic to Alabama, I11: A
‘on Alabama locality for ee! Ha rt’s-tongue, 13
ew taxa and combinations ee perido fie from
‘Gas Mexico, 15: Taxonomy of Thelypter pra
pteris, including Glaphyropteris ( cpa ta) (
Smith, «1B. -E; “Wolford: & J: Li. Co sg
Athyrium fives eastern Tennessee, ate
Sota, E. R. de la Flora de la provincia de Jujuy Republica
Argentina, parte il. Pteridophyta (rev.). 38
Sphuseropteris brunei, 111
Steyermark, J. A. & 0. Huber. Flora del Avila (rev.). 79
ge gece
72: stoma oe 69-72
Supplemental notes on Lesser An weshist oe lophytes
er of eae sibs Steiropteris. inchadlsae cee
98
r aaa f
W. C. (see C. R. Werth)
rool heracleifolia, 27: incisa, 27.
subsp. transiens. 27:
transiens, 27
Thelypteris. = dentata, 75: amir nba he hispidula. 89.
var. hispidula, 89. var. inconstans. ; kunthii, 89: normalis.
7S. 112, tS noveboracnesis, 75. st pina 89: palustis.
tet simulata, 136, 137: el Steiropteris. 98: torresiana, 80.
Pettis
tela —— to the pteriodphyte flora of Escambia County.
Flori
Vittaria, 12: filifolia, 12, 13; graminifolia. 12-14: lineata, 12. 13.
90
Walter, K. S. (
Wendt, uae
the
. D. Gom
pen - some Pleopelis and Polypodium species of
rth, C. . WiC. Taylor: Asplenium x gravesii discovered
in shisha
ford. B. E(s see D. K. Smith)
wardia areolata, 112
ERRATA
Page 52, line 22: For “beaurepairii” read “beaurepairei.”
Page 55, line 39: For “tuckerheimii” read “tuerckheimii.”
Page 61, line 1: For “Petioloa” read “Petiolosa.”
Page 66, line 10: “For beaurepairii” read “beaurepairei.”
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AMERICAN
FERN
JOURNAL
Volume 71
1981
PUBLISHED BY THE AMERICAN FERN SOCIETY
EDITORS
David W. Bierhorst
Gerald J. Gastony
David B. Lellinger
John T. Mickel
MERCURY PRESS, ROCKVILLE, MARYLAND 20852
CONTENTS
Volume 71, Number 1, Pages 1—32, Issued March 30, 1981
Equisetum variegatum and E. X trachyodon in New Jersey JAMES D. MONTGOMERY
Comparative Ecology of Woodsia scopulina
Sporophytes and Gametophytes PAUL J. WATSON AND MARGARITA VAZQUEZ
Scale Insects Feeding on Farinose Species
of Pityrogramma ECKHARD WOLLENWEBER AND VOLKER H. DIETZ
Spore Germination and Young Gametophyte Development
of Botrychium and Ophioglossum in Axenic Culture DEAN P. WHITTIER
New Species of Moonworts, Botrychium subg. Botrychium
(Ophioglossaceae), from North America
W. H. WAGNER, JR. and FLORENCE S. WAGNER
Shorter Note: Range Extensions for Two Lycopods
on Baranof Island, Southeastern Alaska
Pteridophytes for the Flora Mesoamericana
Reviews g;
Volume 71, Number 2, Pages 33—64, Issued June 29, 1981
Azolla filiculoides New to the
Southeastern United States VERNON M. BATES, JR. and EDWARD T. BROWNE, JR.
The Genus Nephrolepis in Florida CLIFTON E. NAUMAN
The Branching Pattern of Hypolepis repens THERESA M. GRUBER
Diplazium japonicum and Selaginella uncinata
Newly Discovered in Georgia WAYNE R. FAIRCLOTH
Notes on Selaginella, with a
New Variety of S. pallescens ROBERT G. STOLZE
Lepisorus kashyapii in the
Western Himalayas S. S. BIR and CHANDER K. SATIJA
Taxonomic Notes on Jamaican Ferns-IlII GEORGE R. PROCTOR
~~ Notes: Notes on North American Lower
ascular Plants—II; Equisetum arvense
Alabama
10
13
48
Volume 71, Number 3, Pages 65—96, Issued September 30, 1981
Arachniodes simplicior New to South Carolina
and the United States JUDITH E. GORDON
A New Isoétes from Jamaica R. JAMES HICKEY
Leaf Turnover Rates and Natural History of the
Central American Tree Fern Alsophila salvinii RALPH L. SEILER
Nomenclatural Notes on Micronesian Ferns F. R. FOSBERG and M.-H. SACHET
x Asplenosorus shawneensis, a New Natural Fern
Hybrid Between Asplenium trichomanes and
Camptosorus rhizophyllus ROBBIN C. MORAN
Notes on North American Ferns DAVID B. LELLINGE
Shorter Notes: Salvinia minima New to Louisiana;
An Unusual Record of Asplenium trichomanes
from Northeastern Florida
Reviews
Suggestions to Contributors
Volume 71, Number 4, Pages 97—124, Issued December 30, 1981
Bog Clubmosses (Lycopodiella) in Kentucky
Chain Ferns of Florida TERRY W. LUCANSKY
Spore Germination Patterns in Anogramma, Bommeria, Gymnopteris,
Hemionitis and Pityrogramma
CLARK S. HUCKABY, R. NAGMANI, and V. RAGHAVAN
Shorter Notes: The Chemoidentity of the Holotype of
Pityrogramma triangularis; A Major Range Extension
for Thelypteris simulata in the Southern Appalachians;
A New Indiana Station for Epiphytic Resurrection Fern
Review
American Fern Journal
Index to Volume 71
Erratum
R
R. CRANFILL
69
101
109
MISSOURI BOTANICAL
APR 44 son]
AMERICAN ows
FERN | aed
J O U R N A @ January—March, 1981
QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY
Equisetum variegatum and E. x trachyodon in New Jersey JAMES D. MONTGOMERY
Comparative Ecology of Woodsia scopulina
Sporophytes and Gametophytes PAUL J. WATSON AND MARGARITA VAZQUEZ = 3
Scale Insects Feeding on Farinose Species
of Pityrogramma ECKHARD WOLLENWEBER AND VOLKER H. DIETZ 10
Spore Germination and Young Gametophyte Development
of Botrychium and Ophioglossum in Axenic Culture DEAN P. WHITTIER = 13
New Species of Moonworts, Botrychium subg. Botrychium
(Ophioglossaceae), from North America a eres :
W. H. WAGNER, JR. and FLORENCE S. WAGNER 20
Shorter Note: Range Extensions for Two Lycopods
on Baranof Island, Southeastern Alaska
Pteridophytes for the Flora Mesoamericana
Reviews
The American Fern Society
Council for 1981
ROBERT M. LLOYD, Dept. of Botany, Ohio University, Athens, OH 45701. President
DEAN P. WHITTIER, Dept. of Biology, Vanderbilt University, Nashville, TN 37235. Vice President
MICHAEL I. COUSENS, Faculty of Biology, University of West Florida, Pensacola, FL 32504
Secretary
JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, TN 37916.
Treasurer
JUDITH E. SKOG, Dept. of Biology, George Mason University, Fairfax, VA 22030.
Records Treasurer
DAVID B. LELLINGER, Smithsonian Institution, Washington, DC 20560. Journal Editor
ALAN R. SMITH, Dept. of Botany, University of California, Berkeley, CA 94720. Memoir Editor
JOHN T. MICKEL, New York Botanical Garden, Bronx, NY 10458. Newsletter Editor
American Fern Journal
EDITOR
DAVID B. LELLINGER Smithsonian Institution, Washington, DC 20560.
ASSOCIATE EDITORS
DAVID W. BIERHORST Rt. 3, Box 188, Picayune, MS 39466.
GERALD J. GASTONY Dept. of ae Indiana Padon Se Bloomington, IN 47401.
JOHN T. MICKEL w York Botanical Garden, Bronx, NY 10458.
The “American Fern Journal” (ISSN 0002-8444) is an illustrated quarterly devoted to the general
study of ferns. It is owned by the American Fern Society, and os at the Smithsonian Institution,
Washington, DC 20560. Second-class postage paid at Washin
Claims for missing issues, made 6 months (domestic) to 2 month (foreign) after the date of issue,
and the matters for publication should be addressed to the
Changes of address, dues, and applications for membership ae be sent to Dr. J. E. Skog. Dept. of
Biology, George Mason University, Fairfax, VA 22030.
Orders for back issues should be addressed to the Treasur
General inquiries concerning ferns should be addressed to ae Secretary.
Subscriptions $9.00 gross, $8.50 net if paid through an agency (agency fee $0.50): sent free to
members of the American Fern Society (annual dues, $8.00; life membership, $160.00).
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pages, $2.00 each; over 80 pages, $2.50 each, plus shipping. Back volumes 1979 et seq. $8.00 each;
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Dr. John T. Mickel, New York Botanical Garden, Bronx, NY 10458, is Librarian. Members
may borrow books at any time, the borrower paying all shipping costs.
Newsletter
Dr. John T. Mickel, New York Botanical Garden, Bronx, NY 10458, is editor of the newsletter
“Fiddlehead Forum.” The editor welcomes contributions from members and non-members, including
miscellaneous notes. offers to shoe or purchase materials, personalia, horticultural notes, and
reviews of non-technical books on fern
Spore Exchange
Mr. Neill D. Hall, 1230 rire 88th Street, Seattle. WA 98115. is Director. Spores exchanged and
collection lists sent on reque
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Gifts and bequests to the Society enable it to expand its services to members and to others interested
in ferns. Botanical books, back issues of the Journal, and cash or other gifts are always welcomed. and
are tax-deductible. Inquiries should be addressed to the Secretary.
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 1 (1981) l
Equisetum variegatum and E. xX trachyodon
in New Jersey
JAMES D. MONTGOMERY*
Equisetum variegatum Schleich., the Variegated Scouring-rush, occurs northward
from the northern tier of states of the United States into Canada, on damp sand or
shores, often on calcareous substrate (Fernald, 1950; Wherry, 1961). This paper
reports the first verified record of E. variegatum from New Jersey and discusses a
dubious record from that state. The hybrid E. X trachyodon A. Br. is also reported
for New Jersey.
Equisetum variegatum was listed for New Jersey by Taylor (1915) and Fernald
(1950), but not by Chrysler and Edwards (1947), Wherry (1961), or Hauke (1963).
The record for New Jersey was apparently based on an undated specimen collected
by C. F. Austin, labeled “Closter, Bergen Co.” Chrysler and Edwards (1947)
disallowed this record, indicating that E. variegatum and E. pratense were mounted
on one sheet and concluding that E. variegatum was mounted by mistake with E.
pratense. Further investigation by the author showed that there are specimens of
many pteridophytes from New Jersey collected by C. F. Austin, usually labeled only
“Closter, Bergen Co.” and without dates or specific locality. A total of 36 species of
the more common northeastern ferns and fern allies are represented at CHRB or NY
by such collections. It is interesting that specimens of plants known to be difficult to
grow or transplant (e.g., Lycopodium spp., Botrychium spp. other than B. dissectum)
are lacking. I suggest, therefore, that at least some of these C. F. Austin collections
represent garden plants which were originally collected at various places in western
New Jersey and probably elsewhere. This conclusion is supported by the fact that, in
addition to the mixed collection of E. variegatum and E. pratense, there is a sheet
of E. fluviatile at NY with labels from both “Closter, Bergen Co.” and “St.
Lawrence Co., NY,” and another undated sheet of FE. pratense labeled “Closter and
Sparta.” I agree with Chrysler and Edwards that the record for E. variegatum from
Closter, Bergen Co. is highly suspect. Other Closter records by this collector
without specific location or date are likewise dubious.
A hybrid involving E. variegatum has been known from New Jersey since at least
1950: E. x trachyodon is the hybrid between E. variegatum and E. hyemale L. As
far as is known from herbarium records, this hybrid was first collected by J. L.
Edwards along the Delaware River, near Flatbrookville, Sussex Co., 28 October
1950 (CHRB, NY).
On 19 August 1977, Vincent Abraitys collected plants of E. variegatum neat
Marksboro, Warren Co. I came across this material in connection with a project to
revise the Chrysler and Edwards book. Material was sent to Dr. Richard Hauke, who
verified the identification.
*Ichthyological Associates, Inc., R. D. 1, Berwick, PA 18603.
Volume 70, number 4, of the JOURNAL was issued December 30, 1980.
9 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
The discovery of E. variegatum in Warren County, New Jersey, represents a
southward range extension for this species. The plant is known in New York from a
collection made in 1960 along the Hudson River in southern Ulster Co. see 7
from central Columbia and Greene Cos. (NYS). The plant is more commo
central New York, and there are many records from the edges of the Adieaeer
(New York State Botanist’s Office, pers. comm.). The distance from the Ulster Co.
site is about 100 km (60 mi); other locations in eastern and central New York are
more than 160 km (100 mi) distant. The only location for EF. variegatum in
Pennsylvania is from Presque Isle, Erie Co., in the extreme northwestern corner of
the state (Wherry, Fogg, & Wahl, 1979).
Equisetum variegatum occurs on the damp shore of a lake underlain by limestone,
a habitat similar to that recorded in herbarium records from New York and Pennsyl-
vania
Mr. Abraitys reports that it is unlikely that plants were present about 1960, so this
is represents a recently established colony. The source of the plants and
means of introduction are unknown, although cultivation can be ruled out. It is
cet aa that both E. variegatum and E. X trachyodon should appear recently in
New Jersey. The localities are separated by approximately 15 km (9 mi). The
appearance of E. x trachyodon is more easily explained since the colony is on the
shore of the Delaware River which extends northward into the range of E.
variegatum in New York. Herbarium records (and observation of the E. variegatum
site) indicate that the hybrid became established separately from and earlier than the
species in New Jersey.
LITERATURE CITED
senior lee A., and J. L. EDWARDS. 1947. The Ferns of New Jersey. Rutgers Univ.
, New Brunswick, NJ.
FERNALD, M. L. 1950. Gray’s Manual of Botany, 8th ed. American Book, New York, NY.
HAUKE, R. L. 1963. A taxonomic monograph of the genus Equisetum subgenus Hippochaete. Nova
—123.
TAYLOR, N. 1915. Flora of the vicinity of New York. Mem. N. Y. Bot. Gard. 5:1—683.
WHERRY, E. T. 1961. The Fern Guide. Doubleday, Garden Cit Xs
OGG, Jr., and H.A. WAHL. 1979. Atlas of the Flora of Pennsylvania. Morris Arbore-
tum, Philadelphia, PA.
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 1 (1981) 3
Comparative Ecology of Woodsia scopulina
Sporophytes and Gametophytes
PAUL J. WATSON and MARGARITA VAZQUEZ*
Woodsia scopulina D. C. Eaton, an obligate rock fern, is widespread in the Rocky
Mountains and has a few disjunct populations in eastern North America. This report
concerns the ecology of W. scopulina gametophytes and sporophytes.
An understanding of the life history of any fern is incomplete as long as the
functioning of its gametophytes remains obscure. Yet, the study of gametophyte
ecology is in its infancy. Adaptations of gametophytes to cold and dessication have
been explored by Pickett (1914). Hill (1971) compared the habitat requirements for
spore germination and gametophyte development for three ferns in Michigan.
Gametophyte population divergence and general ecology have been studied by
Cousens (1979).
Wagner and Sharp (1963) found that free-living Vittaria prothalli occurred in
areas far north of their sporophytes. Since this discovery, several other genera of
tropical ferns have been found to possess gametophytes with geographic ranges much
more extensive than the sporophyte (Wagner & Evers, 1963; Farrar, 1967; McAlpin
& Farrar, 1978). Page (1979) sums up much of the research on fern gametophyte
ecology.
Gametophyte ecology is, of course, microecology. We find this a fascinating and
unusual level at which to study plant ecology, a science in which investigation of
systems at the macro level is the norm.
PROCEDURES
Our study areas consisted of xeromesic to xeric talus slopes and rock outcrops in
the immediate vicinity of Bigfork, Montana. These rocky sites represent typical
Woodsia scopulina habitat, and many hundreds of sporophytes as well as thousands
of gametophytes are found there. Cystopteris fragilis (L.) Bernh. also occurs at the
sites; however, it is only a minor contaminant of the nearly pure W. scopulina fern
communities. Other herbaceous plants are only sparsely distributed on the sites.
Shrubs such as Mountain Spray (Holodiscus discolor (Pursh) Maxim.), Serviceberry
(Amelanchier alnifolia Nutt.), and Rocky Mountain Maple (Acer glabrum Torr. )
occur, especially on the less disturbed sections of the talus slopes. Douglas-fir
(Pseudotsuga menziesii (Mirbel) Franco.) also occurs sparingly. A rich moss flora is
present on the talus and the rock outcrops.
We explored the microhabitats of numerous W. scopulina gametophytes, gameto-
phytes harboring juvenile sporophytes (in which the sporophytes’ leaves were still
dichotomous, not yet resembling those of the mature sporophyte), young, sterile
Sporophytes, and fully-developed, fertile sporophytes. For each of these life cycle
phases, we noted such environmental factors as substrate composition and pH,
sunlight exposure, and nearby plant associates. Consistently occurring differences in
the habitat of each of the four phases were recorded.
*532 University Ave., Missoula, MT 59801.
AMERICAN FERN JOURNAL: VOLUME 71 (1981)
FIG. 1. Talus hollows where Woodsia scopulina gametophytes are found. FIG. 2. Two clumps of soil
illustrating juvenile sporophytes (SP) in association with mosses and gametophytes (GP) growing on
soil,
WATSON & VAZQUEZ: COMPARATIVE ECOLOGY OF WOODSIA 5
Quantitative data on the substrate preferences of gametophytes were gathered by
examining all the cave-like talus hollows on a chosen talus slope. Sixty-one hollows
were inspected. For each, the presence or absence of gametophytes and nature of the
substrate (bare soil, litter on soil, moss and litter on soil, etc.) were recorded. (We
define litter as decomposing but still recognizable plant material upon soil or rock
surfaces.)
In order to compare the habitat preferences of W. scopulina gametophytes and
sporophytes with those of a typical mesophytic forest fern, we familiarized ourselves
with the habitats of the life cycle phases of Athyrium filix-femina (L.) Roth. The
study sites for this fern were located in the moister forested areas of the University
of Montana Biological Station, approximately 12 miles south of Bigfork.
RESULTS
The puberulent, slightly sticky gametophytes of W. scopulina were found only in
cave-like hollows on the talus slopes (Fig. /) and in crevices on rock cliffs and
tables. These talus hollows and rock crevices contain various amounts of soil, bare
rock, mosses, and litter. Gametophytes existed only on bare soil over rock (Fig. 2).
Gametophytes usually were found within talus hollows containing obvious patches of
bare soil (23 out of 29 hollows), but seldom were found in hollows with soil mostly
covered by litter or mosses (5 out of 32 hollows). Gametophytes were never found in
hollows lacking soil. A mere dusting of soil about a millimeter thick was enough to
support gametophytes. Soil pH readings from gametophyte substrates ranged from
6.4 to 7.2.
Gametophytes in the various populations were sparse to dense, and were not
closely associated with other plants, even with the young, leafy moss shoots.
Favorable gameotophyte habitats typically were sheltered from direct sunlight.
However, gametophytes did not grow so far back in hollows and crevices that
illumination was too heavily diminished, even if other environmental conditions
were favorable. Gametophytes were oriented with their apical notches farthest from
the opening of the crevice or hollow. Most gametophytes did not lie flat on the
substrate, but had the apical two-thirds of their thalli slanted steeply upwards (Fig.
3)
Populations of gametophytes giving rise to juvenile sporophytes also grew in the
crevices and hollows, but the soil substrate often was not bare (Fig. 2). This phase
of W. scopulina was usually associated with the moss Brachythecium velutinum
(Hedw.) B.S.G. This small, pleurocarpous moss grew sparingly around gametophytes
with new juvenile sporophytes but more robustly around those with more advanced
juvenile sporophytes having two or three well-developed leaves (in the latter case the
gametophytic tissue was still visable but totally chlorotic). Where B - velutinum grew
densely, a second moss, Encalypta vulgaris Hedw.., often occurred intermixed with
it. Occasionally, a thin layer of litter also covered the soil. Clusters of tiny juve-
nile sporophytes grew from gametophytes positioned in the more illuminated portions
of the microhabitat as, for example, near the front of talus hollows, but still out of
direct sunlight. Those gametophytes furthest back in such a hollow often had few or
no sporophytes.
F AMERICAN FERN JOURNAL: VOLUME 71 (1981)
Young, sterile sporophytes were seen growing only out of talus hollows, rock
crevices, and upon rock tables. When seen growing upon a rock table, examination
of the plants’ bases showed them to be anchored to at least a small crack or other
irregularity in the rock. These sporophytes usually grew in close association with
several mosses such as Dicranum scoparium (L.) Hedw., Rhytidiadelphus triquetrus
(Hedw.) Warnst., Tortula muralis Hedw.. Homalothecium sp., and Brachythecium
sp. Openings that harbored young, sterile sporophytes supported only one
AY
G wiv,
gh. "eich TED ONY f oS
.
ogo
biel 4 :
Aprreen :
AA\SO1L .SBORELING
GAMETOPHYTES Ay @
MMH
FIG. 3. Position of gametophytes and juvenile sporophytes within a typical talus hollow.
Fully developed, fertile sporophytes of W. scopulina were found growing out of
talus hollows, rock crevices, and from irregularities on rock tables and cliffs. Their
fronds always extended completely out of the rock opening of their origin, but grew
in filtered to direct sunlight, depending on the presence of overstory shrubs or trees.
Mature sporophytes had huge systems of fine roots. These roots were much greater
in extent and density than we expected for a fern so small as W. scopulina. The roots
never grew on bare rock. Instead, they grew within a layer of soil over rock and
spread down into even the thinnest layers of soil between rocks. Thick mats of the
same mosses associated with young, sterile sporophytes commonly covered the soil
substrate. Woodsia sporophytes were not associated with thick growths of grasses
and did not occur on sites with a great abundance of deciduous litter. The pH of the
soil substrates we tested ranged from 6.2 to 6.8.
The prothalli of Athyrium filix-femina were found only on bare soil or moist,
heavily rotted wood in shady microhabitats. As was true for W. scopulina gameto-
phytes, A. filix-femina gametophytes did not grow among mosses or upon forest litter.
Sporophytes of A. filix-femina occurred in mesic forest conditions, often beneath
gaps in the tree canopy. Mature sporophytes grew in association with many herbs,
shrubs, and thick layers of mosses.
WATSON & VAZQUEZ: COMPARATIVE ECOLOGY OF WOODSIA 7
DISCUSSION
We conclude that there are at least three salient differences in the ecologies of
Woodsia scopulina gametophytes and sporophytes: (1) Gametophytes become estab-
lished only in secluded microhabitats where sunlight is diffuse for most or all of the
day. Sporophytes, on the contrary, can tolerate exposure to intense direct sunlight for
many hours each day. (2) Competitive abilities of the two generations differ.
Gametophytes cannot coexist with other plant growth, including the mere leafy
shoots of small bryophytes. On the other hand, even the youngest sporophytes are
often surrounded by mosses with no apparent ill effects. Association with mosses
may even benefit sporophytes by reducing desiccation of their roots. Sporophytes
also grow well in close proximity to widely spaced herbaceous and woody
angiosperms. (3) Gametophytes cannot grow upon or under litter, but sporophytes are
not disadvantaged by moderate litter accumulations.
The above conclusions conflict with certain general statements in the literature,
such as that by Nayar and Kaur (1971) who claim that “sporophytes and gameto-
phytes have nearly the same ecological requirements.”
The first difference in ecological requirements mentioned above is a function of
place and, of course, results from sporophytes growing towards sunlight. Gameto-
phytes typically orient towards sunlight, but they do not grow towards it to any extent.
The latter two ecological differences mentioned above are a function of time rather
than place. The microhabitats that provide suitable conditions for gametophytic
growth contain patches of fresh, bare soil in talus hollows and rock crevices. Such
microhabitats are ephemeral and exist due to very recent accumulation of dust and
soil or to soil-churning rock movements. Sporophytes grow in the same places, but
after other plant life has invaded and litter has accumulated.
Although the sporophytes of W. scopulina and A. filix-femina are adapted to different
habitats, their gametophytes exist in strikingly similar habitats. Gametophytes of
both genera grow on bare substrates in relatively moist and shady microhabitats.
Difference in the sporophytes’ yet similarity in the gametophytes’ habitat require-
ments of these two ferns suggests that specialization of the haploid generation of W.
scopulina has lagged behind that of the diploid phase. This observation reinforces
the concept that the evolution of ferns is primarily a diploid affair (Wagner, pers.
comm.). It also is in harmony with the suggestion of Cousens (1979) that a potential
exists for some degree of independent evolution by the two generations. Cousens
suggestion is based upon the observation of Pray (1968) that differences among
populations of Pellaea andromediifolia gametophytes were not correlated with
differences in the sporophyte generation. Woodsia scopulina gametophytes are
evolutionary conservatives, developing only in the most mesic, forest-like
microhabitat available on the otherwise xeric, rocky macrohabitat to which the
sporophyte has become adapted.
8 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
Nonetheless, it may be that Woodsia gametophytes have made modest advances
towards becoming xerophytes. Unicellular glandular hairs and a slightly sticky
surface coating may be desiccation-inhibiting adaptations. (We note however that
Stokey (1951) states that hairs of this type are widely distributed and usually not of
generic significance.) Also, the nonoccurrence of Woodsia gametophytes on rotting
wood in the immediate vicinity of the study sites indicates that spore germination on
this substrate has been selected against. Perhaps the ability to grow on wood has
been traded for some xerophytic adaptation.
Our studies suggest that the frequency and abundance of W. scopulina gameto-
phytes in suitable microsites is greater than that of A. filix-femina gametophytes.
This has interesting implications. Perhaps ferns that do not reproduce extensively by
means of their rhizome system, such as W. scopulina, produce gametophytes of
greater vigor than ferns capable of such asexual reproduction and dispersal, such as
A. filix-femina.
In a study of this type, the tremendous individual mortality due to random
phenomena and intraspecific competition that may take place as plants of a species
struggle to advance from one developmental stage to another becomes highly
evident. For example, only those spores of W. scopulina that happen to land on
properly illuminated bare soil in specially protected areas among rocks will
germinate and develop into gametophytes. Spore wastage must be incredibly high. Of
the gametophytes that do develop, some are not in the right portion of the
microhabitat to give rise to new sporophytes, such as gametophytes found towards
the inner recesses of talus hollows. The threshold light intensity allowing gameto-
phyte development may be too low for the growth of new sporophytes. Alternatively, it
may be that not enough free water reaches the deeper parts of talus hollows to allow
much sperm transfer from antheridia to archegonia. Thus, innumerable gameto-
phytes are also wasted. Further, out of the mats of juvenile sporophytes that arise
from part of a gametophyte population, only one ultimately survives to occupy the
crevice or hollow as a young, sterile and latter as a fertile plant.
In W. scopulina, almost all intraspecific competition takes place between juvenile
sporophytes. New sporophytes promptly kill their parent gametophytes, possibly by
secreting a toxic chemical or by parasitism, thereby avoiding competition from other
sporophytes that otherwise might arise on the same gametophyte. Competition
between juvenile sporophytes of different gametophytes probably is mostly for
available light. The reason most intraspecific competition takes place between
juvenile sporophytes is because of the small size and spotty distribution of
microhabitats suitable to nurture the birth, growth, and continued survival of W.
scopulina sporophytes.
We extend deep graditude to Dr. W. H. Wagner, Jr., whose counsel made this
study possible. Special thanks are also due Dr. C. N. Miller for valuable support
throughout the study. We would like to extend our appreciation to the other readers
of our final report, Dr. D. E. Bilderback, Dr. F. Wagner, and R. Moran. We also
thank R. Hoham for aid in moss identification, Dr. C. A. Speer for providing
darkroom facilities, D. Watson for help in preparing photographs, and the students
and faculty of Yellow Bay Biological Station for their friendly support.
WATSON & VAZQUEZ: COMPARATIVE ECOLOGY OF WOODSIA 9
LITERATURE CITED
COUSENS, M. I. 1979. Gametophyte ontogeny, sex expression, and genetic load as measures of
population divergence in Blechnum spicant. Amer. J. Bot. 66:116—132.
FARRAR, D. R. 1967. Gametophytes of four tropical fern genera reproducing independently of their
sporophytes in the southern Appalachians. Science 155:1266-1267.
HILL, R. H. 1971. Comparative habitat requirements for spore germination and prothallial growth in
three ferns from southern Michigan. Amer. Fern J. 61:171-182.
McALPIN, B. & D. R. FARRAR. 1978. Trichomanes gametophytes in Massachusetts. Amer. Fern J.
68:97-98.
NAYAR, B. K. and S. KAUR. 1971. Gametophytes of homosporus ferns. Bot. Rev. 37:295-396.
PAGE, C. N. 1979. Experimental aspects of fern ecology. In A.F. Dyer (ed.). The Experimental
Biology of Ferns. Academic Press, New Yor :
PICKETT, F. L. 1914. Some ecological adaptations of certain fern prothalli—Camptosorus rhizophyllus
Link., Asplenium platyneuron Oakes. Amer. J. Bot. 1:477-498
PRAY, T. R. 1968. Interpopulational variation in the gametophytes of Pellaea androemedaefolia. Amer.
J. Bot. 51-960.
STOKEY, A. G. 1951. The contribution of the gametophyte to classification of the homosporous ferns.
Phytomorphology 1:39-58.
WAGNER, W. H., Jr, and R. A. EVERS. 1963. Sterile prothallial clones (Trichomanes”) locally
abundant on Illinois sandstones. Amer. J. Bot. 50:623.
_and A. J. SHARP. 1963. A remarkably reduced vascular plant in the United States. Science
142:1483-1484.
7 REVIEW
“FERNS, FERN ALLIES AND CONIFERS OF AUSTRALIA,” by H. T.
Clifford and J. Constantine. xviii + 150 pp. illustr. University of Queensland
Press. 1980. ISBN 0-7022-1447-7. $24.25.— Although this book is subtitled “A
Laboratory Manual,” and does contain illustrations of morphological details, it is a
good introduction to the pteridophyta of Australia, and, by extension, to the Old
World tropics. Two-thirds of the book concerns pteridophyta. For those interested in
identifying Australian ferns and fern allies, there are keys down to species,
interesting and useful generic descriptions with notes on habitats, and tables of
distribution by species within Australia and Tasmania. Species descriptions are
missing, and are not really compensated for in the brief keys to species. Fortunately,
the genera of Australian ferns are diverse and mostly with only a few species, and so
identification is likely to be easy in most cases. References and literature cited, a
table of vegetative characteristics of major vascular plant groups, a list of synonyms
of Australian pteridophyta with the names accepted by the authors, a short glossary,
and an index conclude this useful work —D. B. L.
10 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
Scale Insects Feeding on Farinose Species of Pityrogramma
ECKHARD WOLLENWEBER and VOLKER H. DIETZ*
Several well known species of the fern genus Pityrogramma are called Silverback
Ferns or Goldback Ferns because of the conspicuous white or yellow farinose
deposit on the abaxial surface of their fronds. This material is composed of
flavonoid aglycones, mostly of dihydrochalcones and chalcones (Wollenweber, 1978;
Wollenweber & Dietz, 1980). The correct structural formulae of these compounds
are shown in Fig. /; the formulae in both cited papers are incorrect. These lipophilic
phenolic compounds are secreted by glandular trichomes and form quasi-crystalline
rods or filaments on the surface of the enlarged terminal cell of the capitate glands
(Wollenweber, 1978, pp. 13-15).
R1
ea
|
On OU R! OH
R2
Chalcones { *
1. R'=OCH3,R2=H
2. R'=R*=0CH, OH O
Dihydrochalcones
7 8 OCH, Roe
4. R'=R*=0CH,
FIG. 1. Correct structural formulae of chalcones and dihydrochalcones, which form the major consti-
tuents of the farina on Pityrogramma fronds.
No doubt, glandular trichomes are anatomical features developed for some
definite function, and the natural compounds secreted ought not to be regarded
merely as waste products (cf. Swain, 1977). The farinose indument of Pityrogramma
has been the subject of speculation as to its physiological and ecological function for
about a century. Among the postulated functions are protection of the lower frond
surface and the young spores against wetting on the one hand, and against water loss
by transpiration on the other hand. Antibacterial and antifungal effects of such
*Institut fiir Botanik, Technische Hochschule Darmst: ittspahnstrz - é tadt,
Federal Republic of Germany. armstadt, Schnittspahnstrasse 3, D-6100 Darms
WOLLENWEBER & DIETZ: SCALE INSECTS ON PITYROGRAMMA 1]
phenolic substances have some likelihood. Insect deterrence also has been discussed
(Wollenweber, 1978). Field observations indicate that these ferns are rarely attacked
by feeding insects (L. D. Gomez, pers. comm.). Hoéhlke (1902) favored the latter
function of the farinose deposits because he had observed that “plants in the
greenhouse are free of destructive insects even in the summer.” There is as yet
no evidence for any of the foregoing functions; allelopathy from frond exudates has
been demonstrated (Star, 1980).
FIG. 2. Saissetia on the pinna costa and pinnules of Pityrogramma austroamericana, showing older and
younger glandular trichomes covered with exudate (Leitz Reprovit II). FIG. 3. Close-up of Saissetia
(Cambridge Stereoscan 600), x 30.
For several years, we have been growing plants of Pityrogramma austromaericana,
P. calomelanos, P. chrysophylla, and P. trifoliata in a greenhouse at Darmstadt. We
Saissetia (Coccidae) thrives on our Silverback and Goldback ferns. The insects are
found on the abaxial costae surface of old fronds, and also on the pinnule surface.
The insects seem to be not harmed or irritated by the rich flavonoid deposits, and they
grow and reproduce quite well while sitting in and between the exudate (Figs. 2 and
3). We are able to rid the plants of the insects for several months by removing infested
fronds.
It is not our intent to refute the possibility that flavonoid exudates may function to
deter some insects, even though they do not deter Saissetia. Observations of other
insects, especially in the field, would be welcome. We wish to thank Mrs. R. Heger
12 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
(Darmstadt) for the photograph and Dr. W. Bartholott ‘aie for the SEM
picture. We are grateful to Dr. D. R. Miller of the Dept. of Agriculture
(Beltsville) for determining the genus of the scale insect.
LITERATURE CITED
HOHLKE, F. 1902. Uber die Harzebehilter und die Harzbildung bei den Polypodiaceen und einigen
Phanerogamen. Beih. Bot. Centralbl. 11:8-45
STAR, A.E. 1980. Frond exudate flavonoids as allelopathic agents in Pityrogramma. Bull. Torrey Bot.
Club 107:146-153.
SWAIN, T. 1977. Secondary compounds as protective agents. Ann. Rev. Pl. Physiol. 28:479-S01.
WOLLENWEBER, E. 1978. The distribution and chemical constituents of the farinose exudate in
iaiaiglanig ferns. Amer. Fern J. 68:13-28.
——, and V. H. DIETZ. 1980. Flavonoid patterns in the farina of goldenback and silverback
ferns. Biochem: Syst. Ecol. 8:21-33.
REVIEW
“FERNS AND FERN ALLIES OF KENTUCKY,” by R. Cranfill, Kentucky,
Nat. Pres. Comm. Sci. Techn. Ser. 1. 284 pp. 1980. $4.50.—This new pteridophyte
Flora is a model for what state floristic treatments should be. General topics,
covered in about 35 pages, include collecting history, phytogeography and ecology,
pteridophyte life history, identifying pteridophytes, and a statistical summary of
Kentucky’s pteridophytes (69 species and 10 hybrids in 29 genera are known). The
technical treatment includes a synoptical key to families, a key to genera using fertile
and another using sterile material (the latter a very worthwhile novelty for ecolo-
gists), and treatments of the genera and species, including keys, synonymies, notes,
and illustrations. Although there are no descriptions, the keys are fairly extensive
and very thorough notes help to distinguish critical taxa. Distribution maps grouped
at the back of the book show the location of each species and the herbarium where
vouchers documenting the locations may be found. Seventeen herbaria were consult-
ed, as well as the literature, in compiling the maps. An illustrated glossary,
extensive literature cited, and an index conclude the volume. This book is a must for
anyone studying ferns in the east-central portion of the United States. It is available
rom the Kentucky Nature Preserves Commission, 407 Broadway, Frankfort, KY,
40601.—D.B.L.
