AMERICAN FERN JOURNAL 1976 -7 7 PUBLISHED BY THE AMERICAN FERN SOCIETY EDITORS David W. Bierhorst Gerald J. Gastony David B. Lellinger John T. Mickel MERCURY PRESS, ROCKVILLE, MARYLAND 20852 @issour! BoTANIcAG CONTENTS Volume 66, Number 1, Pages 1-32, Issued April 1, 1976 Spore Morphology of the Hawaiian Genus Sadleria (Blechnaceae) ROBERT M. LLOYD xsomopteris anasilla, a New Fossil Fern Rhizome from the Cretaceous of Maryland JUDITH E. SKOG A New Species of Hymenophyllum from Central America ROBERT G. STOLZE Uncommon Wall Thickenings in the Sieve Cells of Pteris wallichiana J. J. SHAH and M. N. B. NAIR Some Lesser Known Ferns from the Western Himalayas, 1. Cheilanthes anceps var. brevifrons S. P. KHULLAR Shorter Notes: A New Locality for Lycopodium serratum in Mexico; Naturalization of Cyrtomium fortunei in North America; Forked Fronds in Asplenium rhizophyllum; A Note on the Young Fronds Ophioglossum palmatum; The Identity of Polypodium sectum 19 No _ furfuraceum f. pinnatisec Ph, Reviews 28-30 Suggestions to Contributors 32 Volume 66, Number 2, Pages 33-80, Issued July 1, 1976 Edgar Wherry in Pennsylvania JOHN M. FOGG, JR. 33 Perry Creek, Washington, a Fern-watcher’s Eldorado A. R. KRUCKEBERG 39 Origin of the Pteridophyte Flora of the Bahamas, Caicos and Turks Islands DONOVAN S. CORRELL 46 Spore Retention and Release from Overwintering Fern Fronds DONALD R. FARRAR 49 The Distribution S Abundance of Dryopteris in New Jerse JAMES D. MONTGOMERY 53 Cystopteris bulbifera in the Southwestern United States TIMOTHY REEVES 60 Variation in North American Asplenium platyneur W. CARL TAYLOR, ROBERT H. MOHLENBROCK. and FREDDA J. BURTON 63 The Distribution of Dryopteris spinulosa and its Relatives Eastern Canada DONALD M. BRITTON 69 Ferns: Potential In-situ Bioassay Systems fo Aquatic-borne Mutagens EDWARD J. KLEKOWSKI, JR. and DAVID M. POPPEL_ 75 Review 80 Volume 66, Number 3, Pages 81-112, Issued October 1, 1976 Vegetative Propagation in Asplenium exiguum JOHN T. MICKEL Cold a of tle! Ferns n Southeastern Michiga ROYCE H. HILL Variation in Costa Rican Ophioglossum palmatum and Nomenclature of the Species LUIS D. GOMEZ P. Anatomical Studies of the Neotropical Cyatheaceae. I. Alsophila and Nephelea TERRY W. LUCANSKY Prothallus Morphology in some Tectarioid Ferns SURJIT KAUR and SANTHA DEVI The Occurrence of Thelypterin in Ferns G. H. DAVIDONIS Cystopteris fragilis in the Western Himalayas S. S. BIR and CHANDER K. TRIKHA Shorter Notes: Two New Sites for Ceratopteris thalictroides in Texas; Adiantum Capillus- veneris in the Bahama Islands; Vascular Cryptogams at a Site Deglaciated in 1880 Review Volume 66, Number 4, Pages 113-140, Issued December 31, 1976 Re-introduction of Marsilea vestita into Florida DANIEL B. WARD and DAVID W. HALL Diplazium delitescens and the Neotropical Species of Asplenium sect. Hymenasplenium ALAN R. SMITH Diffusive Resistance, Titratable Acidity, and CO2 Fixation in Two Tropical Epiphytic Ferns S. C. WONG and C. S. HEW Comparative Studies in the Biolo ogy of Lyc copodium carolinianum JAMES G. BRUCE Shorter Note: A _— of Ophioglossum vulgatum for North Dak American Fern Journal Index Errata oo _ coo \o oitesess tel Volume 66 F E R N Number 1 J O U be N A January-March, 1976 QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY Spore Morphology of the Hawaiian Genus Sadleria (Blechnaceae) ROBERT M. LLOYD — Loxsomopteris anasilla, a New Fossil Fern Rhizome from the Cretaceous of Maryland JUDITH E. SKOG_ 8 _ in A New Species of Hymenophyllum from Central America ROBERT G. STOLZE Uncommon Wall Thickenings in the Sieve Cells of Pteris wallichiana J.J. SHAH and M. N. B. NAIR _— \o Some Lesser Known Ferns from the Western Himalayas, i 1. Cheilanthes anceps var. brevifrons S.P. KHULLAR 2 ~ Shorter Notes: A New Locality for Lycopodium serratum in Mexico; Naturalization of Cyrtomium fortunei in North America; Forked Fronds in Asplenium rhizophyllum; A Note on the Young Fronds of Ophioglossum palmatum; The Identity of Polypodium furfuraceum f. pinnatisectum Reviews Suggestions to Contributors Missour! BorTanie al APR 18 (976 GARDEN LIBRARY Council for 1976 DAVID W. BIERHORST, Dept. of Botany, University of Massachusetts, Amherst, Mass. 01002. President RICHARD L. HAUKE, Dept. of Botany, University of Rhode Island, Kingston, R.1. 02881. Vice-President TERRY R. WEBSTER, Dept. of Botany, University of Connecticut, Storrs, Conn. 06268. : Secretary JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916. Treasurer DAVID B. LELLINGER, Smithsonian Institution, Washington, D.C. 20560. Editor-in-Chief JOHN T. MICKEL, New York Botanical Garden, Bronx, N.Y. 10458. Newsletter Editor American Fern Journal EDITOR-IN-CHIEF Smithsonian Institution, Washington, D. C. 20560 ASSOCIATE EDITORS DAVID W. BIERHORST ..Dept. of Botany, University of Massachusetts, Amherst, Mass. 01002 GERALD J. GASTONY ....Dept. of Plant Sciences, Indiana University, Bloomington, Ind. 47401 JOHN T. MICKEL New York Botanical Garden, Bronx, New York 10458 DAVID B. LELLINGER The “American Fern Journal”’ is an illustrated quarterly devoted to the general study of ferns. It is owned by the American Fern Society, and published at the Smithsonian Institution, Washington, DC 20560. Second-class postage paid at Washington. Matter for publication and claims for missing issues (made within six months of the date of issue) should be addressed to the Editor-in-Chief. Changes of address, dues, and applications for membership should be sent to Dr. Terry W. Lucansky, Dept. of Botany, University of Florida, Gainesville, Florida 32601. Orders for back issues should be addressed to the Treasurer. General inquiries concerning ferns should be addressed to the Secretary. Subscriptions $8.50 gross, $8.00 net (agency fee $0.50); sent free to members of the American Fern Society (annual dues, $5.00; sustaining membership, $10.00; life membership, $100.00). Extracted offprints, if ordered in advance, will be furnished to authors at cost, plus postage. Back volumes $5.00 to $6.25 each: single back numbers of 64 pages or less, $1.25; 65-80 pages, $2.00 each; over 80 pages, $2.50 each, plus shipping. Ten percent discount on orders of six volumes or more: postage additional. Library and Herbarium W. H. Wagner, Jr., Department of Botany, University of Michigan, Ann Arbor, Michigan Dr. W. 48104, is Librarian and Curator. Members may borrow books and specimens at any time, the borrower paying all shipping costs. Newsletter Dr. John T. Mickel, New York Botanical Garden, Bronx Park, Bronx, New York 10458, is editor dlehead Forum.’ The editor welcomes contributions from members and non- bers, including miscellaneous notes, offers to exchange or purchase materials, personalia, hor- ticultural notes, and reviews of non-technical books on ferns. Spore Exchange Mr. Neill D. Hall, 1230 Northeast 88th Street, Seattle, Washington 98115, is Director. Spores exchanged and collection lists sent on request. Ciitt Ba a th Gifts and Bequests tm b d to others interested e Journal, and cash or other gifts are ddressed to the Secretary. a ~~, fe boc iety b in ferns. Herbarium specimens, botanical books, back issues of th always welcomed, and are tax-deductible. Inquiries should be a AMERICAN FERN JOURNAL: VOLUME 66 NUMBER 1 (1976) l Spore Morphology of the Hawaiian Genus Sadleria (Blechnaceae) ROBERT M, LLOY D*?! The Blechnaceae is a small family of terrestrial ferns which includes about eight genera. The family is characterized by fronds with elongate sori (discrete or forming coenosori) on a secondary vein which runs parallel to the midvein of the pinna. These sori are protected by introrse indusia which open toward the costae. Most of the genera of Blechnaceae have a circumtropical, Southern Hemisphere distribution. The genus Sadleria Kaulf. is one of three fern genera endemic to the Hawaiian Islands. Nine species have been described in the genus, including S. cyatheoides Kaulf., S. pallida Hook. & Arn., S. souleyetiana (Gaud.) Moore, S. squarrosa (Gaud.) Moore, S. polystichoides (Brack.) Heller, S. unisora (Bak.) Robinson, S. hillebrandii Robinson, S. fauriei Copel., and S. rigida Copel. Past taxonomic treatments have recognized a variety of species. Hillebrand (1888) recognized our: $. cyatheoides, §S. pallida, S. souleyetiana, and S. squarrosa, the last species with three varieties. Christensen (1925) recognized seven species, includ- ing in addition to the above S. fauriei, S. rigida, and S. unisora. Copeland (1947) and Stone (1967) mentioned that six or seven species exist in the genus. A modern treatment of the genus is lacking. The species of Sadleria are found in a variety of ecological habitats, from gr lava flows to wet rain forests. The most common species, S. cyatheoides, among the first invaders of new lava flows, but is found as well in nearly Poe Acacia-Metrosideros-Cibotium forest. The remaining species are less common; S. squarrosa, for example, is restricted to wet, dark banks in upland rain forest. Morphologically, the genus is much like Blechnum. However, in contrast to the usually non-arborescent rhizome and pinnatifid to pinnate fronds of most Blechnum species, the rhizome of Sadleria is erect and in most species arbores- cent and the fronds are bipinnatifid to bipinnate. There is little available literature on spore morphology in Sadleria. Previous studies dealing only with spore size and shape include those by Skottsberg (1942), Selling (1946), Carlquist (1966), and Holbrook-Walker and Lloyd (1973). The following study was undertaken to document more fully spore morphology in this unusual genus, utilizing the scanning electron microscope, to see if this feature could provide insights into the taxonomic relationships of the species. *Department of Botany, Ohio University, Athens, OH 4 ' Appreciation i is expressed to the curators of the nc jolt within for lending the specimens o which this study is based. I would also like to acknowledge Dr. Otto Degener for his helpful ganiaas he and for making available to me his specimens of the genus for study. This work was supported by NSF staid t GB-36923. Volume 65, pie 4. of the JOURNAL was issued December 31, 1975. 