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 1 (1981) 13
Spore Germination and Young Gametophyte Development
of Botrychium and Ophioglossum in Axenic Culture
DEAN P. WHITTIER*
Spores of the Ophioglossaceae have been sown by numerous investigators, but few
have succeeded in germinating them (Boullard, 1963). Du Buysson (1889) reported
germinating the spores of Ophioglossum vulgatum and Botrychium ternatum. The
spores of B. virginianum and three tropical species of Ophioglossum (O.
moluccanum, O. pendulum, and O. intermedium) were germinated by Campbell
(1895, 1907). Recent studies have shown that spores of B. dissectum f. obliquum
(Whittier, 1972) and B. multifidum (Gifford & Brandon, 1978) can germinate in
axenic culture.
Spores sown in axenic culture required darkness, took weeks to germinate, and, at
least for B. dissectum, produced a three-dimensional gametophyte immediately after
germination (Whittier, 1973; Gifford & Brandon, 1978). None of the earlier reports
by Campbell and du Buysson gave any indication that darkness was required for
germination. In soil cultures, O. moluccanum, O. vulgatum, and B. ternatum
germinated in a few days, which is a period comparable to that for spores of the
Polypodiaceae (Campbell, 1907; du Buysson, 1889). Young gametophytes of O.
vulgatum and B. ternatum were illustrated by du Buysson (1889) with filamentous
and two-dimensional growth habits which are similar to those for polypodiaceous
gametophytes. Consequently, the present study of additional species was carried out
in axenic culture to determine how the requirements for germination and the type of
early gametophyte development for these species compared with the results from
previous studies.
MATERIALS AND METHODS
Spores of eight Ophioglossaceae species, Botrychium biternatum ( Sav.) Underw.,
B. dissectum Spreng. var. dissectum, B. lunarioides (Michx.) Swartz, B. matri-
cariifolium A. Braun, B. virginianum (L.) Swartz, Ophioglossum engelmannii
Prantl, and O. vulgatum var. pycnostichum Fern. and var. pseudopodum (Blake)
Farw., were used in this study. With the exception of the spores of B. matricartifolium
and O. vulgatum var. pseudopodum, which were from Ontario, the spores were
collected in the southeastern United States. Voucher specimens are on deposit in the
Vanderbilt University Herbarium (VDB).
The techniques of Whittier (1973) were employed. The spores were sown on ?
ml of nutrient medium in culture tubes with a diameter of 20 mm. The tubes had
screw caps which were tightened to reduce moisture loss. The nutrient medium was
composed of Knudson’s solution of mineral salts, minor elements, FeEDTA, and
0.6% agar (Whittier, 1973). The medium was supplemented with 0.5% sucrose and
had a pH of 6.3. The spores were cultured at 24+1°C in the light at an intensity of
1400 lux from cool white fluorescent lamps or in darkness. For some treatments, the
cultures were maintained at 3+1°C in a cold room.
*Dept. of Biology, Vanderbilt University, Nashville, TN 37235.
14 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
For the first eight weeks after sowing, the spores were removed weekly from at
least six cultures of each species and examined for germination. After eight weeks,
the cultures were sampled at irregular intervals. The percentage of germination was
determined after examining 1000 or more spores.
Early gametophyte stages of development were cleared and stained with
acetocarmine-chloral hydrate according to Edwards and Miller (1972). The cleared
and stained gametophytes were drawn with a camera lucida for study. No effort was
made to indicate either the walls of cells lying behind illustrated cells or the spore
coat if one was present. However, the nuclei of cells positioned behind other cells
are denoted by dotted lines.
OBSERVATIONS
A few of the spores of most species studied germinated in the dark on the nutrient
medium at 24°C (Table 1). Lengthening the dark periods for those spores which
germinated under these conditions increased the amount of their germination. Even
after extended periods of time, the spores did not germinate in illuminated cultures.
TABLE |. GERMINATION OF SPORES OF THE OPHIOGLOSSACAE “ AXENIC CULTU
S
pecies Weeks in darkness Percent germination
Botrychium biternatum ~ 0.5
B. dissectum var. dissectum 8 0.1
B. lunarioides 3 0.7
B. matricariifolium 8 0.5
B. virginianum be 0
Ophioglossum engelmannii! 52 0
O. vulgatum var. pseudopodum 16 0.7
0
O. vulgatum var. pycnostichum 52
'See text as conditions which promote germina’
fy th of time in darkness bette: Ber ipation varies considerably. The fastest
pais was three weeks for B. lunarioides. The spores of O. vulgatum var.
pseudopodum took the longest time to germinate at 24°C; although the cultures were
not sampled on a weekly basis after eight weeks, initial gametophyte stages were
found in the cultures of this species after 16 weeks in the dark, which indicated that
the spores had germinated recently. Thus, under these conditions, spores of the
Ophioglossaceae do not have identical dark requirements for germination.
Sowing the spores in darkness at 24° did not promote germination in B.
virginianum, O. vulgatum var. pycnostichum, or O. engelmannii. However, 0.1% of
the spores of O. engelmannii germinated in the dark when maintained at 24°C for
three months, followed by 3°C for three months, and returned to 24°C for six
months. Germination did not occur during the cold treatment; consequently the final
warm period was necessary for germination. When the first warm period was omitted
and the spores were placed directly into the cold (3°C) for three months, no
germination occurred. Other modifications of the warm and cold temperature regime
are now under investigation in an attempt to increase the amount of germination.
Although successful for O. engelmannii, the regime of warm and cold temperatures
in the dark failed to bring about germination in B. virginianum and O. vulgatum vat.
pycnostichum.
D. P. WHITTIER: BOTRYCHIUM AND OPHIOGLOSSUM IN AXENIC CULTURE 15
The early developmental stages of gametophytes were studied in the five species
which germinated in the dark at 24°C (Table 1). Spores of these species exhibited
similar early development. The spore coat cracked open at the triradiate ridge and
the spore divided transversely to its polar axis (Figs. /, 14, 20 and 27), producing
a distal cell (away from the triradiate ridge) and a somewhat smaller proximal cell
(near the triradiate ridge). The proximal cell enlarged, forcing the three lobes of the
FS" (\B> fo
oS
BO
oe
wy
Is
=)
G
FIGS. 1-35. Stages in the early development of Botrychium and Ophioglossum gametophytes, x 275.
The bottom cell in Figs. 1-33 is the proximal cell. FIGS. 1-6. B. matricariifolium. FIGS. 7-13. B.
dissectum var. dissectum. FIGS. 14-19. B. lunarioides. FIGS. 20-26. B. biternatum. FIGS. 27-35.
O. vulgatum var. pseudopodum.
cracked spore coat apart, and bulged out of the spore coat. The distal cell, sch
remained inside the spore coat for a longer period of time, continued to divide. The
second division was longitudinal and divided the distal cell into two nearly equal
cells (Figs. 2, 7, 15, 21 and 28). The third division, which was either longitudinal
or transverse, occurred in one of the two distal cells (Figs. 3, 8, 16, and 29). The
fourth division occurred in the other cell at the distal end. Usually the plane of
division was perpendicular to the plane of the third division (Figs. 10, 17, 22, and
30) but occasionally its plane was almost the same as the plane of the third division
(Figs. 4 and 9). The development to the 5-celled gametophyte usually occured inside
the spore coat. Usually the gametophytes lost the spore coat by the 10-celled stage,
although it was seen occasionally on gametophytes with more —
16 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
The exact pattern of the later divisions was not followed for two reasons: there was
more variation in the sequence of the later divisions (Figs. 5, 6, 11, 12, 18, 23, 24,
and 25) and the three dimensional form of the young prothalli made it difficult to
follow the divisions. However, a small, more or less spherical or globular gameto-
phyte formed as a result of these later divisions (Figs. 13, 19, 33, and 36). The
mature gametophytes with morphology typical of the species developed from the
globular stage.
The early sequence of divisions in the gametophytes of O. vulgatum var.
pseudopodum was similar to that for the species of Botrychium. However, at about
the 7-celled stage (Fig. 32), the Ophioglossum gametophytes appeared to have an
apical cell. Side and apical views of 10-celled gametophytes showed definite apical
cells (Figs. 33 and 34). Older gametophytes had a typical apical cell with three
cutting faces (Fig. 35). Botrychium gametophytes of similar sizes or cell numbers
had no recognizable apical cells (Figs. 1/3 and 26).
FIG. 36. Young globular gametophyte of O. engelmannii, x 275.
The young gametophytes of O. vulgatum var. pseudopodum were larger than those
of Botrychium because the individual cells of the Ophioglossum gametophytes were
larger. More than likely, the larger cell size in Ophioglossum gametophytes is
controlled by the same factor which caused the spores of O. vulgatum vat.
pseudopodum to be larger than those of the Botrychium species. The average long
dimension for O. vulgatum var. pseudopodum spores was 43.8 ym; Botrychium
spore averages ranged from 31.0 to 34.9 um. The spores of O. vulgatum vat.
pseudopodum were not exceptional; those of the other Ophioglossum species studied
also were larger, with their longest dimensions averaging greater than 40 pm.
DISCUSSION AND CONCLUSIONS
The dormancy of those Botrychium and Ophioglossum spores which germinated is
broken by culturing them in the dark. Thus, the findings of Whittier (1973) and
Gifford and Brandon (1978) are corroborated by the results of the present study.
However, not all the spores germinate after a dark period at 24°C. In O. engelmannii,
germination occurs after a sequence of warm and cold periods in the dark. Although
the germination in O. engelmannii is limited and does not increase with extensions
of the final warm/dark period, the sequence of warm/dark, cold/dark, and warm/dark
periods is the only treatment which promoted germination. It remains to be
¢
D. P. WHITTIER: BOTRYCHIUM AND OPHIOGLOSSUM IN AXENIC CULTURE 17
determined if increasing the duration of the first warm/dark period and/or the
cold/dark period would increase germination in O. engelmannii. In B. virginianum
and O. vulgatum var. pycnostichum spores, there is no germination after a dar
period of 52 weeks at 24°C or following a sequence of warm and cold periods, as for
O. engelmannii. Although the dormancy of most species’ spores is broken by a dark
period, spores of O. engelmannii, O. vulgatum var. pycnostichum, and B. vir-
ginianum demonstrate that either a longer period in the dark at 24°C or a more
elaborate treatment is necessary for germination.
Campbell (1895, 1907) germinated spores of four species of Ophioglossaceae,
including those of B. virginianum, on soil or humus. These spores must have
germinated in the light because Campbell noted whether or not there was a trace of
chlorophyll in the young gametophytes. In three of the species, germination was
slow (taking a month or more), but the spores of O. moluccanum (now O.
petiolatum) from Java germinated in three days. Sussman (1965) suggested that
Campbell possibly used old spores which had lost their dormancy. At least for the
Ophioglossum species, this seems unlikely because Campbell ( 1907) discussed
collecting and sowing the spores in a relatively short time. The germination of these
Ophigglossum spores in the light is contrary to the findings for axenic culture. In
Lycopodium, on the other hand, the speed of germination varies considerably
(Barrows, 1935). Some of the more tropical species of Lycopodium tend to
germinate rapidly and on the surface of the soil, compared to the slower underground
germination of the more temperate species. Possibly a similar situation exists with
Ophioglossum, with the spores of the tropical species germinating more rapidly and
in the presence of light.
The spores of B. virginianum which germinated for Campbell (1895) but which
failed to germinate in axenic culture present a different problem. These spores are
from a temperate zone species in which dormancy is not broken in axenic culture.
Although the age of the spores used by Campbell is unknown, their age may not be
that significant because spores of B. virginianum up to two years old failed to
germinate in axenic culture. More time may be necessary for germination; Campbell
(1895), for instance, found early stages of germination after 18 months. Jeffrey
(1897), after failing to get germination after 18 months, suggested that the warmer
climate in which Campbell worked may have been a factor. Increasing the duration
of the experiments and employing higher temperatures along with other variations
are now being tested on the spores of B. virginianum. ae
The earliest stages of gametophyte development for the five taxa studied in detail
are consistent with the reports of Campbell (1895, 1907). The gametophytes in
axenic culture never produced a filament or plate of cells. Within a few divisions
and usually before the young gametophyte had completely broken out of the spore
Coat, a 3-dimensional growth pattern was established. Although Campbell (1907)
reported that the proximal cell usually divided in the early stages of the species he
Observed, the proximal cell remained undivided in the young gametophytes of the
present study. However, there are indications that the proximal cell may divide at
later stages in the development of the gametophytes in axenic culture. The undivided
proximal cell in the young gametophytes and the total lack of chlorophyll (because
18 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
the gametophytes in axenic culture were grown in total darkness) are essentially the
only differences found between the gametophytes from axenic culture and those
observed by Campbell (1895, 1907). All the species that have been studied have a
similar pattern of early development which produces a small, globular gametophyte.
Campbell (1907) suggested that an apical cell was established very early in
Ophioglossum gametophytes. In the present study, O. vulgatum, which has a
prominent apical cell in the mature gametophyte (Bruchmann, 1904), set off an
apical cell at possibly the sixth division. In slightly older gametophytes (10 or 11
cells), a characteristic apical cell with three cutting-faces was present. Campbell
(1907) observed the earliest stages of apical cell formation in side view, but
apparently he did not obtain apical views to confirm the presence of a typical apical
cell. No apical cell is found in the young gametophytes of Botrychium, which
probably is not surprising because the mature gametophytes of Botrychium are
generally without a prominent apical cell or sometimes even without a recognizable
apical cell (Bierhorst, 1958).
The report of du Buysson (1889) on spore germination and early gametophyte
growth of B. ternatum and O. vulgatum must be considered separately because of
his unusual observations. He reported germination times comparable to that for
spores of the Polypodiaceae, gametophytes with filamentous and two-dimensional
growth patterns, and young gametophytes with large amounts of chlorophyll. The
characters described by du Buysson (1889) are typical for the Polypodiaceae, but not
for the Ophioglossaceae. The possibility that spores of the Ophioglossaceae he sowed
never germinated and that stray spores of the Polypodiaceae (he was also studying
the development of other fern gametophytes) germinated instead would explain his
results. It is probably best to question or ignore the observations of du Buysson until
they have been repeated.
Some differences exist between the observations of Campbell and those from
axenic culture. Whether these differences are related to the source of the spores
(tropical as opposed to temperate) or to environmental variations (soil as opposed to
axenic culture) remains to be investigated. Nevertheless, the studies in axenic
culture basically substantiate Campbell’s descriptions of the early development of
gametophytes of the Ophioglossaceae.
I want to thank Mr. Alan Anderson (University of Guelph), Dr. Robert Kral
(Vanderbilt University), and Dr. R. Dale Thomas (Northeastern Louisiana Universi-
ty) for supplying spores of O. vulgatum var. pseudopodum, B. biternatum, and B.
lunarioides, respectively; Dr. W. H. Wagner, Jr. (University of Michigan) for
confirming the identification of B. biternatum; and the Vanderbilt University
Research Council for support of this research.
LITERATURE CITED
BARROWS, F. L. 1935 Propagation of Lycopodium. I. Spores, cuttings, bulbils. Contrib. Boyce
Thompson Inst. 7:267-294.
BIERHORST, D. W. 1958. Observations on the gametophytes of Botrychium virginianum and B.
dissectum. Amer. J. Bot. 45:1—9.
BOULLARD, B. 1963. Le gametophyte des Ophioglossacées. Considérations biologiques. Bull. Soc. *
Linn. Normandie 4:8 1-97.
D. P. WHITTIER: BOTRYCHIUM AND OPHIOGLOSSUM IN AXENIC CULTURE 19
ogee H. 1904. ae das Prothallium und die Keimpflanze von Ophioglossum vulgatum L.
, Zeit. 62:227-2
PAYSON. i du 1889. Monographs des cryptogames d’Europe. II. Filicinées. Rev. Sci. Bourbonnais
Cent. France 2:153—16
CAMPBELL, D. H 1895. The ee and Development of Mosses and Ferns, Ist ed. Macmillan,
dyin ork.
——— . Studies on the Ophioglossaceae. Ann. Jardin Bot. Buitenzorg, Il, 6:1
Trey yy E. and J. H. MILLER. 1972. Growth regulation by ethylene in fern 2p one Ill.
Inhibition a spore germination. Amer. J. Bot. 59:458—465.
GIFFORD, E. M., Jr. and D. D. BRANDON. 1978. Gametophytes of Botrychium multifidum as grown
n axenic clue Amer. Fern J. 68:71-75.
JEFFREY. E. C. 1897. The gametophyte of Botrychium virginianum. Trans. Roy. Canadian Inst.
—2 i
SUSSMAN, e S. 1965. Physiology of dormancy and germination in the propagules of i i
plants. Pp. 931- eo in W. Ruhland (ed.). Encyclopedia of Plant Physiology, vol. 15, pt.
Spri sateen 8
WHITTIER, D. P. 1972. igen! i of Botrychium dissectum as grown in sterile culture. Bot. Gaz.
133;336—339.
. 1973. The effect of light and other factors on spore germination in Botrychium dissectum.
Canad. J. Bot. 51:1791-1794.
20 AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 1 (1981)
New Species of Moonworts, Botrychium subg. Botrychium
(Ophioglossaceae), from North America
W. H. WAGNER, JR. and FLORENCE S. WAGNER*
Since the genus Botrychium was last revised (Clausen, 1938), much research on it
has been done, and many of our taxonomic ideas concerning the use of infraspecific
categories (subspecies, variety, form) have changed (Wagner, 1960). Many
Botrychium species can be distinguished only by subtle differences, and there is
often a high level of variability, even in single populations. In the littke Moonworts,
Botrychium subg. Botrychium, taxonomic difficulties are exacerbated by the small
size of the plants and the tendency for the segments to become folded when pressed.
To gain an idea of the distinctness of the species, careful field studies of sizeable
populations are needed, and the specimens must be pressed so as to spread out the
segments. Finding these usually rare and local plants requires “a labor of love.”
Together with colleagues and students, we have spent an extaordinary amount of
time exploring for Moonworts over the past several decades. Some of the fruits of
this quest are described here.
In making our taxonomic judgments, we have adhered to a strict requirement of
consistency. Plants from one locality to another must display the same characteris-
tics. Likewise, the differences from other species must be consistent; there should
be no connecting forms. We have emphasized the ability of two critical taxa to occur
together in the same habitats without intergrading. Mixed populations have thus been
of value in making decisions. We also have used the method of mutual associations—the
ability of two taxa to maintain their differences in their separate habitats, even
though their mutually associated relatives stay alike. Thus if taxon A grows with
taxon C in one habitat, and taxon B grows with C in another, and if taxon C remains
uniform in both habitats, then the differences between A and B are probably
genetically fixed.
The four new Moonworts described here have been studied in wild populations,
and in all cases the evidence upholds their validity as discrete species. We treat each
only briefly, but we hope that the description and comments will give other workers
a preliminary idea of their distinctions and also will lead to finding new populations.
More detailed reports on all of these plants will be made in the future.
The measurements are based upon the best-developed and most distinctive
specimens from each of several collections. Small specimens may be hard to
identify, and all of these species include tiny fertile individuals less than 3 cm tall.
Spores were mounted in Hoyer’s solution (Beeks, 1955) and measured along the
largest diameter. The ranges represent the averages of several different collections.
*Herbarium and Department of Botany, University of Michigan, Ann Arbor, MI 48109.
WAGNER & WAGNER: NEW SPECIES OF MOONWORTS 2]
B D
FIG. 1. Silhouette drawings of four new species of moonworts. A. Botrychium crenulatum. B. B.
paradoxum. C. B. mormo. D. B. montanum.
Botrychium crenulatum W. H. Wagner, sp. nov. Figs. 1A, 2.
Folium 10 (6—16) cm altum, herbaceum, flavo-virens; segmentum sterile dispositum
altum in axe, stipite 5 (1-17) mm longo, lamina ovato-oblonga vel lineari-oblonga, 2
(1.5-6.5) cm longa, 1.2 (8-18) cm lata, divisionibus lateralibus 3 (2-5) paribus,
cuneatis, remotis, 6 (3-12) mm longis, 5 (3-12) mm latis; margines crenulati, et
interdum paucincisi.
Leaf 10 (6-16) cm tall, texture herbaceous, color (living and freshly pressed)
yellowish green; common stalk 0.60 (0.50—0.80) of the total leaf length; sterile
segment stalk 5 (1-17) mm long, the blade (pressed) ovate-oblong to linear-oblong, 2
(0.5-6.5) cm long, 1.2 (0.8—1.8) cm wide; segments 3 (2-5) pairs, wedge-shaped, the
lower and upper margins of the medial pinnae an angle of 70° (20°—110°), their outer
edges separated by 80 (30—120)% of the width of the pinnae, often narrowly cuneate
toward the base, the largest pinnae 6 (3-12) mm long, 5 (3-12) mm wide, with 15
(10-25) veins reaching the distal margin; margins crenulate (entire to crenate or
dentate), and in very large specimens sometimes with I—5 incisions 1/5 (up to 1/2) of
the pinna length; fertile segment 4.5 (2.5-9.5) cm tall, the sporangial branches
confined mainly to the upper 1/2 (1/3-2/3), without major proximal sporangial
branches; spores 45 (40—48) pm in diam.; chromosomes 2n= 90.
TYPE: Mt. Baden-Powell Trail, Hamell Springs, San Gabriel Mountains, Los
Angeles Co., California, 7745 ft., L. L. Kiefer 1488 (MICH; isotype UCLA).
9 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
OTHER COLLECTIONS:
CALIFORNIA: San Bernardino Co.: San Bernardino Mts.: Santa Ana River: Big Meadows, Hilend
373 (LA), Bluff Lake Meadow, | specimen with B. simplex, Hilend 238 (LA), S Fork of Santa Ana
River, with B. simplex, Munz 6164 (DS). San Gabriel Mts.: N of Baldy Notch, headwaters of Coldwater
Fork of Lytle Creek, Kiefer 1489 (LA). San Gorgonio Wild Area: S Fork of Santa Ana River, ca.
halfway between Poopout Hill and Slushy Meadows, Kiefer 815 (LA). South Mountain: Fern Canyon,
branch of Mill Creek near Mill Creek Falls, opposite Vivian Creek Falls, Robertson in 1907 (DS).
Tulare Co.: Southern Sierra Nevada, ridge between forks of Monache Creek. SW of Olancha Fedak,
Munz 15333 (DS).
3 » *
/ | >
FIG. 2. Botrychium crenulatum from the type locality in the San Gabriel Mts., CA. A. View of living
plant, showing outline of fertile segment. B. Same, showing spacing and shape of sterile divisions. Leaf
8 cm tal
One might be tempted to treat this as a geographical subspecies of the worldwide
and variable Common Moonwort, B. /unaria L. However, there are several reasons
why we cannot do this. There are no intermediate populations, nor any evidence of
geographical transition. The new species is approximately equally distinct
morphologically from B. /unaria and the endemic North American tetraploid species
B. minganense Vict. (cf. Wagner & Lord, 1956). Botrychium crenulatum (Figs. 1A
and 2) may be distinguished from B. lunaria by its herbaceous (not fleshy) texture,
yellow (not dark-green) color, higher placement of the sterile segment on the leaf,
stalked rather than sessile sterile portion, an average of three pinna pairs (rather than
five), much more remote, smaller, and narrower segments, with fewer veinlets
WAGNER & WAGNER: NEW SPECIES OF MOONWORTS 23
terminating along the distal margin’, and the prevalence of crenulate, as opposed
to nearly entire, outer margins (except in B. lunaria f. subincisum (Roeper) Milde,
with the distal margins shallowly toothed, and f. incisum (Milde) Holmberg, with
pinnae more or less deeply cut; both usually are highly sporadic). The spores of B.
crenulatum average 6 wm larger in diameter than those of B. lunaria.
Superficially, B. crenulatum resembles B. minganense more than B. lunaria
because of the over-all proportions of the leaf and its parts. Several of the collections
have been identified as B. lunaria var. minganense (Vict.) Clausen. However, B.
crenulatum differs from B. minganense in its more delicate texture, the more abrupt
reduction of the apex, with fewer, coarser, and more angular, rather than more
rous, smaller, and rounder ultimate and penultimate segments, in the mostl
crenulate rather than mostly entire distal pinna margins, in the apex of the sterile
segment in the leaf primordium for the following year overtopping the fertile and
clasping it (in contrast to equally and paralleling it), and in the chromosome number
of 2n=90 rather than 2n = 180.
Whether the plant described here is the same as the one named B. lunaria subsp.
occidentalis by Léve et al. (1971) is questionable. They distinguish it from typical
B. lunaria in being smaller and having a yellower color. However, the type locality,
above Graymont in Clear Creek Co., Colorado, is remote from the known range of
B. crenulatum. We know of no localities in Colorado or its surrounding states where
B. crenulatum has been found. The specimens were probably depauperate sun-forms
of B. lunaria, judging from the characters given. Crenulations are not mentioned.
This Southwestern Moonwort is very rare, judging from the few collections that
exist. It may occur locally, however, in large numbers. Most of the specimens come
from southern California, especially San Bernardino County, at altitudes averaging
around 2500 (2100-3400) m. The fern extends in the mountains of California as far
north as Butte County, where we discovered it in 1949 in company with Dr. E. B.
Copeland. Possibly B. crenulatum has a wider range than given here. Plants we
previously identified as the South American B. dusenii (Christ) Alston from various
western states may prove to be variations of B. crenulatum. Also, in Oregon and
Montana, moonworts with extremely narrow segment divisions may be conspecific
or closely related.
Botrychium crenulatum grows in the drier places of damp meadows, boggy
areas, and marshes, either on hillsides or flat lands where there are wet banks or
springy spots. The plants are rooted in tussocks or “rises” around isolated trees or
shrubs, or in depressions that dry out during the summer, or at the edges of marshes.
They may occur either in sun or shade, but evidently prefer partial shade.
The associated genera that have been recorded include woody plants Pinus and
Salix, and herbs Dodecatheon, Hypericum, Liparis, Mimulus, Smilacina, and
Veratrum. Various sedges and grasses under which these moonworts grow can make
detecting the plants very difficult. Associated species of grapeferns include com-
monly B. simplex, and rarely B. multifidum.
‘Some collections (e.g., Wagner 4609, Johnston 2080) have relatively wider segments, with angles
approaching those of B. lunaria.
24 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
Botrychium paradoxum W. H. Wagner, sp. nov. Figs. 1B, 3.
lateral branches are sessile or nearly so and short, usually only 1-3 mm long;
sporangia mostly 2 or 3 (1-12) per lateral branch. Spores 40 (36-43) wm in dia-
meter. Chromosome number unknown.
ad
FIG. 3. Botrychium paradoxum. A. Small form of deep shade. Marias Pass, MT. Leaf 3 cm tall.
Large form of open meadows from the type locality at Storm Lake, MT. Leaf 15 cm tall.
TYPE: Storm Lake, ca. 6 mi S of Georgetown Lake, Deerlodge National Forest,
Flint Ridge Mountains, Deerlodge Co., Montana, Wagner 80/28 (MICH; 9 plants
observed).
OTHER COLLECTIONS:
CANADA: Alberta: Waterton Lakes National Park, 1 mi NW of Red Rock, Blair & Nagy 1280 (LEA: 6
plants observed). U.S.A.: Montana: Glacier Co.: 2 mi W of N end of Swiftcurrent Lake, Many Glaciers Trail
to Fishermancap Lake above Wilbur Creek, Wagner 78547 (MICH: 2 plants). Ca. | mi W of Marias Pass,
Pondera and Flathead county line, Wagner 78528, 80/17. (MICH; 45 plants).
WAGNER & WAGNER: NEW SPECIES OF MOONWORTS 25
This extraordinary Moonwort is surely one of the most peculiar ferns in North
America. It shows modifications of the leaf as profound as, for example, those in
Schizaea pusilla (Schizaeaceae). Two species of Ophioglossum lacking sterile
segments have been described: O. kawamurae Tagawa from Osumi Province, Japan,
and O. lineare Schlechter & Brause from New Guinea. These plants are
morphologically different from Botrychium paradoxum, however, in that the sterile
segment has been lost rather than transformed. The Sumatran O. simplex Ridley has
vestiges of the sterile segment (Tagawa, 1939; see also Sahashi, 1980). The fact that
bladeless species of the sister genus Ophioglossum have been reported points up the
fact that these are strongly mycorrhizal plants that may lose much or all of the
photosynthetic lamina. In Botrychium we can find a progression from forms like B.
lanceolatum (Gmel.) Angstr. to such extremes as B. montanum and B. mormo with
the ample blade replaced by a narrow, much reduced blade. In B. paradoxum, the
sterile lamina is not really lost, but has become converted to a second fertile
segment. The two fertile segments are both apparently positively phototropic and
negatively geotropic, so that they are held erect and parallel. In other words, what
might be an expected loss of the sterile segment, as illustrated partially in B.
montanum, B. mormo, and Ophioglossum simplex, and completely in O. kawamurae
and O. lineare, does not take place.
In regard to the western flowering plant, the Leafless Wintergreen, Pyrola aphylla
Smith, Camp (1940) wrote that “...the Pyrolaceae are on the physiological
borderline between autophytism and parasitism,” and he showed that several species
of wintergreens were capable of developing aphyllous forms. Similar physiological
conditions no doubt prevail also in the Ophioglossaceae. All of the fronds we have
studied are without laminae. It is interesting to note, however, that there is a
distinctive Moonwort, the sterile segments of which have remote and narrow
divisions, that has been found at two of the four Botrychium paradoxum localities.
This may represent the atavistic form of B. paradoxum, but our evidence for this 1s
tenuous; it may be merely a coincidence. eee
The four localities for B. paradoxum are all near the Continental Divide,
extending 200 miles (325 km), and the altitudes range from ca. 1700 m to 2500 m.
Plants of this species are very scarce. In spite of intensive searches by many
people, the numbers of specimens observed are low, as noted in the citations of
collections above. The plants occur in diverse habitats with two very different
extremes—open, sunny meadows and closed, shaded fireweed clones. The Waterton
Lakes specimens were taken in a moist drainage area on a grassy slope and those
from Storm Lake in meadows only a few feet from the shore. The Marias Pass
Population grows in black muck and rotting plant materials under a dense cover.
Prominent woody plants growing near Botrychium paradoxum include Pinus
contorta, Picea englemannii, Abies lyallit, Potentilla fruticosa, and shrubby species
of Salix. The herbaceous vegetation is very different between the open and closed
closed sites are dominated by Epilobium angustifolium (Fireweed), Geum
macrophyllum, Heracleum lanatum, Osmorhiza occidentalis, and Senecio foetidus,
% AMERICAN FERN JOURNAL: VOLUME 71 (1981)
as in the habitat near Marias Pass. Weeds are found in small numbers (Taraxacum
officinale and Bromus inermis). Our largest specimens of Botrychium paradoxum
were found in open sites, where they reached 15 cm in height with stalks 2.0 mm in
diameter (Fig. 3A). The heavily shaded plants of B. paradoxum are very delicate,
reaching only 7 cm in height with stalks 0.5 mm or less in diameter (Fig. 3B). Some
individuals reached only 3 cm in height and had fertile branches only 1-2 mm tall
with only | or 2 sporangia each. The latter appeared to be very young juveniles.
In the Fireweed habitats, plants are easily found by parting the large herbs and
searching the soil below. But in the open meadows, the plants are challenging to find
because of close interlacing with other vegetation and the presence of “look-alikes.”
The inflorescences of various small grasses, sedges, and plantains may superficially
resemble the fertile segments of B. paradoxum. The inflorescences of the tiny
knotweed, Polygonum (Bistorta) viviparum are especially troublesome.
Botrychium mormo W. H. Wagner. sp. nov. Figs. 1C, 4, 5.
Folium 8.6 (7—12.5) cm altum, succulentissimum, flavo-virens, nitidum;
segmentum sterile lineare, 2 (1.3-4.1) cm longum, 5 (3-7) mm latum, stipite |
(0.51.6) cm longo, lobis 2 (1-3) paribus, acutis vel truncatis, marginibus distalibus
integris vel parum crenatis, nec acute dentatis nec irregulariter laceratis; segmentum
fertile 4.5 (2.5-7.5) cm altum, plerumque in parte tertia proximali ramosum
Gametophyte commonly persisting at the bases of even the largest plants; leaf 8.6
(7-12.5) cm tall, very succulent, yellow-green, shiny; the common stalk making up
50 (20-70)% of the total length; sterile segment linear, 2 (1.3-4.1) cm long, 5
(3-7) mm wide, the stalk 1 (0.5—1.6) cm long; lobes 2 (1-3) pairs, round-pointed to
truncate, the distal margins entire or shallowly crenate, not sharply dentate or
irregularly lacerate, and with no tendency for exaggerated basal lobes; the tip
usually with 2—4 angular triangular or squarish lobes; fertile segment 4.5 (2.4—7.5)
cm tall, commonly branched in the lower third, the branches 1/3 to 2/3 as long as
the main axis; sporangia large, sunken, not opening until late September and
October, the aperture narrow, only 15—30°; spores 49 (45-53) wm in diameter;
chromosomes 2n= 90.
TYPE: Rich woods on W side of route 9, 8.15 mi N of Bagley City Park,
Clearwater Co., Minnesota, Wagner 79314 (MICH).
OTHER COLLECTIONS:
MICHIGAN: Alger Co.: Grand Sable Lake, Hagenah 2550 (BLH). Gorge at Sable Falls, near Grand
Marais, Hall & Hagenah 83] (BLH). Cheboygan Co.: Witkout definite locality, Hagenah 25/1] (BLH).
0.5 mi WSW of Riggsville Corners, Wagner 8062 (BLH). Chippewa Co.: E of Trout Lake, Hagenah
3480 (BLH). Dickinson Co.: 3 mi N of Norway, Hagenah 2920 (BLH). Luce Co.: N of Hendriks
Quarry, Hagenah 3205 (BLH). Otsego Co.: Without definite locality, Hagenah 2717 (BLH). MINNE-
SOTA: Becker Co.: N side of Route 143, NE of Twin Island Lake, Wagner 79327 (MICH). Beltrami
Co.: S and N of Route 1, ca. 1.5 mi E of Clearwater Co. line. Wagner 73196 (MICH). Cass CoN
side of Leech Lake, Ottertail Peninsula, Trana 756/5 (MIN). Clearwater Co.; U. of Minn. Biol.
Station, Lake Itasca, Bearpaw Point, Wagner 73104 (MICH: | specimen mixed with B. minganense),
Rosendahl! 5929 (MIN). Lake Itasca, Garrison Point, Wagner 7329] (MICH). Route 200, ca. 1/3 mi
of Wild Rice River, Wagner 73127 (MICH). Mahnomen Co.: E side of county road 4, | mi N of Route
200, Waterway 122 (WIS). WISCONSIN: Ashland Co.: Chippewa Twp.. Sec. 10, Peck 79-590
(UWL), Moran 917 (MIL). Forest Co.: N of Route 8, 2 mi W of Crandon, Wagner in 1951 (MICH).
Iron Co.: 3.5 mi NW of Hurley, R. M. Tryon 4023 (WIS). Wood Co.: Arpin, Goess! 3001 (MIL).
WAGNER & WAGNER: NEW SPECIES OF MOONWORTS
FIG. 4. Botrychium mormo, Mature plants. A. Leaf 8 cm tall. Note low branching of fertile segment.
B. Leaf 7 cm tall. Note form of segment divisions. Lake Itasca, MN.
FIG. 5. Botrychium mormo. Young plants 1-1.5 cm tall. Note highly reduced sterile and
; N
segments at tip of petiole, with only 2-4 sporangia. Lake Itasca, MN.
28 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
We have studied this Moonwort for three decades and have determined that it is
totally consistent. Among the many curious features of B. mormo, we include the
tendency for persistence of the gametophyte even on full-sized sporophytes, the
narrow range, the habitat in mesic forest, and the disappearance of the plants during
drought periods. Figure 4 shows well developed, mature specimens and Fig. 5 shows
the rudimentary, immature specimens. The extremely “chubby” form is nicely
illustrated by Peck (1980).
This, together with the following species, may constitute a distinct section of
subg. Botrychium. Although some small leaves may resemble juvenile or shade
specimens of B. simplex E. Hitchc., the fully developed leaves show important
differences. Mature individuals have the sterile segment higher on the axis, unlike
mature B. simplex (“Blade inserted almost basally or toward the middle of the
plant,” Clausen, 1938). The sterile segment has lower divisions approximately equal
to those above, but in B. simplex these are normally enlarged in mature individuals,
in the fullest development producing a ternate blade. The strongly truncate and
adnate lateral lobes of the two new species differ from the rounded lobes and
spatulate to lunate, more or less stalked pinnae of B. simplex. (Indeed, many
medium and large specimens of B. simplex have middle and upper lateral divisions
so flabellate as to resemble B. /unaria). The tip of the sterile segment is usually
toothed to deeply cleft, unlike the mainly undivided and entire tip of B. simplex.
The usual habitats of B. mormo and B. montanum are shaded forest floors under
mature trees, rather than those of B. simplex, which are open, marshy places,
meadows, and edges of wet woodland pond shores.
On the campus of the Lake Itasca Biological Station in Minnesota, sizeable
populations of both B. mormo and B. simplex are found within a quarter of a mile of
each other. Class studies have revealed over twenty differences between these
species.
Botrychium mormo is very difficult to find and apparently very rare. It if does
occur in areas other than Michigan, Wisconsin, and Minnesota, it has been
overlooked. Our students have made extensive searches for the plants in many
localities. We estimate that only one in fifty seemingly suitable sites have yielded
specimens.
The plant grows in rich leaf mold in Basswood (Tilia americana)—Sugar Maple
(Acer saccharum) forests. East of Marquette, Michigan, these dominants are joined
by Beech (Fagus grandifolia). The little plants push their way through the leaf litter
or simply lie under the litter, failing to appear at all. The goblinish appearance and
behavior of this odd plant has inspired the English name of Little Goblin and the
scientific epithet mormo. Often very small plants less than 1.5 cm tall (Fig. 5) will
dominate a population. In drought years, even large plants may fail to send up a leaf,
or one has to scrape off the litter to find any plants, these whitish and apparently
lacking chlorophyll. It is not clear why as many as half the individuals, including
specimens of all sizes, retain gametophytes at their bases. These are readily detected
as swollen, brown masses 1-6 mm long, protruding among the roots.
The grapeferns that most commonly grow with or near B. mormo are B.
virginianum and B. minganense, and more rarely B. lunaria, B. lanceolatum, B.
WAGNER & WAGNER: NEW SPECIES OF MOONWORTS 29
matricariifolium, B. dissectum, and B. multifidum. None of these other species
shows the peculiarities of B. mormo such as the persistent gametophytes, extremely
succulent texture, and peculiar shiny yellow color. We conclude, therefore, that the
habitat is not responsible for its distinctive characteristics. They are evidently
genetically fixed.
Botrychium montanum W.H. Wagner, sp. nov. Figs. 1D, 6.
Folium 8.7 (4-12.5) cm altum, herbaceum, glaucum, hebes; segmentum sterile
FIG. 6. Botrychium montanum from the type locality at Swan Valley, MT. Largest plants 8 cm tall. Note
Shape of divisions of sterile segments.
Leaf 8.7 (412.5) cm tall, herbaceous, glaucous, dull; the common stalk making
up 60 (40-90) per cent of the total length; sterile segment oblong to linear, 2.1
(0.7-4.0) cm long, 5.5 (2-9) mm wide, the stalk 0.7 (0.3-1.5) cm long: lobes 3
(1-6) pairs, irregular, narrow and pointed, oblong, square, often grouped or
confluent, the distal margins irregularly toothed to lacerate, frequently with ew
teeth, basal lobes lacking tendency for enlargement, the blade tip usually with 2-
30 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
TYPE: Crane Mountain Rd. (Route 498), 3.6 mi S of junction with Ferndale
Road, Swan Valley, Lake Co, Montana, cedar swamp forest, Wagner 80/10 (MICH).
OTHER COLLECTIONS:
MONTANA: Flathead Co.: Glacier National Park, Johns Lake, Wagner 78510 (MICH). S end of
Lake McDonald, N of Apgar, Wagner 78522 (MICH). Avalanche Creek, Wagner 78532 (MICH). Lake
Co.: Station Creek, near Flathead Lake, Wagner 78534 (MICH). W side of Cedar Bay Road, Wagner
80114 (MICH). Soup Creek, | mi E of Route 83, Wagner 80121 (MICH).
Botrychium montanum differs from B. mormo in characters that are especially evident
in living plants (Fig. 6) The leaf is herbaceous (not succulent), glaucous (not eee
green), and dull (not shiny). The position of the sterile segment along the vertical axis
tends to be higher, and the fertile segment only rarely branches near the base. The
cutting of the sterile segment is most distinctive: the lobes are more deeply cut and
irregular in pattern, with some narrow and some broad (due to grouping or confluence),
and most have sinuses of various sizes. Sporangial dehiscence takes place a couple of
months earlier than in B. mormo, and the sporangial valves open wider. We were unable
to 88 persistent gametophytes in.B. montanum as we did in B. mormo
rychium montanum is relatively easy to find, and we are surprised at the paucity of
pone collections. A single locality may have hundreds of plants in a small area (Fig.
6). It is most abundant in moist, springy Western Red Cedar (Thuja plicata) forests. The
species may grow in habitats which are quite different, however, as along grassy trail
edges (Logan Pass and Many Glaciers, Glacier National Park). Although all of the
known localities for B. mormo are at less than 1500 ft (450 m) altitude, B. montanum
grows at altitudes from 3200 ft (970 m) at Swan Lake to 6000 ft (1800 m) at Logan Pass.
e fern may occur in pure stands, but generally it is associated with B. virginianum
and more rarely with B. lunaria, B. lanceolatum, B. boreale, and B. minganense.
We acknowledge the support of NSF Grant DEB 800 55 36, “Evolution and
Systematics of the Grapefern genus Botrychium.” William R. Anderson kindly translated
the Latin diagnoses. A large part of the information we have about the species described
here was obtained by our students in Pteridology courses at the Lake Itasca Biological
Station, Minnesota, and the Flathead Lake Biological Station, Montana.
LITERATURE CITED
BEEKS, R. M. 1955. Improvements in the squash technique for plant eigen Aliso 3:131-133.
CAMP, W. H. 1940. Aphyllous forms in Pyrola. Bull. Torrey Bot. Club 67:453—465.
CLAUSEN, R. T. 1938. A Monograph of the Ophioglossaceae. Mem ater a Club 19(2):1-177.
LOVE, A., D. LOVE, and B. M. KAPOOR. 1971. Cytotaxonomy of a century of Rocky Mountain
orophytes. Arctic and Alpine Res. 3:139-165.
PECK, J. H. 1980. Discovery of the Goblin Fern in Wisconsin. Bull. Bot. Club Wisconsin 12:2-4 +
cover illustration.
SAHASHI, N. 1980. Morphological and taxonomical studies on Ophioglossales in Japan and the
se regions (4). Comparative morphology of spores of some species in Ophioglossales. J.
Jap. Bot. 55:73-80.
TAGAWA, M. ne Ophioglossum kawamurae Tagawa, a new species from Japan. Acta Phytotax.
Geobot. 8:134—136.
WAGNER, W. H., Jr. 1960. Evergreen grapeferns and the ie of infraspecific categories as used
in North American pteridophytes. Amer. Fern J. 50:32—4
, and L. P. LORD. 1956. The morphological and cytological distinctness of Botrychium
minganense and B. lunaria in Michigan. Bull. Torrey Bot. Club 83:261—280.
_~
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 1 (1981) 31
SHORTER NOTE
RANGE EXTENSIONS FOR TWO LYCOPODS ON BARANOF ISLAND,
SOUTHEASTERN ALASKA.—Early in 1976, I discovered a colony of Tree
Clubmoss (Lycopodium dendroideum Michx.) at Thimbleberry Bay, about 5 km
southeast of Sitka, Alaska (57°02’N, 135°16’W). Sitka is located on the outer coast
of Baranof Island, one of the larger islands of southeastern Alaska. The small,
vigorous colony of about five plants grows on a moss-covered rock bench about | m
above the extreme high tide line. This collection re-extends the range of the species
in southeastern Alaska about 200 km to the northwest. It is the first authenticated
report of L. dendroideum from the large outer islands. The voucher collected in 1979
(Muller 2854) is in the herbarium at the University of Alaska Museum, Fairbanks,
Alaska (ALA).
During the early 1840’s Eduard Blashke, a physician and botanist stationed in
Sitka, found L. dendroideum. Franz Joseph Ruprecht, curator of the herbarium at St.
Petersburg, Russia, published Blashke’s sighting. However, in 1941, Eric Hulten
expressed doubts about the century-old report of L. dendroideum at Sitka (Hulten,
E. 1941. Flora of Alaska and Yukon, |. Lunds Univ. Arsskr. N. F., avd. 2, vol. 37.
pp. 69-70).
The species occurs at widely scattered localities in the main part of Alaska and is
known to grow in the Alaska panhandle (southeastern Alaska) south and east from
Wrangell. A previously unreported collection of L. dendroideum was made in June
of 1973 at Thomas Bay east of Petersburg, a northward range extension of 65 km
within the Alaska panhandle. The specimen (Robuck 1403) is in the herbarium of
the U.S.D.A. Forest Service, Forestry Sciences Laboratory, Juneau, Alaska.
In October 1980, several colonies of the Bog Clubmoss (Lycopodium inundatum
L.) were found about 25 km south of Sitka, 1/3 km northwest of Big Bay at the
southeastern end of an unnamed lake system (56°49°N, 135°21’W). These plants
were growing on a thin muskeg mat overlying granitic rock at the edge of a shallow
lake. Vouchers (Muller 4181) are at ALA and the University of Washington
herbarium (WTU). In Alaska, the plant is known to occur along lake shores and in
muskeg areas in the vicinity of Ketchikan and Wrangell. The Big Bay collection is a
range extension of about 200 km to the northwest. This is the first time L.
inundatum has been reported from the large outer islands of the Alaska panhandle.
Since southeastern Alaska is sparsely settled and travel is difficult and expensive,
a lack of botanical information is not surprising. More range extensions can be
expected as botanists investigate southeastern Alaska more thoroughly.—Mary Clay
Muller, Chatham Area, Tongass National Forest, P.O. Box 1980, Sitka, AK 99835.
AMERICAN FERN JOURNAL: VOLUME 71 (1981)
PTERIDOPHYTES FOR THE FLORA MESOAMERICANA
The Flora Mesoamericana Project, run by a consortium of museums, aims at
producing a comprehensive vascular plant flora covering southern Mexico (Chiapas,
Tabasco, Quintana Roo, and Yucatan) south to the Panama/Colombia_ border.
Although pteridophytes initially were to be excluded, a decision was made in
November 1980 to include them. The volume on Pteridophyta, scheduled to appear
around 1986, will Se edited by Ramén Riba and Luis Diego Gémez. Pteridologists
interested in contributing generic or family treatments are urged to contact the
editors for general information. Please write to Flora Day eee
Museo Nacional de Costa Rica, Apartado 749, San José, Costa Rica,
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QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY
Azolla filiculoides New to the
Southeastern United States VERNON M. BATES, JR. and EDWARD T. BROWNE, JR.
The Genus Nephrolepis in Florida CLIFTON E. NAUMAN
The Branching Pattern of Hypolepis repens THERESA M. GRUBER
Diplazium japonicum and Selaginella uncinata
Newly Discovered in Georgia WAYNE R. FAIRCLOTH
Notes on Selaginella, with a
New Variety of S. pallescens ROBERT G. STOLZE
Lepisorus kashyapii in the
Western Himalayas S. S. BIR and CHANDER K. SATIJA
Taxonomic Notes on Jamaican Ferns-IIl GEORGE R. PROCTOR
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Council for 1981
ROBERT M. LLOYD, Dept. of Botany, Ohio University, Athens, OH 45701. President
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American Fern Journal
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AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 2 (1981) 33
Azolla filiculoides New to the Southeastern United States!
VERNON M. BATES, JR.* and EDWARD T. BROWNE, JR.**
When studying specimens of Azolla, one is often confronted with a large number
of sterile collections that cannot be identified with confidence. Although Svenson
(1944), in his monograph of the New World species of Azolla, based his classifica-
tion on characteristics of the infrequently occurring sporocarps and paid little
attention to vegetative characters, we believe that megaspore ornamentation pro-
vides the only suitable means of identification.
Through SEM examination of its megaspores, we have confirmed the identifica-
tion of a collection of A. filiculoides Lam. from Georgia. The collection was made
15 April 1956 from plants floating in a large fresh-water lake at the north end of
Sapelo Isaland, McIntosh Co., Georgia (Wilbur H. Duncan 1 9969, with William P.
Adams and Clyde Connell, TEX). Apparently this is a new record for the
southeastern United States. We found that the megaspore ornamentation agreed very
closely with Svenson’s (1944) line drawings and description; the “raised, hexagonal
markings” reported by Svenson actually are localized masses of fused excrescences
which form a flattened to slightly convex reticulum (Fig. /). Although there has
been no previous ultrastructural study of Azolla megaspores from North America,
recent descriptions of European material support our identification (Pieterse, de
Lange & van Vliet, 1977: Martin, 1978).
Traditionally, Azolla in the southeastern United States has been thought to be A.
caroliniana Willd. Megaspores apparently never have been described for this species
in North America (Svenson, 1944; Correll & Correll, 1975). Di Fulvio (1961)
illustrated a cross-section of the perispore of an A. caroliniana megaspore, but the
source of her material and the voucher documenting it are not stated. Unfortunately,
so little is known about the cytology of Azolla that it cannot be said whether or not
A. caroliniana is a sterile hybrid; it has been reported as 2n=48 in Europe (Love,
Love & Pichi Sermolli, 1977, p. 373). Finding consistently sterile material over the
entire range of a species does, however, raise the possibility that the material is
hybrid or otherwise incapable of forming spores, perhaps as a result of aneuploidy.
Svenson reported the distribution of A. filiculoides as widely scattered along the
Pacific Coast from Alaska southward into Washington, Oregon, California, and
Mexico. Although he believed that this species was naturalized in the New England
states, he never attempted to explain two New York collections of A. filiculoides
which may not have been the result of artificial introductions: Riverhead, Long
Island, Suffolk Co., 20 Aug 1938, Muenscher & Curtiss 6647 (US) and Oak
Orchard, Orleans Co., Aug 1876, Herb. N. L. Britton 5. n. (NY).
If, as some have suggested (Smith, 1955), Azolla is disseminated by adhering to
the feet of migratory waterfowl, it seems strange that A. caroliniana and A.
* Data Communications Corp., 3000 Directors Row, Memphis, TN 38131.
** Dept. of Biology, Memphis State University, Memphis, TN 38152.
E Portion of a Master of Science thesis submitted to the Department of
sity, 1980.
Volume 71, number 1, of the JOURNAL was issued March 30, 1981.
Biology, Memphis State Univer-
34 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
filiculoides would be limited to the Atlantic and Pacific coasts, respectively. In light
of the large number of sterile collections of Azolla from the eastern United States
which cannot be identified with certainty, it seems quite possible that A. filiculoides
is more widely distributed in the eastern United States than has been thought. Our
discovery of A. filiculoides in Georgia, considered with those specimens already
known from New York, supports the hypothesis that this species may be widely
distributed along the Atlantic coast. Unfortunately, this cannot be confirmed until
more fertile collections are observed or until vegetative differences can be found to
distinguish the various species of Azolla.
. 1. Scanning electron micrograph showing characteristic features of the megaspore of Azolla
filiculoides. Proximal to the girdle (g), elevated masses of excrescences form a distinct reticulum (r).
We wish to express our appreciation to Dr. Lewis B. Coons and Mrs. Naomi
Roberts for their kind assistance in obtaining the micrograph. We are also especially
indebted to Dr. Billie L. Turner, curator of the Herbarium, University of Texas at
Austin, who loaned us collections of Azolla to examine. In addition, we are very
grateful to Dr. W. Carl Taylor, curator of the Vascular Herbarium, Milwaukee Public
Museum, for reading the manuscript and making valuable suggestions.
LITERATURE CITED
CORRELL, D. S. and H. B. CORRELL. 1972. Aquatic and Wetland Plants of the Southwestern United
States. Environmental Protection Agency, Washington, DC. :
DI FULVIO, T. M. 1961. Sobre el episporio de las especies Americanas de Azolla con especial
referencia a A. mexicana Presl. Kurtziana 1:299-302.
OVE, A, D. LOVE, and R. E. G. PICHI SERMOLLI. 1977. Cytotaxonomical Atlas of the
Pteridophyta. J. Cramer, Vaduz.
MARTIN, A. R. H. 1976. Some structures in Azolla megaspores and an anomalous form. Rev.
Paleobot. Palynol. 21:141-169.
PIETERSE, A. H., L. de LANGE, and J. P. van VLIET. 1977. A comparative study of Azolla in the
Netherlands. Acta Bot. Neerl. 26:433—449.
SMITH, G. M. 1955. Cryptogamic Botany, vol. II. Bryophytes and Pteridophytes, ed. 2. McGraw-Hill,
New York.
SVENSON, H. K. 1944. The new world species of Azolla. Amer. Fern J. 34:69-84.
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 2 (1981) 35
The Genus Nephrolepis in Florida
CLIFTON E. NAUMAN*
The recent addition of N. multiflora (Roxb.) Jarrett ex Morton to the Florida flora
(Gillis & Proctor, 1975) and the description of a new hybrid species (Nauman,
1979a) has created a need for a revised treatment of the genus in Florida.
This treatment is based on a study of more than 120 morphological characters
involving more than 1000 specimens. A transplant experiment and ecological
observations (Nauman, 1979b) also aided in interpreting morphological variability.
his study was based in part on a Masters thesis, and I would like to express my
appreciation to D. F. Austin (Florida Atlantic University) for his guidance during the
course of study and for advice and assistance in the gathering of data. Appreciation
is also given to the curators and staff of the following herbaria for the loan o
materials and types: B, BM, BR, F, FAU, FLAS, FSU, FTG, GH, K, MA, NY, P,
PENN, PH, S, TENN, UC, US, USF, UWFP.
KEY TO THE FLORIDA SPECIES OF NEPHROLEPIS
1. Adaxial costa surface of poe pinnae glabrous (sometimes with a few scales); indusium reniform,
hippocrepiform, or lunate, .2 mm or more wide.
2. Pinnae falcate with acute to attenuate tips; plants never bearing tubers; rachis ere appearing
concolorous or obscurely bicolorous . exaltata
2. Pinnae not falcate or slightly so with blunt tips; plants sometimes bearing oe "chs scales
distinctly bicolorous (pale with a dark point of attachment) 2. N. cordifolia
1. Adaxial costa surface of medial pinnae sparsely to ad covered with short erect trichomes (often
also with scales); indusium orbicular, ca. 1.0 mm
Basal portions of mature stipes covered with jeg brown, appressed scales with a margins.
4. N. multiflora
3. Basal portions of mature stipes not covered with dark brown, appressed scales, but often with a
few loose, reddish to light brown scales or the scales absent.
4. Adaxial costa surface sparsely = the trichomes 0.4 mm long; pinnae ered falcate;
blade length/width ratio 6.8-17.9, mean 9.5 5.N. X averyi
4. Adaxial costa surface densely sinocen to tomentose (rarely glabrous), trichomes 0.3 mm
long; pinnae not falcate or slightly so; blade length/width ratio 3.8-7.8, mean 5.1.
1. N. biserrata
1. Nephrolepis biserrata (Swartz) Schott, Gen. Fil. text to pl. 3. 1834.
Aspidium biserratum Swartz, J. Bot. Schrad. 1800(2):32. 1802. TYPE: Mauritius. Groendal, (S-Hb.
wartz !).
Nephrodium biserratum (Swartz) Presl, Rel. Haenk. 1:31. 1825.
Hypopeltis biserrata (Swartz) Bory in Bél. Voy. Indes. a Bot. 2:65. 1833.
Lepidoneuron biserratum (Swartz) Fée, Gen. Fil. 301. ;
Nephrolepis exaltata var. biserrata (Swartz) Baker in sii Fl. Brasil. a 493. 1870.
Nephrolepis hirsutula 8 biserrata (Swartz) Kuntze, Rev. Gen. Pl. 2:816. _
Aspidium acutum Schkuhr, Krypt. Gew. 32, pl. 31. 1806. TYPE: =ge wal collector not state
(Hb. Breyne presumably destroyed).
Nephrodium acutum (Schkuhr) Presl, Rel. Haenk. 1:31. 1825.
Nephrolepis acuta (Schkuhr) Presl, Tent. Pterid. 79. 1836.
_Nephrolepis hirsutula « acuta (Schkuhr) Kuntze, Rev. Gen. Pl. 2:816. 1891.
“Department of Biological Sciences, Florida Atlantic University, Boca Raton, FL 33431.
AMERICAN FERN JOURNAL: VOLUME 71 (1981)
N. biserrata N. cordifolia
N. exaltata N. multiflora
200 miles
N.x% averyi
~
FIG. 1. County distributions of the species of Nephrolepis in Florida.
C. £. NAUMAN: NEPHROLEPIS IN FLORIDA 37
DISTRIBUTION AND HABITAT: This is a species of swamps and wet ham-
mocks in which it is usually terrestrial, but may be epiphytic or epipetric. The
general distribution is pantropical; in Florida it is largely restricted to Florida’s
tropical fringe (Fig. /). Distribution in Florida appears to be limited by the plant’s
ability to tolerate cold temperatures as revealed by transplant experiments.
Nephrolepis biserrata is best distinguished from the other Florida Nephrolepis
species by its size, hyaline margined scales on the rhizome and croziers, pubescent
pinnae, and orbicular indusia. The species is closest to N. multiflora in indumentum
and sori, to N. X averyi in size, and to N. cordifolia in spores.
REPRESENTATIVE SPECIMENS:
Broward Co.: Durand 58, 64, 72 (FAU); Leeds 342 (NY, PH); Moldenke 483 (NY, PH, US); Nauman
& Nauman 380 (FAU); R. P. St. John 1751 (FLAS). Collier Co.: Correll 6081 (FLAS), Diddle 698
(FLAS); Evans (TENN); Lakela et al. 27996 (USF); Nauman & Austin 555 (FAU). Dade Co.: Avery &
Loope 1918 (FAU); Britton 428 (F, NY); Buswell (FAU); Carter 204 (PH); Correll 5897 (GH, US);
Eaton (F); Long et al. 1941 (NY, USF); Munroe (GH, NY, UC); Safford & Mosier 105 (US); Small &
Mosier 5890 (NY, US). Highlands Co.: Garrett 60 (FLAS); McFarlin 10714 (GH). Monroe Co.:
Delchamps & Wherry (PH); Lakela & Almeda 30526 (USF). Palm Beach Co.: Durand 25, 80 (FAV);
Hill 152 (NY); McJunkin (FAV).
2. Nephrolepis cordifolia (L.) Presl, Tent. Pterid. 79. 1836.
Polypodium cordifolium L., Sp. Pl. 2:1089. 1753. TYPE: Petiver, Pterigr. Amer. f. /, f. //. 1712 (').
The Petiver plate is supposedly a copy of Plumier, Tract. Fil. Amer. 1. 7/. 1705 (!). The Plumier plate is
a poor drawing of what might be a species of Nephrolepis. Until plants can be examined from the area
where Plumier obtained his material, application of the epithet N. cordifolia will be problematical.
Aspidium cordifolium (L.) Swartz, J. Bot. Schrad. 1800(2):32. 1802.
Aspidium tuberosum Bory ex Willd., Sp. Pl. ed. 4, 5:234. 1810. TYPE: “Bourbon sur les arbres, No.
111, Bory de St. Vincent” (B-Hb. Willd. 19759 photo FAU !, photo GH !; isotypes P photo FAU !, FI,
not seen).
Nephrodium tuberosum (Bory ex Willd.) Desv., Mém. Soc. Linn., Paris 6:252. 1827.
Nephrolepis tuberosa (Bory ex Willd.) Presl, Tent. Pterid. 79. 1836.
Nephrolepis cordifolia var. tuberosa (Bory ex Willd.) Baker in Mart. Fl. Brasil. (1)2:491. 1870.
Nephrolepis exaltata B tuberosa (Bory ex Willd.) Kuntze, Rev. Gen. Pl. 2:816. 1891 .
DISTRIBUTION AND HABITAT: Whether or not this species is native to
Florida is uncertain. Wherry (1964) considered it possibly native to the southern-
most portions of the State. Though widespread, the plants are almost entirely
persistent from cultivation in dumps and at abandoned homesites. I know of only
one site where the plants may have colonized without the help of man. This is on the
Sturrock estate in West Palm Beach. The plants were reported to have been blown in
by a hurricane in the late 1940’s (Sturrock, pers. comm.). This species’ distribution
is scattered in Florida and doesn’t conform to any apparent trends in temperature
extremes (Fig. ]). The general distribution is possibly worldwide in the tropics and
subtropics, also in Japan and New Zealand.
Tubers are the most distinctive feature of this species, although tuberless colonies
are frequent throughout Florida. Whether tuber production, or the lack of it, is
controlled by environmental or genetic factors is at present uncertain. There appears
to be a correlation between the substrate in which the plant is growing and tuber
production. In the Florida populations, tuber production seems restricted to plants
growing in humus and has not been seen in epiphytic plants or plants growing in
38 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
drier sites. These observations suggest some specific soil and moisture requirements
for tuber production.
The distinctly bicolorous rachis scales are as diagnostic as tuber- production and
may be used to distinguish this species from any of the other Florida species, even in
the absence of other key features..
REPRESENTATIVE SPECIMENS:
Brevard Co.: Hollister (US); Shuey M1084 (USF). Broward Co.: McCart & Snyder 9523 (FAV).
Citrus Co.: R. P. St. John 197 (FLAS). Dade Co.: Evans (TENN); Hardy 5 (PH). Duval Co.: Darling
(US). Escambia Co.: Burkhalter 5919 (UWFP). Hernando Co.: Cooley et al. 8307 (GH, USF); Mickel
et al. 1782 (UC); Moldenke & Moldenke 29488 (US). Highlands Co.: McFarlin 8957 (FLAS).
Hillsborough Co.: Evans 2303 (TENN); Mickel et al. 1789 (US); Scudder 441 (FAU). Leon Co.:
Hume (FLAS). Marion Co.: Mickel et al. 1742 (UC). Martin Co.: Austin et al. 6469 (FAU); Orange
Co.: Githens 2579 (PH); Medgser (UC). Palm Beach Co.: Austin (FAU); Baker (FAU); Cassen 89
(USF); Hill 149 (NY); Nauman et al. 218, 410 (FAU); Wilhelm 121 (FAU). Pasco Co.: Carpenter (GH).
Pinellas Co.: Genelle & Fleming 2508 (USF); Scudder 436 (FAU). Polk Co.: Cooley 11783 (USF);
3. Nephrolepis exaltata (L.) Schott, Gen. Fil. text to pl. 3. 1834.
Polypodium exaltatum L. Syst. Nat. ed. 10(2):1326. 1759. TYPE: Sloane, Jam Voy. 1. 31, 1107,
which is based on a specimen communicated to Sloane by Dr. Sherard from Jamaica (BM not seen,
photo FAU).
Aspidium exaltatum (L.) Swartz, J. Bot. Schrad. 1800(2):32. 1802.
Nephrodium exaltatum (L.) R. Brown, Prodr. Fl. Nov. Holl. 1:148. 1810.
DISTRIBUTION AND HABITAT: This is the most common of the Florida
species. It is found in a wide variety of habitats, such as tropical hammocks, . low
hammocks, and swamps. Frequently N. exaltata is found as an epiphyte on Sabal
palmetto or species of Quercus, but it also may occur epipetrically or terrestrially. In
Florida, N. exaltata is found from Dade and Monroe to Duval Counties (Fig. /).
Occasionally it occurs farther north, but only in cultivation. The plants are most
common south of Lake Okeechobee, becoming occasional to rare northward. The
general distribution has been traditionally construed as pantropical, but studies by
Proctor (1977) and Stolze (pers. comm.) have implied that N. exaltata may have a
more restricted range than previously thought. Examination of herbarium specimens
shows that a large proportion of plants identified as N. exaltata is actually N.
multiflora, N. biserrata, N. cordifolia, or N. rivularis (Vahl) Mettenius. The
misapplication of the epithet exaltata is so widespread that a thorough monographic
study will be necessary to determine the actual range of this species. Authentic
x phar are known from Florida, Jamaica (the type locality), the Antilles,
10802 (NY). Broward Co.: Durand 15, 73 (FAU); Hopkins 31 (FAU): Janda 29 (FAU); Nauman 826
(FAU). Charlotte Co.: Ward A-69 (FLAS). Citrus Co.: E. P. St. John (FLAS). Collier Co.: Austin &
Austin 6669 (FAU); Austin el al. 6763 (FAU); Clewell 286 (FSU); Cooley 786 (USF); Evans (TENN);
Eyles & Eyles 8235 (GH); Hitchcock (F); Lakela & Almeda 2994] (USF), Long et al. 2393 (USF);
Nauman et al. 328 (FAU); Sturtevant 44 (FLAS, US). Dade Co.: Garber (F, NY); Hill 3079 (FTG);
Moldenke 436 (NY); Small 7384 (NY, TENN); Tracy 9135 (F, NY, PENN, US). Duval Co.: Calkine
(F). Glades Co.: Ward 1-10 (FLAS). Hardee Co.: Kirk (FLAS). Hendry Co.: Jennings (USF); Ward
C. E. NAUMAN: NEPHROLEPIS IN FLORIDA 39
et al. 2396 (FLAS, FSU). Hernando Co.: E. P. St. John (FLAS). Highlands Co.: Evans 2292
(TENN); Lakela 26793 (USF); Mickel et al. 1815 (FSU, UC); Nauman 90 (FAU); Porter & Porter
10618 (NY, UC); Wilbur & Webster 2610 (GH, NY, US). Hillsborough Co.: Mickel et al. 1789. (UC).
Indian River Co.: Small 8849 (NY). Lake Co.: Nash 1288 (F, GH, NY, PH, UC, US); Underwood (F,
GH). Co.: Brumbach 5348 (FAU, FLAS); Correll 5919 (GH); Hitchcock 542 (F, GH. US):
Standley 124 (F, GH, NY). Leon Co.: Jackson (FSU). Martin Co.: Nauman 527 (FAU); Nauman &
Tatje 520 (FAU); Popenoe & Popenoe 688 (FTG). Monroe Co.: Long et al. 1703 (USF). Okeechobee
Co.: McCart 10287 (FAU). Osceola Co.: Mearns 3] (US). Palm Beach Co.: Austin 6636 (FAU):
Cooley et al. 4877 (GH, USF); Durand 77, 81 (FAU); Hitchcock 2402 (F); Kral 5680 (FSU); Meagher
895 (FTG); Nauman et al. 702. (FAU); Randolph 140 (GH); Underwood 2219 (NY). Pasco Co.:
Underwood 1931 (NY). Pinellas Co.: Curtiss 3764 (F, NY, UC); Genelle & Fleming 1640 (USF);
Ralphs 750 (F); Scudder 104 (FAU); Thorne 10315 (UC). Polk Co.: Jennings & Jennings (USF); Smith
(US); Wherry (PH). Saint Lucie Co.: Austin et al. 6454 (FAU); Leeds 344 (PH); Small & Matthaus
9640 (NY). Sarasota Co.: Smith (PH). Seminole Co.: Lambert 18 (PH); Nauman et al. 293 (FAV).
Sumter Co.: E. P. St. John (FLAS, NY); Smith (US). Suwannee Co.: Leonard 6667 (FSU).
4. Nephrolepis multiflora (Roxb.) Jarrett ex Morton, Contrib. U. S. Natl.
. 1974.
Davallia multiflora Roxb., Calcutta J. Nat. Hist. 4:515, t. xxxi left hand. 1844. LECTOTYPE: India,
Roxburgh (BR not found, fide Lawalrée in litt., fragment US!), chosen by Morton.
DISTRIBUTION AND HABITAT: Occasional in disturbed sites, usually near
canals and other bodies of water in loose, well drained soil, frequently in full sun.
Scattered in southern Florida, but reaching as far north as Pinellas and Hillsborough
Counties (Fig. 1). Like N. biserrata, the distribution appears limited by tolerance to
cold temperatures. A native of the Old World tropics, N. mulitflora is widely
naturalized in the New World. The species was first reported for Dade County,
Florida by Gillis and Proctor (1975). Though the data are inconclusive, this species
seems to have arrived in Florida in the late 1940’s or 1950’s somewhere in Lee
County and is actively spreading. This conclusion is based on the dates of collection
of this species throughout the State, but may be biased by the uneven collecting of
certain areas. Nephrolepis multiflora was reported by Proctor (1977) to occur in the
Bahamas and the Antilles. I have seen specimens from these areas, as well as
Central America, Brazil, and Venezuela.
REPRESENTATIVE SPECIMENS: :
Broward Co.: Avery & McPherson 1327 (USF); Lakela & Long 1598 (USF). Collier Co.: Austin et
al. 6767 (FAU); Correll & Popenoe 47290 (FTG); Lakela & Almeda 29988 (GH, USF); Lassiter-et al. 9
(USF). Dade Co.: Avery 1329 (FLAS, USF); Evans (TENN); Gillis 10856 (FTG). Hillsborough Co.:
Long et al. 2942 (USF): Scudder 439 (FAU); Shuey (USF). Lee Co.: Austin 6625 (FAU); Brumbach
8743 (NY, US); Cooley 2548 (FLAS, GH, NY, US). Martin Co.: McCart 10406 (FAU, FLAS);
Nauman & Tatje 258 (FAU); Nauman et al. 200 (FAU). Monroe Co.: Avery (FLAS). Palm Beach Co.:
Brawner (FAU); Durand 5, 85 (FAU); Nauman 312 (FAU); Nauman & Austin 181 (FAU); Nauman et al.
).
5. Nephrolepis x averyi Nauman, Amer. Fern J. 69:69. 1979.
TYPE: Fakahatchee Strand off West Grade, 50 ft E of Indian Mound Slough
Bridge, Collier Co., Florida, 29 Jan 1979, Nauman et al. 635 (US; isotypes FAU,
AS, GH, MSC, NY). oe
DISTRIBUTION AND HABITAT: Terrestrial, epiphytic, or epipetric in ham-
mocks and swamps. Known to occur only with its putative parents, N. biserrata and
40 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
N. exaltata, in mixed colonies, in Florida south of Lake Okeechobee to as far north
as Pinellas County (Fig. /). Distribution outside Florida is uncertain; one specimen
seen from Jamaica.
Nephrolepis X averyi is best distinguished from N. biserrata by its falcate pinnae
and narrower fronds, and from N. exaltata by its larger size and lightly pubescent
adaxial costa surface.
PRESENTATIVE SPECIMENS:
Co.: Durand 4, 49, 56, 67, 70 (FAU); Nauman 318, 434 (FAU); Nauman et al. 647 (FAV).
Collier Co.: Beck 2027 (FSU); Field & Lazon (US); Fischer 8 (US); Howell 855 (US); Nauman et al.
631. (FAU). Dade Co.: Curtiss 5460 (FLAS, UC, US); Eaton 99] (F); Garber (F, FLAS, PH); Hardy
10 (PH); Lakela 31449 (USF), Small 7384 (NY); Tatnall 805 (PH). Manatee Co.: Cuthbert (FLAS).
Palm Beach Co.: Stevens (FAU). Pinellas Co.: Bebb (F). Polk Co.: White (FLAS).
SPECIES AND FORMS EXCLUDED
Several cultivated forms of Nephrolepis are occasionally found in Florida. These
forms are usually in cultivation or are persistent from cultivation. The distribution is
scattered from Dade to Duval Counties. Forms represented in the herbaria are: N.
exaltata cv. ‘Bostoniensis,’ cv. ‘Elegantissima,’ Cv. ‘Florida Ruffles,’ and cv. ‘M. P.
ills’; N. falcata f. furcans (Moore in Nicholson) Proctor [=N. biserrata cv.
‘Furcans’]; and N. hirsutula cv. ‘Superba.’
Nephrolepis pectinata (Willd.) Schott was reported by Wherry (1964) for southern
Florida. I have seen no specimens of this species in the herbaria or field. It is
doubtful that this species exists in Florida.
LITERATURE CITED
GILLIS, W. T. and G. R. PROCTOR. 1975. Additions and corrections to the Bahama flora—II. Sida
6:52-62
NAUMAN, C. E. 1979a. A new Nephrolepis hybrid from Florida. Amer. Fern J. 69:65—70.
_______. 1979b. The genus Nephrolepis in Florida. Unpublished Master's Thesis, Florida Atlantic
University. 126 pp.
PROCTOR, G. R. 1977. Pteridophyta. In R. A. Howard, A Flora of the Lesser Antilles. Vol. II]. Arnold
Arboretum, Harvard University, Jamaica Plain, MA.
WHERRY, E. T. 1964. The Southern Fern Guide. Doubleday, Garden City, NY.
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 2 (1981) 4]
The Branching Pattern of Hypolepis repens
THERESA M. GRUBER*
Hypolepis, a pantropical genus of the Dennstaedtiaceae, is a terrestrial fern with
usually large leaves and rather slender, long-creeping stems bearing trichomes and
long, fibrous roots. Hypolepis repens (L.) Presl may form large colonies, and a
single plant was measured in Puerto Rico with 42 meters of stem (R. and A.
Tryon, pers. comm.). The leaves may be up to three meters or more long and often
are partially supported by surrounding vegetation. Roots occur along the entire length
of the stem, but are especially numerous at the leaf bases.
The morphology of Hypolepis repens and other species has been studied by
Gwynne-Vaughan (1903), Bower (1923), Troop and Mickel (1968), and Imaichi
and Nishida (1973). These treatments are limited, however, to describing the vas-
cular system and analyzing single branch units. This paper considers the branching
pattern of the stem and its components of entire plants of H. repens and the
development and adaptive significance of the pattern.