2 AMERICAN FERN JOURNAL: VOLUME 66 (1976) Sadleria spores. FIG. 1.8. hillebrandii, St. John et al. 18447 (MICH), x 1200. FIG. 2.8. hillebrandii, surface view, St. John et al. 18447 (MICH). x 3000. FIG. 3. S. Piste te showing broken, multilayered perispore, Degener 18502 (GH), x os a 4. S$. cyatheoides , surface view, Degener 18502 (GH), x 5000. FIG. 5. S$. souleyetiana, St. John 24727 (MICH). x 1100. FIG. souleyetiana, surface view, St. John 24727 aie x 6000. R. M. LLOYD: SPORE MORPHOLOGY OF SADLERIA 3 MATERIALS AND METHODS Spores were obtained from herbarium specimens of most taxa of Sadleria. Untreated spores were mounted on specimen stubs with double-stick tape, coated with gold approximately 10 nm thick, and observed at 20 kv accelerating voltage with a Hitachi HHS-2R scanning electron microscope. Spore sizes were obtained from spores mounted in diaphane and observed with light microscopy. The voucher specimens are: S. cyatheoides: Hawaii: St. John & Catto 18423 (MICH). Lanai: St. John & Eames 18752 (MICH). Maui: Degener 18502 (GH). Oahu: Copeland s. n. (MICH), Fosberg 13309 (MICH). S. hillebrandii: Hawaii: St. John et al. 18447 (MICH). Maui: Bonsey s. n. 13 Aug 1951 (MICH). ? S. cyatheoides < hillebrandii: Hawaii: Degener 18494y (GH) S. pallida: Oahu: Degener 10329 (MICH), chic de 13665 (MICH). S. oe ee Maui: St. John 24725, 24727 (M ‘ : Lanai: Munro 940 (NY). Molokai: a & Anderson 1696 (MICH). Oahu: Grether & Washer Soe (US), Wagner 5756 (MICH S. unisora: Kauai: Heller 2863 (GH). RESULTS All Sadleria spores are bilateral, monolete, somewhat concave-convex in lat- eral view, and ovate in polar view. The external spore wall appears to be com- posed of several distinct layers. Light microscopy and scanning electron micro- scope studies indicate that the innermost layer in all species is smooth and without ornamentation and probably represents the exospore (Figs. 3 and /4). On top of this innermost layer are found one, two, or possibly three additional layers of spore wall, which in most species are loosely attached and easily separated from the exospore (Figs. 3 and /4). Since the outer layers are easily fractured and are not tightly affixed to the inner exospore, it is most likely that they represent a perispore, although it has not been possible to demonstrate that they are deposited from without the spore. However, in this paper the innermost smooth layer will be referred to as the exospore and the outer loosely attached layers as the perispore. Spore sizes without perispore in S. cyatheoides, S. pallida, S. hillebrandii, and S. souleyetiana are similar: 42.0-70.0 (mean = 53.5) wm long and 26.0- 45.0 (mean = 35.4) wm wide. Spores of S. squarrosa and S. unisora are significantly larger, however: 58.0-81.0 (mean = 67.5) wm long and 37.0-56.0 (mean = 45.2) wm wide. Spores of S. hillebrandii are illustrated in Figs. 1, 2, and /3. The exospore surface is smooth. The perispore appears to consist of a single layer; however, some spores possess what appear to be remnants of an inner layer similar to that of spores of S. cyatheoides. The perispores are 2.4-4.4 (7.5) wm thick and are multitiered as in S. cyatheoides (Fig. 13). Generally, the surface of the perispore appears to be composed of a series of thin, irregularly sized and shaped plates. This surface pattern overlaps a substructure of small, crowded, irregularly oriented rods, seen in cross-section in Fig. 13. The inner surface of the perispore is minutely roughened. The spores of S. pallida that were examined were found to be similar in nearly all respects to those of S. mere Spores of S. cyatheoides are illustrated in Figs. 3, 4, and /2. The exospore surface is smooth (Fig. 3). The perispore consists of at bat two different lami- AMERICAN FERN JOURNAL: VOLUME 66 (1976) aria aie FIG. 7. S. unisora, Heller 2863 (GH). x 850. CHG 8:6 Cinisora wirtate VIEW, ar Ze , ¥ m4 iy. SY . mee s ( id 8000. FIG. 9. S. squarrosa, Grether & Wagner 3924 (US), x 1000. FIG. 10. S. a, Surtace view, Gre ye & erence ; ae - aE 4 a . elas nce ; ibe ther & Wagner 3924 (US). x 5000. FIG. 11.S. squarrosa, Surface view, ~) R. M. LLOYD: SPORE MORPHOLOGY OF SADLERIA ) nated layers, an inner layer 0.5-1.0 um thick, closely appressed to the exospore, and an outer layer. Light microscopy indicates that the total perispore thickness can be up to 10 um. The outer surface of the outer layer is more or less smooth, with widely scattered and irregular perforations. An irregular deposition pattern with occasional areas showing the underlying rod pattern can be seen in Fig. 4. Connecting the outer and inner surfaces is a series of vertical rods (Fig. /2). The inner surface of the outer perispore layer appears to be smooth. The inner layer of perispore is thin and roughened on the external surface (Fig. /2). In some spores there is evidence that a third perispore layer exists, internal to the above two layers and overlying the exospore. This layer appears to be very membraneous and less ornamented than the layers above. Spores of S. souleyetiana are illustrated in Figs. 5,6, and /4. The exospore is smooth. The perispore in surface view is highly irregular and tuberculate (Fig. 5). The external surface is composed of an irregular and incomplete matrix, through which project numerous, irregularly oriented rods (Fig. 6). In some specimens, the matrix is nearly lacking. In cross-section, the perispore is seen to be composed of a single layer, with each tubercle enclosing a large lumen (Fig. /4). The interior of the perispore is composed of densely packed irregularly oriented and anas- tomosing rods. The outer surface of the perispore is variable in different spores. In some spores, the surface consists of irregular deposits between the papillate reticulum (Fig. 6); in other spores the deposits are lacking and the surface has a reticulate, areolate-papillate pattern. The perispores are 2.5-5.2 (13.3) wm thick. Spores of S. squarrosa are illustrated in Figs. 9-11. They are more variable in surface pattern than are those of the other species investigated here. The peri- spore is apparently more tightly affixed to the exospore, as fractured walls were not observed. In the spores of some plants, the perispore is constructed of irregu- larly sized and shaped anastomosing rods arranged in a reticulum (Fig. 11). Spores of S. unisora are similar (Figs. 7, 8). In other plants, the reticulate pattern of rods and lumina is more minute and difficult to see (Figs. 9, /0). Frequently, there are regular deposits of matrix between rods and overlying much of the surface. Spores from freshly collected specimens of S$. squarrosa are brownish, but in much of the herbarium material examined the spores appear white, indicating perhaps that some change has occurred in the surface structure or that the mate- rial has been bleached out. Hillebrand (1888) indicated that spores of this species were pale, ‘‘. . . at first enveloped by a dense layer of soft clavate papillae, which disappear with age, leaving only a rough surface.”” Fresh spores have not been examined under the scanning electron microscope in this study. DISCUSSION Details of the spore wall that have been discovered using the scanning electron microscope have provided important features which may be used taxonomically to help delimit taxa in Sadleria. Four perispore types are present and correspond to S. hillebrandii, S. cyatheoides, S. souleyetiana, and S. squarrosa. The spores of S. pallida are almost identical to those of S. hillebrandii; although the former species is recognizable on other morphological grounds (Degener & Degener, 6 AMERICAN FERN JOURNAL: VOLUME 66 (1976) 14 Sadleria spores. FIG. 12. §. cvatheoides, cross-section of « ngs perispore (a) and surface of inner perispore layer (b), Degener 18502 (GH), x 6300. FIG. 13. S. hillebrandii, cross-section of perispore (arrow indicates inner surface), St. John et al. 18447 ole H), x 5000. FIG. 14. S. souleyetiana, cross-section of perispore, St. John 24727 (MICH), x 5000. R. M. LLOYD: SPORE MORPHOLOGY OF SADLERIA 7 1974), spore structure indicates a very close relationship to S. hillebrandii. Spores of S. unisora are nearly identical to spores of some plants of §. sguarrosa, rein- forcing the hypothesis that S. wnisora is at best only an isolated form of S. sqguar- rosa. The morphological features of §. squarrosa make it one of the most distinct and easily characterized species in the genus. The spore variation in this species may be due to ontogenetic processes or may indicate divergence in various popu- lations. Spore diversity in Sadleria could lead to a hypothesis that the genus is polyphyletic. However, the spores do have several features in common: a peri- spore which is composed of small rods which are arranged either in a regular reticulations (in §. squarrosa and S. unisora) or are more irregularly arranged (in the remaining taxa). The basic difference between spores, with the exception of the laminate perispore of S. cyatheoides, is the outer surface pattern of the peri- spore itself. The multi-layered perispore of S. cyatheoides is apparently unique to the genus, although fractured spores of S. squarrosa have not been observed. It can be hypothesized that the basic spore type in the genus is that of S. cyatheoides. From this type, by loss of the inner perispore layer and by progres- sive loss of outer surface deposits, the surface pattern of §. squarrosa spores has been produced. Investigation of spores from plants thought by Degener (pers. comm.) to be hybrids between S. cyatheoides and S. hillebrandii reveals perispore features which are intermediate between the two species. The outer perispore surface of individual spores exhibits both the irregular plates of S. hillebrandii, as well as the smooth surface of §. cyatheoides spores, thus supporting Degener’s conclusions. LITERATURE CITED CARLQUIST, S. 1966. The biota a ser distance dispersal. III. Loss of dispersibility in the Hawaiian flora. Brittonia 18: CHRISTENSEN, C. 1925. Revised i * ese Pteridophyta. B. P. Bishop Mus. Bull. 25; 1-30. COPELAND, E. B. 1947. Genera Filicum. Chronica Botanica, Waltham, Mass. DEGENER, O. and I. DEGENER. 1974. Sadleria about Kilauea, wail Hawaiian Bot. Soc. Newsletter 13(5): 21-2 HILLEBRAND, W. 1888. Feces of ge Hawaiian Islands. C. Winter, Heidelb HOLBROOK-WALKER, S. and R. M. LLOYD. 1973. Reproductive Panay and gametophyte morphology of the Hawaiian hal genus ante (Blechnaceae) relative to habitat diversity and propensity for colonization. Bot. J. Linn. Soc. 67: 157-174. SELLING, O. H. 1946. Studies in Hawaiian colle statistics. I. The spores of the Hawaiian pteridophytes. B. P. Bishop Mus. Spec. Pub. 37: 1-87. 1. /-7. SKOTTSBERG, C. 1942. Vascular sie from the Hawaiian Islands. III. Pteridophytes collected during the Hawaiian bog survey 1938. Acta Horti Gotob. 15: 35-148 STONE, B. C. 1967. A review of the dated genera of Hawaiian plants. Bot. Rev. 33: 216-251. 8 AMERICAN FERN JOURNAL: VOLUME 66 NUMBER 1 (1976) Loxsomopteris anasilla, a New Fossil Fern Rhizome from the Cretaceous of Maryland UDI E-SKOG* A fusinized fern rhizome from the Lower Cretaceous beds of the Patuxent Formation (Potomac Group) in College Park, Maryland, is here described as a new genus and species. This rhizome is the first from a locality that was uncov- ered after the flooding of tropical storm Agnes in 1972. Locality and stratigraphy.—In June 1972, flooding that followed a tropical storm uncovered Cretaceous clays containing plant remains in College Park, Maryland, near Washington, D.C. The predominantly grey to dark grey color, sandy lithol- ogy, and mica content of the outcrop, as well as its geographic position to the west of overlying Arundel and Patapsco strata, indicate its placement in the Patuxent Formation (L.. J. Hickey, pers. comm.), the lowest of the three of the Potomac Group. A preliminary examination of a pollen preparation from the fossil-bearing bed is consistent with its placement in Zone I of the Brenner (1963) and Doyle (1969) classifications. The age of this zone is probably Aptian, but may range into the Barremian (Fig. /). Materials and methods.— Many fossils of gymnosperms and ferns corresponding to forms described by Fontaine (1889) and Berry (1911) occur in the clay lens, mainly as small fragments. Although some fragmentation probably occurred prior to fossilization, numerous planes of slippage (slickensides) in the clay are evi- dence of internal motion subsequent to burial. The clay material containing the plants was maintained in a wet condition until further preparation was possible. It was bulk macerated in hydrofluoric acid. Larger plant fragments were sieved out: then the smaller fragments were sorted under a dissecting microscope. Some of the remaining residue was centrifuged and prepared for pollen and spore analysis. Fragments for sectioning were embedded in plastic, either in Epon 812 (50% A and 50% B mixture) or in Spurr Firm Embedding Medium. Sections were cut on a rotary microtome and mounted on microscope slides in Canada balsam. Observations and photographs were made using a Wild M-5 microscope and a Nikon S-Kt microscope with a combination of transmitted and reflected light. Photographs utilized Kodak Plus-X Pan film. Description.—The well-preserved rhizome is approximately 2 cm long and 5 mm in diameter, with several roots (2.6 mm in diam.), two nodes, and a covering of hairs (some pointed and some blunt) that is more dense at the nodes. The car- bonized remains of the rhizome are preserved as fusain (Schopf, pers. comm.) of a type not derived through fire (Schopf, 1975). The cell walls are opaque, brittle, Department of Biology, George Mason University, Fairfax, VA 1 A os . . . . . -, o4 thi author is deeply indebted to Dr. Francis M. Hueber, Division of Paleobotany, Smithsonian i itution, for support and encouragement in all phases of the study, to Dr. Leo J. Hickey of the same fol bovis one on the ae deposits in the area, and to the Division of Paleobotany / adoratory space for the research. She also thanks Charles é iS aSSIS- tance with the photographs. TTT Tee ee *Research Associate, Division of Paleobotany, Smithsonian Institution, Washington, DC 20560 and 22030. —. oe ace in ee J. E. SKOG: LOXSOMOPTERIS ANASILLA FROM MARYLAND 9 glistening, and show smooth, conchoidal fractures, corresponding to Schopf's (1948) description of fusinized plant tissues. During the dehydration process for embedding in plastic, many of the brittle hairs lost their pointed tips, but the multicellular, bulbous bases remained and can be seen in section (Fig. 3). Some sections show the uniseriate tips of the hairs (Fig. 5). The cells of the hairs measure 16-33 wm in diameter. In section the rhizome is solenostelic with a sclerotic pith (cells 16-33 wm in diam.). The xylem is well preserved and is composed of scalariform tracheids (Fig. 4), protoxylem tracheids 12-17 wm in diam., and metaxylem tracheids 18-30 um in diam. Maturation of the xylem is exarch with the protoxylem surrounding FIG. 1. Stratigraphic relations of the Potomac Group in Maryland. Stages! Formational Equivalents' Pollen Zones' Time 2 Maryland New Jersey Seeinning af ataee 3 (millions of years ago)* 7) & Raritan Fm. IV Cenomanian Se ‘a z 2 Bi Ill 5 **Raritan”’ Fm. lic Upper ——— Sal dil IIB ; A 2 Albian Middle Patapsco Fm. S zs 2 oO o z IIA g reer 3 3 RE nba EOE Se ago 5 Lower 2 Arundel Clay o : ies ~ Aptian : Patuxent Fm. 3 Barremian —_—- Fe — - 1Correlations of units from Doyle et al. (1976). 2 Approximate dates from Dickinson and Rich (1972). this tissue (Fig. 4). The phloem (cells 5-8 w~m in diam.) surrounds the xylem and contains parenchyma cells 7-12 wm in diam. There is an endodermis of rectangu- lar cells 8-12 xm in diam. around the stele, although this tissue is poorly preserved and difficult to distinguish (Fig. 4). The cortex is composed of an inner layer of mixed sclerenchyma (cells 13-23 »m in diam.) and parenchyma (cells 26-46 «#m in diam.) and an outer layer of dense sclerenchyma (Fig. 2). The epidermis is a layer of isodiametric cells with hairs arising from them. The roots appear to arise from the protoxylem area and become diarch as they pass through the cortex (Fig. 2). Petiole traces are poorly preserved, but appear C-shaped and produce a gap In the stele as they are formed. No petiole traces are preserved outside the cortex. 