MATERIALS AND METHODS
Stems of Hypolepis repens were studied in a Liquidambar cloud forest located
about 12 km south of Misantla, Veracruz, Mexico at an altitude of 1400 meters. A
large plant (Fig. 7) was excavated and exposed in situ; it was measured, mapped,
and portions were taken for dissection. The plant was growing on a 35—45° slope in
loose humus in association with herbaceous angiosperms. The stem length totaled
30 m and occupied an area of 13 m?. In clear weather, most of the colony received
sunlight between 10:00 AM and 1:00 PM, despite the surrounding tree canopy. A
smaller plant (Fig. 2) was collected intact from a nearby wooded area. It was
growing in loose humus on a slope of about 30° around a tree base nearly a meter in
diameter.
The vascular system of the stems was studied by dissection and the anatomy of the
branch unit by means of a cinematographic record of the surface between serial
slices (Tomlinson, 1971).
OBSERVATIONS
The pattern of stem growth, with its branches, buds, and leaves is shown in Figs.
I and 2. An analysis of these diagrams revealed a surprising regularity of pattern,
which is illustrated schematically in Fig. 3. The following regular components of the
whole stem and leaf system were noted.
(1) Each bifurcation of the stem (Fig. 3, axis 1) produces a leaf (Fig. 3, axis 2)
and a continuing stem (Fig. 3, axis 1). Mes
The bifurcation may be described as dichotomous since the division of the parent
Stele into stem and leaf is equal (Fig. 4). However, this designation may be
inappropriate because dichotomous branching usually refers to the production of two
*6206 Braidwood Drive, San Antonio, TX 78249.
AMERICAN FERN JOURNAL: VOLUME 71 (1981)
TR yo
Je /! >
T/
7 TR
TR
uphill
downhill
i
O
y @: Dead
o: Matur mo
©: Infant l
cnaesaine t tuned
Oo: Crozier
~~ ud or stem apex ]
1 METER ; TR : Truncat
pan
Dead leaf
roe panel
nfant lea
cagatains ges we
Crozier
eoo®
Bud or stem apex
Truncate
Aborted Leaf
Decayed but continuous
; at
2
FIG. 1. Diagrammatic sketch of Hypolepis repens, mapped in situ. FIG. 2. Diagrammatic sketch of
Hypolepis repens, collected intact and later mapped.
1 METER
T. M. GRUBER: BRANCHING PATTERN OF HYPOLEPIS REPENS 43
initially equivalent organs rather than a stem and a leaf and because it is not known
whether the bifurcation arises from a division of the apical cell.
Sections of the vasculature illustrated in Fig. 4 agree with the observations of
Gwynne-Vaughan (1903) and Troop and Mickel (1968), except that the leaf petioles
arise on the lateral rather than the upper side of the stem.
(2) Leaves are produced successively on alternate sides of the stem. The simplest
assumption for descriptive purposes is that the system is monopodial, that is, the leaf
is an appendage of the stem axis.
(3) The petiole (Fig. 3, axis 2) bears 1-4 buds (Fig. 3, axes 3-6) which may
develop into stems (Fig. 3, axes 3, 4) that function like the stem from which their
parent leaf originated.
An analysis of the number of buds on the petioles showed that of a total of 41
petioles on the plants in Figs. / and 2 in condition to study, two had no buds, five
had one bud, ten had two, 22 had three, and two had four buds. The third and fourth
buds, when present, were borne in the region of the petiole where it became erect
and they grew down toward the ground, perhaps providing support for the leaf.
(4) The proximal bud that develops into a branch (Fig. 3, axis 3) is always on the
side of the petiole (Fig. 3, axis 2) opposite to the continuing main axis (Fig. 3,
axis 1); successive buds that may develop into branches (Fig. 3, axes 4-6) are
alternate beyond the proximal bud.
It follows that the second bud is always on the side of the petiole nearest to the
continuing main axis.
(5) When the proximal bud develops into a stem (Fig.-3, axis 3), the symmetry of
its resulting next higher-order branch unit (Fig. 3, leaf 7 and petiolar buds 8, 9) is
identical to that of the unit (Fig. 3, leaf 2 and petiolar buds 3-6) from which the
branch originated. ;
(6) When the second bud develops into a stem (Fig. 3, axis 4), the symmetry of its
resulting next higher-order branch unit (Fig. 3, leaf 10 and petiolar buds 11, 12) is
the mirror image of that unit (Fig. 3, leaf 2 and petiolar buds 3-6) from which the
branch originated. ;
(7) All stems developing beyond the first branch unit repeat the patterns laid down
in (1) through (6).
The distance between leaves and the angle between bifurcations of axes were both
rather variable.
The spacing of the leaves varied widely in each of the plants investigated. The
small plant (Fig. 2), which was growing in a heavily shaded locality, had the leaves
an average of 19.5 cm apart, with two 28 cm apart. The large plant (Fig. /),
which was growing in a somewhat exposed site, had the leaves an average of 33.9
cm apart and two were as much as 68 cm apart. Apparently the distance between
leaves is influenced by the immediate environment and is not a fixed characteristic
of the branching pattern. : .
The angle between bifurcations of the stem and leaf approximates 60 - In a tota
of 17 branch units of the small plant (Fig. 2), the angle was 40 in two units, 50° in
four, 55° in one, 60° in eight and 90° in two (an average of 59°). The angle between
the petiole and its first bud averaged 63°, for the nine that were measurable.
AMERICAN FERN JOURNAL: VOLUME 71 (1981)
A
666
0000000000000
0000000000000
Bud
nh
Leaf
= Fn
Bud \ :
: ae mxAy
G. 3. The Branch Complex. Branch order is represented by the different patterned axes: main axis,
solid; first-order, solid dots; second-order, stripes; third-order, open circles, fourth-order, stippled. The
individual axes are numbered according to their ontogenetic appearance. An axis terminated by a
circular tip represents a leaf; a triangle represents a stem apex. _ 4. Serial sections of the vasculature
of the branching unit of Hypolepis repens, from a cinematographic record.
T. M. GRUBER: BRANCHING PATTERN OF HYPOLEPIS REPENS 45
DISCUSSION
Ecological Implications. — Bell and Tomlinson (1980) noted the infrequency of
adequately detailed accounts of basic branching patterns in rhizomatous systems,
especially among the ferns. Ferns are ideal for this kind of study because they have
no secondary cambial growth.
A fern with a creeping, branching stem, as exemplified by Hypolepis repens, has
important advantages over its relatives which have short-creeping or erect stems. An
environment such as the Liquidambar cloud forest presents a substantial limitation
to gametophytes in the form of competition for open space. An elongate, branching
stem provides a means of vegetative propagation which alleviates the necessity for
local propagation through sexual reproduction. Interestingly enough, not a single
fertile frond was found during several days of mapping and excavating.
In addition, a regular method of branching enables the plant to survive destruction
of one or more stem apices. When growth along one axis is unsuccessful, that shoot
may abort without the loss of the entire plant. The closely related H. punctata Mett.
ex Kuhn responds to the destruction of the leaf apical cell by the development of the
most proximal bud into a leaf or shoot (traumatic reiteration, Imaichi and Nishida,
1973). Hypolepis repens probably responds in a similar manner, as indicated by the
development of the proximal bud in two out of four aborted leaves (Fig. 2).
A branched network with bifurcation angles of 60° tends to form hexagons, a
pattern which covers a maximum amount of surface area with a minimum path
length (Stevens, 1974). This patterned branching enables the plant to make a
systematic exploration of the local environment with a minimal energy output. This
same pattern enables the plant to reenter territory, over a span of time, which
previously proved favorable to growth.
A shoot might be expected to be sufficiently plastic to respond to an especially
favorable environment. This theory of “adaptive reiteration under supra-optimal
conditions” is difficult to prove, as was pointed out by Bell and Tomlinson (1980).
The difficulty lies in there being no established basis for quantification. A favorable
environment could exist in time, such as especially favorable weather during a
growing season, or in space, such as nutrient-rich soil or good light availability.
Adaptive reiteration could refer to the production of a maximum amount of
photosynthetic material within a limited amount of time or space, exceeding that
observed to be average. Hypolepis repens appears to support this theory, as indicated
by the inordinate number of croziers within the area delineated by broken lines in
Fig. 1. Furthermore, within this same region, there are three examples of advanced
development of the second of the second-order branch (as in Fig. 3, axis 4), a
relatively rare occurrence in the two plants.
These ecological advantages and the predictability of the morphology of H gd
repens suggest that this particular pteridophyte has genetically fixed growth an
branching patterns.
Development. — It is interesting to spe
ment for a branching system as well defined a
examination of the rhizome raises the question 0
culate on the possible mode of develop-
s that of H. repens. A superficial
f whether branching in this species
46 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
is sympodial, monopodial, or dichotomous. If the first were the case, the question
of development would be simplified from a descriptive point of view; all the leaves
would be terminal and all branches could be produced in the same way. However, the
anatomy of H. repens and the developmental studies of the closely related H.
punctata by Imaichi and Nishida (1973) indicate otherwise. Instead, there are two
types of branches in a branch complex: (1) the stem (Fig. 3, axis 1) bifurcates to
produce a leaf (Fig. 3, axis 2) and a continuing stem; and (2) and one to four buds (Fig.
1, axes 3-6) branch off from the base of the leaf (Fig. 3, axis 2).
The first type of branching may be dichotomous in the strictest sense, that is, by a
division of the apical cell of the parent stem axis (Fig. 3, axis 1). The equal
distribution of the vasculature between the daughter stem and leaf favors this
explanation (see Fig. 4). On the other hand, the two daughter products are not the
same. This might result from a lateral mode of branching, that is, one where
meristematic tissue proximal to the apical cone of the continuing stem differentiates
into and terminates as a leaf. —
The methods by which the buds branch from the leaf are somewhat less clearly
defined. Bower (1923) theorized that the formation of extra-axillary buds in H.
repens represented a modification of dichotomous branching of the rhizome system,
that, in effect, the basal bud (Fig. 3, axis 3) bore the leaf (Fig. 3, axis 2). The
regular pattern of buds on the leaf base strengthened his argument that these buds
were part of a regular branching system and therefore did not develop. adventitiously.
Imaichi and Nishida’s (1973) decapitation experiments and ontogenetical observa-
tions of the related H. punctata demonstrated that the bud meristem is formed after
the leaf is established, making Bower’s theory untenable.
In his developmental studies of Onoclea sensibilis, Dryopteris aristata, and D.
filix-mas, Wardlaw (1943) demonstrated that adventitious buds occur only in specific
positions corresponding to those occupied by detached meristems. These superficial
bud meristems develop from the region of the shoot apical meristem not involved in
the development of leaves. In the case of D. filix-mas, every bud initially occupies
an axillary position. It may then become separated from the leaf with which it
originally was in an axillary relationship by displacement onto the enlarging base of
another leaf which was lateral to it in the earlier developmental phase. Since the bud
develops more slowly than the leaf, its vasculature tends to become joined with that
of the leaf it had been carried up on, rather than with that of the stem from which it
originated. In summary, Wardlaw (1943) demonstrated that each bud of D. filix-mas,
despite its position on the petiole of an adult leaf, occupies an approximately
axillary position on the shoot at the time of its formation. Wardlaw concluded that
extra-axillary buds in other fern species might originate in a similar fashion.
It is possible that some interaction of hormones produced by the petiolar roots
and/or developing frond may trigger the differentiation of meristematic cells pro-
duced by the apical cone of the leaf. Only a thorough investigation of the
developmental process will provide the insight needed to explain the origin of the
interesting branching pattern of Hypolepis repens.
Field work in Veracruz, Mexico was supported by the Atkins Garden Fund,
Harvard University, and assistance in Mexico was obtained from the Instituto de
T. M. GRUBER: BRANCHING PATTERN OF HYPOLEPIS REPENS 47
Biologia, Universidad Nacional Autonoma de México. I am grateful for the help and
encouragement of ching 2 Rolla Tryon and Dr. Alice Tryon and especially for the
guidance of Professor P. B. Tomlinson in preparation of the film. Appreciation is
also extended to Scott Clempson, Kathleen Buckley, David Karachuk and the staff of
the Gray Herbarium and Arnold Arboretum Library.
LITERATURE CITED
BELL, A. D. and P. B. TOMLINSON, 1980. Adaptive architecture in rhizomatous plants. Bot. J.
Linn. Soc. 80:125—160.
BOWER, F. O. 1923. The Ferns, Vol. I. University Press, Cambridge.
GWYNNE-VAUGHAN, D. T. 1903. Observations on the anatomy of solenostelic ferns. Ann. Bot.
:689-742.
IMAICHI, R. and M. NISHIDA. 1973. Studies on the extra-axillary buds of Hypolepis punctata. J.
Jap. Bot. 48:268-279.
STEVENS, P. S. 1974. Patterns in Nature. Little, Brown, Boston.
TOMLINSON, P. B. 1971. Monocotyledons—Towards an understanding of their morphology and
peri In R. D. Preston (ed.) Advances in Botanical Research, vol. 3. Academic Press,
TROOP, J. ra aa Yes 8 ae 1968. Petiolar shoots in the Dennstaedtioid and related ferns.
Amer. Fern J. 58:64—7
WARDLAW, C. W., 1943. aA and analytical studies of pteridophytes, II. Ann. Bot. n. s.,
7:349-377.
48 AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 2 (1981)
Diplazium japonicum and Selaginella uncinata
Newly Discovered in Georgia
WAYNE R. FAIRCLOTH*
The Japanese Twin-sorus Fern, Diplazium japonicum (Thunb.) Bedd., has been
reported as escaped from cultivation in Gadsden County, Florida since 1957
(Wherry, 1964), and recently Short (1980) published an account of its discovery in
Lee County, Alabama. This species subsequently has been found in three widely
disjunct locations in Georgia (Fig. /).
In late summer of 1980, my wife and I led a fern field trip in central south
Georgia for the Georgia Botanical Society. During the field trip, Marge White and
Frieda Polsfuss (fern hobbyists from middle Georgia) showed several of us some
pressed fronds of a fern which Mrs. White had collected in Houston County,
Georgia. Upon superficial examination, most of us identified the plant as a form of
Athyrium, although the possibility of its being a Diplazium was suggested. A frond
was given to Lloyd H. Snyder, Jr. (a fern hobbyist from Atlanta), who discovered the
same fern beneath a highway bridge in Berrien County, Georgia the following day.
Specimens from Houston and Berrien Counties were shown to Dr. Murray Evans of
the University of Tennessee, who confirmed the identity of both as Diplazium
japonicum.
The two sites were rechecked in October to determine the extent and condition of
the colonies. The team of White, Polsfuss, and Snyder visited the Houston County
site, which is located in dense, deciduous woods between Hatcher Road and Fagin
Mill Road southwest of Warner Robins. Two groups of approximately a dozen plants
each about 1000 m apart were found along the edge of a small stream in association
with Asplenium platyneuron, Athyrium asplenioides, Onoclea sensibilis, Osmunda
cinnamomea, O. regalis, Polystichum acrostichoides, and Thelypteris torresiana.
My wife and I found the Diplazium in Berrien County growing beneath a slough
bridge of the Withlacoochee River, west of Nashville on Georgia Highway 125.
Thirteen plants bearing fertile fronds were clustered together in an area of three
square meters. Eighty-two sporelings were widely scattered beneath the bridge,
indicating that the colony was successfully reproducing and expanding. Other ferns
found beneath the bridge were: Asplenium platyneuron, Lorinseria areolata,
Lygodium japonicum, Onoclea sensibilis, Ophioglossum petiolatum, Osmunda
regalis, Pteridium aquilinum, Thelypteris dentata, T. kunthii, T. torresiana, and
Woodwardia virginica.
On 2 January 1981, I discovered an immense colony of the Japanese Twin-sorus
Fern on the Farmer’s Branch prong of Sofkee Creek in Grady County, Georgia.
Hundreds of plants were growing thickly on both banks of this spring-fed stream for
a distance of 28 m; in addition, scattered plants were found downstream for a
distance of approximately 120 m. Most of the fertile plants were robust, with fronds
*Department of Biology, Valdosta State College, Valdosta, GA 31601.
W. R. FAIRCLOTH: DIPLAZIUM AND SELAGINELLA IN GEORGIA 49
commonly as long as 65 cm. The only ferns growing in association with D.
japonicum at this site were Dryopteris ludoviciana, Lorinseria areolata, and Osmun-
da cinnamomea.
Houston County is located in the Upper Coastal Plain Province, almost in the
center of the state and very near the Fall Line (the junction with the Piedmont
Province). Geographically, it is similar to the Fall Line location in Lee County,
Alabama. Grady County also is in the Upper Coastal Plain Province, but is about
FIG. 1. Distribution of Diplazium japonicum (circles) and Selaginella uncinata (triangles) in Alabama,
Florida, and Georgia.
140 miles to the southwest and borders Gadsden County, Florida. Berrien County is
located in the Lower Coastal Plain Province, a region which differs from the Upper
both in elevation and in having predominantly sandy soils. Unlike the rolling, well
drained Upper Coastal Plain, the Lower is very flat, with numerous ponds, vast
Swamps, and broad river systems. ae
Not only do these new locations extend the range of Diplazium japonicum
Significantly, the species is a new addition to the vascular flora of Georgia. Recent
papers by Bruce, Jones, and Coile (1980) and by Duncan and Kartesz (1981) do not
include D. japonicum as a component of Georgia’s flora.
50 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
Specimens from the three locations (Berrien County, Snyder 697, Faircloth 8518;
Houston County, Snyder 757; Grady County, Faircloth 8521) are on deposit in the
herbaria at the University of Georgia (GA) and Valdosta State College (VSC).
Another addition to the vascular flora of Georgia is Selaginella uncinata (Desv.)
Spring. The location of an unfamiliar Spike-moss in Decatur County was first
mentioned to me by Angus Gholson, U. S. Army Corps of Engineers, Resource
Manager for Lake Seminole, Chattahoochee, Florida. My wife and I visited the site
in late March, 1980. We found an extensive colony of the Blue Spike-moss growing
luxuriantly on the banks of a small stream in a ravine immediately north of Greenshade
Cemetery between Faceville and Fowltown (Fig. /).
Shoot development at that season of the year was strictly sterile because a late
freeze on 1 March 1980 had killed the aerial shoots back to ground level. There was
evidence of abundant strobili development the preceeding year, although I could find
no microspores or macrospores that had been retained. My identification, based
upon sterile material, was confirmed by Dr. John T. Mickel (pers. comm.). Fertile
material was collected later in the summer; specimens (Faircloth 8531) are on
deposit in the Valdosta State College Herbarium.
Naturalized Selaginella uncinata, a native of China, has been known from Florida
and some of the Gulf Coast states for more than 20 years (Brown & Correll, 1942;
Lakela & Long, 1976). With the return of horticultural interest in hanging baskets, it
has lately been cultivated and widely sold as a hanging basket plant under the names
of Rainbow-fern and Parlor-moss. Its iridescent, blue-green foliage and arching-
trailing growth habit make it a choice plant for this purpose. The origin of the colony
in Decatur County is puzzling. There are no homesites nearby, and its location in
close proximity to a cemetery appears coincidental because the plant is not likely to
be used either as a potted plant or in other types of floral arrangements for cemetery
ornamentation. Mr. Gholson’s familiarity with the site indicates that the colony is at
least four years old but judging from its size, it is perhaps as much as 8 to 10 years
old.
I wish to thank the following people for their help in field work and in providing
information for this report: Juanita N. Faircloth, Marge White, Frieda Polsfuss,
Leslie Garland, Lloyd H. Snyder, Jr., and Angus Gholson.
LITERATURE CITED
BROWN, J. W. and D. S. CORRELL. 1942. Ferns and Fern Allies of Louisiana. Louisiana State
University Press, Baton Rouge,
G. JO
BRUCE, J. oe NES, Jr, and N. C. COILE. 1980. The Pteridophytes of
Geo; sae Castanea 45:185-193.
DUNCAN, W. and J. T. KARTESZ. 1981. The Vascular Flora of Georgia; An Annotated
oe University of Georgia Press, Athens, GA.
LAKELA, O. and R. W. LONG. 1976. Ferns of Florida; An Illustrated Manual and Identification
Gade Banyan Books, Miami, FL.
SHORT, J. W. 1980. Diplazium japonicum New to Alabama. Amer. Fern. J. 70-111.
WHERRY, E. T. 1964. The Southern Fern Guide. Doubleday, Garden City, NY.
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 2 (1981) 51
Notes on Selaginella, with a New Variety of S. pallescens
ROBERT G. STOLZE*
One of the most variable neotropical species in the genus Selaginella is S.
pallescens (Presl) Spring. It may grow on soil or rocks, in sun or in deep shade,
from sea level to over 3000 m elevation, and it occurs in Mexico, Central America,
Cuba, Jamaica, and parts of South America. The species belongs to the heterophyl-
lous subg. Stachygynandrum, which is characterized by the stems (at least distally)
and the vegetative leaves of branches having 2 rows of smaller, usually appressed,
median leaves and 2 rows of larger, spreading lateral ones. Contrasting with this is
subg. Selaginella, with plants homophyllous throughout, i.e., leaves are borne on all
sides of the stem and branches and all are appressed for most of their length.
Subgenus Stachygynandrum is sharply divided into two groups, one having stems
articulate, or at least constricted at or near the nodes and here usually discolored,
and with rhizophores produced dorsally. The other group, containing S. pallescens,
has stems neither articulate nor with discolored or constricted nodes, and with
thizophores produced ventrally, Selaginella pallescens and its nearest relatives
commonly have the stems densely caespitose, often forming rosettes. They tend to
curl inward when dry, then uncurl again when moisture is introduced, thus giving
rise to the common name “Resurrection Plant.”
Taxonomy of the entire S. pallescens complex in the neotropics needs careful
re-examination. There are several species which I feel are not truly distinct, and yet
there are some hitherto unrecognized variants, which perhaps should be formally
treated as varieties or forms. Some very closely related species are: S. cuspidata
(Link) Link and var. elongata Spring, S. harrisii Underw. & Hieron., S.
microdendron Bak., S. millspaughii Hieron., and S. pulcherrima Liebm. During a
study of the genus for the “Ferns and Fern Allies of Guatemala,” it appeared to me
that most of these species might better be included under S. pallescens, for
whatever differences were noted by previous authors appear to be thoroughly
inconsistent. On the other hand, some new features have come to light which seem
Significant and consistent enough to indicate recognition of some specimens at the
varietal level.
In his study of the spores of heterophyllous Selaginellae, Hellwig (Ann. Mo. Bot.
Gard. 56:444—464. 1969) annotated a number of specimens in various herbaria as S.
pallescens, “red-stemmed variant.” Some other minor features have been discovered
to be consistent with stem color on these specimens, and the combination of
characters support a decision to name the following new variety.
Selaginella pallescens (Presl) Spring var. acutifolia Stolze, var. nov.
arietas haec a varietate typica /S. pallescens (Presl) Spring var. pallescens]
differt caulibus rubellis, foliis lateralibus acutis (mec acuminatis nec aristatis), et
interdum pallide roseo-tinctis, et foliis medianis acutis (nec acuminatis nec aristatis).
TYPE: Rocky hills near Santa Rosalia, 2 mi south of Zacapa, alt. 200 m; Depto.
Zacapa, Guatemala, 1939, Steyermark 29293 (F).
*Department of Botany, Field Museum of Natural History, Chicago, IL 60605.
52 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
In forests or wooded ravines; commonly on rocks, cliffs, or rock outcrops, from
sea level to 1100 m; Guatemala, Honduras, El Salvador, Nicaragua, and Costa Rica.
Plants epipetric or terrestrial; stems reddish, at least at base; lateral leaves acute,
rarely subacute, sometimes a few of the older ones becoming streaked or tinged with
pale red; median leaves commonly acute at apex.
SELECTED SPECIMENS EXAMINED:
GUATEMALA: Chiquimula: Damp thicket along road between Chiquimula and Zacapa, 400-600
m, Standley 74517 (F, US). Escuintla: Medio Monte, Palén, Mario Dary Rivera 755 (F). Jutiapa:
Quebrada near Mangoy, 550 m, L. O. Williams 1420] (F). HONDURAS: Choluteca: Moist bank
above San Antonio de Flores, 30 m, Williams & Molina 16714 (F). Morazan: Sabana Grande, 1100 m,
J. desided ip 3260 (F, UC, US). El Paraiso: Drainage of Rio Yeguare, 600 m, Molina 4014 (F,
US). EL SALVADOR: Chalatenango: Hills outside San José Cancasque, 400 m, Seiler 365 (F). San
sees Riverside, Canton San Antonio Chavez, 300 m, Seiler 54] (F). La Unién: Humus, woods N of
La Union, Morrison & Beetle 8761 (F, US). NICARAGUA: Leon: Canyon of Rio Sinecapa, near Santa
Rosa, 200 m, Williams & Molina 42442 (F). Nueva Segovia: Ravine W of Ocotal, Seymour 840 (F).
COSTA RICA: Guanacaste: Margen rocosa del Rio Recreo, 80 m, Jiménez 1172 (F).
e€ most conspicuous difference between this and the typical variety is the
reddish coloration present in the plants (a phenomenon not uncommon in the.
subgenus). The stems are always reddish at base, and the color often extends
halfway to the apex. Also, as the lateral leaves begin to age, some become tinged
with red. Typical S. pallescens has stems pale greenish to stramineous throughout;
and if lateral leaves turn color with age, it is to a dull whitish or tawny hue.
Most median and lateral leaves in var. acutifolia are never more than acute, whereas in
var. pallescens the leaves are acuminate or even aristate. The new variety prefers low
altitudes from sea level to 800(1000) m, and is most common in rocky habitats. The
typical variety is found occasionally near sea level, but most frequently occurs
between 800 and 2500 m. It, too, is found in rocky situations, but seems equally at
home on the forest floor.
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 2 (1981) 53
Lepisorus kashyapii in the Western Himalayas
S. S. BIR and CHANDER K. SATIJA*
Lepisorus kashyapii (Mehra) Mehra is a polymorphic species closely related to
L. excavatus (Bory) Ching. The two species are known to hybridize in the Himalayas
(Bir & Trikha, 1969, p. 271). The latter species was studied in detail by Bir and
Trikha (1974).
These two species of Lepisorus are more easily separated in the field than in the
herbarium. The rhizomes of L. kashyapii are loosely attached to trees or rocks by
long, straight roots, whereas L. excavatus has rather tightly adherent rhizomes.
Lepisorus kashyapii laminae are thicker and have more obscure veins, and their color
is paler yellow-green. According to a note by R. R. Stewart in the U.S. National
Herbarium, L. kashyapii has quickly deciduous, dark brown, contorted hairs on the
abaxial surface. Unfortunately, these characteristics are mostly difficult to observe in
herbarium material. The rhizome scales of the two species are very similar.
Although there is some variation, the sori of L. kashyapii are more round and only
slightly immersed in the laminae, whereas those of L. excavatus are oval and more
deeply immersed. Lepisorus kashyapii tends to dry brown, whereas L. excavatus
usually dries green. These characteristics will aid in separating the two species in
the herbarium.
During a study of various collections of Lepisorus from the Himalayas, L.
kashyapii was found to be morphologically very interesting. Some specimens from
various localities around Nainital, a popular health resort located in the northwestern
Himalayas, exhibit variations from typical L. kashyapii that proved to be two new
varieties. These, along with var. kashyapii, are distinguished in the following key.
KEY TO THE VARIETIES OF LEPISORUS KASHYAPII
. Fronds linear to linear-lanceolate, 7.5—11.5 cm long, 0.5—1 cm wide; rhizome scales long-acuminate at
3. L. kashyapii var. minor
ea
the apex . 2
Finsae narrowly to sometimes broadly elliptic-lanceolate, (12)15—35(42) cm long, 1.5-3.5(5.5) cm
wide; rhizome scales acute to acuminate at the apex.
2. Laminae bright brown on drying; rhizome scales peltate-lanceolate, dark brown, strongly clathrate
in the center, yellowish and not clathrate at the margins, especially around the peltate base, the
margins contorted and finely toothed when young, usually worn away and merely erose in age,
sporangial paraphyses all peltate, clathrate .........-.-.+-ssessseeseeeeees Lak, kashyapii var. kashyapii
. Laminae dull brown on drying; rhizome scales narrowly ovate-lanceolate, yellowish-brown through-
out, clathrate in the center, less so at the margins, the margins erose and hyaline; sporangial paraphyses
hair-like as well as peltate, clathrate ............-:sssesseesseeeeenersseeseen es 2. L. kashyapii var. major
1. Lepisorus kashyapii (Mehra) Mehra in Bir, Res. Bull. Panjab Univ. n.s.,
13:23. 1962, var. kashyapii.
Polypodium satslinagh arte tee Univ. Publ. 24, f. 5. 1939. TYPE: Not stated; a lectotype
should be chosen from Mehra’s material at Panjab University, Lahore (LAH).
Pleopeltis kashyapii (Mehra) Alston & Bonner, Candollea 15:208. 1956.
Rhizomes long-creeping, 3-5 mm in diam.; rhizome scales ig abel pe
central portion clathrate, brown to dark brown in mass, the marginal porti
—
nN
*Department of Botany, Punjabi University, Patiala 147002, India.
AMERICAN FERN JOURNAL: VOLUME 71 (1981)
005mm ;
figs.4,5
Ff etagd ds Gege >
sO. 24 e,,
Cer
boat ar
y 49,¢8. “MN j
i] a%, 4
MERE »_Smm
lig Pr) figs.2,7
BIR & SATUA: LEPISORUS IN THE WESTERN HIMALAYAS 55
scarcely or not clathrate, yellowish, broader around the peltate base than toward the
apex, the margins contorted, finely toothed, usually worn away and merely erose in
age. Stipes 2-5 cm long, yellowish, bearing a few, usually somewhat contorted
scales. Laminae narrowly to sometimes broadly aes -lanceolate, (12)15—35(40) cm
long, 1.5—3.5(5.5) cm wide, bright brown on drying, acute above an acuminate bas
acuminate at the apex, entire or slightly wavy along the margin; sporangia protected
by subpersistent, clathrate, peltate paraphyses; spores reniform or oval, plane to
concavo-convex, (idtie minutely verrucose, ca. 42-60 wm long, 30-50 pm wide;
mber n=36.
OTHER CI TATIONS: Mehra & Bir, Res. Bull. Panjab Univ. n.s., 15:168. 1964;
Bir & Trikha, Bull. Bot. Surv. India 11:271-273. 1969 [1971].
ILLUSTRATIONS: Mehra (1939, pp. 24-25, fig. 5a—g); Bir & Trikha (1969, pp.
271-273, figs. 41-45).
SPECIMENS EXAMINED:
INDIA: Himachal Pradesh: Simla: Near Shali Peak, 2400 m, Sept 1969, Bir 1046 (PAN); Sanjouli,
2100 m, Aug 1969, Bir (PAN), Sept 1958, Bir (PAN); Chharbara, 2400 m, Aug 1960, Bir (PAN).
Dalhousie: Satdhara, 2000 m, Sept 1968, Trikha 106 (PUN). Uttar Pradesh: Mussoorie: 1800 m, Aug
1959, Bir (PAN); Nag Tiba, 2400 m, Sept 1947, Fleming 82 (US); Lal Tiba, 2400 m, Sept 1968, Bir 44
(PUN), 2250 m, Sept 1968, Trikha 1070 (PUN). Nainital: Laria Kanta, 2400 m, Sept 1967, Bir (PUN);
Tiffon Top, 2100 m, July 1971, Trikha 1901 (PUN); Khurpatal, Sariyatal, 1600 m, July 1971, Trikha
1904 (PUN). West Bengal: Darjeeling: Senchal forest, 1500 m, July 1957 Malhotra 744 (PAN);
Manibhangang—-Tonglu Road, 2400 m, July 1957, Bir 774 (PAN), 2000 m, July 1957, Bir (PAN); Near
Dingle Kothi, 1800 m, July 1969, Tritha 1025 (PUN); Senchal Lake, 2400 m, Aug 1969, Trikha 1030
(PUN); without definite locality, Thomson (US).
NEPAL: Shotibas, 3000 m, Oct 1958, Polunin, Sykes & Williams 5568 (US); Garpung Kholen, 3000 m,
Sept 1962, Polunin, Sykes & Williams 5410 (US).
2. Lepisorus kashyapii var. major, Bir & Trikha, var. nov. Figs. 1-5.
Squamae rhizomatis ovatae, acuminatae, flavescenti-brunneae, margine eroso,
kg lamina lanceolata, 29-42 cm longa, 2.7-3.5 cm lata, statu sicco obscure
brunnea; sporangia paraphysibus biformibus, unis, umbelliformibus, parietibus
cellularum comparate tenuibus, alteris bicellulatis, uniseriatis, piliformibus; sporae
verrucosae vel tuberculatae 55-67 um longae, 33-55 pm latae; chromosomatum
numerus n=36.
TYPE: Cheena Peak, Nainital, Uttar Pradesh, India, epiphyte, 2400 m, July
1971, Trikha 1906 (PUN 1240; isotypes PUN 1241, 1242).
PARATYPE: Tiffon ae Nainital, Uttar Pradesh, India, lithophyte, 2100 m, July
1971, Trikha 1907 (PUN 1357).
3. sami kashyapii var. minor Bir & Trikha, var. nov. Figs. 6-8.
quamae rhizomatis ovatae, longe acuminatae, brunnescenti-atrae, margine eroso;
lamina Hipearié vel lanceolato-linearis, 7.5-11.5 cm longa 0.5—1 cm lata, statu a
laete fusca; sporangia paraphysibus solum umbelliformibus, clathratis, parietibus
cellularum fortibus, Seas: sporae verrucosae, 50-67 jm longae, 38-50 jum latae;
chromosomatum numerus
FIGS. 1-5. Holotype of Lepisorus kashyapii var. major (Trikha 1906, PUN). FIG. 1. es FIG. 2.
Rhizome scale. FIG. 3. Peltate sporangial paraphysis. FIG. 4. Hair-like paraphysis. Fl ae .
FIGS. 6-9. Holotype of Lepisorus kashyapii var. minor (Trikha 1096, PUN). FIG. 6. Habit
Scale from rhizome apex. FIG. 8. Peltate iolawaiat paraphysis. FIG. 9. Spore.
56 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
TYPE: Tiffon Top, Nainital, Uttar Pradesh, India, epiphyte, 2100 m, July 1971,
Trikha 1096 (PUN 1175; isotype PUN 1243).
Our grateful thanks are due to Prof. K. U. Kramer (Zurich) for the Latin
diagnoses, to Dr. D. B. Lellinger for advice on taxonomic matters, and to the
keepers of the cited herbaria for the loan of material.
LITERATURE CITED
ALSTON, A. H. G. & C. E. B. BONNER. 1956. Résultats des expéditions scientifiques genevoises au
Népal en 1952 et 1954 (Partie botanique), 5.—Pteridophyta. Candollea 15:193-220
BIR, S. a and C. K. TRIKHA. 1969. Taxonomic revision of the polypodiaceous genera of India—IV.
Polypodium lineare complex and allied species. Bull. Bot. Surv. India 11:260—276. [Pub-
lished in 1971].
, and C. K. TRIKHA. 1974. Taxonomic revision of the polypodiaceous genera of India—VI.
Legis orus excavatus group. Amer. Fern J. 64:49-63.
MEHRA, P. N. 1939. Ferns of Mussoorie. Panjab Univ. (Lahore) Publ. 29 pp.
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 2 (1981) 37
Taxonomic Notes on Jamaican Ferns—III
GEORGE R. PROCTOR*
This paper continues my series concerning the taxonomy of the ferns of Jamaica
(see Proctor, 1965, 1968).
The genus Thelypteris is the largest and one of the most complex genera of ferns
in Jamaica, when construed in the broad sense, with a total of 59 known species. In
recent years, this world-wide taxonomic group has received intensive scrutiny,
especially by R. E. Holttum (Old World) and A. R. Smith (New World). The former
has presided over the disintegration of the genus into numerous small “splinter”
genera, a process already initiated by Ching and others; Smith has developed an
integrated classification exercising the concept of subgenera and sections. The
present writer prefers the latter approach.
The first of Smith’s subgenera is Amauropelta. This is the largest and most
difficult group of Thelypteris species occurring in Jamaica, with a total of 24
species, nine of them believed to be endemic. Amauropelta was classified by Smith
(1974) in nine sections, all but one of which are represented in Jamaica. The
position of the Jamaican species in these sections may be summarized as follows:
2. Phacelothrix: thomsonii.
3. Uncinella: negligens, oligocarpa, germaniana, linkiana, gracilis, heteroclita.
4. Amauropelta: firma, basiattenuata, ba bisii, trelawniensis, randallii, sancta, nockiana,
underwoodiana, harrisii, gracilenta, resinifera vats. resinifera and caribaea.