10 AMERICAN FERN JOURNAL: VOLUME 66 (1976) 3 . we ed FIGS. 2-5. Anatomical details of Loxsomopteris anasilla. FUG. 2. Cross section of rhizome (USNM 208539 c- 1), < 50. FIG. 3. Cross section of outer cortex and epidermis with multicellular bases of hairs, x 150. FIG. 4. Cross section of portion of stele, x 600. (USNM 208539 c-10). « 50. The abbreviations are: e endodermal cells, h uniseriate portion of hair, p parenchyma in the cortex, px protoxylem, r ikea of metaxylem tracheids. FIG. 5. Cross section of part of rhizome root production, and s scalariform J. E. SKOG: LOXSOMOPTERIS ANASILLA FROM MARYLAND 1] Loxsomopteris J. E. Skog, gen. nov. Fern rhizome covered with bristle-like hairs, these multicellular at the base and tapering to a uniseriate tip, with a sclerotic pith, solenostele, exarch xylem mat- uration, and a cortex of mixed parenchyma and sclerenchyma (Fig. 2). TYPE: Loxsomopteris anasilla J. E. Skog. Loxsomopteris anasilla J. E. Skog, sp. nov. Figs. 2-5. Rhizome dorsiventral, terete in cross section, solenostelic, covered with bristle-like hairs; hairs multicellular at the base, tapering to a uniseriate tip; outer cortex sclerenchymatous, inner cortex of sclerenchyma and parenchyma; endodermis present; xylem of scalariform tracheids, maturation exarch with pro- toxylem area around the periphery of the tissue; phloem with sieve elements and parenchyma surrounding the xylem; pith sclerotic; petiole traces C-shaped; roots diarch. TYPE: United States National Museum 208539, a, b, and c 1-21, series of mounted slides, all deposited in the Paleobotany Collections, U. S. National Museum of Natural History. TYPE LOCALITY: Paint Branch, 39°00’ N Lat., 76°56’ W Long., on the creek bank 500 ft NW of the intersection Greenbelt Road and Route US-1, Col- lege Park, Maryland, U. S. A.; USNM locality 14212. STRATIGRAPHY: Lower Cretaceous Barremian or Aptian, Potomac Group, Patuxent Formation. DERIVATION OF THE EPITHET: From the Greek andsillos, meaning with bristling hairs. The amphiphloic siphonostele, exarch maturation of the xylem, scalariform tracheids, sclerenchymatous pith, mixed cortex, size of the stem, and relative age relate L. anasilla to the form genus Solenostelopteris Kershaw (1910). Based upon her description and a reexamination of the type material in the British Museum (Natural History) Paleobotany Collection (Fig. 8), Loxsomopteris differs from Solenostelopteris in cortical arrangement of sclerenchyma and parenchyma, presence of epidermis, age, locality, and size. The outer cortex and epidermis of S. japonica are not present, and so there is no indication of hairs on the type species. Kershaw (1910) suggested possible relationships with the Davallieae and, in particular, the extant fern genus Microlepia. Her opinion was based on the distribution of sclerenchyma in the cortex, the arrangement of the xylem and phloem, and the marginal thickening of the xylem in forming the leaf gap. A second species of Solenostelopteris was described by Ogura (1930) from the Upper Cretaceous of Japan as S. loxsomoides. This species and L. anasilla ap- pear to be closely related, particularly in the possession of hairs on the epidermis. The hairs of §. loxsomoides are conical, multicellular, and arise from the outer cortex, which is composed of large, thin-walled cells. Ogura’s diagram and illustration together do appear to show this sort of structure; however, the ques- tion arises as to whether the protrusions are hairs. The epidermis is not clear from the illustration, and if these are truly hairs it is unlikely that the cortex would participate as extensively in their formation as has been illustrated. Thus, al- though S. loxsomoides may very well belong in the new genus Loxsomopteris , Mt AMERICAN FERN JOURNAL: VOLUME 66 (1976) a Ae ¥ ‘ee ee te ooo es = Sty 3 : “ase te Sees 2 ~ s: s* wet *, ee : wy antes ee. %s .: : “ore : Sree Pay te st & = a ee. ~< - RAR, Neng Roe ‘ ms . ‘ %: ers : .% 33 ‘ bee eS _— > R ‘wee i *, bt ms wet OC at x ~ " oF zs <. Po < * oF. ago, ati * ss pesescee ee ie: e mts «®, a Se ‘a? @ee % ‘Ay yer CS iris @ * we ba o> os ve. ee. ee Oy i alt * BEES ,4 ott ee Lin one +6) 2 re ee is * oy tbe a pthaeete 2 < ides by terete. a er ae “A YW ~S ' + FIGS. 6-8. Anatomical detalis of Loxsomopsis and Solenostelopteris. FIG. 6. Cross section of L. costaricensis (Mickel 3001, US), x 11. FIG. 7. Cross secti aa with the pith in the lower portion of the photograph, x 5 G. 8. Cross section of the type of S. japonica, specimen v-28872a (Stopes no. 1Y A-21a), x 8. Same as Kershaw (1910. fig. 3); photograph courtesy of the British Museum (Natural History). The abbreviations are: e = endodermal cells; h = multicellular base of hair; oc = sclerenchym atous outer cortex, px = protoxylem, r = production of root, and s = scalariform thickening of metaxylem tracheids. on of portion of stele of L. costaricensis, FIG J. E. SKOG: LOXSOMOPTERIS ANASILLA FROM MARYLAND 13 seems advisable to let it remain in the form genus Solenostelopteris until the type specimen has been reexamined. Loxsomopteris differs in having hairs with pointed tips arising from the epidermis, a sclerotic outer cortex, and is terete in cross section rather that elliptic. A further distinction between S. japonica and L. anasilla is that the inner cortex is composed of thick-walled cells in the former. gura suggested a close relationship with Loxsoma (Loxsomaceae). reflected in the specific epithet, based upon the indument and stelar pattern. Two other species, S. nipanica and S. sahnii, were described by Vishnu-Mittre (1958) of Jurassic age from India, but neither of these was described as having hairs on the epidermis. On the basis of locality, age, sclerotic outer cortex, and the characteristic hairs on the epidermis, the specimen from Maryland is described as a new genus in the family Loxsomaceae. The systematic position of Loxsomopteris is difficult to determine because of the lack of attached fronds. Further work on the pinnules found in the same deposit is continuing, and may eventually suggest a more certain relationship. On the basis of the epidermal appendages, relationship with Loxsomaceae is clearly suggested. Within that family, Loxsomopteris anasilla can probably be compared most closely with Loxsomopsis costaricensis. The hairs of the latter species are multicellular at the base tapering to uniseriate tips, and the points may break off leaving some hairs blunt with only the bulbous bases remain- ing (Fig. 6). Anatomically, the fossil is strikingly similar to the living fern (Fig. 7) in the solenostele, sclerenchymatous pith, mixed cortex with the outer layer of sclerenchyma, exarch maturation of the xylem with protoxylem areas around the xylem, similar production of diarch roots, and C-shaped petiole trace at its point of origin. However, because no attachment to fronds is yet available, one must not ignore other possible living relatives of Loxsomopteris, although the similarities are not as close to any other genus of extant ferns which has a soleno- stelic vascular system. According to Gwynne- Vaughan (1903, p. 727), the exarch protoxylem seems to limit comparison to Loxsoma, Dicksonia | Dennstaedtia | apiifolia, Davallia platyphylla, D. speluncae, D. hirta, or D. marginalis. The genus Davallia can be eliminated from comparison because of the presence of paleae instead of hairs, and Dennstaedtia does not possess the bulbous-based hairs (Bower, 1926). In Loxsomaceae the characteristic islets of parenchyma in the cortex (Gwynne-Vaughan, 1901) are further evidence of the affinities Lox- somopteris anasilla has with this family. The new fossil fern rhizome indicates that the anatomical characteristics typical of the family Loxsomacae apparently were present in the Lower Cretaceous. However, there is no spore record of this family—or the available spores may not be distinctive enough to determine family relationships. Brenner (1963, p. 31) attributes fern spores of this age to the Matoniaceae, Cyatheaceae, Gleicheniaceae, Schizaeaceae, and Osmundaceae. He interprets the paleoecol- ogy of the region as a warm-temperate, broadleaf evergreen rainforest and suggests that these fern families are not inconsistent with this environment. Doyle (1969), on the other hand, suggests the possible presence of Cyatheaceae, Schizaeaceae, and Gleicheniaceae in the Lower Cretaceous, but attributes all 14 AMERICAN FERN JOURNAL: VOLUME 66 (1976) other spores to ‘‘groups of less certain affinities,’ an interpretation which is probably more accurate because trilete spores are found in many families of pteridophytes. Information from the leaf and pinnule types of the Lower Creta- ceous must be interpreted with caution, since most of the work on these beds by Fontaine (1889) and Berry (1911) was based upon compression fossils without critical comparative study. Details of leaf anatomy have not yet been investigated, but further work in progress will likely yield a reinterpretation of the groups present. Previous descriptions of these fossil beds in the Potomac Group (Fontaine, 1889; Berry, 1911) have dealt mainly with compression material. Only the fern see from the Patapsco Formation has been found as a petrifaction (Berry, 1911, p. 295). This rhizome of L. anasilla is therefore interesting, as it is the ne fossil fern stem from the Lower Cretaceous yet found in Maryland. That the most closely related species are from the Upper Cretaceous of Japan indicates further evidence of the relationship of eastern Asiatic and eastern North American floras since the Cretaceous. Li (1952) indicates that there are few extant ferns that show this relationship (Camptosorus, Osmunda, Onoclea), possibly because of the production of spores rather than seeds as a dispersal mechanism. Correlation may prove to be closer when ages of Cretaceous continental beds are better defined for Asia and America. LITERATURE CITED BERRY, E. W. 1911. Pteridophyta and Gymnospermae in Clark, W. B. et al. Maryland Geological Survey: Lower Cretaceous. Johns Hopkins Press, Baltim BOWER, F. O. 1926. The dermal appendages of the ferns. Ann. oe; 40: 479-490. BRENNER, G. J. 1963. The spores and ei of the Potomac Group of Maryland. Bull. Maryland Dept. Geol. Mines, Water Res. 27:1-215. DICKINSON, W. R. and E. I. RICH. os Petrologic intervals and petrofacies in the ha Valley Sequence, Sacramento Valley, California. Bull. Geol. Soc. Amer. 83: 3007-302 DOYLE, J. A. 1969. Cretaceous angiosperm pollen of the Atlantic coastal plain and its ee significance. J. Arnold Arb. 50: 1-35. » M. VAN CAMPO and B. LUGARDON. 1976. Observations on exine structure of Eucommiidites and Lower Cretaceous angiosperm pollen. Pollen & Spores 18: (in press). FONTAINE, W. M. 1889. The Potomac or Younger Mesozoic flora. U.S. Geol. Surv. Monogr. 15: i-xiv, 1-377, t. i-clxxx. Government Printing Office, Washington, GWYNNE-VAUGHAN, D. T. 1901. Observations on the anatomy of sdlenostelic ferns. I. Lox- gs Ann. Bot. 14: 71-98. ————.. 1903. Observations on the anatomy of solenostelic ape Part II. Ann. Bot. 17: 689-742. cass E. M. 1910. A fossil solenostelic fern. Ann. Bot. 683-691. CET 2 lone Floristic ee between eastern Asia ot eastern North America. Trans. . Phil. Soc , 42: 371-429. Obuee. Y. inn On ay OE and affinities of some Cretaceous plants from Hokkaido. J. Fac. Sci., Imp. Univ. Tokyo, III, 2: 381-413, pl. XVITI-XXI. SCHOPF, J. = Rin Variable coalification: the processes involved in coal formation. Econ. Geol. ———_.. - Modes of fossil preservation. Rev. Paleobot. Palynol. 20: 27-5 VISHNU- witli 1958. Studies on the fossil flora of Nipania, Rajm ih | Series, India- Pteridophyta, and general observations on Nipania fossil flora. The Palaeobotanist 7: 47-66. pean ROEM sinas AMERICAN FERN JOURNAL: VOLUME 66 NUMBER 1 (1976) 15 A New Species of Hymenophyllum from Central America ROBERT G. STOLZE” The Filmy Fern genus Hymenophyllum includes an interesting and closely knit complex of species, treated by C. V. Morton (1947) as sect. Sphaerocionium, subsect. Lanata. Morton (1968) revised the classification of Hymenophyllaceae and placed this group in subg. and sect. Sphaerocionium, subsect. Hirsuta, The latter is distinguished by the following characteristics: segment margins entire rather than toothed; blades bearing stellate trichomes on the leaf surface between, as well as on, the veins; and veins lacking accessory wings not in the same plane as the leaf. The group contains 20-25 species, which occur chiefly in the neo- tropics. While studying subsect. Hirsuta for “‘The Flora of Guatemala,” | came upon a new species of Hymenophyllum represented by specimens previously undeter- mined or misidentified as H. sieberi (Presl) v. d. Bosch or H. trapezoidale Liebm. A few of those determined as H. sieberi had been annotated by Morton; perhaps he (1947, p. 180) was referring to these when he said, “Some of the Guatemalan specimens are more or less aberrant.” Hymenophyllum crassipetiolatum Stolze, sp. nov. Figs. 1, 3. Rhizoma repens; folia indeterminita, 12-42 cm longa, 3-8(10) cm lata: petioli 4-11(16) cm longi, (0.5)0.6-0.9 mm crassi, non alati; laminae anguste lanceolatae vel ovatae, rhachidibus late alatis, rarius ad basin non alatis; pinnae plerumque bipinnatifidae, late alatae; pinnulae pinnatifidae, pinnulae apicales bifidae vel simplices; segmenta ultima integra; trichomata in venis et in superficiebus foliorum stellata, sessilia, trichomata marginalia radiis 4-6 plerumque adpressis et versus apicem segmenti flexis; indusia saepe latiora quam longa, trichomatibus simplicibus vel bifurcatis instructa. ‘ TYPE: Slopes of Volcan Gemelos, Dept. Zacapa, Guatemala, 1942, Steyer- mark 43302 (US; isotypes F, GH). Pendent from tree trunks, or growing on moist banks, in deep shade in cloud forests, 1,250-3,300 m. Known from Mexico (Chiapas), Guatemala, Honduras, and El Salvador. : Rhizome wiry, long-creeping, provided with simple, reddish to light brown trichomes; leaves subdistant on the rhizome, indeterminate, pendent, mature ones 12-42 cm long, 3-8(10) cm wide: petiole 4-11(16) cm long, (0.5)0.6-0.9 mm in diameter, nonalate (although the basal pinna sometimes short decurrent), sparsely to abundantly provided with simple, bifid, or (mostly) stalked, stellate trichomes; lamina ovate or, more commonly, narrow-lanceolate, not reduced at base, or the lower 1-4 pairs of pinnae somewhat shorter; rachis broadly alate throughout, or nonalate at the base, the wings plane to slightly crispate, sparsely or abundantly provided with sessile or subsessile, stellate trichomes; pinnae commonly bipin- natifid, the costae broadly alate; pinna segments 4-12 pairs, the larger ones deeply pinnatifid, the apical ones bifid to simple, ultimate segments plane or slightly “Department of Botany, Field Museum of Natural History, Chicago, IL 60605. AMERICAN FERN JOURNAL: VOLUME 66 (1976) CHICAGO NATURAL HISTORY MUSEUM Lil: << SReneeenliaeeeenennae ee centimeters I 2 PLANTS OF GUATEMALA ‘ Beapedition, 1961-52 43302 Sieera de Las Minas Hymenophylius Pendent from mossy bluffs, banks, and at base ] of tree trunks. HOLOTYPDS Common type neer summit. Field Museum of Nanural History ss HYMENOPHYLLUM CRASSIPETIOLATUM Stolze apa: middle and upper south-facing slopes of Voledin Gemelon, alt. 2,100-3,200 m. hie hi 1975 JOLIAN A. STRYEOMARK genes FIG. | Holotype of Hy : oO ? } ‘rassipeti ype of Hymenophyllum | latum, Stevermark 43302 (US), R. G. STOLZE: A NEW CENTRAL AMERICAN HYMENOPHYLLUM 17 undulate, entire, veins and leaf surfaces sparsely to abundantly provided with sessile-stellate trichomes; marginal trichomes abundant, sessile-stellate (rarely minutely stalked), most of the 4-6 stout rays commonly appressed and directed toward the segment tip; indusium as broad or broader than the ultimate segment, not or scarcely immersed in the segment tip, the valves as broad or broader than long, their margins provided with simple or bifurcate trichomes. SELECTED SPECIMENS EXAMINED: MEXICO: Chiapas: Matuda 5233 (F). GUATEMALA: Alta Verapaz: Finca Los Alpes, ca. 4000 ft Wilson 352 (F). Baja Verapaz: Sierra de las Minas ca. 5 km S of Purulha, 1600 m, L. O. Williams et al. 41976 (F). Chiquimula: Cerro Brujo, SE of Concepcion de las Minas, 1700-2000 m, Steyermark 31034 (F, GH, US). El Progreso: Between Finca Piamonte and the summit of Volcan Sta. Luisa, 2400-3333 m, Steyermark 43546 (F, US). Zacapa: Between Loma El Picacho and Cerro de Monos, 2000-2600 m, Steyermark 42795 (F, US). HONDURAS: Morazan: Dry slopes of Mt. San Juancito, near El Rosario, 1400 m, Molina 23402 (F). Ocotepéque: Mt. Cocal, 20 km NW of Ocotepéque, 1800 m, Molina 22100 (F). EL SALVADOR: Chalatenango: East slope of Los Esemiles, ca. 2430 m, Tucker /062a (F). Santa Ana: Montana Montecristo, 2100 m, Molina & Molina 12488 (F). The comparatively thick petiole of this species is one of its most distinctive features. The petiole diameter of most Hymenophyllum species is less than 0.5 mm, and frequently is only 0.1-0.2 mm. Noteworthy also is its luxuriant, highly dissected leaf blade, which often attains lengths of over 40 cm. In larger primary segments (pinnules) of most pinnae, the veins are pinnately arranged, so that the pinnae are essentially bipinnatifid. (In more distal pinnae the primary segments are merely bifid or simple.) Some other characteristics in H. crassipetiolatum are subtle, but important. The indusial valves vary somewhat in dimension, occasionally they are suborbicu- lar or sometimes slightly longer than broad, but most commonly the valves of mature indusia are broader than long. The stellate trichomes of the segment mar- gins are sessile (rarely very short-stalked), with 4-6 stout rays closely appressed along the margin and most of them bent toward the segment tip (Fig. 3). The marginal stellate trichomes of many similar species appear much more delicate and have long slender stalks with filiform rays that spread in a more random pattern. (Fig. 2). ; In Mexico and Central America, the affinities of H. crassipetiolatum are with H. sieberi. The two may be readily separated by a number of characteristics, including those in the following key: Petioles (0.5)0.6-0.9 mm in diam, not alate; pinnae essentially bipinnatifid; marginal trichomes stout, stellate, sessile, the rays