5. Blennocaulon: cheilanthoides.
6. Pachyrhachis: pachyrhachis, malangae vat. sitiorum.
7. Lepidoneuron: rudis.
8. Blepharitheca: concinna.
9. Apelta: (not represented in Jamaica).
Three of the above species are new to science and are described herein, along
with a new minor form of 7. rudis; four other names represent new combinations
requiring validation. The localities of these taxa are shown in Fig. /. The writer is
grateful to Dr. John Mickel for helping to locate, at the New York Botanical Garden,
types of species described by Jenman.
Thelypteris decrescens Proctor, sp. nov. he,
Subg. Amauropelta, sect. Adenophyllum. Ex affinitate 7. pilosulae a qua stipitibus
multo brevioribus, densissime minutissimeque stipitato-glandulosis, glabratis vel
parce pilosulis; laminis minoribus, utrinque, sed plerumque subtus abundanter
glandulosis, glandulis stipitatis, flavis resinaceis differt.
hizome stout, erect, its scales yellow-brown, lance-attenuate, the margins
subentire with a few minute colorless stipitate glands. Stipe very short, 3-4 cm
long, scaly at base, puberulous, and, to ether with the rhachis, mec): and
minutely stipitate-glandular, the glands colorless; also bearing few to many lon
soft, pluricellular hairs. Blades narrowly elliptic to oblanceolate, 35-50 cm long,
*Harvard University Herbaria, 22 Divinity Ave., Cambridge, MA 02138.
58 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
10-15 cm broad at or above the middle, markedly decrescent downward with 7 pairs
or more of reduced pinnae; longest pinnae oblong-linear, sessile, acuminate, up to 7
cm long, 1-1.7 cm wide at the base, with up to 20 pairs of oblong or narrowly
deltate- oblong segments, these blunt at the apex and | .5— 2.5 mm wide, the margins
flat or very narrowly reflexed; veins 5-8 pairs, simple, scarcely prominulous but
with slightly enlarged tips (seen from the upper side). Sori supramedial, round;
indusium delicately reniform, densely glandular (the glands minute, globular, and
ales to stipitate), soon withering. Rhachis and other vascular parts densely
pubescent on the upper (adaxial) side with whitish pluricellular hairs; similar hairs
less dense beneath; all parts on both sides, but more abundantly beneath, beset with
stipitate yellow-resinous glands
TYPE: Upper west slope of Blue Mt. Peak, Parish of St. Thomas, Jamaica,
6500-7325 ft (1981-2233 m), L. M. Underwood 1513, 11-12 Feb 1903 (NY).
Paratype from lower western ridge of Blue Mt. Peak, 5500-6318 ft (1700-1950 m),
4-9 July 1926, W. R. Maxon 10025, (NY, US).
This is the first species of sect. Adenophyllum to be reported from the West
Indies. Most of the other species of this section are South American, but the closely
related Thelypteris pilosula (Mett.) Tryon has an extensive range from southern
Mexico to Peru, and has been reported from Jamaica and Hispaniola.
Thelypteris negligens (Jenm.) Proctor, comb. nov.
Nephrodium negligens Jenm. Bull. Bot. Dept. Jamaica, n.s. 3:21. 1896. TYPE: Jamaica, without
exact locality, Jenman s.n., (NY).
bag ns ote abe naga Proctor, sp. nov.
uropelta, sect. Amauropelta. Ex affintate T. balbisii a quo laminis
ne Selon thachidi et costis laminarum supra sulcatis earum marginibus
pilis incurvatis multicellularibus ca. 0.2 mm longis; venis 9-11 paribus differt.
Rhizome decumbent-ascending or suberect, clothed at the apex with dark brown,
lustrous, pabraie lance-attenuate scales 3-4 mm long. Fronds few, erect-arching,
up to 65 cm long; stipes 4-8 cm long, deciduously scaly toward the base, minutely
stipitate-glandular throughout and lightly clothed with small, curved, pluricellular
hairs. Blades lanceolate, 45-60 cm long and up to 18 cm broad below the middle,
rather abruptly narrowed at the base, acuminate at the apex. Rhachis yellowish-
rown, together with the costae densely clothed just inside the adaxial groove with
short (ca. 0.2 mm long), stiffly incurved, pluricellular hairs; underside of the
pegs minutely stipitate- ore and sparingly clothed with a few long, transpar-
t, septate hairs up to 1.5 mm long. Pinnae mostly at right-angles to the rhachis,
Bae the lower reduced ones somewhat reflexed, the largest linear- to narro owly
deltate-oblong, 1.5—-2 cm bi at the base, sessile, acuminate, deeply pinnatifid,
with up to 23 pairs of segments; very small, brown aerophores present at abaxial
base of costae. Segments Sbtong. subfalcate, 2.5—-3.5 mm wide and not over 6 mm
long, subacute at the apex, the margins strigillose-ciliate, and with 9-11 pairs of
simple veins. Veins lightly strigillose on upper (adaxial) side; veins and tissue
beneath with small, erect, unicellular, straight hairs and numerous sessile, reddish-
G. R. PROCTOR: NOTES ON JAMAICAN FERNS-III 59
resinous glands. Sori medial to supramedial; indusium erect, glabrous, densely
resinous-glandular; sporangia glabrous.
TYPE: 1 mile N of Spring Garden, Parish of Trelawny, Jamaica, 1500-1700 ft
(457-518 m), 2 Mar 1978, G. R. Proctor 37704 (IJ).
This species is known only from the type specimen. It appears to be related to
Thelypteris balbisii (Spreng.) Ching, but differs in the nature of its indument, in the
deflexed lower pinnae, and in the more oblique segments with fewer veins. From the
related T. randallii Maxon & Morton ex Morton it differs in the much thicker
thachis clothed on the sides of the adaxial groove with stiff, incurved hairs, and in
having sessile, reddish-resinous glands only beneath. These three species and the
next (7. harrisii) differ from the rest of sect. Amauropelta in having pluricellular
hairs; in T. balbisii such hairs, however, may be present or absent.
JAMAICA
FIG 1. Known localities of cited species and varieties of Thelypteris. In order from west to east ( left to
right): T. trelawniensis, T. resinifera var. caribaea, T. harrisii and T. malangae vat. sitiorum, i.
negligens, and T. decrescens.
Thelypteris harrisii Proctor, sp. nov.
tdi kaulfussii sensu Jenm. Ferns Brit. W. Ind. & Guiana 210. 1908, non Hook., 1862.
Jenman believed that this plant had been treated by Grisebach (Fl. Brit. West Ind. 691. 1864) as a
variety of what is now called Thelypteris oligocarpa, and rightly rejected this assignment. However,
taking up the name kaulfussii, he resurrected an epithet whose antecedents had been hopelessly
confused by Hooker (Sp. Fil. 4:97. 1862), and which cannot be applied to the present pie =
ubg. Amauropelta, sect. Amauropelta. Ex affinitate T. underwoo coger q y
rachide dense pilosa, pilis ca. 1 mm longis vel longioribus; indusiis pilis se
f
nnae, the lowest often being mere auricles; rhe
throughout and also soft-pilose with long, spreading, whit
mm long or longer. Pinnae mostly linear-ob
1.5-1.8 cm broad above the sessile base; costae and other va
60 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
throughout with soft hairs; tissue freely resinous-glandular beneath and nearly
glabrous. Segments close, strongly oblique, oblong, acute, up to 3 mm broad, the
margins narrowly reflexed; veins 7-9 pairs, the lower ones often forked, all strongly
prominulous on the upper (adaxial) side. Sori supramedial; indusium relatively
large, round-reniform, densely long-pilose and ciliate, but without glands, decidu-
ous; sporangia glabrous.
TYPE: Moody’s Gap, border of Parishes of St. Andrew and Portland, Jamaica,
ca. 3000 ft (914 m), 22 Oct 1898, W. Harris 7430, (IJ; isotypes BM nae
Paratype from same locality, 13 Feb 1900, W. N. Clute 173 (NY).
Thelypteris harrisii seems to be related to T. underwoodiana (Maxon) Ching, but
clearly differs in its much longer stipes, denser, longer, and softer pilosity, and in
having the indusium pilose but without glands, instead of ciliate and resinous-
glandular.
Thelypteris gracilenta (Jenm.) Proctor, comb. nov.
Polypodium gracilentum Jenm. Bull. Bot. Dept. Jamaica, n.s. 4:129. 1897. TYPE: Jamaica, without
exact locality, Jenman s. n. (N.Y.)
is species somewhat resembles a large. T. gracilis in general appearance, but
markedly differs from that species in details of the indument, sorus, and indusium.
It is not clear why Jenman placed it in Polypodium, in view of its evident indusium,
unless through faulty observation. Also, he stated that it is “common from 3,500 to
5,000 ft. altitude, in grass by the sides of open shallow streams and in similar wet
exposed places.” The total lack of any subsequent collections indicates, however,
that it must in fact be very rare, and its continued existence needs confirmation.
Thelypteris resinifera var. caribaea (Jenm.) Proctor, comb. & stat. nov.
Nephrodium caribaeum Jenm., J. Bot. Brit. For. 24:270. 1886. TYPE: North slopes of Mt. Diablo,
Parish of St. Ann, Jamaica. Sherring s. n., (K photo US; isotypes IJ. US).
Dryopteris caribaea (Jenm.) C. Chr., Ind. Fil. 257. 1905.
Thelypteris caribaea (Jenm.) Morton, Amer. Fern J. 53:65. 1963.
In recognizing this plant as a distinct species, Morton stressed characters which
do not provide clear and sharp differentiation from 7. resinifera, but which in each
case are more a question of degree (e.g., relative cell width vs. cell length in the
clathrate rhizome scales, relative hairiness of the indusium). Although the ensemble
of differences suggests a recognizable local variant of T. resinifera, more can hardly
be said until living plants are rediscovered.
—
Thelypteris malangae var. sitorium (Jenm.). Proctor, comb. nov.
Nephrodium jenmanii var. sitiorum Jenm. J. Bot. Brit. For. 17:261. 1879. TYPE: Jamaica, without
exact locality, Jenman 38, in 1878 (K; isotype US).
Nephrodium conterminum sensu Jenm. Bull. Bot. Dept. Jamaica n.s. 3:45. 1896, non Aspidium
conterminum Willd. in L., 1810.
Dryopteris consanguinea var. aequalis C. Chr. Smiths. Misc. Coll. 52:380. 1909. TYPE: Second
Breakfast Spring, Parish of St. Andrew, Jamaica, W. R. Maxon 997 (US) (= Underwood 2131, NY).
Differs from typical 7. malangae of Hispaniola in having narrower, more tapering
pinnae (mostly 1.5—-1.8 cm wide vs. usually over 2 cm wide), the segments usually
distinctly crenulate and relatively shorter and broader, and in the presence on
G. R. PROCTOR: NOTES ON JAMAICAN FERNS-—III 61
Jamaican plants of widely scattered, very minute, colorless, stipitate glands,
especially in the adaxial grooves of the rhachis and costae. In addition, the sori of
var. malangae are approximately medial, whereas those of var. sitiorium are
submarginal.
Thelypteris rudis f. cristata Proctor, f. nov.
A forma typica marginibus pinnarum apicem versus integerrimis, apice ipso
cristato-laciniato differt.
Differs from the typical form in having the distal part of the pinnae entire, at the
end expanding into a cristate-laciniate apex.
PE: Jamaica, without definite locality or collector, J. P. 1232-a (K; isotype
IJ). This plant was gathered in 1885, probably by J. H. Hart or one of his
colleagues in the then Botanical Department of Jamaica.
LITERATURE CITED
PROCTOR, G. R. 1965. Taxonomic notes on Jamaican ferns. Brit. Fern Gaz. 9: Sie
—. 1968. Taxonomic notes on Jamaican ferns—II. Brit. Fern Gaz. 10:21-25,
SMITH, A. R. 1974. A revised classification of Thelypteris subg. eamiaabe "Amer. Fern
J. 64:83-95.
62 AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 2 (1981)
SHORTER NOTES
NOTES ON NORTH AMERICAN LOWER VASCULAR PLANTS-II.—Field
work in Arizona and examination of herbarium specimens at ASU, LL, and NY have
revealed several new state records for Arizona and various states in northern Mexico.
Also presented here are range extensions for two species within states where
previously reported, one for Arkansas and one for Arizona. I am especially grateful
to have been able to examine the numerous fine collections of Marshall C. Johnston
and his former students, Thomas L. Wendt and Fernando Chiang C.
A second locality in Arkansas is now known for Cheilanthes eatonii Baker in
Hook. & Baker. The collection data are: Benton Co., Arkansas, E. N. Plank s. n. in
1899 (NY). The species was first reported in Arkansas in Baxter Co., by W. C.
Taylor & D. Demaree (Rhodora 81:514. 1979). They reported it as C. castanea
Maxon, which I do not recognize as distinct from C. eatonii [A monograph of the
fern genus Cheilanthes section Physapteris (Adiantaceae), Ph.D. Dissertation Ari-
zona State University, 1979].
Cheilanthes < parishii Davenp. (pro sp.) has been found new to Arizona. The
collection data are: Dushey Canyon, Harquahala Mts., Maricopa Co., Arizona,
growing near C. covillei Maxon and C. parryi (D. C. Eaton) Domin, desert scrub
vegetation with Saguaro, Ocotillo and Jojoba, igneous substrate, 3000 ft elevation,
Reeves 7127 (ASU). Reported previously from three localities in southern California
by A. R. Smith (Madrofio 22:377. 1974). I agree with Smith that this is a sterile
hybrid between the species listed above. A single plant was found at this site where
the presumed parents are abundant.
The first collection from Chihuahua, Mexico for Notholaena bryopoda Maxon has
been made. The collection data are: Sierra del Roque, N of Julimes and N and NW
of Rancho el Sauz, 28°39’—28°41'N, 105°20’18”—105°20'30"W, 1450-2150 m ele-
vation, matorral desertico con espinos laterales, steep slopes of limestone mountains,
limestone gravel, with Acacia neovernicosa, Dasylirion, Agave lecheguilla, Fouqueria
splendens, and Parthenium incanum, M. D. Johnston et al. 12314 (LL). Previously
known from Coahuila and Nuevo Leén, where it occurs on gypsum, according to
R. M. Tryon (Contr. Gray Herb. 179:77. 1956).
Notholaena greggii (Kuhn) Maxon is now known from Nuevo Leén. The collec-
tion data are: Sierra Madre Oriental, Nuevo Leén, Mexico, calcite and limestone
hills beyond Pablillo toward Santa Clara, 15 mi SW of Galeana, scattered on bank of
calcite, C. H. & M. T. Muller 1080 (LL). Previously known from Texas, Chihua-
hua, Coahuila, and Durango (Tryon, 1956, p. 76).
The first collection of Notholaena neglecta Maxon in Nuevo Leon has been made.
The collection data are: Minas “Manto Blanco” y “Sabana Blanca” just N of the
Canon de Potrerillos, Nuevo Leén, Mexico, 26°04’N, 100°45’W, 950-1000 m
elevation, crasi-rosulifolios espinos, limestone ridge, gypsiferous clay loam, with
Agave lecheguilla, Hechtia, Fouqueria, Larrea, and Opuntia rufida, M. C. Johnston
et al. 10248C sae Previously known in Texas, Arizona, Chihuahua, and Coahuila
(Tryon, 1956, p. 76).
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 2 (1981) 63
Notholaena parvifolia Tryon has been collected in San Luis Potosi. The collection
data are: Estaci6n Microondas “Pastoriza” about 22 km S of Matehuala, San Luis
Potosi, Mexico, 23°25'05"—23°25'25"N, 100°38’50’—100°39'00"W, 1550-1650 m
elevation, crasi-rosulifolio espinoso, a few patches of matorral, limestone hills,
calcareous gravelly soil, with Orthosphenia mexicana, Cnidoscolus sp., Eysenhardtia
sp., and Agave lecheguilla, M.C. Johnston et al. 11111B (LL). Previously known
from New Mexico, Texas, Chihuahua, Coahuila, Nuevo Leén, Tamaulipas and
Zacatecas (Tryon, 1956, p. 99).
The first record for Pellaea intermedia Mett. ex Kuhn in San Luis Potosi has the
following collection data: 1 km by winding road below and W of Real de Catorce,
on road to Estacién Catorce, above Socavon La Purisima, San Luis Potosi, Mexico,
23°41'40"N, 100°53’50”W, 2400-2450 m elevation, crasi-rosulifolio espinoso, badly
disturbed agriculturally, very steep canyon slopes of metamorphic rock, with Agave
spp. and Opuntia spp., M. C. Johnston et al. 11070A (LL). Known previously from
Arizona, New Mexico, Texas, Chihuahua, Sonora, Coahuila, Nuevo Leon and
Zacatecas, according to A. F. Tryon. (Ann. Missouri Bot. Gard. 44:179. 1957).
Pitryogramma triangularis (Kaulf.) Maxon var. triangularis is now known to
occur in Arizona. The collection data are: Frehner Canyon, Virgin Mountains,
Mohave Co., Arizona, very scarce, steep rocky N slope, granite, with Pinon,
Quercus turbinella, Ephedra, Galium, Stipa, Sitanion, and Thamnosma, 5000 ft
elevation, R. Gierisch 4598 (ASU). This variety was previously known from
southern British Columbia, Washington, Oregon, California, Baja California, south-
ern Nevada, and southwestern Utah, according to K. S. Alt and V. Grant (Brittonia
12:155. 1960). Variety maxonii Weath. occurs in central and southern Arizona. The
locality reported here for variety triangularis is in the extreme northwestern corner
of Arizona.
The first collection of Polypodium glycyrrhiza D. C. Eaton in Arizona has been
made. The collection data are: Devil’s Chasm, Sierra Ancha, Gila Co., Arizona,
narrow, deep gorge, one large patch ca. 6 X 6 ft on cliff, ca. 5000 ft elevation, B.
Warner s.n., 10 Jan 1979 (ASU). Known previously from Kamtchatka, the Aleutian
Islands, Alaska and coastal British Columbia, Washington, Oregon, and California
(south to central part of state), according to R. M. Lloyd and F. A. Lang (Brit. Fern
Gaz. 9:171. 1964). The specimen examined has the “sweet” rhizome and free
venation of P. glycyrrhiza, in contrast to the acrid rhizome and usually anastomosed
venation of P. californicum Kaulf. The material does not resemble P. hesperium
Maxon, which is rather widely distributed in Arizona. The presence of P. glycyrrhiza
in Arizona brings to three the number of predominantly Pacific Coast ferns found
disjunctly in central Arizona. The other two are Dryopteris arguta (Kaulf.) Watt. and
Woodwardia fimbriata J. E. Smith in Rees. na ae
Selaginella eremophila Maxon is now known from several additional localities in
southwestern Arizona. The collection data are: Sierra Estrella Regional Park,
Maricopa Co., Arizona, W-facing wash on west face of Squaw Tit, moist desert
under rocks with Penstemon antirrhinoides, Salvia mohavensis, Notholaena standleyi,
and Castilleja lanata, 2400 ft elevation, E. & M. Sundell 177 (ASU); West side of
White Tank Mountains, Maricopa Co., Arizona, slope leading up to light 1, lower
64 AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 2 (1981)
sonoran desert slope, at base of cliff, D. Keil 4088 (ASU); 7 mi S of Buckeye on
U.S. 80, 2.4 mi along Buckeye Recreation Area Park road, on S-facing slope, under
rocky ledge, 1189 ft elevation, A. Pierce 2/1 p. p. (a small fragment is apparently
S. arizonica Maxon); Cabeza Prieta Game Range, Yuma Co., Arizona, S4, T14S,
RI5W, ca. 6 mi NW of Tule Well, E-facing slope of low mountain, E. Lehto et al.
23548 (ASU, NY, US). This species previously was known in Arizona from a single
collection near Tinajas Altas, Yuma Co., collected by Jaeger in 1934, according to
R. Tryon (Ann. Missouri Bot. Gard. 42:80. 1955). All of the cited collections
previously had been identified as S. arizonica, which is more common in Arizona
than is S. eremophila. So far as I know at present, the two species occur together
only at the Buckeye locality. I have seen material of S. arizonica from the White
Tank Mountains, but from a different locality than the one cited above for S.
eremophila. Selaginella eremophila is otherwise known from southern California
and Baja California (Tryon, 1955).
Field work in Arizona was supported by National Science Foundation Dissertation
Improvement Grant 77-00182 to Dr. D. J. Pinkava and the author. I thank the
curators of the cited herbaria for permission to examine their collections.—Timothy
Reeves, Biological Science Center, Boston University, Boston, MA 02215.
EQUISETUM ARVENSE IN ALABAMA.—The Common Horsetail, Equisetum
arvense L., has been reported from Alabama by various authors from Small (Ferns
of the Southeastern States, 1938) to Evans in Radford, Ahles, and Bell (Manual of
the Vascular Flora of the Carolinas, 1968), none of whom gave any indication of
locality. According to Dean (Ferns of Alabama, 1969, p. 153), “A large colony was
discovered in Marengo County by Dr. R. M. Harper.” This was puzzling because
Marengo County is in the coastal plain of southwestern Alabama, but E. arvense is a
northern plant reaching its southern limit in Alabama. One would expect the plant to
be present only in the northern part of the state; indeed, Wheatstone and Atkinson
(Castanea 44:1—8. 1979) found it in Morgan County in central northern Alabama in
1974. It also has been found recently in Calhoun County (R. R. Haynes, UNA).
While boating on the Black Warrior River in Greene County in October 1978,
I was quite surprised to find a thriving colony of E. arvense on the west bank of the
river. This locality is in the coastal plain about 20 miles upstream from Demopolis,
which is in Marengo County. Therefore, this find lent credence to the report from
that county. The Greene County colony was much larger and denser than the Morgan
County population. It was growing in damp, partially shaded sand at the base of a
wet chalk bluff, just above the normal level of the river (Short 1183, AUA, and
duplicates to be distributed). On a visit to the U.S. National Herbarium in April,
1980, I examined a specimen of E. arvense (R. M. Harper 121, US) collected on
October 11, 1908 in Marengo County on the bank of the Tombigbee River
about 10 miles downstream from Demopolis. The plants were growing in sand at the
base of a wet chalk bluff. This evidently is the collection referred to by earlier
authors.—John W. Short, 905 McKinley Ave., Auburn, AL 36830.
BRITISH PTERIDOLOGICAL SOCIETY
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residents should apply to Dr. J. Skog, Biology Department, George Mason
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G. Coke, Robin Hill, Stinchcombe, Dursley, Gloucestershire, England.
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AMERICAN Boe
FERN Aas
J O U R NA L July-September, 1981
QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY
Arachniodes simplicior New to South Carolina
and the United States JUDITH E. GORDON 65
A New Isoétes from Jamaica R. JAMES HICKEY 69
Leaf Turnover Rates and Natural History of the
Central American Tree Fern Alsophila salvinii RALPH L. SEILER 75
Nomenclatural Notes on Micronesian Ferns F. R. FOSBERG and M.-H. SACHET 82
x Asplenosorus shawneensis, a New Natural Fern
Hybrid Between Asplenium trichomanes and :
Camptosorus rhizophyllus ROBBIN C. MORAN 85
Notes on North American Ferns DAVID B,. LELLINGER 90
Shorter Notes: Salvinia minima New to Louisiana;
An Unusual Record of Asplenium trichomanes
from Northeastern Florida 95
Reviews 68, 8&4
S i : : ss aici 06
uggestions to Contributors +
MISSOURI ome
ocT 17 s196f"
GARE WT
The American Fern Society
Council for 1981
ROBERT M. LLOYD, Dept. of Botany, Ohio University, Athens, OH 4570 President
DEAN P. WHITTIER, Dept. of Biology, Vanderbilt University, Nashville. Ane 47235. -Vice Pah i!
MICHAEL I. COUSENS, Faculty of Biology, University of West Florida, Pensacola. FL 325
eee
JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, TN 37916.
Treasurer
JUDITH E. SKOG, Dept. of Biology, George Mason University, Fairfax, VA 22030.
Records Treasurer
DAVID B. LELLINGER, Smithsonian Institution, Washington, DC 20560. Journal Editor
ALAN R. SMITH, Dept. of Botany, University of California, ih CA 94720. Memoir Editor
JOHN T. MICKEL, New York Botanical Garden, Bronx, NY 104 Newsletter Editor
American Fern Journal
EDITOR
DAVID B. LELLINGER Smithsonian Institution, Washington, DC 20560.
ASSOCIATE EDITORS
DAVID W. BIERHORST Rt. 3, Box 188, Picayune, MS 39466.
GERALD J. GASTONY Dept. of Biology, Indiana University, Bloomington. IN 47401.
JOHN T. MICKEL New York Botanical Garden. Bronx, NY 10458.
The “American Fern Journal” (ISSN 0002-8444) is an illustrated quarterly devoted to the general
study of ferns. It is owned by the American Fern Society, and or: at the Smithsonian Institution,
Washington, DC 20560. Second-class postage paid at Washin
Claims for missing issues, made 6 months (domestic) to ie suis (foreign) -after the date of issue,
and the matters for a should be addressed to the Editor.
Changes of address, dues, and applications for membership should be sent to Dr. J. E. Skog. Dept. of
Biology, George se University, Fairfax, VA 22030.
Orders for back issues should be addressed to the Treasure
General inquiries oe ferns should be addressed to a Secretary.
Subscriptions $9.00 gross, $8.50 net if paid through an agency (agency fee $0. os sent free to
members of the Aenchiah Fern Society (annual dues, $8.00; life membership. $160.
Back volumes 1910-1978 $5.00 to $6.25 each; single back numbers of 64 pages or 68 $1.25: 65-80
pages, $2.00 each; over 80 pages, $2.50 each, plus shipping. Back volumes 1979 et seq. $8.00 each;
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Library
Dr. John T. Mickel, New York Botanical Garden, Bronx, NY 10458. is Librarian. Members
may borrow books at any time. the borrower paying all shipping costs.
Newsletter
John T. Mickel, New York Botanical Garden, Bronx, NY 10458. is editor of the newsletter
“Fiddlehead Forum.” The editor welcomes contributions from members and non-members, including
miscellaneous notes, offers to exchange or purchase materials, personalia. horticultural notes, and
reviews of non-technical books on ferns.
Spore Exchange
Mr. Neill D. Hall, 1230 Northeast 88th Street, Seattle, WA 98115, is Director. Spores exchanged and
collection lists sent on request.
Gifts and Bequests
Gifts and bequests to the Society enable it to expand its services to members and to others interested
in ferns. Botanical books, back issues of the Journal, and cash or other gifts are always welcomed, and
are tax-deductible. Inquiries should be addressed to the Secretary.
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 3 (1981) 65
Arachniodes simplicior New to South Carolina
and the United States
JUDITH E. GORDON*
During the past year, a population of about 100 plants of Arachniodes simplicior
(Mak.) Ohwi was found along nearly 100 m of the bank of a small, unnamed
tributary of the Savannah River in North Augusta, South Carolina. The species was
identified using Ching’s (1934) treatment of Asiatic species of Rumohra Raddi, a
genus now correctly designated Arachniodes Blume, as explained by Tindale (1961).
The identification was confirmed by the United States National Herbarium. Speci-
mens from the American population are described as ee (see also Figs. 1, 2):
Rhizome moderately creeping, commonly S—8 mm in diameter, densely covered
with tan to cinnamon-brown, papery scales, these long triangular with auricular-
clasping bases, 0.5-2.5 mm wide, 3-8 mm long, the tips one cell wide and 2-4
cells long, the margins entire except for | "3 trichomes, 3—5 cells lo geod near scale
bases and a few basal teeth A ose from outward distensions of two cell end
junctures. Stipes commonly 0.5—4 cm apart, 15—46 cm long, mm diane: at
the base, often equal to or longer than aie laminae, sub-terete, latened on the upper
surface and grooved toward the apex, tan at the base, pale green above, aging to a
straw-yellow color, somewhat scabrous from small preiberences on ica scattered
scales are borne, the scales more oe sehr ae tibet the stipe ,2-I12m
long, and similar to those of the rhizome ex cept more elongate and hocolale: brown,
~ Len consisting of 2 large and 3 small fanilles at the base, fe he to 3 at the
Rachis unwinged, pale green, the upper surface grooved, the scales like those
of the stipe except smaller. Costae yellow-green, with grooves continuous with Spa
o me rachis, the scales smaller and more abruptly ogee Laminae ts
26-34 cm wide, deltoid- Sonal 2—3-pinnate, coriaceous, Tare ial, the
mata surfaces glossy dark green with yellow- as Be along the costes and
adjacent bases of pinnules, the abaxial surfaces uniformly pale green and not glossy,
the apex gradually acuminate. Pinnae generally |-pinnate, 3—4 Bette pairs below
i i m long cm
the terminal pinna, 1-6 cm distant, at 30—45° to the rachis, 8-1 ,
wide at the base except the basal pair, 2-pinnate with the base 8-10 cm wide, the
petiolules 1-6 mm long, the apices gradually acuminate, the pinnules and venation
anadromically arated on all pinnae. Pinnules, except the lower pairs on the basal
pinna pair, 16—22 alternate pairs, asymmetrically ovate-auriculate with an acroscopic
basal lobe, 13-22 mm long, 5-10 mm wide, the petiolules narrowly whe gies sessile
or up to | mm long, the apices sie, the margins with spines up to 1 mm long,
with a few trichomes along the larger veins; basal pinnule pair on the nee pinna
pair basiscopically produced with the upper about /- ys the length of the lower, the
latter 6-10 cm long, the ultim ee segments resembling the pinnules of the 2-pinnate
pinnae. Pintiale venation free, semi-dichotomous, not extending to the margins, the
first branch arising anadromically. Sori dorsal, globular, arranged in a single row on
each side of the pinnule main vein, terminal on smaller lateral veins, somewhat
closer to the main vein than the margin. Indusia 0.5-1.0 mm in diameter,
slightly darker brown in the center, glabrous, the margins entire, persisting for about
two months before being shed. Sporangia with stalks composed of three rows of
*Department of Biology, Augusta College, Augusta, GA 30910.
Volume 71, number 2, of the JOURNAL, was issued June 29, 1981.
66 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
FIGS. 1 and 2. Photographs of Arachniodes simplicior. FIG. 1. Frond as seen in natural habitat, FIG. 2.
Lowermost basal pinna, dorsal view.
J. E. GORDON: ARACHNIODES NEW TO THE UNITED STATES 67
cells, 0.3-0.4 mm long, englandular, the annulus of 13-16 thickened cells. Spores
bilateral, dark brown, rugose-reticulate, 4144ym X 28—-324m. Chromosome
number 2n = 164 (Léve et al., 1977), undetermined for this population.
The unusual nature of the habitat deserves further mention. The area, although
within the city limits, is accessible only by foot and is characterized by steep,
southwest-facing cliffs and heavy undergrowth within southern hardwood forest that
probably has not been logged. The area surrounding the creek is dominated by
Fagus grandifolia with a scattering of Celtis occidentalis and various species of
Quercus. The understory is dominated by a species of Aesculus, apparently a hybrid
between A. pavia and A. sylvatica. Polystichum acrostichoides (Michx.) Schott plants
grow intermixed with the population of A. simplicior, but are less numerous. A
small population (ca. eight plants) of Pteris multifida Poir., a naturalized species, 1s
also present along the creek bank. The site has the following coordinates: 33°30’ 11”
N Lat., 81°58'54” W Long., 42-53 m elev. This places the site along the western
edge of the Hammond Hills subdivision.
The area is seldom visited by hikers, but there have been two major disturbances
within the last 50-55 years according to city officials. About 1927-29, the Georgia
and Florida Railroad extended a line northward through the area. The rail bed was
built with fill (source unknown) rising about seven meters above the creek bed. the
creek water being piped through at the base. Plants of A. simplicior grow within a
meter of the pipe. Use of the rail line was discontinued about 1954, at which time
the city laid a sanitary trunk sewer line through the area. The sewer pipe, which is
elevated about seven meters, parallels the old rail bed and is downstream about 60
m. If the population were present in 1954, it would have been disturbed by the
construction of the sewer line.
Observations of greenhouse and field plants show a growth pattern based on two
new fronds produced yearly, usually in April in the North Augusta area. The
persistence of the old frond bases permits a rough estimate of individual rhizome
age, obtained by counting old bases and dividing by two. Ten rhizomes were
examined; ages ranged from 2-15 years, using this method of calculation. Consider-
ing the likely decay of older rhizome portions, the population size, and possible
establishment from a spore source, it appears that the population is at least 20-25
years old.
There are several possible spore or rhizome sources in the vicinity, including
several nurseries, although none is within a mile radius of the site. One of the
nurseries reported selling A. simplicior about three years ago, but not prior to that
time nor within the last year. Another source of spores or rhizomes would include
plants purchased by residents of the Hammond Hills subdivision, which was
established about 1955. The probability of someone having actually planted the fern
at the site is highly unlikely since access to the area is difficult. Based on the
available information, I believe the population was probably established from a spore
source shortly after the construction of the sewer line in 1954. Further studies
centering on chromosome counts and gametophyte developmental stages are antici-
pated.
68 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
LITERATURE CITED
CHING, R. C. 1934. A revision of the compound leaved Polysticha and other related species in the
continental Asia including Japan and Formosa. Sinensia 5:23-91.
LOVE. pt D. LOVE, and R. E. G. PICHI SERMOLLI. 1977. Cytotaxonomical Atlas of the
erido’ saa rene Vaduz.
TINDALE, M. D. 1961. Aspidiaceae in nals of South Eastern Australia. Contr. N. S. W. Natl.
erb. Ben Leong 208-21 1:1
REVIEW
VASCULAR PLANTS OF CONTINENTAL NORTHWEST TERRITORIES,
CANADA, by A. Erling Porsild and William J. Cody. 1980. 667 pp. National
Museums of Canada Publ. Div., Ottawa, Canada K1A OM8. ISBN 0-660-001 19-5.
Can$80.00.—This admirable volume provides descriptions, keys, habitat data, and
maps for 1113 species of vascular plants which occur in the region immediately
north of British Columbia, Alberta, and Saskatchewan. Information also is included
on a number of species expected by the authors to occur in the region. There is an
excellent survey of the history of collection in the region and a worthwhile bibliogra-
h
y.
The pteridophytes are treated as 40 species, one subspecies, and one variety
present, with four more species expected. Nearly half are fern allies. There are no
great surprises in this subarctic flora; no hybrids or taxonomic novelties were noted.
The limestone plants are of interest.
I found the distribution maps to be curious and unexpected. They include Alaska
and the Aleutians to the west, Greenland and Iceland to the east, and Chicago to the
south. Thus, they are too small for detailed distributions within the region covered
by the book. They do nothing to support the six phytogeographic zones designated
by the authors, and certainly do not give a clear picture of the tree line, areas well
collected, or other extrapolations one usually extracts from dot maps.
Although we are warned (p. 3) that the maps are “in no way complete” for the
Canadian distributions and are meant to show a “broad picture,” they have serious
shortcomings. A person interested in Newfoundland might conclude that Woodsia
alpina or Phegopteris connectilis does not occur there. Also, no fiddleheads in New
Brunswick where they are canned in quantity? More serious are the maps which
record more than the taxon being treated, e.g., Botrychium lanceolatum versus
subsp. lanceolatum, Gymnocarpium dryopteris versus subsp. disjunctum, and Wood-
sia oregana versus W. cathcartiana.
I think it is a great pity that the authors fall back on names such as Dryopteris
disjuncta for Gymnocarpium dryopteris and D. phegopteris for Phegopteris
connectilis. 1 do not like Polypodium vulgare subsp. virginianum for P. virginianum
or Dryopteris dilatata (the common tetraploid of Europe) for D. expansa, but these
are newer and more problematical changes
All in all, this book is a most useful compendium of the pteridophytes of a broad
area of boreal, sub-arctic and arctic Canada and will be a constant source of
reference for anyone interested in this part of the world.—D. M. Britton, Dept. of
Botany and Genetics, University of Guelph, Guelph, Ont. NIG 2W1, Canada.
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 3 (1981) 69
A New Isoétes from Jamaica
R. JAMES HICKEY*
The genus /soétes L. is poorly represented in the Caribbean, with only two
species reported to date. /soétes cubana Baker collections made by Wright in the
western, lowlands of Cuba. To my knowledge, there have been no subsequent
collections of this species from Cuba, although certain collections from Belize
strongly resemble the Cuban material and probably are conspecific with it. /soétes
tuerckheimii Brause is known from several collections made in the Cordillera
Central in the Dominican Republic at altitudes of 2200-2900 meters. Thanks to the
generosity of Dr. George Proctor, who kindly supplied spirit and herbarium material,
I am able to report the discovery of a third Caribbean species which is endemic to
Jamaica.