mostly appressed and bent toward the segment tip; indusial Fed ‘ 5(0- trichomes simple to forked; indusia mostly broader than long; plants growing at 1,250 3,300 ie LTT > SERRE cane ecie aD serene Shere aOT SUES) aking ar veh usetanne tines emer eee H. crassipetiolatum Petioles 0.3-0.5 mm in diam., alate at the apex or in the upper half; pinnae essentially pinnatifid: marginal trichomes delicate, stellate, stalked, the rays spreading; indusial trichomes pues - stellate; indusia mostly longer than broad; plants growing at 950- 1,600 m altitude... H. sieberi Although H. crassipetiolatum frequently has been identified in herbaria as H. trapezoidale, the two belong to different subsections. The blades of the latter are glabrous between the veins and the margins, the rachises are peerage portion of the blade, and the marginal trichomes are simple, bifurcate, or stalked- Stellate with 3 delicate rays. 18 AMERICAN FERN JOURNAL: VOLUME 66 (1976) Three South American species perhaps are more closely related to H. cras- sipetiolatum than any of the Central American species: H. lindenii Hook., H. interruptum Kunze, and H. plumieri Hook. & Grev. Hymenophyllum lindenii is one of the few with petioles nearly a full millimeter in diameter, but the rachis is nonalate in the lower portion of the blade and indusial trichomes are predomi- nantly stalked-stellate, whereas those of H. crassipetiolatum are simple or bifur- cate. Both H. plumieri and H. interruptum have more delicate petioles and their blades are less highly dissected. The latter has groups of fertile pinnae often separated by several sterile ones, and H. plumieri has the petiole alate at the apex. ee 3 FIGS. 2-3. Trichomes on segment margins of Hymenophyllum. FIG. 2. Hymenophyllum sieberi. FIG. 3. Hymenophyllum crassipetiolatum. In spite of Morton’s past work with the family and genus, Hymenophyllum still poses many interesting problems in taxonomy. The plants are small and often inconspicuous, and grow primarily in the deep, dense forests, so undoubtedly there will be new discoveries as additional collections are made. LITERATURE CITED MORTON, C.V. 1947. The American species of Hymenophyllum, section Sphaerocionium. Contr. U.S. Nat. Herb. 29: 139-201. . 1968. The genera, subgenera, and sections of the Hymenophyllaceae. Contr. U.S. Nat. Herb. 38: 153-214. AMERICAN FERN JOURNAL: VOLUME 66 NUMBER 1 (1976) 19 Uncommon Wall Thickenings in the Sieve Cells of Pteris wallichiana J. J. SHAH and M. N. B. NAIR*:! Pteris wallichiana Agardh was collected from Kerala, India, fixed in FAA, and processed for microtomy by conventional methods (Sass, 1958). Sections were stained following Cheadle, Gifford and Esau (1953). In addition, I2KI and H2SO4 was used for cellulose, the PAS reaction for insoluble polysaccharides (Jensen, 1962), and Toluidine blue ‘O’ for cell walls (O’Brien, Feder & McCully, 1964). During our study of phloem structure in Preris wallichiana we observed un- common papillose thickenings on the walls of sieve cells, rarely in the rhizome and frequently in the rachis. These sieve cells were randomly distributed and generally belonged to the metaphloem. Sieve cells with this thickening did not appear to be structurally different from other sieve cells. The thickenings project 2.4-10.4 wm from the sieve cell wall into the cell lumen (Figs. /-2). Phloem parenchyma cells having a common wall with sieve cells rarely show this type of thickening (Fig. 4). The thickening may be on only one side of the common wall or it may be on both sides of the wall of two adjacent sieve cells. Occasionally small fissures, some of which may be artifacts, appear in the papillose thickenings, which cause the thickening to appear to consist of different parts (Figs. 5-8). The thickenings are neither tyloses nor, apparently, a reaction to fungal infection. The thickenings are PAS-positive and give a confirmatory test for cellulose. They stain purple red with toluidine blue ‘O’, a staining reaction similar to the remaining part of the sieve cell wall. A membraneous lining is sometimes observed over the thickenings (Fig. 3). Similar wall thickenings have not been reported in sieve elements of other fern species. Warrington (1970) reported papillose cellulose thickenings of cell walls in the cortical cells of rhizome of Geocaulon lividum (Santalaceae), a dicotyledon- ous plant, but these thickenings were always in pairs on common cell walls. LITERATURE CITED CHEADLE, V. I., E. M. GIFFORD, Jr. and K. ESAU. 1953. A staining combination for phloem and contiguous tissues. Stain Technol. 28: 49-53. JENSEN, W. A. 1962. Botanical Histochemistry. W. H. Freeman, London. O'BRIEN. T. P.. N. FEDER and M. E. McCULLY. 1964. Polychromatic staining of plant cell wall by Toluidine blue ‘O’. Protoplasma 59: 367-373. SASS, J. E. 1958. Botanical Microtechnique. Iowa State College Press, Ame WARRINGTON, P. D. 1970. Cell wall thickening in Geocaulon lividum (S Bot. 48: 1677-1679. §.. antalaceae). Canad. J. *Department of Botany, Sardar Patel University, Vallabh Vidyanagar 388120, India. "This work was fied by grant GF-36747 from the National Science Foundation, U.S.A. 20 AMERICAN FERN JOURNAL: VOLUME 66 (1976) 24. um [ oe EE a FIGS. 1-4. Rachis phloem of Preris wallichiana. FIG. 1. Transection showing sieve cells with and without wall thickenings. FIG. 2. Longisection of a portion of a sieve cell showing a thickening. FIG. 3. Longisection of a portion of a sieve cell showing a thickening with membranous lining. FIG. 4. Longisection of a portion of a sieve cell and a phloem parenchyma cell showing paired thickening along the common wall. P = phloem parenchyma, SC = sieve cell. FIGS. 5-8. Longisections of portions of sieve cells showing a variety of fissures in the thickenings. AMERICAN FERN JOURNAL: VOLUME 66 NUMBER 1 (1976) 21 Some Lesser Known Ferns from the Western Himalayas, 1. Cheilanthes anceps var. brevifrons S. P. RHULLAR”* Nearly a century ago Blanford (1886) recognized two new species in the Cheilanthes farinosa (Forsk.) Kaulf. complex: C. anceps Blanf. and C. grisea Blanf. Later, Blanford (1888) reduced these species to varieties of C. farinosa. Hope (1901), Alston (1956), and Verma (1964) considered these taxa to be species. Panigrahi (1962) regarded the tetraploid occurring in Ceylon as Aleuritopteris anceps (Blanf.) Panigr. and the diploid as A. grisea (Blanf.) Panigr. Hope (1900, p. 250) apparently was the first to point out that C. anceps had two forms. His specimens, from near Simla in the western Himalayas, although col- lected at the same time as Blanford’s, differed in being smaller, more delicate, and sometimes in having apparently concolorous rhizome scales. This led Hope to comment that his specimens seemed near to C. farinosa var. grisea (Blanf.) Blanf. These small specimens cannot be C. grisea, however, because that species has a naked rachis, always has concolorous, brown scales that are restricted to the stipe bases, and has a crenate or irregularly lobed indusial margin. Cheilanthes anceps, on the other hand, is characterized by its lanceolate to oblong-lanceolate fronds, scaly stipes and rachises, and lacerate indusial margins. Sufficient morphological and cytological differences occur between the two types of C. anceps to require their separation into varieties, but I hesitate to assign the rank of species to the two because of certain resemblances between them. In both, bicolorous scales are present on the stipe and rachis, the fronds are more or less lanceolate, and the indusial margin is always lacerate. Cheilanthes anceps Blanf. var. anceps. oe Fig. 1. Fronds 25-48 cm long; stipes 10-20 cm long, 0.75-1.5 mm in diam., with a sulcate stele, approximately equalling the laminae, dark chestnut to almost black, glossy, the stipe scales linear-lanceolate, bicolorous, dark with pale margins; rachis scales similar; laminae 10-24 cm long, (2.5) 3-4.5 cm wide, lanceolate to oblong-lanceolate, thin, not compact, heavily white-waxy beneath, the lowest 2 or more pinna pairs subequal, rather distant; indusia narrow, toothed or lacerate, n=29 in material from Darjeeling, eastern Himalayas (Verma in Mehra, 1961). SPECIMENS CITED: INDIA: Himachal Pradesh: Simla Hills, 650 m, 15 Sept 1960, S. S. Bir (PAN 4266-69). Uttar Pradesh: Mussoorie, Jamna Bridge, 600 m, 27 Aug 1959, S. S. Bir (PAN 2639-40); Nainital, Kathogodam Road, 1200 m, 5 Oct 1975, S. P. Khullar 24 (PAN). West Bengal: Darjeeling, Manjitar- Teesta Road, Aug 1951, S. C. Verma (PAN 3776-3779). : This variety occurs at 600-1800 m altitude in the western Himalayas and at about 150-450 m altitude in the eastern Himalayas. *Department of Botany, Panjab University, Chandigarh 160014, India. 22 AMERICAN FERN JOURNAL: VOLUME 66 — 1976) HER GARIU™M afer Boss . fhe bet tates Ley OS PANJAB UNIVERS:TY FIG. 1. Cheilanthes anceps Blanf. var. anceps (S. S. Bir, PAN 2639). S. P. KHULLAR: FERNS FROM THE WESTERN HIMALAYAS, 1 23 r. brevifrons (S. P. gesagt meee §977). FIG. FIGS. Det: Holotype . Cheilanthes anceps va a eas Entire specimen. FIG. 3. Indusial lobe sate highly lacerate margin. Stipe scale, x 20. 