Isoétes jamaicensis Hickey, sp. nov. Figs. 1-4, 6, and 8
Radices dichotome ramosae. Cormus trilobatus, 1-2 cm diametro. Folia 20-55
(X =38), 8-20 cm longa, 0.5—0.8 mm lata in medio, attenuatissima, recurvata.
Stomata et fasciculi fibrosi peripherales praesentes. Velum parvum. Margines ligulae
hyalini, ephemeri; area centralis triangularis, attenuata versus apicem. Megasporae
albidae, 320-440 (X=398) pm, tuberculatae. Microsporae cinereae, ellipticae,
30-40 (x = 36) ym longae, 22.5-32.5 (X= 27.3) um latae, cum papillis cavis.
TYPE: In mud of drying seasonal rain pools, ca. 350 ft, Harris Savanna,
Clarendon Parish, Jamaica, 26 Nov 1974, G. R. Proctor 34357 (1).
PARATYPE: Harris Savanna, Clarendon Parish, Jamaica, 26 Nov 1974, G. R.
Proctor 34358 (IJ).
In habit and spore morphology, /. jamaicensis most closely resembles /.
montezumae Eaton of Mexico (see Table | for comparison with other Mexican and
Caribbean species). Although the distinctness of the latter species from /. mexicana
Underw. has been questioned (e.g., Reed, 1953; Pfeiffer, 1922), preliminary
examination of numerous Mexican collections, including type specimens, suggests
that hybridization, possibly involving the formation of partially fertile hybrids, has
caused the confusion between these species. In light of this possibility, /. montezumae
certainly merits provisional specific status until my more detailed study of these
species and /. pringlei Underw. is completed.
Isoétes jamaicensis and I. montezumae differ in several respects, of which the
more salient are discussed here. The corm of /. jamaicensis is distinctly three-lobed,
whereas in the latter it is deeply bilobed. The membranaceous margin at the base of
the leaves is less pronounced in /. jamaicensis than in /. montezumae. In the former,
it is 1-2 mm wide and merges into the leaf proper 5-10 mm above the sporangium;
in the latter, the margin is 1.5-2 mm wide and extends 10-20 mm above the
sporangium. Both species have similar velum coverage of the sporangium. Eaton
(1897) reported a very narrow velum in megasporophylIs and virtually none in
microsporophylls of /. montezumae. In I. jamaicensis, there 1s little dimorphism in
*Biological Sciences Group, U-43, University of Connecticut, Storrs, CT 06268.
TABLE |. COMPARISON OF ec sca the AND CARIBBEAN ISOETES SPECIES. !
Character
Habitat
Pana igi
Corm
Leaf a
Leaf length (cm)
Leaf diameter (mm)
Stomates
Fibrous strands
Megaspore ornamentation
Perispore strands
Megaspore size (wm)
Microspore color
Microspore ornamentation
Microspore length £m
Microspore width £m
+
>
white
tuberculate
nsis
I, mexicana
amphibious
1830-2150
grey-white
h
wn
smooth-echinate
25-38
25-33
I. montezumae
amphib-terr
2
8-20
8-14
0.6-1.2
+
+
>
white
tuberculate
+ united
350-510
ash-grey
papillate
23-28
I. pringlei
amphibious
A
10-20
16-25
0.5-1
+
>lA
white
cristate-echinate
'Data obtained from original descriptions, Pfeiffer’s monograph (1922), and personal observations.
I. cubana
amphibious
>50
rudimentary
grey-white
tuberculate
all united
290-400
fawn
echinate-papillate
25-33
20-25
1. tuerckheimii
amphib-aquat
2200-2900
(1861) TZ SWATOA “IWNUNO! NY34 NYOMI
R. J. HICKEY: NEW ISOETES FROM JAMAICA 71
velum coverage between the megasporohylls and the microsporophylls: most of the
leaves show some velum coverage (Fig. / and 2), although in a single microsporo-
phyll of the type collection there was no appreciable velum development. Ligule
characteristics show considerable variability due to the ephemeral nature of the
hyaline margin and due to the delicate nature of the central region, which deterio-
rates rapidly with age (Figs. / and 2). The sporangium wall is unspotted and
consists of thin-walled cells in /. jamaicensis (Fig. 3), just as it does in many
specimens of J. montezumae.
The tuberculate megaspores of both species are quite similar (Figs. 4 and 5).
However, /. montezumae has consistently larger equatorial ridges and is more
variable in the extent of ornamentation; the spores range from distinctly to indistinctly
tuberculate. /soétes jamaicensis is more consistent in possessing well developed
tubercles. Specimens of /. montezumae also show variation in tubercle distribution.
On the distal surface, the tubercles diminish in size and increase in frequency close
to the equatorial ridge. The tubercles of the proximal surface are always less well
developed. In /. jamaicensis, on the other hand, the tubercles are uniformly
developed and distributed.
The tubercles of both species consist of solid masses of perispore material
extending outward from the spore surface (Figs. 6 and 7). In both species, cords of
perispore material radiate out from the tubercles, giving the spores a cobwebby
appearance. While superficially similar, a closer examination of the spores and
tubercles shows some distinct morphological differences. In /. montezumae, the
perispore layer between the tubercles consists of very slender, interwoven strands
which form a mat-like surface (Fig. 7, lower right). These small strands merge and
two groups of them twist together to form the cords which can be seen radiating out
from the tubercles. The cords in /. jamaicensis are also composed of smaller
strands, but they are larger than those in /. montezumae, parallel, and are never
interwoven. Between the tubercles, these small strands are isolated from one another
and do not form any sort of solid structure.
The microspores of /. jamaicensis are ornamented with numerous, conical,
hollow papillae (Fig. 8). In some spores of the type collection, the papillae are quite
pointed, approaching echinate projections in form, and are occasionally branched
apically. The bases of the papillae are confluent. In /. montezumae (Fig. 9), the
papillae are sparse, rounded, and give no indication of being hollow (as evidenced
by their invariably unbroken appearance). The papillae are quite distant, and
between them the microspores are covered with a finely granular perispore. Unlike I.
jamaicensis, the equatorial ridges are visible in /. montezumae when the micro-
spores are viewed from the side. Both species have a prominent proximal suture.
The holotype of /. montezumae (Pringle 3459, MO) shows no indication of spore
abortion, nor do either of the collections of /. jamaicensis. Spore abortion is not
uncommon in many of the Mexican collections of /soétes.
Isoétes jamaicensis inhabits seasonal rain pools in open xerophytic scrub of the
Harris Savanna. This region receives 35-40 inches of rain annually during seasonal
rainfalls. However, according to Dr. Proctor, the pools containing I. jJamaicensis
form only once every four or five years, when the rains are particularly heavy.
AMERICAN FERN JOURNAL: VOLUME 71 (1981)
FIGS. |I-3. Leaf morphology of the type of /svétes jamaicensis (Proctor 34357, \J). FIG. 1.
Megasporophyll showing incomplete velum and degraded ligule, x 9.6. FIG. 2. Microsporophyll from a
more central region of the corm showing velum and ligule which still retains a portion of the apex of the
central region, x 9.6. FIG. 3. Cellular detail from outer sporangial wall of a megasporangium, x 396.
All drawings made with a camera lucida drawing tube.
R. J. HICKEY: NEW ISOETES FROM JAMAICA 73
FI 4-9. peels of the types of /soétes jamaicensis (Proctor 34357, IJ) and J. montezumae (Pringle
IGS.
3459, MO). FIG. 4. Megaspore of /. jamaicensis, equatorial view,
F Megaspore of /. jamaicensis, close up of a partially
FIG. 7. gen of J. montezumae, close up of a
_ 8. Microspore of /. jamaicensis, equatorial
95
x 125. FIG. 5. Megaspore of /.
montezumae, distal view, * 125
developed tubercle, distal surface. 2500.
partially splape er tubercle, distal surface, < 2500.
view, X 1665. FIG. 9. Microspore of /. montezumae, near proximal view,
74 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
Whether the corms break their dormancy in drier years is not known, but observa-
tions on other species of similar habitats suggest that they may not (Hall, 1971).
Considering the unusual habitat and apparent sporadic growth of this plant, it is not
surprising that it has eluded detection this long. It will not be surprising if this species
is collected in other areas of Jamaica or perhaps on some of the other islands of the
Greater Antilles.
LITERATURE CITED
EATON. A. A. 1897. A new quillwort from Mexico (1. seaegn Fern Bull. 5:25.
HALL, J. B. 1971. Observations on Isoetes in Ghana. Bot. J. Linn. Soc. 64:117-139.
PFEIFFER. N. E. 1922. Monograph of the Isoétaceae. a ar Bec Gard. 9:79-232, pl. 12-19.
REED, C. F. 1953. Index Isoétales. Bol. Soc. Brot. 27:5
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 3 (1981) 75
Leaf Turnover Rates and Natural History of
the Central American Tree Fern Alsophila salvinii
RALPH CL. SEILER*
There are few studies of growth rates of tropical trees. Moreover. the growth of
tree ferns in their native habitats has seldom, if ever, been studied because of their
slow growth and the need to make observations over a long period of time. As a
Peace Corps volunteer in El Salvador I had the chance to study the growth of the
cloud forest tree fern Alsophila salvinii Hooker (Cyatheaceae).
Bosque Montecristo is a cloud forest that covers about 15 km* and is located at
the common borders of El Salvador, Guatemala, and Honduras. Approximately 60
percent of the forest is in El Salvador and has been set aside as a national park and
wildlife preserve. The highest point in the forest, the summit of Cerro Montecristo,
has an altitude of 2414 meters. Bosque Montecristo receives an average of 2250 mm
of rain annually. Fog drip during the night deposits about the same amount of water
on the forest (Reyna, 1979). March and April are the driest months. The rainy
season begins in mid-May or early June and continues until December.
Bosque Montecristo is a subtropical, lower montane, very humid forest according
to the Holdridge life zone classification (Holdridge. 1975). The cloud forest proper
begins at ca. 2100 m and extends to ca. 2350 m, where it is replaced by an
ericaceous shrub association on the mountain summits. The ecotone between the
cloud forest and the pine-cypress association below occurs between 1900 and 2100
meters. Secondary forest is found in the whole zone wherever there has been logging
(Reyna, 1979).
The majority of the trees in the cloud forest are evergreen. The emergent trees in
the forest canopy are principally oaks, which are thickly covered with epiphytes
(Reyna, 1979). Tree trunks near the forest floor support lush growths of mosses,
filmy ferns, and many other ferns such as Asplenium, Grammitis, and Elaphoglos-
sum, Reyna (1979) reports 175 species of trees in the Montecristo area and 71
species from the cloud forest proper. In the area where the present study was
undertaken three tree ferns are common: Dicksonia gigantea Karst.. Trichopteris
schiedeana (Presl) Tryon, and Alsophila salvinii Hooker.
Alsophila salvinii is a cloud forest tree fern known from southern Mexico,
Guatemala, Honduras, and El Salvador (Stolze, 1976). It is one of the most
conspicuous ferns in Bosque Montecristo and is the most common tree fern. In very
wet parts of the forest, A. salvinii forms large, dense thickets, often in areas where a
large tree or branch has fallen. The thicket shown in Fig. / is brightly lit during the
morning from holes in the forest canopy. The thicket appears to be perpetuating
itself, as there are hundreds of very young tree ferns present with trunks as yet
undeveloped. Fronds of these are only about 30 cm long. whereas fronds of adults
are about 2.5 m long. ;
Alsophila salvinii is also found in relatively dry, open secondary forest at ca. 2050
meters in Bosque Montecristo. The ferns in the secondary forest don’t form dense
*3977 S. 775 W., Bountiful, UT 84010.
FIG. 1. The thicket of tree ferns in
ae Sk ee
which the study was undertaken. FIG. 2. Alsophila salvinii showing persistent stipes a
nd
=! 3
rachises of dead fronds.
(1861) TZ SWMIOA “IWNYNOF NY34 NVOINIWY
R. L. SEILER: LEAF TURNOVER OF ALSOPHILA SALVINII 77
thickets, are less common, and in general appear less robust than the ferns of the
cloud forest proper. Young tree ferns are not seen in the secondary forest. The
absence of young tree ferns in the secondary forest might be partially due to grazing
by cattle; cattle from a nearby pasture have been observed in the forest eating
fiddleheads from the crowns of adult tree ferns.
The fronds of A. salvinii are tripinnate, and when the tronds die only the
secondary pinnules fall; the stipe and main rachis remain attached to the trunk for
several years (Fig. 2). The base of the stipe is closely appressed to the trunk for
about 15 cm and then arches out to support the frond. Eventually the dead rachis
and most of-the stipe will break off, but the appressed stipe bases remain attached to
the trunk. The bases decay slowly and for many years will hold water either from the
roots or captured from rainfall and condensation. The outer trunk of A. salvinii is
not thickly covered with adventitious roots as it is in some tree ferns such as
Dicksonia gigantea.
The goals of this study were to investigate frond production rates, death rates, and
lifespan; frond phenology; the relation between trunk length and tree age: the effects
of water stored in the appressed stipe bases on frond production and death rates: and
the factors affecting tree mortality.
METHODS
In late June 1978, twenty-eight mature, healthy trees of Alsophila salvinii: were
chosen from a dense thicket (Fig. /) in very wet forest at 2300 m altitude on Cerro
Montecristo and were tagged with forester’s flagging tape. All fiddleheads on these
trees were separately tagged and their subsequent development followed. All dead
fronds were cut from the tree ferns, but the appressed stipe bases were left intact.
The mean trunk length of the 28 selected ferns was 1.18 m (s=0.47). The terns
were randomly divided into equal control and experimental groups. Holes were cut
in the bottoms of the dead stipe bases of the trees in the experimental group with a
pocket knife so that they could no longer hold water.
At irregular intervals over the next two years, eleven visits were made to Bosque
Montecristo to observe the tree ferns. All mature, immature, and dead fronds
present on each fern were counted. Fiddleheads and young fronds in which the
pinnules had not fully expanded were considered immature. All living fronds with
fully expanded pinnules were considered mature. Because dead fronds were removed
at the beginning of the study, the ferns had no dead fronds at the first observation.
Obtaining an accurate count of the number of fronds that died over the period of the
study would have been difficult if dead fronds fell without leaving a trace; this was
not a problem, however, because dead fronds remained attached to the trunk.
RESULTS AND DISCUSSION
Means of mature, immature, and dead fronds per tree in the control group during
the two year period from late June 1978 to May 1980 are shown in Fig. 3. There is a
rapid flush of fronds produced starting at the end of the dry season in early May.
The immature fronds of A. salvinii do not develop synchronously. The first
fiddlehead of the season may be nearing maturity as the last fiddlehead just barely
CVG)
| (14) (12)
(13) (10)
(13)
C13}. C08} (13)
(14)
HME MATURE FRONDS
JE IMMATURE FRONDS
2K DEAD FRONDS
MEAN WKUMBER OF MATURE, IMMATURE AMD DEAD FROMDS PER TREE
: ;
i
24 /
/
/
/
: /
/
/
tbl as
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T T T T t TT 1 Sah cus T qT T 1 & i T T T T 7 af Fa oa T z
J A $ 0 N D J F M A M J J A 5 0 N 1) J F M A M J
MONTH
1978 ‘Beet 1979 et 1980. ————-——»)
_The mean number of mature, immature, and dead fronds per tree of Alsophilia salvinii in the control group during the period June 1978 to May
G.
1980. eo bars indicate 95% contidence limits for the means; number of individuals is shown in parentheses.
(1861) TZ SWNIOA “IWNYNOF NY34 NVOIMSWY
R. L. SEILER: LEAF TURNOVER OF ALSOPHILA SALVINII 19
protrudes from the crown. The asynchronous production of fronds by A. salvinii is
not typical of all tree ferns; in some tree ferns such as Dicksonia gigantea, all the
fiddleheads produced during one growing season develop and are mature nearly
synchronously (personal observation).
Throughout the year, even during the dry season, fiddleheads are present on a few
ferns, but these normally just barely protrude above the trunk apex. In February and
April 1979, several very young fiddleheads were tagged and the time required for
their maturation was observed. Fiddleheads initiated in February, just before the
driest part of the year, required 3-4 months to mature, and one of the tagged
* fiddleheads died during this period. Many of the pinnae of the surviving fronds were
undeveloped, dead, and dried out when observed in April. Fiddleheads tagged in
April 1979, at the end of the dry season, required 2-3 months to mature and all
survived.
Frond death rate fluctuates seasonally (Fig. 3). Few fronds died during the dry
part of the year (January to June). Nearly dead fronds with only a few pinnules
remaining survived this entire period. In June, at about the time the pinnules of the
fronds initiated in April began to expand, the frond death rate sharply accelerated
and was quite constant from June to January. However, the end of the period of rapid
frond death does not coincide with the time the fiddleheads were finally mature.
On | April 1979, the ferns averaged 1.7 dead fronds per tree; one year later they
averaged 4.7 dead fronds per tree (data interpolated from Fig. 3). This indicates an
average frond death rate of three fronds per year per tree. During this same period,
there was no net gain or loss of mature fronds, which together with the calculated
death rate, indicates an average frond production of three fronds per tree per year.
Since the number of mature fronds per fern averages about six, the death rate of
three fronds annually implies a frond lifespan of about two years. The period trom
April to April was used to measure net changes in the number of mature and dead
fronds because April is in the middle of a 3—4 month period when there are no rapid
changes in the number of mature and dead fronds.
The lives of the 27 fiddleheads tagged in late June 1978 were followed for two
years. By October 1978 all the tagged fiddleheads were mature. In February 1979,
about half the fronds had begun to drop some pinnules. In July 1979, one year after
the fronds were initiated, all those not killed by catastrophe were alive, although
nearly all were dropping pinnules. By December 1979, one of the fronds was dead,
but at the last observation in May 1980 (22 months after the fronds were initiated)
no more had died. It is probable that the fiddleheads tagged in June 1978 died
during the 1980 wet season.
If the number of fronds produced by a given length of trunk is constant, the age of
a tree fern can be estimated. To determine the number of fronds produced in 50 cm
of trunk, ten ferns not previously included in the study were selected from the same
thicket. The number of stipe bases between 20 and 70 cm below the crown was
counted on the ten tree ferns; the mean was 18.1 fronds/ 50 cm (s= 1.59). Since
about three fronds are produced annually, about 12 years are needed to grow a meter
of trunk under the wet conditions of the primary forest. The growth rate may be
different in the drier secondary forest. The exact age of a tree fern cannot be
80 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
calculated from its trunk length because neither the time needed for establishment
nor the rate of frond production in very young individuals is known.
The lifespan of some of the tree ferns may be in excess of 50 years. In the
primary forest, one vigorous fern with a trunk 4.6 meters long was observed;
assuming a growth of | meter/I2 years, the plant was about 55 years old. Such
longevity apparently is not common. Of 92 healthy adult tree ferns measured in both
the primary and secondary forests, only one other trunk over 4 meters long was
noted and the third largest was only 3.3 m long. The median length of the 92 tree
trunks was |.3 meters.
One of the most common natural catastrophes in the forest is the falling of large
epiphyte-covered trees or their branches. Winds in the forest can reach 80 kph
(Reyna, 1979). In February 1979 a large oak branch fell at the edge of the thicket
and completely crushed and broke one of the tagged tree ferns: although it did no
harm to the trunk of another fern, all the fronds except one immature frond were
broken off.
The ferns seem to be able to withstand extraordinary damage without being
killed. Few of the tree ferns in the thicket have erect trunks, most are inclined away
from the vertical, and some nearly lie on the ground. Probably this is caused by
falling branches or trees knocking the ferns over. The ferns recover from being
knocked over by turning the crown and growing upright (Fig. /). One fern was seen
which had had its direction of growth radically changed at least twice, and possibly
three times, with no apparent ill effects.
Water is retained in the persistent petiole bases of the ferns during the entire wet
season and part of the dry season. It was originally thought that this might be
advantageous to the ferns during the dry season, perhaps providing additional water
trapped from rain or condensation. Observations indicate otherwise; the tree ferns
which could not store water in their punctured petiole bases did not differ
significantly from the control in number of fronds produced annually or in the frond
death rate. The results might have been different in the drier secondary forest, where
the ferns are less vigorous.
Possible reasons for tree fern mortality were investigated. Falling branches kill
some trees; nevertheless, being knocked over or the loss of fronds may not kill the
fern. The tree that lost all except one immature frond subsequently developed
normal fronds. No sign of insect damage to fronds was ever observed. Two fronds
on one tree fern were stricken by a blight; normal fiddleheads developed into twisted,
stunted fronds with only a few functional pinnules. This did not kill the fern during
the period of the study, and no other fronds seen were affected by a similar blight.
Relatively few dead tree ferns were seen, as the soft wood of the trunk probably
decays rapidly in the wet conditions of the forest. Nevertheless, several dead
individuals were observed in which the trunk diameter was sharply constricted at the
crown. Living trees with this syndrome were easily recognized by their extremely
small fronds, which usually were only 40-50 cm long. The crowns of several of
these living trees were examined by splitting them open; there was no evidence of
rot or decay in the crown or of damage to the base or roots of the trees. It may be
that the constricted trunk and the very small fronds are symptoms of senescence in
R. L. SEILER: LEAF TURNOVER OF ALSOPHILA SALVINII 81
the trees. Further research on these ferns could provide evidence supporting the
hypothesis that death from old age is fairly common in the population. Data
supporting the hypothesis would include a relatively high frequency of ferns with
this syndrome occuring in the population and evidence that these trees are old
compared with the population median.
ACKNOWLEDGMENTS
Agradezco sinceramente a la Unidad de Parques Nacionales y Vida Silvestre del
Ministerio de Agricultura y Ganaderia y a la Fundacion Freund por el apoyo
recibido para la realizaci6n de este estudio. Expreso mi aprecio a la Lic. Maria
Luisa Reyna V. por su valiosa ayuda con la descripcion del bosque y ademas a la
Lic. Kathy DeRiemer, Lic. Dennis Witsberger y Senor Gabriel Calderon Hernandez
quienes hicieron las observaciones en la parte final del estudio por encontrarme
afuera de El Salvador. Asimismo a los Sefores Amadeo, Julio y Ricardo Martinez.
encargados de los jardines de Los Planes de Montecristo; al Senor Juan Escobar,
motorista de Parques Nacionales; y al Senior Pedro Hernandez y los demas vigilantes
por su plena cooperacion en el trabajo del campo.
LITERATURE CITED
HOLDRIDGE, L. R. (Consultor). 1975. Zonas de Vida Ecoldgica de El Salvador: informe preparado
r la Organizacién de la Naciones Unidas para la agricultura y la alimentacion.
PNUD/FAO/ELS/73/004, Documento de trabajo No. 6. San Salvador, EI Salvador.
REYNA V., M. L. 1979. Vegetacién Arborea del Bosque Nebuloso de Montecristo. Thesis (Licenciatura
en Biologia), Facultad de Ciencias y Humanidades, Universidad Nacional de El Salvador. San
Salvador, El Salvador. 177 pp.
STOLZE. R. 1976. Ferns and Fern Allies of Guatemala. Part 1. Ophioglossaceae through Cyatheaceae.
Fieldiana Bot. 39:1—130
82 AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 3 (1981)
Nomenclatural Notes on Micronesian Ferns
F. R. FOSBERG and M.-H. SACHET*
Three problems concerning the names of ferns found in Micronesia have arisen
during the preparation of a geographical checklist of Micronesian pteridophytes. Our
solutions to these problems are as follows. In addition, a new synonymy is given for
the western Pacific species Thelypteris immersa.
List gris guamensis CHont: ) Fosb. & Sachet, comb.
O77 ay PE23 mi-S o hte at. on Agat-Umatac —
m, “Marin ana Is Sands by ssnull stream. Grether 4354 (BISH not seen; isotype US). PARATYP
Ercthce 4385 (US
Since we ie the genus Thelypteris Schmidel in a broad sense, including such
segregates as Christella Léveillé, it is necessary to provide the appropriate combina-
tion for C. guamensis in Thelypteris. Wagner and Grether (Occ. Pap. Bishop Mus.
19:56. 1948) list as Cyclosorus dentatus (Forsk.) Ching the two collections cited by
Holttum (Grether 4384, 4385). We could not match these exactly in the consider-
able series of specimens from Guam labelled 7. dentata or T. parasitica. The
thickened, reddish, glandular hairs scattered over the underside of the lamina, the
pubescence of two ranges of length, the broad fronds with the lowest pinnae only
slightly or not reduced, and the short-creeping rhizome would seem to place T.
guamensis between T. dentata and T. parasitica. We follow Holttum in recognizing
T. guamensis as a separate species, but with some misgivings. The Guam plants of
this affinity do not sort well into three, and even less well into two, populations.
BE 9
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Thelypteris immersa (Blume) Ching, Bull. Fan Mem. Inst. Biol. 6:306. 1936.
Aspidium immersum gon Enum. Pl. Javae 156. 1828. TYPE: Gaenaeng Parang, Java. Blume (L
not seen, Morton photo 116
Drvopteris immersa oe ca Kuntze, Rev. Gen. PI. 2:313. 1891; Hosokawa. Trans. Nat. Hist. Soc.
Formosa 28:147. |
eatin en see Hosokawa, Trans. Nat. Hist. Soc. Formosa 32:285. 1942. TYPE: Peliliu
Island, Palau, Hosokawa 922/ (TAI not seen),
Amphineuron immersum (Blume) Holtt. in Nayar & Kaur, Comp. Bedd. Handb. 203. 1974.
Thelypteris peliliuensis Fosb. Smiths. Contr. Bot. 45:4. 1980, nom. nov. for G. palauensis
Hosokawa, non T. palauensis (Hosokawa) Reed.
Professor R. E. Holttum (in /itt. 1 Mar 1981) informed us that he studied the
type of G. palauensis and found it to be his Amphineuron immersum, a widely
distributed East Asian and Western Pacific species. Hosokawa himself had reported
this earlier as Dryopteris immersa. Since we regard Amphineuron as a part o
Thelypteris, T. peliliuensis must go into synonymy under 7. immersa.
Trichomanes falsinervulosum (Nishida) Fosb., Smiths. Contr. Bot. 45:4. 1980.
Mic Secs falsinervulosum Nishida, J. Jap. Bot. 32:156. 1957. TYPE: Aimiriik. Babeldaob. Palau.
8 Sept . Tsuyama (TI not seen).
This. . was made after checking recent indices and publications on
co of Botany, National Museum of Natural History, Smithsonian Institution, Washington,
FOSBERG & SACHET: NOMENCLATURAL NOTES ON MICRONESIAN FERNS 83
Malesian and Micronesian ferns. After its publication, Dr. Lellinger called our
attention to C. V. Morton’s combination 7. “falsivenulosum” (Nishida) Morton
(Contr. U. S. Natl. Herb. 38:192. 1974), based on Microgonium “falsivenulosum™
Nishida (J. Jap. Bot. 32:156. 1957). We find no such name published by Nishida:
because the references are identical, it seems likely that Morton misread the epithet
or unintentionally changed the epithet to one that meant exactly the same thing.
If this were an obvious typographical error or a simple orthographic error that
involved only a connecting vowel or similar change, it could be regarded under
ICBN Art. 73 as a correctible error. However, when the change produces a different
Latin root which could properly be used in this position in an epithet, it seems most
advisable to treat the two as different epithets and to index the resulting combina-
tions as different names. This will save future workers from rechecking to account
for the difference. The present wording of Art. 73 does not make it clear where the
line should be drawn between correctible and non-correctible errors. Therefore.
one’s best judgment must be followed. In this case we choose to use the 1980
combination.
Trichomanes motleyi (v. d. Bosch) v. d. Bosch, Nederl. Kruid. Arch. 5(2):145.
1861.
Microgonium motleyi v. d. Bosch, Hym. Jay. 5. t. 1. f. /. 2. 1861. SYNTYPE: “Hab. Insula Borneo
(pr. Laboan), Motley No. 203 (comm. ill. W. J. Hooker)” (L not seen).
SYNTYPE CITED WITH T. MOTLEYI: “Hab. Ins. Borneo (pr. Laboan) Motley (H. Hook.)” (L not
seen).
In verifying the original publication of this species, a Curious circumstance was
noticed. Van den Bosch published M. motleyi and T. motleyi independently in
different publications in the same year, each with a slightly different description and
statement of type material, and each with no reference to the other.
No firm evidence is available to us as to which publication is earlier, other than
that the “Hymenophyllaceae Javanicae” has an introduction dated “Mart. 1860” and
a prefatory note in the other publication is dated “Aug. 1860.”
Presumably Motley’s two sheets are different sendings of the same material,
although they could be different collections from the same place. Since van den
Bosch published the two species only a few months apart, it is likely that he had
both sheets at hand while both publications were being prepared. Therefore, both
collections should be considered as syntypes. In response to an inquiry directed to
the Leiden Herbarium concerning the Motley specimens, G. J. de Joncheere
informed us (in litt. 17 Oct 1980) that one sheet is labelled in van den Bosch’s hand
“Trichomanes motleyi v. d. B. from Laboan (Borneo) no. 203 Mr. Motley ad
truncos arborum.” This sheet bears two small specimens which are, according to de
Joncheere, apparently the plants from which the excellent illustrations in the
“Hymenophyllaceae Javanicae” were made. Attached to the other sheet 1s a descrip-
tion written by van den Bosch which is similar to but not identical with the two
published descriptions, which are themselves similar but not identical. Interestingly,
none of the annotations on either of these sheets mentions Microgonium; all refer to
Trichomanes motleyi. We here designate the sheet with the description as the
lectotype of M. motleyi.
84 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
As to the exact status of the two names, we can do no better than to quote from de
Joncheere’s letter:
From the above you will see that indeed the names Microgonium motleyi and Trichomanes
motleyi are independently published, strictly speaking. However, as the dual names originated
at about the same time from the same man from the same type and were apparently just the
result of a wavering mind, one cannot regard these names as truthfully unconnected: on the
contrary these publications are very much connected and should in my opinion be treated as
the publication of a new species and a new combination. depending on what exact date of
publication of the V. D. B. articles in question can eventually be ascertained.
We are prepared to accept this well considered opinion. Since the scanty evidence
from the introductory statements indicates that M. motleyi may have appeared
earlier, we treat the 7richomanes as a transfer. Copeland (Phil. J. Sci. 51:201—202.
1933; 67:62. 1938), in his major works on Hymenophyllaceae, used both names
without raising any question of the propriety of their publication.
his species was reported as Microgonium motleyi by Ito (Bot. Mag. Tokyo
67:219. 1954) from Aimiriik, Babeldaob, Palau, Caroline Islands, Okabe /6 (TI not
seen).
REVIEW
“SINOPSIS DE LAS ESPECIES DE LYCOPODIUM L. (LYCOPODIACEAE
PTERIDOPHYTA) DE LA SECCION CRASSISTACHYS HERTER,” by Cristina
H. Rolleri. Revista del Museo de La Plata, n.s. 13:61-114. 1981.—1In the first half
of this century, Herter and Nessel published several papers and one book (“Die
Barlappgewachse”) which thoroughly confounded the taxonomy and nomenclature
of Lycopodium. So complete was the chaos introduced by these authors, especially
in the tropical members of the genus, that subsequent botanists often have avoided
studying them. The synopsis now published by Dr. Rolleri goes a long way toward
resolving the problems that were introduced in sect. Crassistachys, a largely
terrestrial group of mostly Andean species. Dr. Rolleri has studied the anatomy and
morphology of the plants in great detail, which is evident from her key to the 56
species of the section. A synonymy, statement of habitat and range, and brief
description are given for each species. A rather long list of excluded names, many of
which are of uncertain application because the author was not able to obtain type
material, indicates that there is yet more work which must be done before the
taxonomy and nomenclature of this section of Lycopodium can be completely
understood, but the progress in the present synopsis is very great and will be most
helpful to all who must deal with these confusing plants.—D. B. L.
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 3 (1981) 85
x Asplenosorus shawneensis, a New Natural Fern Hybrid
Between Asplenium trichomanes
and Camptosorus rhizophyllus
ROBBIN C. MORAN*
The Appalachian spleenworts constitute one of the most interesting and diverse
groups of ferns in North America. They are popular with amateur fern growers due
to their beauty and ease of culture. To professional botanists, the study of these
small ferns has contributed to advances in cytology, chemical systematics, reticulate
evolution, hybridization, and ecology. Knowledge of the Appalachian Spleenwort
complex is still far from complete: indeed, new hybrid ferns and cytogenetic races
have only recently been described. The purpose of this paper is to describe a new,
naturally occurring spleenwort hybrid.
In October, 1979, while botanizing in the Shawnee Hills of southern Illinois. I
discovered one plant of a hybrid between Asplenium trichomanes and Camptosorus
rhizophyllus. The hybrid was found on a north-facing crevice in a sandstone canyon
where a large number of the parent species were present. The elongate-triangular
shape of the blade at once suggested C. rhizophyllus; however, the base of the blade
was cut into opposite, pinnate lobes that looked very much like enlarged basal pinnae
of A. trichomanes. The stipe resembled that of A. trichomanes due to its narrow
diameter and dark brown color that extended a short distance up the rachis (Figs. /
and 3). Furthermore, the frond venation was both dichotomously free branched. as
in A. trichomanes, and anastomosing, as in C. rhizophyllus (Fig. 3). The suspected
hybrid was removed and brought to Southern I[Ilinois University. where it was grown
for further stud
To gain further evidence of the plant’s hybrid nature, alpha and beta esterase
isoenzymes were studied using starch gel electrophoresis in the suspected hybrid and
the putative parents. It was hypothesized that the suspected hybrid would show
isoenzyme banding patterns present in both parents and perhaps intermediate bands
as well.
The electrophoretic system followed was that of Steiner and Johnson (1973). Their
instructions for specific esterase stains and gel and electrode buffer were used
exclusively. Only mature blade tissue was ground. This eliminated variation in the
results which could be caused by using fronds of different ages. Plants of A.
trichomanes and C. rhizophyllus used for the analysis were collected at the same site
where the hybrid was found. Electrophoresis was run until a standard marker dye had
migrated about 3.5 cm. After staining the starch gel. it was possible to differentiate
the beta esterase enzymes that stained pink—purple from the alpha esterases that
stained black (Brewer, 1970, p. 88). The resulting zymograms were wrapped with
clear plastic wrap, stored in a refrigerator at 1° C, and later photographed with
Ektachrome film. Several replicates were made to confirm the validity of the results.
*Department of Botany, Southern Illinois University. Carbondale. IL 62901. Present ron Illinois
Natural History Survey, Natural Resources Bldg., 60 )7 E. Peabody Dr.. Champaign. IL 6182¢
AMERICAN FERN JOURNAL: VOLUME 71 (1981)
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* i
.
4
ight)
e. unreduced spore mother cells (ri
<
oO
4
&
FIG. |. Mature fronds cultivated from the type of x Asplenosorus shawneensis (Moran 1269. MICH).
S. Wagner.
nosorus Shawneensis showing 16 lar
and 64 aborted spores (left). Photo by F.
FIG. 2. Sporangia of x Asple
R. C. MORAN: x ASPLENOSORUS SHAWNEENSIS, A NATURAL HYBRID 87
The hybrid combines the esterase isoenzyme banding patterns of A. trichomanes
and C. rhizophyllus (Fig. 4). Isoenzyme band numbers seven and nine. present in C.
rhizophyllus, were the only parental bands undetected in the hybrid: they may have
been present in the hybrid but simply not seen due to their extreme faintness.
Isoenzyme band number eight, present in A. trichomanes and the hybrid. is distinct
from all other bands because it is a purple-staining beta esterase band. All other
bands are black-staining alpha esterases. No intermediate bands were observed. The
hybrid esterase isoenzyme banding patterns provide strong evidence that
trichomanes and C. rhizophyllus are indeed the parents.
F rhizophyllus
trichomanes
shawneensis
~
FIG. 3. Frond outlines and venation patterns drawn from cleared fronds of * Asplenosorus shawneensis
and its parents.
Esterase zymograms of two other common spleenworts in southern Illinois,
x Asplenosorus pinnatifidus and Asplenium platyneuron, were made in order to
ascertain if they were involved in the hybrid’s formation (Fig. 4). Both species
produced banding patterns suggesting that they were not involved.
x Asplenosorus shawneensis R. C. Moran, hybr. nov. Figs. 1 and 3
(L.) Link. Rhizoma breve, reptans; squamae nigricantes, clathratae. Frondes
u
humilo-patentes, caespitosae, sempervirentes, usque ad IS cm longae. Stipites
filiformes, atrobrunnei, nitidi, usque ad 4 cm longi. Lamina leviter coriacea,
usque ad I cm latae, marginibus non profunde crenatis. Rachis atrobrunnea usque
ad 7 cm ad basim, viridis ad apicem. Sori usque ad 4 mm longi.
88 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
TYPE: East of Devil’s Kitchen Lake, Williamson County, Illinois, T10S, RIE,
sec. 15. in crevice of a shaded, north-facing sandstone canyon, with Asplenium
trichomanes and Camptosorus rhizophyllus growing abundantly nearby, 21 O
979, Robbin C. Moran 1269 (MICH
Observations of the hybrid’s chromosomes showed 72 univalents at early meiotic
metaphase. Thus, x Asplenosorus shawneensis is an allodiploid with the genomic
constitution R'T2, with no pairing between members of the two different genomes
(R and T are the respective genomes and superscripts the pairing controls). The
spores of XA. shawneensis vary between 64 aborted spores and 16 unreduced
spores per sporangium (Fig. 2).
=
4r
SLPS
oT Se
rs sitetetats
oO
2 eaoet
Ti
1 —
merece
: —
trichomanes hybrid rhizophyllum = pinnatifidum = platyneuron
FIG. 4. Esterase isoenzyme banding patterns obtained with starch gel electrophoresis in five spleen-
worts, Relative darkness of the bands is indicated by cross-hatching. Band 8 is a beta esterase: all other
bands are alpha esterases.
The following key has been provided to distinguish x A. shawneensis from the
similar
Index to Volume 71 an? eg
PAN 3 122
Erratum
The American Fern Society
Council for 1981
ROBERT M. LLOYD, Dept. of Botany, Ohio University, Athens, OH 45701. President
DEAN P. WHITTIER, Dept. of Biology, Vanderbilt University, Nashville, TN 37235. Vice President
MICHAEL I. COUSENS, Faculty of Biology, University of West Florida, Pensacola, FL 32504.
Secretary
JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, TN 37916.
Treasurer
JUDITH E. SKOG, Dept. of Biology, George Mason University, Fairfax, VA 22030.
Records Treasurer
DAVID B. LELLINGER, Smithsonian Institution, Washington, DC 20560. Journal Editor
ALAN R. SMITH, Dept. of Botany, University of California, Berkeley, CA 94720. Memoir Editor
JOHN T. MICKEL, New York Botanical Garden, Bronx, NY 10458. Newsletter Editor
American Fern Journal
EDITOR
DAVID B. LELLINGER Smithsonian Institution, Washington, DC 20560.
ASSOCIATE EDITORS
DAVID W. BIERHORST Rt. 3, Box 188, Picayune, MS 39466.
GERALD J. GASTONY Dept. of Biology, Indiana University, Bloomington, IN 47401.
JOHN T. MICKEL New York Botanical Garden, Bronx, NY 10458.
The “American Fern Journal” (ISSN 0002-8444) is an illustrated quarterly devoted to the general
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Changes of address, dues, and applications for membership should be sent to Dr. J. E. Skog, Dept. of
es George Mason University, Fairfax, VA 22030.
rs for back issues should be midressed to the Treasure
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Dr. John T. Mickel, New York Botanical Garden, Bronx, NY 10458, is Librarian. Members
may borrow books at any time, the borrower paying all shipping costs.
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Dr. John T. Mickel, New York Botanical Garden, Bronx, NY 10458, is editor of the newsletter
“Fiddlehead Forum.” The editor welcomes contributions from members and non-members, including
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Spore Exchange
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AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 4 (1981) 97
Bog Clubmosses (Lycopodiella) in Kentucky
R. CRANFILL*
Distributed largely on the Atlantic and Gulf Coastal Plains, the southern members
of Lycopodiella Holub (= Lycopodium subg. Lepidotis (Pal. Beauv.) Baker) are
rare and local in the interior of the continent. Inland occurrences are scattered as far
north as the middle Mississippi Valley, and often are markedly disjunct from
populations on the coastal plain. The purposes of this paper are to record recent
observations on this genus in Kentucky, to give some explanation of its recent
appearance and spread, to describe a new hybrid, and to make a series of new
combinations.
t is now apparent that the genus Lycopodium in the broad sense may be divided
into several smaller, more natural genera (Beitel & Bruce, 1980). To facilitate the
use of the segregate genus Lycopodiella, the following new hybrid and combinations
are necessary.
Lycopodiella x brucei Cranfill, hybr. nov.
Planta hybrida inter L. appressam et L. prostratam intermedia, a L. appresso
microphyllis ciliatis et sporophyllis patentibus, a L. prostrato rhizomatibus crassiore
et strobilis angustiore, minus quam 10 mm latis differt; sporae non abortivae.
TYPE: Borrow pit ca. 200 m ENE of road KY-280, 0.6 mi from its junction with
road KY-121, Calloway County, Kentucky, 15 May 1975, J.G. Bruce 76006 (MICH).
Named in honor of James G. Bruce, III.
Lycopodiella alopecuroides (L.) Cranfill, comb. nov.
Lycopodium alopecuroides L. Sp. P|. 2:1102. 1753.
Lycopodiella appressa (Chapman) Cranfill, comb. & stat. nov.
Lycopodium inundatum var. appressum Chapman, Bot. Gaz. 3:21. 1878.
Lycopodiella < copelandii (Eiger) Cranfill, comb. nov.
Lycopodium X copelandii Eiger, Biol. Rev. City Coll. New York 18:21. 1956.
Lycopodium inundatum var. elongatum Chapman, FI. So. States, ed. 2:671. 1883.
Lycopodiella prostrata (Harper) Cranfill, comb. nov.
Lycopodium prostratum Harper, Bull. Torrey Bot. Club 33:229. 1906. os
_ Although much has been written about the Lycopodiella population in Kentucky
(Johnson & McCoy, 1975; Bruce 1975; Cranfill, 1980), confusion still exists over
the number and kind of taxa present. Johnson and McCoy reported L. appressa and
L. prostrata from a large gravel pit in eastern Calloway County. Bruce’s (1975)
extensive survey of the North American members of the genus includes comments on
the Kentucky material, which he concluded is comprised of L. appressa, ae
copelandii (L. alopecuroides appressa), and L. X brucei (as L. appressa X
*Division of Biological Sciences, University of Michigan, Ann Arbor, MI 48109. Present address:
Department of Botany, University of California, Berkeley, CA 94720.
Volume 71, number 3, of the JOURNAL was issued September 30, 1981.
98 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
prostrata). For want of authentic L. x brucei, | admitted only the first two species
to the flora (Cranfill, 1980). In an attempt to resolve these differences, I undertook
fieldwork in Calloway County during the summers of 1979 and 1980. I recorded the
topographic and ecological features of each site where Lycopodiella was found and
made representative collections of each population for further analysis. Results of
these studies are reported below.
Reconnaissance on 10 Aug 1979 in the vicinity of the gravel pit where Johnson
and McCoy had collected (Site 1) revealed three additional populations (see Fig. /)
of L. appressa (Site 2, Cranfill 4730; Site 3, Cranfill 4731; Site 4, Cranfill 4738a).
All sites are scattered along the bases of Cretaceous ridges composed of sand and
partially consolidated gravel and are characterized by low-lying flats of sands with
virtually no contamination from silt or organic matter. These plants seem especially
sensitive to siltation, for no Bog Clubmoss sites have been found where siltation had
occurred. Ground water in the area is soft and acidic and lies at or just below the
surface of the flats for much of the growing season. The original pit is the largest
and best developed of the four sites, has been abandoned for some time, and
contains the most extensive populations of the clubmosses. Lycopodiella appressa is
abundant, occupying nearly all the moist, open areas, whereas L. X copelandii is
confined to the wettest spots, often around pools. The moss Aulacomnium palustre
is an associate of both species, and Cladonia and Leucobryum albidum are scattered
about on dryer hummocks. Vascular plant associates include species with decidedly
southern affinities, e.g., Boltonia asteroides, Gymnopogon ambiguus, Scirpus
koilolepis, Spiranthes odorata, Vaccinium atrococcum, Woodwardia areolata, and
FIG. 1. Portion of south-central Calloway County, Kentucky, showing extant sites of Lycopodiella.
R. CRANFILL: BOG CLUBMOSSES IN KENTUCKY 99
Xyris torta. Pinus taeda is naturalized along the banks above the pit. Sites 2-4 have
been disturbed recently and contain only small populations of L. appressa. Few
species other than the clubmosses have become established along the roadside
ditches and sandy flats, although Boltonia was seen at Site 2.
Winter and early spring visits reveal almost complete dieback of the clubmosses
during the winter. Bruce (1975) reports the formation of “tubers” in L. alopecuroides
and L. prostrata which serve as food storage organs for the following spring. I
observed a similar situation in L. appressa at Site 1. Growth is initiated from the tips
of these organs in early April.
From these observations, it is evident that any management programs designed to
protect these stations must (1) maintain the open nature of the site by controlling the
invasion of woody vegetation, (2) attempt to maintain a high water table, which is
important not only for reproduction but also perhaps in moderating the harsh
winters, and (3) minimize siltation, which appears to have adverse effects on these
ants.
Although I have been unable to confirm the presence of L. X brucei at any of the
sites out of ca. 100 individuals that I collected from all populations, recently |
received herbarium material of this species (Bruce 76006) from Dr. J. G. Bruce.
Since L. prostrata is the most southern of the species in the complex, it is plausible
that it and its hybrids are less winter hardy than other taxa in the pits. Therefore, a
combination of harsh winters in the late 1970’s and competition with other
clubmosses may be responsible for the hybrid’s disappearance.
On the other hand, hybrids which occur without one or both parents often have
been explained by “long distance hybridization,” which may apply in this case. In
Lycopodiella, hybrids between parents with the same chromosome number exhibit
normal pairing of chromosomes at meiosis, and produce what appear to be
functional spores (Bruce, 1975). If the spores be viable, the occurrence of isolated
hybrids may represent the introduction of single spores from hybrid plants. Such a
mechanism for hybrid reproduction and dispersal would be novel among the
pteridophytes, which generally produce infertile hybrids that reproduce largely
vegetatively.
Although the bog clubmosses may have been present in the area for some time,
the evidence points to a relatively recent introduction. In Calloway County, these
plants occur in situations which have been created by man’s activities in the recent
past. This also appears to be the situation with populations in west Tennessee.
Natural habitat suitable to Lycopodiella is very limited in the northern portion of the
Mississippi Embayment and is occupied quickly by the adjacent floodplain forests. It
seems likely that these clubmosses have migrated from their metropolis to the south
by hopping from one gravel pit to another along the Cretaceous hills in western
Kentucky and west Tennessee. If migration is occurring still, close inspection of
suitable sites to the north may reveal the presence of these taxa on Cretaceous
outcrops in southern Illinois.
I am indebted to M. E. Medley, Laurina Lyle, and D. M. Johnson for assistance
in the field, to D. M. Johnson and J. M. Beitel for reading and commenting on the
manuscript, and especially to J. G. Bruce, III who helped me in various ways.
100 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
LITERATURE CITED
BEITEL. J. and J. G. BRUCE, III. 1980. Generic concepts in the Lycopodiaceae. Abstract. Bot. Soc.
r. Misc. Ser. Publ. TEL.
BRUCE, J. G., Ill. 1975. Systematics and morphology of subgenus Lepidotis of the genus Lycopodium.
Unpublished Ph.D. dissertation, University of Michigan, Ann Arbor, MI. 15
CRANFILL, R. 1980. The Ferns and Fern Allies of Kentucky. Ky. Nat. Pres. Comm. Monogr. Tech.
Ser. 1:1-284.
JOHNSON. R. G. and T. N. McCOY. 1975. Some Lycopodiums new to western Kentucky. Amer. Fern J.
65:29
REVIEW
FERNS AND FERN ALLIES in the “FLORA OF BAJA CALIFORNIA” by
Ira L. Wiggins, viii + 1025 pp. Stanford University Press, 1980. ISBN
0-8047-1016-3. $65.00—This is the first publication to treat all the known ferns and
fern allies of Baja California, Mexico. Sixty-five species and an additional five
varieties are included in the treatment, which appears on pp. 51-71 of the book.
Descriptions are provided for families and genera. The species and varieties are
delimited through rather detailed keys which include comments on the species’
habitat, distribution in Baja California, and overall oe Line drawings
illustrate one member of each genus treated. These are the work of the author in
many instances; others are taken from L. R. Abrams’ “lllustrated Flora of the
Pacific Coast States” (1923). The illustrations are nae of very high quality and
are detailed enough to give an accurate representation of the species illustrated. Two
exceptions are the illustrations of Woodsia plummerae and Equisetum laevigatum,
which are not very useful below the genus level.
The taxonomy used is conservative and traditional in treatment of families (e.g.,
Polypodiaceae s. lat. is used) and species and varieties. Three examples of outdated
nomenclature to be corrected are: Pteridium entry corrected to Pteridium aquilinum
(L.) Kuhn var. pubescens Underw.; Anemia anthriscifolia corrected to A. tomentosa
(Savigny) Swartz var. mexicana (Presl) Mickel; and Pellaea longimucronata cor-
rected to P. truncata Goodd. | have noted several cases of misspelling or incorrect
citation of authors. The overall distributions given for the species vary considerably
in completeness and accuracy. Several species should be added to the list of ferns
known in Baja California (based upon recent literature reports): Bommeria pedata
(Swartz) Fourn., Pellaea skinneri — and Pellaea seemannii Hook. (both are
Cheilanthes, pers. obs. and R. M. Tryon, pers. comm., but there are not available
names in Cheilanthes), Cheilanthes den Hook., and C. wootonii Maxon. In
addition, I have seen specimens of C. eatonii Baker in Hook. & Baker from Baja
California.
This treatment of the ferns and fern allies of the exciting botanical region of Baja
California is most welcome, as is the treatment of the entire vascular flora. Dr.
Wiggins is to be commended for producing such a monumental work.—Timothy
Reeves, Biological Science Center, Boston University, Boston, MA 0221
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 4 (1981) 101
Chain Ferns of Florida
TERRY W. LUCANSKY*
Woodwardia is a genus of rather large, terrestrial ferns with mostly ascending to
erect rhizomes. The species native in the United States, however, typically have
creeping rhizomes. The genus includes 11 or 12 species, is worldwide in its
distribution, and. is one of a comparatively few genera characterized by boreal
distribution (Copeland, 1947). Three species of Woodwardia, commonly known as
chain ferns, occur in Florida. Woodwardia virginica and W. areolata are native
species which occur primarily on the Atlantic Coastal Plain and extend from Florida
to Texas and northward to Nova Scotia. Scattered inland localities also exist for both
species. Woodwardia radicans is native to the Old World, but is cultivated and has
reportedly escaped in peninsular Florida (Small, 1938). All three species possess
distinctive leaf venation, with elongate areoles arranged in chain-like fashion along
the costae and/or costules. In W. areolata, an extensive network of areoles extends
to the margins of the leaf, whereas in W. virginica and W. radicans the veins are
simple and forked beyond the areoles and are free at the margins. Elongate sori also
are arranged in chain-like fashion along the costae and/or costules in each species.
Woodwardia virginica, the Virginia or Giant Chain Fern, and Woodwardia
radicans, the European Chain Fern, were originally placed in the genus Blechnum
(Linnaeus, 1771), although Smith (1793) later transferred them to his genus
Woodwardia, with W. radicans the type species. Subsequently, Pres! (1851)
founded the genus Anchistea for W. virginica, although only the glandular indusia
and the presence of a single row of areoles distinguish this species from the other
species of Woodwardia (Morton & Neidorf, 1956). Today both species are typically
included in the genus Woodwardia (Copeland, 1947; Wherry, 1964; Fernald, 1970;
Lakela & Long, 1976). Some workers, however, still recognize the genus Anchistea
(Radford et al., 1964; Small, 1938; McVaugh & Pyron, 1951).
Woodwardia areolata, the Net-vein or Dwarf Chain Fern, was first named
Acrostichum areolatum (Linnaeus, 1753). Smith (1793) included the species in
Woodwardia (as W. angustifolia), although it was not until much later that Moore
(1857) called the species W. areolata. Today some botanists (Fernald, 1970; Lakela
and Long, 1976) use this name. Pres! (1851), however, established the genus
Lorinseria, and many taxonomists (Copeland, 1947; Small, 1938; McVaugh and
Pyron, 1951) consider L. areolata to be the correct name for the species. Wherry
(1964) felt that the completely areolate venation and marked dimorphism of this
species justified its segregation into a separate genus.
Much morphological data are available for these species (Shaver, 1954; Wherry,
1964: Small, 1938; Fernald, 1970), but comparative anatomical data are almost
totally lacking. In this study, the anatomy of W. virginica and W. radicans is
compared with that of W. areolata, and these data are correlated with the two
taxonomic systems currently in use for these species.
Plant materials of W. virginica and W. areolata were collected in Alachua County,
Florida. Plants of W. radicans were obtained from the Strybling Arboretum in San
*Department of Botany, University of Florida, Gainesville, FL 32611.
102 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
Francisco, California. The plant parts were killed and fixed in formalin-acetic
acid-alcohol (FAA), dehydrated in a tertiary-butyl alcohol series and infiltrated with
Tissuemat (Johansen, 1940). Sections (8 4m) were stained with a safranin-fast green
series and photographed with a 35 mm Zeiss C35 camera. Habit photographs were
taken with a 35 mm single-lens reflex camera.
FIG. 1. Prostrate underground rhizomes of W. areolata (above), x 0.4 and W. virginica (below), 0.5
Note adventitious roots. FIG. 2. Erect stem of W. radicans, X0.9. Note large mass of adventitious
roots.
RESULTS AND DISCUSSION
Both Woodwardia virginica and W. areolata occur in wet pinelands, bogs,
marshes, and alongside streams and roadside ditches, whereas W. radicans is
cultivated in fern gardens and has reportedly escaped to swamps and hammocks in
the state. Woodwardia areolata and W. radicans thrive in partial shade around the
bases of trees, but W. virginica does best in sunny locations and can tolerate a
lower degree of acidity in the soil (Cody, 1963; Wherry, 1964)
Both W. virginica and W. areolata have a branched, creeping rhizome (Fig. /),
whereas W. radicans has an ascending to erect stem (Fig. 2). The stems of W.
virginica (x = 10 mm) and W. radicans (x = 18 mm) are stouter than the rhizome
of W. areolata (x = 5 mm) and more deeply buried in the soil. Golden brown,
elongate scales with either blunt, rounded, or acuminate apices occur on the rhizome
and petiole bases of W. virginica and W. radicans. Pale brown, cordate scales with
acuminate apices are found on the rhizome and petiole bases of W. areolata. Shaver
(1954) reported brown-black, oblong scales with acute apices in W. virginica and
brown, ovate scales on the rhizome of W. areolata. I found small, light-brown,
subcordate to cordate scales with acuminate apices abaxially on or proximal to the
T. W. LUCANSKY: CHAIN FERNS OF FLORIDA 103
rachis and midveins of all three species. Woodwardia virginica has rather large,
subcoriaceous, pinnate leaves, whereas W. radicans possesses large, coriaceous,
pinnate-pinnatifid leaves with scaly buds produced on the rachis at the bases of the
upper pinnae. Woodwardia areolata, however, has marked dimorphism, with the
taller fertile leaves typically found in the summer and fall, distinct from the
pinnatifid sterile leaves.
Stem transections of the three species reveal similar anatomical features, although
differences are noted (Fig. 3-5). A single-layered epidermis composed of elongate,
bulbous, or irregularly shaped, thick-walled cells is partially sloughed off in mature
sporophytes of both W. virginica and W. areolata, whereas the epidermal layer is
typically intact and composed of small, thick-walled cells filled with tanniferous
substances in W. radicans. A hypodermis composed of two zones occurs in the three
species, although the outer zone may be sloughed off in W. virginica and W.
radicans. In W. areolata, the outer zone consists of irregularly shaped, thick-walled
parenchyma cells (Fig. 6), whereas in W. virginica and W. radicans the parenchyma
cells of this zone are thicker-walled and more heavily lignified. The inner zone in all
species is composed of sclerified, thick-walled parenchyma cells that are filled with
tannins or contain starch grains (Fig. 6), although the zone is less extensive in W.
areolata than in the other two species.
In all three species, the bulk of the stem is composed of ground tissue (Figs.
3-5). The cortical region is distinguishable from the pith region primarily by the
position of the meristeles. The cortex is composed primarily of irregularly shaped,
thin-walled parenchyma cells filled with tanniferous substances, starch grains, or
both. The pith region is composed of large, thin-walled parenchyma cells filled
primarily with tannins, although starch grains are occasionally found in these cells.
In W. virginica, thick-walled parenchyma cells occasionally comprise this region. In
both W. virginica and W. radicans, certain cells in the cortical and pith regions
enlarge singly or in groups, and tanniferous substances partially or totally fill the
lumens of these cells (Fig. 7). These cell contents are abundant in W. radicans,
especially around the meristeles, but are lacking in W. areolata. Large intercellular
spaces occur in the cortical and pith regions of W. virginica and W. areolata and
possibly serve as an adaptation to a marsh habitat. In W. radicans, very small
intercellular spaces are occasionally noted in the ground tissues.
The stelar pattern in all three species is a dictyostele with overlapping leaf gaps
and variously-sized meristeles (Figs. 3-5). Each meristele is an amphicribral bundle
delimited by an endodermis with distinct Casparian strips (Fig. 8). A pericycle of
thin-walled parenchyma cells (1- 4 layers) filled with tannins completely encircles
the primary phloem. The latter tissue is composed of sieve cells and phloem
parenchyma, although protophloem and metaphloem are typically indistinguishable
in mature stems. The primary xylem of all species is composed primarily of
scalariform-pitted tracheids, although spiral and annular-thickened protoxylem ele-
ments do occur. Interspersed among the xylary elements are thin-walled parenchyma
cells filled with tannins. Xylem maturation is mesarch.
In all three species, the sterile leaves exhibit similar anatomical features (Figs.
9-11), whereas the fertile leaf of W. areolata usually is reduced, although it may be
104
AMERICAN FERN JOURNAL: VOLUME 71 (1981)
FIG. 3. Stem transection of W. nee x10. Note meristeles. FIG. 4. Stem transection of W.
radicans, X10. Note dictyostele. FIG. 5. Stem transection of W. areolata, x 10. Note meristeles a
dictyostelic pattern. FIG Two-zon hs hypodermis of W. areolata, x2 FAG: Teste seninestils
substances in pith region of W. radicans, x91
. 8. Individual meristele of W. areolata,
Pid. 9. Transection of leaf of W. areolata, x 245.
extensive ae pesge layer. 10. Transection of leaf of W.
ote tannin-filled palisade layer.. FIG. . Transection of leaf of W. radicans, x 295.
Note extensive tannin-filled wee layer. The pomhens are: abe = abaxial epidermis, ade = adaxial
epidermis, c = cortex, e = endodermis, g = gap, h= hypodermis, iz = inner zone, m= meristele, oz = outer
zone, p=pith, pe= ewe ph= coldiens pl =palisade layer, s = stomate, sm=spongy mesophyll,
t=tannins, and x= xylem.
ote amphicribral arrangement of vascular tissue
Note reduced palisade layer an
virginica, X 279
T. W. LUCANSKY: CHAIN FERNS OF FLORIDA 105
partially sterile (Halsted, 1899) or even may resemble a fertile leaf (Waters, 1903).
In each of the three species, the single-layered adaxial epidermis is composed of
variously sized cells, is covered with a thin cuticle, and lacks stomates. The
mesophyll is differentiated into palisade and spongy mesophyll layers, although the
palisade layer is much reduced and less extensive in W. areolata than in the other
two species (Figs. 9-11). The palisade layer is composed of loosely arranged,
irregularly shaped chlorenchyma cells in W. areolata; a more compact and tannin-
filled palisade layer characterizes W. virginica and W. radicans. Although Payne and
Peterson (1973) noted an abaxial hypodermis in the leaves of W. virginica, none was
found in the present study (Fig. 10). The spongy mesophyll in all three species
consists of loosely-arranged chlorenchyma cells with numerous, large intercellular
spaces. This layer is much more extensive and constitutes a greater proportion of the
leaf in W. areolata than in the other two species (Figs. 9-//). In W. radicans, the
spongy mesophyll layer is greatly reduced and contains small intercellular spaces.
Depending upon the species, the spongy mesophyll cells are either larger (W.
areolata) or smaller (W. virginica and W. radicans) than the cells of the palisade
layer. The chlorenchyma cells of the mesophyll in the latter two species are typically
filled with tannins, whereas this ergastic substance is less evident or lacking in the
mesophyll cells of W. areolata (Figs. 9-11). In all species, the abaxial epidermis is
composed of variously shaped cells and possesses anomocytic stomates (Fig. 12).
Typically two or three epidermal cells abut the guard cells, with one cell nearly
surrounding the stomatal apparatus. Chloroplasts are infrequently noted in the
adaxial epidermis of W. areolata, but do not occur in the epidermal layers of the
other two species. The presence of chloroplasts in the epidermal cells of land plants
is correlated with a deep-shade habitat (Sculthorpe, 1967), and their occurrence in W.
areolata may be an adaptation to the shaded habitat of this species. The mid-vein
(costa) in all three species is surrounded by a bundle sheath of thick-walled
parenchyma cells and is delimited by a single-layered endodermis with distinct
Casparian strips (Figs. 13 and 14). The innermost layer of the sheath may have
tanniferous substances in W. virginica and W. radicans, whereas these cell contents
are lacking in W. areolata. A pericycle composed of thin-walled parenchyma cells
encircles the vascular tissues. The primary phloem is composed of sieve cells and
phloem parenchyma. The primary xylem is U- or V-shaped and consists primarily of
scalariform-pitted metaxylem elements. Protoxylem is generally restricted to a
median position in the concavity of the xylary elements.
The fertile leaves of W. areolata show a much reduced structure; no differentiation
of the mesophyll occurs, and only a few small intercellular spaces are noted.
Stomates are lacking in both epidermal layers of a fertile leaf.
Ground tissue comprises the bulk of the petiole base in the three species (Figs.
15-17). This tissue is aerenchymatous and consists of either thick-walled (W.
virginica) or thin-walled (W. areolata) parenchyma cells with numerous intercellular
spaces. In W. radicans, this tissue consists of thin-walled parenchyma cells or is
composed of two zones in the larger petioles (Fig. 17). The outer zone consists of
thick-walled parenchyma cells, and the inner zone is composed of thin-walled
parenchyma cells. Numerous, small intercellular spaces occur in these zones, and
106 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
the cells are filled with tannins. Occasionally the parenchyma cells of the ground
tissues in W. radicans enlarge and possess tanniferous substances that partially or
totally fill the cell lumen. A single-layered epidermis characterizes the petiole and a
hypodermis is found in all three species (Figs. /5—/7). The latter tissue is much
more extensive in W. radicans than in the other two species.
The number of vascular strands at the base of the petiole differs in each species
In W. virginica, typically five small spherical strands and two large U- or V-shaped
strands occur (Fig. /6), although a total of five or six strands has been noted.
12. Anomocytic stomates of W. virginica, x 225. FIG. 13. Transection of midvein (costa) of W.
‘ . Transection of midvein (costa) of W. virginica, x 100.
substances in bundle s heath cells: FIG? 1S,
small, spherical abaxial b ede ie
Note Saas ak
Transection of petiole base of W. areolata, x21. 2 two
FIG. 16. Transection of petiole base of W. virginica, x2 yao a ee
Transection of petiole base of W. eri x11. FIG. 18. Transection of adventitious root of W.
virginica, X97. Note diarch pattern. FIG. 19. Shoot meristem of W. areolata, x2 IG. 20. Shoot
of
meristem of W. virginica, x 295. Note pamia apical cell. The abbreviations are: aoe apical cell,
bs=bundle sheath, gt=ground tissu ner cortex, mx =metaxylem
oc =outer cortex,
ph = phloem, ps = petiole strand, px = hein s=stomate, x = xylem.
T. W. LUCANSKY: CHAIN FERNS OF FLORIDA 107
Waters (1903) reported seven oval bundles in the petioles of W. virginica. In W.
radicans, four or five petiole strands are noted in the petiole base (Fig. /7). Two or
three small, spherical to oval bundles occur abaxially and two large crescent-shaped
strands are found adaxially in the petiole (Fig. 17). In W. areolata, typically two
large petiole strands are found in a median position in the petiole base, although
occasionally four strands occur at a comparable level in the petiole (Fig. /5). Waters
(1903), however, reported only two oval petiole bundles in W. angustifolia (W.
areolata). When four strands are present, they include two large, median, crescent-
shaped strands and two very small, spherical strands located proximal to the
hypodermis on the abaxial surface of the petiole. The small abaxial strands originate
from the division of the larger strands in the petiole base. In both W. virginica and
W. radicans, small abaxial strands also arise from the division of the two large
strands in the base of the petiole. In all species, a bundle sheath composed of large
parenchyma cells with tanniferous substances surrounds each petiole strand. Cellular
composition and arrangement of the stelar tissues is similar to the midrib (rachis) of
a leaf.
Transections of the adventitious roots of all three species show similar anatomical
features (Fig. 18). The epidermis is typically sloughed off in mature roots, and the
outer cortex, which is composed of irregularly shaped, thick-walled parenchyma
cells, forms the outer boundary of the organ. The inner cortex consists of isodiamet-
ric, sclerified, thicker-walled parenchyma cells that may be filled with tanniferous
substances. A single-layered endodermis with Casparian strips delimits the stele. A
pericycle composed of 1-3 layers of thin-walled parenchyma cells surrounds the
vascular tissue. The primary phloem consists of sieve cells and phloem parenchyma,
and the primary xylem is composed primarily of scalariform-pitted metaxylem and
some spiral and transitional (reticulate-scalariform) protoxylem. In all three species,
the primary xylem is diarch with exarch maturation (Fig. 18), although a triarch
pattern is infrequently noted in W. radicans.
In the species studied, the shoot apical meristem consists of a highly-vacuolated
pyramidal apical cell averaging 50 x 80 ym (Figs. 19-20). Recent derivatives of
the apical cell constitute the promeristem and comprise two groups of cells, the
surface and subsurface zones. The promeristem in W. areolata is slightly dome-
shaped (Fig. 19), whereas this tissue has a pronounced dome shape in W. virginica
and W. radicans (Fig. 20). The surface zone in these three species consists of large,
vacuolated, rectilinear cells and a few isodiametric cells on the periphery. McAlpin
and White (1974) found a similar arrangement of the superficial cells in the genera
Dryopteris and Quercifilix. The subsurface zone in W. areolata is distinct and
consists of small, isodiametric cells (Fig. 19), whereas this zone is indistinct in the
other two species. ;
Comparative data indicate a close relationship between W. virginica, W. radicans,
and W. areolata. Although the marked dimorphism and venation of the leaves justify
the segregation of Lorinseria areolata from the genus Woodwardia, chromosome
numbers (Wherry, 1964) and spore morphology (McVaugh, 1935; Hires, 1965)
favor the retention of these species in the same genus. Comparative anatomical data
also support the placement of all three species in the genus Woodwardia.
108 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
Grateful acknowledgment is made to Messrs. Alan Jones and James Neumann for
their technical assistance in this study.
LITERATURE CITED
CODY, W. J. 1963. Woodwardia in Canada. Amer. Fern J. :
COPELAND, E. B. 1947. Genera Filicum. Chronica wats imal A:
pone M. L. 1970. Gray’s Manual of Botany. 8th corrected ed. D. Van Nostrand, Co., New York,
pepe B. D. 1899. Partial sterility of fertile Woodwardia fronds. Plant World 2:55—
HIRES, C. S. 1965. Spores. Ferns. Microscopic Illusions Analyzed, vol. 1. Mistaire kdaesee
Millburn, NJ.
JOHANSEN, D. A. 1940. Plant Microtechnique. McGraw-Hill, New York, AM
LAKELA, O. and R. LONG. 1976. Ferns of Florida. Banyan Books, Miam ;
LINNAEUS, C. 1753. Species Plantarum, vol. 2. Reprint ed. 1959. Ray ee, London.
———. 177]. Mantissa Plantarum Altera. Reprint ed. 1961. Hafner, New York, NY
McALPIN, s and R. WHITE. 1974. Shoot organization in the Filicales: the promeristeti. Amer. J.
Bot. 61:562-579.
McVAUGH, R. 1935. Studies on the spores of some northeastern ferns. Amer. Fern. J. 25:73-85.
d J. PYRON. 1951. Ferns of Georgia. Reprint ed. 1968. Univ. Georgia Press, Athens,
GA.
MOORE, T. 1857-62. Index Lome William Pamplin, London
MORTON, C. V. and C. NEIDORF. 1956. The Virginia Chain- fern. Amer. Fern. J.
PAYNE, W. W. and K. M. PereRseny 1973. Observations of the hypodermises of pitas Asher. Fern.
34-42.
PRESL, K. B. 1851. = forage — Abhandl. K. Boehm, Gesell. Wiss. V, 6:361-624.
RADFORD, m E., H. E. AHLES, and C. R. BELL. 1964. Guide to the Vascular Flora of the
Carolinas. The hee paces Univ. of North Carolina, Chapel Hill, NC.
SCULTHORPE. C. D. 1967. The Biology of Aquatic Vascular Plants. Edward Arnold, London
SHAVER, J. M. 1954. Ferns of Tennessee. Bureau of Publications, Geo. Peabody College for Teacher’:
Nashville.
SMALL, J. K. 1938. Fert of the Southeastern States. Reprint ed. 1964. Hafner, New York, NY.
SMITH, J. E. 1793. —— Botanicum. Mém. Acad. Turin. 5:401—413.
WATERS, C. E. 1903. Ferns. Henry Holt, New York, NY.
WHERRY, E. T. 1964. The powals Fern Guide. Doubleday, Garden City, NY.
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 4 (1981) 109
Spore Germination Patterns in Anogramma, Bommeria,
Gymnopteris, Hemionitis and Pityrogramma
CLARK S. HUCKABY*, R. NAGMANI** and V. RAGHAVAN**
Based on the morphological characters of the sporophyte and gametophyte,
palynological evidence, and spore germination patterns seen in whole mount
preparations, the genera Anogramma, Bommeria, Gymnopteris, Hemionitis, and
Pityrogramma have been assigned to different families by fern taxonomists. Bower
(1928) included these genera in the group designated as gymnogrammoid ferns,
whereas Christensen (1938) placed all except Gymnopteris in the polypodiaceous
group. Copeland (1947), however, regarded these genera as part of the complex
group of pteroid ferns. Holttum (1949) followed Bower in his conception of
gymnogrammoid ferns and combined the gymnogrammoid and vittarioid ferns of
Bower into the family Adiantaceae. Nayar (1970) placed Anogramma in the
Adiantaceae, Bommeria, Hemionitis, and Gymnopteris in the Cheilanthaceae and
Pityrogramma in the Pteridaceae, whereas Crabbe et al. (1975) treated all five
genera as members of the Adiantaceae. According to Haufler and Gastony (1978),
gametophytes of Bommeria and Hemionitis responded differently from those of
Anogramma and Pityrogramma to antheridiogen A, the male sex hormone isolated
from the gametophytes of Pteridium aquilinum, to merit their further subgrouping.
Previous light microscopic studies (Endress, 1974; Raghavan & Huckaby, 1980;
Huckaby & Raghavan, 1981a; Rutter & Raghavan, 1978) on the orientation of the
initial cell divisions during germination of spores of certain ferns using techniques
generally employed in electron microscopy (fixation in an aldehyde fixative and
embedding in glycol methacrylate plastic) have contradicted earlier work on spores
of the same ferns based on whole mount preparations. Comparative studies of spore
germination in Anemia, Lygodium, and Mohria (Schizaeaceae) (Raghavan &
Huckaby, 1980) and in Cyathea and Dicksonia (Huckaby & Raghavan, 1981a) have
also demonstrated the value of the early division sequence during spore germination
as a stable taxonomic criterion in defining affinities of problematic genera. In view
of the disagreement among pteridologists on the relationships of Anogramma,
Bommeria, Gymnopteris, Hemionitis, and Pityrogramma, the present work is aimed
at evaluating the importance of spore germination patterns in these genera as studied
by modern histological techniques and in the scanning electron microscope (SEM)
as a new source of taxonomic evidence. a
Relatively little information is available in the literature on the initial cell division
patterns during germination of spores of Anogramma, Bommeria, Gymnopteris,
*Department of Biology, Syracuse University, Syracuse, NY 13210.