24 AMERICAN FERN JOURNAL: VOLUME 66 (1976) Cheilanthes anceps var. brevifrons Khullar, C. ancepiti var. anc oe statura cai sete tere stipitibus tides, laminis crassiusculis et chromosomatum numero ert. Fronds (4)9-12.5(27) ses a stipes (2)3.6-4. 7(10) cm long, 0.3-0.8 mm in iam., with a terete stele, generally shorter than or equalling the laminae, light to dark brown, glossy; stipe scales narrowly tities r-lan ge — me OUS, abu ndant, bicolorous, brown at the center, paler towards the ore distinctly bicolorous near the stipe base than near the apex; ens sealee Seite a smaller; laminae 5.5-8.0 cm lo =e 3-3.5 cm wide, thick, compact, more or less oblong- lanceolate, the a aia acuminate, the basal pinna sale shorter than the supra babel pair, the basal bocce teal pair divergent and nearly parallel to the stipe; pinnae subopposite, divergent, the lower 2 or 3 suprabasal pairs equal and distant compared with those in the proximal half of the lamina; pinnules Aine. gril ing, lobed; indusia narrow, generally regularly lobed, not ee en ss rown, margin deeply and regularly lobed with long, marginal teeth; s br ae globose, the exine with reticulate ickennics appearing like mar ms picicctone 50. 70 um Xx 45-65 um; n=30 in material from the western Himalayas (Khullar Mehra, 1973). HOLOTYPE: Taradevi, Simla, Himachal Pradesh, India, 1800 m, July 1967, et Saree a Rea 5977). PARAT INDIA: ibis ee Simla Hills, Kasauli, 1800 m, July 1967, S. P. Khullar (PAN 6053-6068, 6492); Tuti Kundi, 1500 m, 18 Aug 1960, S. S. Bir (PAN 3908- 10); Dalhousie, Bathri, 1350 m, 16 Aug 1964, M. Golaknath (PAN 5166, 5291, 5530). Uttar Pradesh: Ranikhet, 1800 m, Sept 1972, S. P. Khullar (PAN 22). This variety is quite common between 1300 and 1800 m altitude in the western Himalayas LITERATURE CITED ALSTON, A. H. G. 1956. The subdivision of the Polypodiaceae. Taxon 5: 23-2 BLANFORD, H. F. 1886. The silver ferns of Simla and their allies. J. Simla Nat. Hist. Soe, 1213-22. . 1888. A list of the ferns of Simla in the N. W. Himalaya between levels of 4,500 and 10,000 fe et. J. Asiatic Soc. Bengal 57: 294-315. HOPE, C. W. 1900. The ferns of northwestern India. J. Bombay Nat. Hist. Soc. 13: 236-251. KHULLAR, S. P. and P. N. MEHRA. 1973. Cytotaxonomy of W. Himalayan ferns, I. Schizaea- ceous series. Res. Bull. Panjab Univ., n.s., 23: 189-204. MEHRA, P. ¥ 1961. Chromosome numbers in Himalayan ferns. Res. Bull. Panjab Univ., n.s., 12: 139-1 PANIGR AT G. 1961. A note on Aleuritopteris grisea (BI.) Panigrahi comb. nov. and A. anceps (BI.) Panigraphi comb. nov. Bull. Bot. Surv. India 2: 321-322. VERMA, S. C. 1964. Cytotaxonomic investigations on some E. Himalayan pteroids. Ph. D. thesis, Panjab University, Chandigarh, India SHORTER NOTES 25 SHORTER NOTES A NEW LOCALITY FOR LYCOPODIUM SERRATUM IN MEXICO.—Until recently L. serratum had been collected only once in Mexico. Liebmann found it in the State of Oaxaca during the last century and published it as a new species, L. sargassifolium. His species has proven to be the same as L. serratum, which is distributed primarily in the Old World: temperate Asia, the Zonda Islands, New Caledonia, and Hawaii. It is scarcely represented in the New World: Cuba, Hispaniola, and Mexico. This is another example of an Asian-American disjunct distribution; the major range of L. serratum and the entire ranges of the species allied to it are in the Old World. The second known Mexican collection of L. serratum was made by A. J. Sharp and Blanca Pérez Garcia in June 1973 at a locality 6 km southwest of Tianguis- tengo, Municipio of Zacualtipan, in the State of Hidalgo, at an altitude of 2000 m. The plants were found in a subdeciduous forest with Liquidambar styraciflua, Ternstroemia latifolia, Alchornea latifolia, Alnus, and Quercus. The specimens of this collection were found in two small patches about 500 m apart, both near a small stream. Because of the dichotomous branching and the prostrate habit of the older stems, the plants have a circular outline; four series of dichotomies can be noted in the larger specimens. The leaves are in several spiral rows and charac- teristically have their margins irregularly toothed. The sporophylls are similar to the sterile leaves or are just a little smaller. Although strobili are not formed, there is a slight concentration of sporophylls along the stem.—Ramon Riba and Blanca Pérez. Garcia, Universidad Auténoma Metropolitana -Iztapalapa, Apartado Postal 55-535, México, D. F., México, and Martha Pérez Garcia, Instituto de Biologia-Botdnica, Universidad Nacional Auténoma de Mexico, Apartado Postal 70-233, México, D. F., México. NATURALIZATION OF CYRTOMIUM FORTUNE! IN NORTH AMERI- CA.—The fern Cyrtomium fortunei Smith, a native of southeastern China, has become established in Charleston, South Carolina as an apparent escape from cultivation. This record presently represents the only known naturalization rane species in North America. Specimens were first noticed in 1973, growing In as- sociation with Cyrtomium falcatum (L. fil.) Presl, on a moist, north-facing brick wall in a cemetery in downtown Charleston. By 1975 the ferns had increased to a colony of about seventy plants, and a collection was made (MacDougal 184 and 186). The probable parent plant is growing nearby with nursery-raised C. fal- catum, and was evidently brought to the graveyard with them. | sega In the living state this species is easily distinguished from C. faleatum, ae it resembles, by the dull green, not glossy, upper side of the fronds. The pinnae wt more numerous and smaller, lanceolate or oblong, 5-8 x 1-2.5 cm, acuminate, and have finely serrate margins. 26 AMERICAN FERN JOURNAL: VOLUME 66 (1976) It is especially interesting to record this naturalization in view of the success that C. falcatum has had in the Charleston area following its naturalization in the early 1940’s; see K. W. Hunt’s ‘‘Ferns of the vicinity of Charleston, S.C.” (Charleston Museum Leaflet No. 17). Voucher specimens of C. fortunei have been deposited in several herbaria, including those of the New York Botanical Garden, The Smithsonian Institution, and The Charleston Museum. I wish to thank Dr. David B. Lellinger and Albert Giraldi of The U.S. National Herbarium for identifying this fern——John M. MacDougal, 1 New Town Lane, Charleston, SC 29407. FORKED FRONDS IN ASPLENIUM RHIZOPHYLLUM.—Forking of fern fronds is a common occurrence, but has not been reported for Asplenium rhizophyllum L. Buzz Darby, an amateur naturalist of Springfield, Missouri, has discovered several colonies of A. rhizophyllum in Newton County, Arkansas, in which some plants had a forked blade. The identity of two such colonies growing IG. I. Asplenium rhizophyllum with blade forked at base (Key 1238, SMS). FIG. 2. A. rhizophyllum with blade forked (Key 1239, SMS) along a creek bank on moss-covered sandstone boulders has been confirmed and vouchers made (Key 1238, 1239, SMS). Colonies with similarly forked blades have been found on limestone outcroppings in a mesophytic forest in Montauk State Park, Dent County, Missouri (Maupin 1145, SMS) and in a similar habitat near Eminence in Shannon County, Missouri, (Key 1282, SMS). My son has discovered a fourth location on limestone near the opening of the Ozark Under- ground Laboratory Cave in the xerophytic, forested hills of Taney County, Mis- souri. SHORTER NOTES 27 In some plants, the stipe of the forked frond is unusually long (8 cm) and is triangular (1.5-2 mm on each side). Forking occurs 5 mm above the cordate base of the doubled, sorus-bearing blade (Fig. 1). The Dent County fronds are similar to this, although several immature plants have small blades with shallowly or deeply notched, rounded tips, rather than attenuate tips. Another frond is forked 3 cm above the base of the blade, and is immature, lacking sori (Fig. 2). Because only a single forked frond is present on most plants, it is unlikely that forking is genetically controlled. Trauma to the tips of young fronds, probably by insects, seems to be the most likely cause, but against this is the Maupin specimen in which the forking occurs at the upper end of the stipe, rather than in the blade itself. Studies are underway to determine the influence of blade-tip trauma on the development of immature fronds. The discovery of forked A. rhizophyllum fronds in distinctly different habitats and localities makes it quite likely that forking is of more than occasional occurrence. I suspect that close observation would reveal more plants with forked fronds, which are almost impossible to see at a glance and often require several minutes of search, even when one knows the exact area to be searched.—James S. Key, Department of Life Sciences, Southwest Missouri State University, Springfield, MO 65802. A NOTE ON THE YOUNG FRONDS OF OPHIOGLOSSUM PALMATUM.