**Department of Botany, The Ohio State University, Columbus, OH 43210.
AMERICAN FERN JOURNAL: VOLUME 71 (1981)
cell,
HUCKABY ET AL.: SPORE GERMINATION PATTERNS 111
Hemionitis and Pityrogramma. Rao (1949) and Nayar (1956, 1962), who followed
the early development of H. arifolia gametophytes, make no reference to the spore
germination. Nayar’s (1964) study of P. calomelanos and P. chrysophylla gameto-
phytes also lacks details of spore germination. In a study of G. vestita gametophytes,
Kaur (1972) states, without giving any details, that spore germination is of the
Vittaria type described earlier by Nayar and Kaur (1968). In this type of germination,
a rhizoid is cut off at the proximal pole of the spore by a wall perpendicular to the
polar axis. The protonemal cell is formed by a subsequent division of the large distal
cell perpendicular to the first division wall. Elongation of the rhizoid parallel to the
polar axis of the spore and that of the protonemal cell along the equatorial plane are
also characteristic of this type of germination. In their review of the gametophytes of
homosporous ferns, Nayar and Kaur (1971) make statements which imply that the
germination pattern of Anogramma, Bommeria, Gymnopteris, Hemionitis, and
Pityrogramma spores follows the Vittaria type,
Based on whole mount observations, Haufler (1979) recently showed that spores
of Bommeria hispida germinated by a division wall perpendicular to the polar axis,
the small proximal cell differentiating into the rhizoid; origin of the protonemal cell
was not traced in this work. However, in a proportion of spores of B. subpaleacea,
the small (proximal?) cell formed from the first division is believed to yield the
protonemal cell, the rhizoid being derived by a division of the same cell as the
protonemal. According to Baroutsis (unpublished work cited by Haufler, 1979 and
pers. comm.), who followed in whole mounts the pattern of cell division during
germination of spores of several species of Anogramma, the first division of the
spore to form the rhizoid is oblique or nearly parallel to the polar axis. According to
this investigator, in A. osteniana, the protonemal cell sometimes appeared to form
before the rhizoid.
MATERIAL AND METHODS
Spores of Anogramma chaerophylla, Bommeria pedata, B. ehrenbergiana, B.
hispida, B. subpaleacea, Gymnopteris rufa, Hemionitis arifolia, H. palmata, H.
pedata, Pityrogramma calomelanos, and P. chrysophylla used in this work were
obtained from various sources! and stored at 5°C until used. To follow cell division
pattern during germination, spores were sown on the surface of 10 ml modified
Knop’s liquid medium (Raghavan, 1965) contained in 5 cm diameter Petri dishes
and allowed to imbibe in the dark for 48 hr. They were then irradiated continuously
with red light or exposed to weak fluorescent light during a photoperiod of ca. 12 hr
as described earlier (Huckaby & Raghavan, 1981a, b). Samples were collected at
intervals of 12-24 hr during an experimental period of 6 days, fixed in 10% acrolein,
'Spores were obtained from the following sources: A. chaerophylla, B. pedata and H. pedata from
Dr. J. T. Mickel, New York Botanical Garden, Bronx, New York; B. ehrenbergiana, B. hispida, B.
subpaleacea, and H. palmata from Dr. C. H. Haufler, University of Kansas, Lawrence, Kansas; H.
arifolia and P. calomelanos from Dr. C. N. Page, Royal Botanic Gardens, Edinburgh, Scotland; G. rufa
and P. chrysophylla from Dr. T. Walker, Royal Botanic Gardens, Kew, England.
112 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
FIGS. 5-8. Germination of Bommeria spores. FIG. 5. B. pedata section owe the rupture of the
the
exine; the spore nucleus is in mitosis. FIG. 6. B. ehrenbergiana section showing the formation of
rhizoid initial (arrow). oe a B. pie section showing the rhizoid initial (r). Arrow points to the
nucleus of the distal cell. . 8. B. pedata section a the protonemal cell (p); (r) is part of the
rhizoid. The nucleus of e pee cell (arrow) is in divisio
HUCKABY ET AL.: SPORE GERMINATION PATTERNS 113
and embedded in glycol methacrylate according to routine procedures previously
used (Raghavan, 1976, 1977; Raghavan & Huckaby, 1980; Huckaby & Raghavan,
1981a, b; Rutter & Raghavan, 1978). Sections cut at 7 pm thickness on a rotary
microtome equipped with a steel knife were stained in 0.05% toluidine blue,
mounted in Euparal, and examined in the light microscope.
For examination in SEM, spores were fixed in 70% ethanol-acetic acid (3:1) at
4°C, dehydrated in ascending series of ethanol and subjected to critical point drying.
They were subsequently mounted on SEM specimen mounts using double-stick
cellophane tape, vaccuum-coated with gold, and examined in a Hitachi S-500
instrument.
SPORE GERMINATION RESULTS
Anogramma chaerophylla.—Spore germination was initiated by the splitting of
the exine at the proximal pole along the lines of the trilete scar after about 36—48 hr
in red light (Fig. 1). Following this, the spore protoplast divided by a wall
perpendicular to the polar axis, resulting in a small proximal cell and a large distal
cell (Fig. 2). The former elongated into the rhizoid, with a corresponding enlarge-
ment of the latter. The distal cell also acquired chloroplasts and appeared outside
through the opening in the exine as a green protrusion, displacing the rhizoid
laterally, so that the wall delimiting the rhizoid from the distal cell appeared parallel
to the polar axis (Fig. 3). The protonemal cell was formed «about 72 hr after
exposure of spores to red light by the division of the distal cell by a wall
perpendicular to the first wall (Fig. 4), although due to the elongation of the distal
cell, the second division wall barely intercepted the first. The division sequences
observed in spores exposed to white light were similar to those seen in red light. In
spores germinated in either light regimen, the protonemal cell grew parallel to the
polar axis of the spore and the rhizoid grew at a right angle to the protonemal cell.
Initial division of the spore protoplast oblique or nearly parallel to the polar axis as
described in this genus by Baroutsis (cited by Haufler, 1979 and pers. comm.) was
not observed in sections made from our sample.
Bommeria ehrenbergiana, B. hispida and B. pedata.—Spores of B.
ehrenbergiana, B. hispida, and B. pedata germinated by cracking of the spore coats
at the proximal pole after exposure to red light for about 48 hr. As in the case of A.
chaerophylla, the first division of the spore protoplast perpendicular to the polar
axis yielded a small proximal rhizoid and a large distal cell (Figs. 5-7). When
spores initially exposed to red light were transferred to white light for 24 hr,
protonemal initiation occurred by the division of the distal cell by a wall perpendicu-
lar to the first. Occasionally after cutting off the protonemal cell, the nucleus of the
distal cell was found to divide again, probably giving rise to a secondary rhizoid
(Fig. 8). Exposure of spores of B. ehrenbergiana and B. pedata to white light or red
light alone resulted in the elongation of the distal cell through the opening in the
exine accompanied by chloroplast accumulation, but its division to form the
protonemal cell was not observed during the experimen
hispida exposed to white light gave rise to the rhizoid and protonemal cell according
114 AMERICAN FERN JOURNAL: VOLUME 71 (1981)
oo TN it
FIGS. 9 and 10. Germination of Bommeria hispida spores. FIG. 9. Section showing the rhizoid (r).
Arrow points to the distal cell enlarging preparatory to division to form the protonemal cell. FIG. 10.
Section showing the distal cell (arrow), partially loose from the wall, after cutting off the protonemal
cell (p); (r) is the rhizoid. FIGS. 11 and 12. Germination of Gymnopteris rufa spores. FIG. 11. Section
showing the first division giving rise to a small proximal rhizoid initial (r) and a large distal cell (dc).
Arrow points to the division wall. FIG. 12. Section showing the elongation of the distal cell (dc) and the
migration of its nucleus (arrow) to the tip.
HUCKABY ET AL.: SPORE GERMINATION PATTERNS 115
to the same division sequence observed in the other two species. In spores of all
three species examined, due to lateral expansion of the distal cell, the orientation of
the wall delimiting the rhizoid was changed so that the wall appeared parallel to the
polar axis of the spore (Figs. 9 and 10).
In spores of Bommeria examined by us, the orientation of the initial division wall
yielding the rhizoid is similar to that described by Haufler (1979) in B. hispida.
However, a sufficient quantity of spores of B. subpaleacea was not available to us to
confirm or refute Haufler’s (1979) observation that in some spores of this species,
the proximal cell arising out of the first division gave rise to the protonemal cell.
ymnopteris rufa.—The first division, which occurred in spores exposed to red
or white light regimens for 72-96 hr, was perpendicular to the polar axis and yielded
a small proximal cell and a large distal cell (Fig. //). This was the only division
observed in the germinating spore during the experimental period. As the proximal
cell differentiated into a rhizoid, the distal cell elongated, became chlorophyllous
and appeared outside as a green cell. Later the nucleus moved from the basal part of
the cell enclosed within the exine to its exposed tip (Fig. /2). A transverse division
occurred at the tip of the distal cell during its further growth in red or white light. A
similar pattern of cell disposition has been described in germinating spores of Preris
vittata (Raghavan, 1977).
Hemionitis arifolia, H. palmata and H. pedata.—Spores of all three species
followed a uniform pattern of germination in which the first division of the spore
protoplast by a wall perpendicular to the polar axis gave rise to a small proximal cell
and a large distal cell. As seen in other genera, the small cell differentiated into the
rhizoid (Figs. 13 and 14). Rhizoid initiation occurred after exposure of fully imbibed
spores to a red or a white light regime for 48 hr and was preceded by the opening of
the exine at the trilete mark. Formation of the rhizoid was followed by enlargement
of the distal cell through the opening in the exine and its division by a wall
perpendicular to the first to give rise to the protonemal cell (Fig. 15). As seen in
SEM preparations (Fig. 16), expansion of the distal cell resulted in the displacement
of the wall separating the rhizoid from the distal cell to a plane parallel to the polar
axis of the spore. In spores of H. palmata grown in red or white light regimes, the
division of the distal cell to form the protonemal cell was not observed during the
experimental period. :
Pityrogramma calomelanos and P. chrysophylla.—Spores of both species are
characterized by the presence of equatorial flanges of sporoderm material which can
be used as a marker for the equatorial plane. Spores responded to white or red light
regimes in about 24 hr by the cracking of the spore walls at the trilete mark and by
the emergence of the rhizoid (Figs. /7 and /8). The latter was traced to the proximal
cell formed by the division of the spore protoplast by a wall perpendicular to the
polar axis (Fig. 19). Following rhizoid formation, the distal cell expanded laterally
and elongated through the opening in the exine parallel to the polar axis, displacing
the rhizoid in a plane parallel to the equatorial axis (Fig. 20). The division of the
distal cell to form the protonemal cell was delayed until the nucleus migrated from
the base of the cell to its tip, and occurred by a wall perpendicular to the first
division wall.
AMERICAN FERN JOURNAL: VOLUME 71 (1981)
116
13 50um
hy
Pr 16
FIGS. 13-16. Germination of Hemionitis spores. FIG. 13. Section of H. pedata showing the first
division to form the rhizoid initial (r) and a distal cell. Small arrow points to the nucleus of the distal
cell; part of the distal cell has become loose from the spore wall (large arrow). FIG. 14. Section of H.
arifolia showing the rhizoid (r) and the undivided distal cell; arrow points to the nucleus of the distal
cell. FIG. 15. Section of H. pedata showing the protonemal cell (p) and rhizoid (r). White arrow points
to the wall delimiting the rhizoid; black arrow points to the wall delimiting the protonemal cell. FIG. 16.
Scanning electron micrograph of H. pedata showing the displacement of the rhizoid wall (arrow): (r) is
the rhizoi
HUCKABY ET AL.: SPORE GERMINATION PATTERNS 117
DISCUSSION
Planes of cell division during germination of Anogramma, Bommeria, Gymno-
pteris, Hemionitis, and Pityrogramma spores described here follow the general
pattern of the Vittaria type of Nayar and Kaur (1968). The spore is divided by two
walls, the first one perpendicular to its polar axis yielding a rhizoid and the second,
perpendicular to the first, giving rise to the protonemal cell. However, due to the
displacement of the rhizoid initial by the expansion of the distal cell, the rhizoid and
protonemal cell have been found to elongate in planes opposite to those described by
Nayar and Kaur (1968). The only difference noted between the initial division
patterns of spores of different genera is in the timing of the second division under
the experimental conditions employed. As seen in spores of Gymnopteris rufa, the
distal cell apparently functions as the protonemal cell due to a delay in the second
division, giving the impression that a single division of the spore protoplast is
sufficient to give rise to a functional gametophyte. However, the fact that under
extended periods in both red and white light regimes, division of the spore
protoplast follows the Vittaria type, reinforces the stability of this character in the
early gametophyte development of ferns. On the basis of our observations, we
believe that the report of Baroutsis (cited by Haufler, 1979 and pers. comm.) on the
formation of the rhizoid in spores of several species of Anogramma by a wall
oriented obliquely or nearly parallel to the polar axis is due to failure to identify the
first division wall as soon as it is formed. As seen in Figs. 5, 6, //, 13, and 19, this
wall appears before or immediately after the exine is ruptured and by the time the
wall is visible in whole mounts, the distal cell would have expanded, displacing the
original wall and giving the false impression of its occurrence parallel to the polar
axis (Figs. 3, 9, and /5).
If the cell division pattern during spore germination can be considered a stable
character for taxonomic purposes, along with other features of the gametophyte and
of the sporophyte, the uniformity in the pattern observed in the five genera
investigated here tends to support their assignment to a single family, Adiantaceae, as
done by Crabbe et al. (1975). Since no distinctive variants of the germination pattern
were seen between groups of genera, a further subgrouping separating Bommeria
and Hemionitis from Anogramma and Pityrogramma as suggested by Haufler and
A comparison between the cell division patterns observed in sectioned spores of
certain genera of Schizaeaceae investigated earlier (Raghavan, 1976; Raghavan &
According to Nayar (1970), Vittaria type germination is evolutionarily more ad-
vanced than the pattern observed in the Schizaeaceae, designated as the Anemia type
ore germination based on whole mount prepara-
patterns of cell division during sp m
here is an increasing body of opinion (Holttum,
tions may be overstated. However, t
118
AMERICAN FERN JOURNAL: VOLUME 71 (1981)
19 50um
FIGS. 17-20. Germination of Pityrogramma calomelanos spores. FIGS. 17 and 18. Scanning electron
micrographs showing the trilete mark and e
flange. j
2g of rhizoid (r). Arrows point to the ee
19. Section showing the first division. Arrow points to the rhizoid initial. FIG. 20.
Scanning electron micrograph showing elongation of the distal cell (de) and displacement of the rhizoid
(r).
HUCKABY ET AL.: SPORE GERMINATION PATTERNS 119
1949; Crabbe et al., 1975) on the possibie origin of adiantoid ferns as a distinct
group from the schizaeaceous stock. Raghavan and Huckaby (1980) have shown that
spores of M. caffrorum follow the same route as the five genera studied here to form
the rhizoid and protonemal cell, exhibiting typical Vittaria type germination. The
existence of similar patterns of division during spore germination in the Adiantaceae
and in a member of the Schizaeaceae might suggest a close relationship between the
two families, but examination of the germination patterns of spores of other species
of Mohria and of Schizaea and Actinostachys is necessary before this evidence can
be used to support the possible origin of adiantoid ferns from a schizaeaceous
ancestry.
This work was ud aos by grant (DEB 78-01297) from the National Science
Foundation to V. Raghava
LITERATURE CITED
BOWER, F. O. 1928. The Ferns (Filicales), vol. III. University Press, Cambridge.
CHRISTENSEN, C. 1938. Filicinae. Jn F. Verdoorn (ed.), Manual of Pteridology. Martinus Nijhoff,
The Hague.
COPELAND, E. B. 1947. Genera Filicum. Chronica Botanica, Waltham, MA.
CRABBE, J. A., A. C. JERMY, and J. T. oar 1975. A new generic sequence for the
a herbarium. Fern. Gaz. 11:141-
ENDRESS, A. G. 1974. Spore germination of cet thalictroides (L.) Brongn. Ann. Bot.
38: a 7-381.
HAUFLER, C. H. 1979. A biosystematic revision of Bommeria. J. Arnold Arbor. 60:445—-4
_ and G. J. GASTONY. 1978. Antheridiogen and the breeding system in the fern genus
Bon meria. Canad. J. Bot. 56:1594—1601.
HOLLTUM, R. E. 1949. The classification of ferns. Biol. Rev. 24:267-296.
HUCKABY, C. S. and V. RAGHAVAN. 1981a. Spore germination patterns in the ferns Cyathea and
Dicksonia. Ann. Bot. 47:397—403.
. and V. RAGHAVAN. 1981b. Germination of the spores of the thelypteroid ferns. Amer. J.
Bot. 68:517-523.
KAUR, S. 1972. Morphology of the prothallus of Gymnopteris vestita. sociopath 22:46-49.
NAYAR, B. K. 1956. Studies in Pteridaceae Il. Hemionitis Linn. J. Indian Bot. ip stage
________. 1962. Ferns of India—V. Hemionitis. Bull. Lucknow Natl. Bot. Gard. 6
. 1964. Some aspects of the morphology of Pityrogramma calomelanos = es aie ‘.
Indian Bot. Soc. 43:203-213.
. 1970. A phylogenetic classification of the homosporous ferns. Taxon 19:229-236.
_ and S. KAUR. 1968. Spore germination in homosporous ferns. J. Palynol. 1:10—26.
a - KAUR. 1971. Gametophytes of homosporous ferns. Bot. Rev. 37:295-396.
RAGHAVAN, V. 1965. Action of purine and | analogs on the growth and differentiation of the
gametophytes of the fern Asplenium nidus. Amer. J. Bot. 52:900-910.
1976. Gibberellic acid-induced germination ie? spores oH Anemia phyllitidis: Nucleic acid and
nade synthesis during germination. Amer. J. Bot. 972.
Cell morphogenesis and macromolecule synthesis during phytochrome-controlled
germination of spores of the fern, Pteris vittata. J. Exp. Bot. 28: 439-456.
_ HUCKABY. 1980. A comparative study of the cell division patterns paring
, and C
ria (Schizaeaceae). Amer.
germination ‘of spores of Anemia, Lygodium and Mohri
67:653-663.
RAO, A. R. 1949. The prothallus of Hemionitis arifolia Sm. Curr. Sci. 18:349-
RUTTER, M. R., and V. RAGHAVAN. 1978. DNA synthesis and cell division during spore germination
in Lygodium japonicum. Ann. Bot. 42:957-965.
120 AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 4 (1981)
SHORTER NOTES
THE CHEMOIDENTITY OF THE HOLOTYPE OF PITYROGRAMMA
TRIANGULARIS.—In recent years, the Pityrogramma triangularis complex has
been the subject of extensive phytochemical studies, for example, by Smith (Bull.
Torrey Bot. Club 107:134—145. 1980 and references therein). The nominal variety
tacitly recognized by Weatherby (Rhodora 22:113-120. 1920), var. triangularis, has
proved to be rather complex in itself with respect to the composition of frond-surface
flavonoids and chromosome numbers. The plants subsumed under var. triangularis
are morphologically very similar, but their extreme chemical differences and distinct
regional distribution suggest that further taxonomic revision within this group may
prove warranted. It is critical, therefore, to establish the chemical identity of the
holotype of var. triangularis, since ultimately the one biological entity identifiable
with the holotype must bear the name var. triangularis.
The holotype, which is in the herbarium of the Berlin Botanical Garden, was
collected by the German poet, botanist, and globetrotter Adelbert von Chamisso. It
bears a note in his handwriting “Gymnogramma triangularis Kaulf. Enum. p. 73,
legit deditque A. v. Chamisso, California.” According to Eaton, quoted by Alt and
Grant (Brittonia 12:153—170. 1960), the specimen was collected in 1816 near San
Francisco. It is filed at Berlin under “Polypodiaceae, Gattung No. 62a. Ceropteris,
Art No. 21 triangularis.”
A minute fragment of the holotype was made available for analysis of its farina;
22 mg of material was rinsed with acetone to dissolve the exuded flavonoids. These
were identified by direct comparison with authentic markers on polyamide-TLC; for
experimental details see Wollenweber, Dietz, Schillo and Schilling (Z. Naturforsch.
35c:685-690. 1980). The major constituent is ceroptin; minor components are
triangularin and another compound which is not yet fully elucidated, “tvt-11,”
according to Dietz (unpubl. dissertation, Darmstadt). This flavonoid pattern is
characteristic of those plants representing the ceroptin chemotype of var. triangularis.
The holotype specimen itself chemically resembles those plants of the diploid
ceroptin type collected by D. M. Smith from Refugio Pass, Santa Barbara County,
California according to Star, Seigler, Mabry and Smith (Biochem. Syst. Ecol.
2:109-112. 1975).
We think it is a most remarkable result that the holotype of P. triangularis can be
equated unambiguously with the typical and well defined ceroptin chemotype of var.
triangularis. This illustrates the powerful role that chemotaxonomy can play in
determining the application of names by the type method. Thanks are due to Dr. D.
Meyer, Berlin, for kindly supplying the fern fragment used in this study.—Eckhard
Wollenweber, Institut fiir Botanik, Technische Hochschule Darmstadt, Schnittspahn-
strasse 3, D-6100 Darmstadt, Federal Republic of Germany and Dale M. Smith,
Department of Biological Sciences, University of California, Santa Barbara, CA
93106.
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 4 (1981) 121
A MAJOR RANGE EXTENSION FOR THELYPTERIS SIMULATA IN
THE SOUTHERN APPALACHIANS.—Recently, while examining fern speci-
mens in the Illinois Natural History Survey Herbarium (ILLS), I found a collection
of Thelypteris simulata (Davenp.) Nieuwl. collected in August 1931 at New Found
Gap, Sevier Co., in eastern Tennessee at the North Carolina border (Pepoon 935).
Herman S. Pepoon was a botanist from Chicago, Illinois and the author of “Flora of
the Chicago Region.” Pepoon himself had identified the specimen as Aspidium
simulatum Davenp., and wrote on the label that the plant occurred at “summit
elevations, damp woods near New Found Gap.” Pepoon deposited several hundred
specimens from eastern Tennessee in ILLS, and it is unlikely that a person of his
competence would have correctly identified this unusual fern and then would have
made a gross error in describing its location. Furthermore, Pepoon’s collecting
notebooks, which are preserved at ILLS, show that he did not collect in west-central
Wisconsin or in the New England states from which this fern presently is known.
Thus, it seems unlikely that the specimen represents a false record due to a label
mix-up.
The specimen is significant because it represents a disjunction of about 400 miles
from the nearest station recorded for this species, which is in northern West Virginia
(Tryon & Tryon, Amer. Fern J. 63:66. 1973). In addition, this find may bear upon
the species distribution during the late Pleistocene; it seems likely that 7. simulata
was present and probably more widespread in the southern Appalachians during
Wisconsinan glacial times, and then gradually became restricted to higher elevations
and latitudes as the climate changed and the glaciers receded during the Holocene.
Although long-distance spore dispersal can never be completely ruled out, it seems
unlikely. Many other pteridophytes show a similar widespread distribution in the
north and become gradually restricted or disjunct at high elevations southward in the
Appalachians, including Botrychium matricariifolium B. multifidum, Dryopteris
campyloptera, Gymnocarpium dryopteris, Lycopodium annotinum, L. selago, and
Phegopteris connectilis. | encourage pteridologists to look for populations of this
interesting species in acid, boggy areas at high elevations in the Great Smoky
Mountains region.—Robbin C. Moran, Herbarium, Illinois Natural History Survey,
Natural Resources Bldg., 607 E. Peabody Dr., Champaign, IL 68201.
A NEW INDIANA STATION FOR EPIPHYTIC RESURRECTION FERN.—
In 1975, the author discovered a large, healthy population of Polypodium
polypodioides (L.) Watt growing approximately 30 feet up on a branch of a dead tree
at Carnes Mill, Crawford County, Indiana. Hemlock (Tsuga canadensis), Yellow
Buckeye (Aesculus octandra), and Goatsbeard (Aruncus dioicus) flourish on the
sandstone cliffs and the tight valley slopes encasing the Little Blue River at this
point. Although commonly found at similar heights in the south, literature research
indicates that this is a relatively rare habitat in the interior northern reaches of this
fern’s range. It is reported by Deam (Flora of Indiana, 1940) from nine counties in
122
AMERICAN FERN JOURNAL: VOLUME 71 NUMBER 4 (1981)
southern Indiana. He stated that he once found it growing in the crotch of a Bur Oak
tree in Wabash County, but that “this is the only specimen I have ever seen growing
on a tree in Indiana, although it is common in this habitat in the south.” Welch in
Lindsey (The Natural Features of Indiana, 1966) stated that she had not seen the
Resurrection Fern growing on a tree in Indiana.—Ronald R. Van Stockum, Jr., 810
Kentucky Ave., Frankfort, KY 40601.
AMERICAN FERN JOURNAL
Manuscripts submitted to the JOURNAL are reviewed for scientific content by
one or more of the editors, and, often, by one or more outside reviewers as well.
During the past year we have received the kind assistance of J. D. Caponetti, A. M.
E
, R. L. Petersen, H. E. Robinson, R. G. §
tolze, W. C. Taylor, R. A. White,
D. P. Whittier, and J. J. Wurdack, to whom we are deeply indebted. We welcome
suggestions of other reviewers.—D.B.L
INDEX TO VOLUME 71
Acrostichum areolatum, 101
Actinostachys. 119
Adiantaceae, 109, 119
Adiantum capillus-veneris, 91
per ore salvinii, 75-79
Amphine , 82: immersum, 82
"101
117; anthriscifolia
Anemia, 109, . 100; tomentosa var. mexicana,
00
erica 109, 111, 117: chaerophylla. 110, 111, 113; osteniana,
Manes 65: simplicior, 65-67
Arachniodes simplicior new to South Carolina and the United
States, 65
— acutum, 35; alpestre, 91; biserratum, 35; conterminum,
60: co sdibolisien, be exaltatum, 38: immersum, 82; simulatum.
121; whereas
gases fe aon -nigrum, 91: x ere 89; ebeneum, 90:
riophyllum, 91; . 90; ruta-muraria,
tts p. trichomanes, 95:
‘ ; trichomanes-ramosum, 91:
rene!
Ac
88; x herb-wagneri, 89; pinnatifidus. 87.
; Shawneensis, 85-8
* Asplenosorus nsevionmsnanee . new — fern hybrid between
hizophy Illus,
Athyrium, 48: alpestr var. ame no subsp. american-
» 91; Reinnasper pe descsithction: ot: filix-femina, 5-8:
Me cum, 91
Azolla, 33; filiculoides, 33; carolinian
Azolla filiculoides new to the vmiionis United States, 33
Bates, V. M., E. T. Browne, Jr. Azolla filiculoides new to the
Da cae Unie States, 33
By, -S.°3: . Satija. Lepisorus kashyapii in the western
Himalayas, i.
aie clubmosses (Lycopodiella) in Kentucky, 9
Bom 1695-499, (E925; 899; pi ee 111-113:
hsp, VW, 113-115; pedata, 100, 111-113; hialnicet: 11,
115
Botrychium, 1, 15, 16, 8. 20, 25, 28; biternatum, 13-15, 18;
bo 5,30:
besperium, sie lavaciacdaiicha: 25, 28, 30, 68,
nari 8, 30, f. incisum, var.
* cbiaiinan. pe
2 30;
—30; multifidum, ;
m, 21, 24- 26: ternatum, 13; simplex, 2, 23, 28, 92;
virgini . 13, 14, 17, 28, 30
= rach pattern of Hypolepis ye 41
. T., Jr. (see V. M. Bates, Jr.
raved si sorus rhizophyllus, 85, 87-89
Chain ferns of Florida, |
Cheil S ag castanea, 62; ni 62, 100; fendleri, 100:
microphylla, 91; hii, 62: , 62: wootonii, 100
The peaei of vi py of | Plate triangularis.
120
Christella, Ph reser s, 82
Clifford, t cole Ferns, fern allies and conifers of
pean ioe”
Cody, W. J. (
. (see A. E. Porsild)
INDEX TO VOLUME 71
Comparative ecology of Woodsia scopulina sporophytes and ga-
metophytes,
Constantine, Ay ee yk i =
Cranfill, diella) in Kentucky, 97; Ferns
= fem alee of Kentucky pe 12
, 109
c
ve
Cy aoa 4
Cystopteris dickinson fs alee 31, 92, var. mackayii, 92.
subsp. tenuifo rusa, 92: reevesiana, 92; tenuis, 92
Diplazium ghee and ‘Sclaginella uncinata newly discovered in
eorgia, 4:
Dryopteris, 107; arist
consanguinea var. aequalis,
ea 68; filix-mas, 46;
phegopteris, 68
Hah 75
Equisetum arvense, be fluviatile, 1; beara ats
1; rachyodon, |, 2; variegatum,
ma,
ata, 46; campyloptera, 121; baea, 60:
60; dilatata, 68: ones neta, 68:
immersa, 82; ludoviciana, 49:
; eat 100:
2
trachyodon in New Jer
Fairc wees ths R. Diplazium japonicum and Sclvginatla: ‘ancinata
Fete oat _ eae of Kentucky (rev.), 12
Ferns, fi f Australia (rev.), 9
Flora of Baja Calife ), 100
Fosberg, F R. & M--H. Sachet. Nomenclatural notes on Micro-
ee sapen seis
hrolepis in Florida, 35
Clapper soy
Gordon, Judith iaahaaiiins simplicior new to South Carolina
and the ne ae 6
Grammitis, 75
Gruber, T. M b f Hypolepis repens, 41
68, 121, subsp. disjunctum, 68
Gy <, +,
Gymn nogrecnea ang, 120
Gymnopteris, 109, , 117; rufa, 11
, 114, 115, 117; vestita, 111
ae 117; ee 115135,-1
16; palmata,
69
an. Spore germination
; Vi ing
mma, Bommeria, Gymnopteris, Hemionitis
ont 70; ne 69-73; mexicana, 69, 70:
.¢ oss gus 70: tuerckheimii, 70
alvinia minima new to Louisiana, 95
Leaf turnover rates and vat iisioey of the Cetera American tree
tm Alsophila orcbees oo
Lepidoneuron biserratu
Lepisorus, 53; e avatus, S38 oe 53, var. kashyapii, 53. var.
. Var. S35
eae kasapi in ak western Himalayas. 53
Lellinger, . Notes on North American Pen 90
Lorinseria me olata, 48, 49, 101
Lucansky, T. W. Chain ferns of Florida pe
Lycopodiella, 97 des, 97. 99: appressa, 97-99: x
brucei, 97-99: x copelandii, 97: prostrata, 97-99
opodium 17, 97; alopecuroides, 97: pees ik x
Lyco}
bopelendii, 97: dendroideum, 31; inundatum, 31, var. appressum.
123
97, var. elongatum, 97; eke Lepidotis, 97: selago, 121
Lygodium, 109, 117; japoni
A major atl extension Pai Thetypeeris simulata in the southern
Appalac hep
Marsilea pone a a3, — 93
Mi rog 29: falsivenulosum. 83: motleyi
Mohria, 109, 117, 119: cafforum, 119
Montgomery, J. D. Equisetum variegatum and E. x trachyodon in
New Jers
sis, a new natural fern
and Cam) tosorus rhizo-
, R. C. XAsplenosorus shawneen
hybrid between nar trichomane
ee
=
=]
a
g
ae
SRO
eo @ +*
nn
3
3
e
ai
Nau . E. The genus Nephrolepis in Florida,
Nem acutum, 35; biserratum, 35; caribaeum, oo conterm
m, 60; exaltatum, 38; ayaa var. sitioram, 60; kaulfussii. $9.
er
pentane 58; tu
Nephrolepis, 35-37, os acuta, 35; x averyi, 35- 39, 40:
biserrata, 35-40, Furcans, 40 sewers 163 var.
tuberosa, 37; exaltata, 35, 36, . bise ]
Bostoniensis, 40, cv. ntissima, 40. cv. tephra s. 4
M. P. s, 40, var. tuberosa, 37: falcata f. asi 40:
utula var. acuta, 35, serrata, 35, cv. Superba, 40:
multiflora, 35-39; pectinata, 40; rivularis, 38; tuberosa, 37
A new BAe station for a tic resurrection fern, 121
A new Isoétes from sa ca,
e — S ie moonwo = Brn subg. Botrychium (Ophio-
olen ssaceae) from sip ae
apace notes on Micronesia ferns, 82
Notes on North American
sa on North American serie vascular plants— ge 2
Notes on Selaginella, with a new variety of S. pal oe
Notholaena bryopoda, 62: greggii, 62: neglecta, = gi 63
O nsibilis, 46, 48
10, 25
be hs singe 13, 14, 16, 17: intermedi-
are, paagtaorantt 13: pendu-
35 vul 13. va
Ma
; 25; ti
lum, 13; petiolatum, cS pe
: pron m, 13-16, 18, var. igen 13, eve 7
mea ; ‘es regalis, 4
ppv ee 7 interm ile 63; longimucronata, 100:
, 100; skinneri, 100
tilis fa
_
Programa 10, a austroamericana, | 1: calomelan-
we me 5 by: cy il, 111, 115, 118: PSOE
63, 12
0, var. maxonii. 63: trifoliata. 11
BF
<=
Polypodiaceae, 13, 100
Polypodium, 60, 93; californicum, 63: coetitotint, so exaltatum.
i : ilentum, 60:
38; glycyrthiza, 63, subg. G le ,
hesperium, 63; kashyapii, 53: subg. arginaria, 93: pectinatum,
93. subg. Pectin: 93: polypodioides, 121. subg Polypodium,
m,
= scouleri, 93; virginianum, 68: vulgare. 93, subsp. virginian-
m, 68
Pol inh tichoides
, 67
aay. Vascular plants of continental
Pa
Porsild, E. & W.
cites territories, da (rev.).
Proct ‘axonomic notes on Jamaican — 57
Pteridium, “100; aq uilinum, 48, 109, var. pubescen’
Pteris multifida, ‘Gr: vittata, 115
Quercifilix, 107
Raghavan, V. (see S. C. Huckaby)
124
Range ~~ for two lycopods on Baranof Island, southeastern
Alaska
Reeves, T. ‘oie on North American lower vascular gt —Il, 62
Reviews: Ferns and fern allies of Kentucky, 12; Ferns, fern allies
and conifers of Australia, 9; Flora of Baja California, 100;
L.
inopsis de las especies Lycopodiu (Lycopodiaceae
Pteridophyta) de la seccién Crassistachys Herter, 84: Vascular
plants of continental a territ Spey 68
ra, a
t, M.-H. (see F. R. Fosberg)
95
ima new to Louisiana, 95
i K. eS. S. Bir
Scale ge Paar on fariose species of Pityrogramma, 10
Schizaea, 119; pusilla, 25
Schizaeaceae ci oh 117, i
Seiler, R. B bs es and natural history of the Central
merican tree fern A sein salvinii, 75
— sh arizonica, 64; cuspidata, 51, var. elongata, 51:
a. 63 i
gynandrum, 51: ae 49, 50
Short, J. W. Equisetum arvense in Alabama, 64
Sinopsis de las especies de Lycopodium L.
ridophy la
( Soho nashud
seccion Crassistachys Herter (rev.
le
Smith, D. M. (
Spore germination and young gametophyte i At of Botry-
chium and Hoglossum it in axenic culture
S rminat! Anogramma, oN Gymno-
rogramma, 109
on Selaginella, with a new variety of S.
yea ret ferns—III, 57
Thelypteris. 57, 59, Adenophyllum, 58: subg. Adeno-
phyllum. 57, 58: sect. "nea 58, 59; whe Amauropelta,
AMERICAN FERN JOURNAL: VOLUME 71 (1981)
57-59; subg. Apelta, 57; balbisii, 585.59; ~~ ——
, 60 a3
57; subg. Blepharitheca, 57; baea, 60: dec
60; dentata, 48, 82: gracilenta, 60: gracilis, 60: ns sis, “s
harrisii, 60; hispidula var. versicolor, 94; immersa, 82
if .
uen
at | 57, 58: ran ea,
. 57, f. crista zt is mulata, 121; torresiana, a8
tela, x 59; subg. ttncinelia. 37: underwoodian
versicolor,
ea 8A: a 82: falsivenulosum, 83; motleyi,
Trichopteris schiedeana, 75
An unusual record of Asplenium trichomanes from northeastern
Van ‘Stock R.
rection fern 542
satus plants of ponienand northwest territories, Canada (rev.),
68
. Jr. A new Indiana station for epiphytic
erage (see P. J. Watson)
ee
—, eae) sr =
Rae Ba: speci moonworts,
ae pas iain ee from North
America, 20
Watson, P. J. & Margarita Vazquez. Comparative ecology of Wood-
sia scopulina ee wR auaesgaaaile
Whittier, D d you ng ipiarieiea-tail develop-
ment of Botrychi 1 Ophi 13
Wiggins, I. L. Flora of Baja Cali 100
ele VOB, ‘Dietz. Scale insects feeding on farinose
a of Ptyrogramma, 10
Wollenwe .&D ~~ Smith. The chemoidentity of the holo-
type o e paid rogramma triangularis,
Woodsia, 8; alpina, 68; jr ace 68; oregana, 68; plummerae,
= bi lin = 3-8
wardia, 101, 107, angustifolia, 101; areolata, 98,
rae ns, im om 105-107; virginica, 48, 101-107
7
-
101-107:
ERRATUM FOR 1980
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