—In his excellent account of the natural history of Ophioglossum palmatum L. in South Florida, Mesler (Amer. Fern J, 65: 33-39. 1974) writes: “The leaves of O. palmatum have been described as palmately or dichotomously lobed, but no ontogenetic evidence has been presented to support either view’’. I presume he refers to the vernating leaves, as adult forms are obviously lobed and often dichotomous in segmentation. I have grown several plants of O. palmatum col- lected by Wagner and Gomez in 1971 north of San José near Vara Blanca, Province of Heredia, Costa Rica, and have been able to observe the emergence of several generations of fronds under greenhouse conditions. The leaves emerge and uncurl (they are not, like the rest of the Ophioglossaceae, circinnate, but erupt from the substratum almost always bent) and are dichotomously bilobed. Some- times, they are spathulate, with a rather truncate apex. The laminae are pinkish green, soft but already fleshy, small and quite out of proportion to the petiole, which is many times longer and robust. My specimens, after removal from the original habitat, were planted in well-drained pots, placed under 50% sunlight and at about 75% relative humidity, with temperatures that averaged 18°C. The fronds were able to grow laxly erect. I agree with Mesler that the establishment of subspecies or varieties based on blade morphology and size is not only inconven- ient, but, I think, quite improper. The variation in leaf size, shape, texture, -_ insertion of the sporophylls on the petioles and laminae, 1s tremendous.—Lu's Diego Gomez P., Herbario Nacional, Museo Nacional de Costa Rica, Apartado 749, San José, Costa Rica. 28 AMERICAN FERN JOURNAL: VOLUME 66 (1976) THE IDENTITY OF POLYPODIUM FURFURACEUM F. PINNATISEC- TUM.—From 1908 to 1910, Alexander Curt Brade and his brother Alfred col- lected ferns in Costa Rica. On April 10, 1908 they visited La Carpintera, a series of hills between the cities of San José and Cartago. There they gathered, among other things, a specimen determined by A. C. Brade as Polypodium furfuraceum Schlecht. & Cham. that was unusual in having 16-21 lower pairs of pinnae pin- natisect, rather than entire, as is usual in P. furfuraceum. Many years later A. C. Brade described this peculiar plant as P. furfuraceum f. pinnatisectum: ** Differt a forma typica pinnis ex parte pinnatisectis; pinnis 12-16 infimis pinnatisectis, utrinque cum 3-7 segmentis; segmentae usque ad 7 mm longae.’’ The type specimen label also reads in Brade’s hand ‘‘(? P. lindenianum Kze.),’ the name of yet another Polypodium with pinnatisect fronds. However, Brade’s specimen is in fact the natural hybrid P. friedrichsthalianum Kunze x furfuraceum. Both parents are very common in the type locality and vicinity, where natural hybrid, P. x aspidiolepis Baker (P. friedrichsthalianum x thys- sanolepis A. Braun), also occurs. The discovery of the hybrid nature of Brade’s plant requires a change in its nomenclatural status: Polypodium pinnatisectum (Brade) L. D. Gomez, comb. nov. Polypodium furfuraceum f. pinnatisectum Brade, Bradea 1: 16, f. 5. 1969, as ‘‘pinnatisecta.”’ TY PE: La Carpintera, Pcia. Cartago, Costa Rica, 1800 m, 10 Apr 1908, 4. & A. C. Brade 16 (HB not seen). In addition, I have seen one other specimen: Monte de la Cruz, Pcia. Heredia, Costa Rica, 1800 m, A. Jiménez 208 pro parte (CR 37368). In view of the apparent facility with which P. friedrichsthalianum hybridizes with other species of Polypodium, it seems necessary to determine whether P. lindenianum Kunze might also be a hybrid involving P. friedrichsthalianum.— Luis D. Gomez P.., Herbario Nacional, Museo Nacional de Costa Rica, Apartado 749, San José, Costa Rica. REVIEWS COMMON FERNS OF LUQUILLO FOREST, PUERTO RICO,” by Angela Kay Kepler, 1975. Spanish and English editions published by Inter American University Press, P.O. Box 1298, Hato Rey, PR 00919. $5.00 paperback, $15.00 hardcover.— Designed as a popular reference, this book covers most of the ferns likely to be found by those who hike the trails of Luquillo Forest and should be a useful field guide to them. Although the 8% by 11 inch size is not convenient for carrying into the field, the book is useful for quick identifications of ferns because each is illustrated and described clearly. There are keys utilizing soral characters and leaf shapes and a complete list of the ferns and fern-allies in the Luquillo Forest by the author and R. Woodbury. The explanation of terms and general introduction to ferns should be useful to those unfamiliar with ferns who wish to use the book. REVIEWS 29 Some aspects of the book are annoying to serious students of ferns. The author uses the unnecessary, rather amateurish terms fruitdot and fruitcover to refer to the sorus and indusium, in the descriptions the term ecology is used when habitat is meant, the illustrations are more artistic than scientific (although diagnostic, nonetheless), no index to species or genera is included, and the plants are listed in the distribution list by common names so that someone unfamiliar with the com- mon names adopted or invented by the author is at a disadvantage is finding plants quickly. Some statements (such as the ferns are not differentiated into species as juveniles) have to be reinterpreted by the reader (meaning one cannot identify juveniles to species). The scientific names are based on W. R. Maxon’s classifica- tion in *‘Scientific Survey of Porto Rico and the Virgin Islands*’ (1926) and are somewhat outdated, but are useful in further checking with Maxon’s book. The book includes a map of trails in the forest and a distribution list of the ferns and fern allies to be found along the trails. Considering the book as a whole, it should prove valuable to all visitors to Luquillo Forest and to pteridologists who wish to make convenient identifications of the common pteridophytes of that area.—J. E. Skog, Department of Biology, George Mason University, Fairfax, VA 22030. **TRICHOMANES,”’ Philip. J. Sci. 51: 119-280, pl. 1-61. 1933; *“HYMENOPHYLLUM,” Philip. J. Sci. 64: 1-188, pl. 1-89. 1937; “GENERA HYMENOPHYLLACEARUM,” Philip. J. Sci. 67: 1-110, pl. 1-11. 1938, by E. B. Copeland, reprinted 1975 in one volume by Otto Koeltz, P. O. Box 129, D-624 Koenigstein, W. Germany. DM200.—Of the major botanical monographs pub- lished in the pre-World War II Philippine Journal of Science, only two have not been or are not about to be superceded. The 1930's depression which reduced subscription sales, and the War in which all back issues were destroyed, combined to make their reprinting a necessity. Otto Koeltz has obliged with both Bartram's 1939 **Mosses of the Philippines’’ and Copeland’s three-part study of the Hymenophyllaceae. i Copeland’s trilogy is close to his final word on the subject. ‘*Trichomanes (1933) dealt with 113 Old World species including Cardiomanes, and ‘‘Hymenophyllum’”’ (1937) with 127 species; ** Genera Hymenophyllacearum (1938) apportioned these and many New World species into 33 genera with the contention that the two customary ones were polyphyletic. To these, Copeland added only a short paper with notes and novelties from New Guinea (Philip. J. Sci. 73: 457-469, pl. 1-4. 1941), a 34th genus based on a New Caledonian oddity (Gen. Fil. 36. 1947), a treatment of the Philippine species (Fern Fl. Philip. 1: 46-82. 1958), and minor scattered miscellany. Thus, since 1938 botanists have been able to choose from two generic treatments, both presented by a single author: a Hymenophyllaceae either with Over 30 genera, or with 2 main genera plus one to several isolated minigenera._ While Copeland’s taxonomy and nomenclature are subject to considerable ee sion, of major lasting value in his work are the numerous magnificent detaile illustrations, ‘‘Hymenophyllum’’ and ‘tGenera Hymenophyllacearum were 30 AMERICAN FERN JOURNAL: VOLUME 66 (1976) illustrated by the two Filipino artists, Leopoldo A. Alicbusan and Esteban P. Borbe, who worked with Copeland for 14 months in his private herbarium then located at Los Banos, Laguna, Luzon. The microscopic details were drawn with camera lucida from slides prepared by Copeland. Mr. Borbe has told me that Copeland demanded precise reproduction without stylizing, and eventually came to trust the skills of his artists so completely that he did not always compare the completed drawings with the specimens. Each drawing was made on separate paper; the final lay-out of the plates was Copeland’s. Alicbusan survived the infamous ** Death March”’ upon the fall of Bataan in 1942, and after the War made a career in the military, meeting a soldier’s death in 1954. Borbe is still working from retirement as an artist. The reprint is handsomely bound with good quality paper. The single volume is one-half the weight and thickness of the three originals if separately bound. Page size is reduced, but only by the eliminating of the unnecessarily large margins. Unlike those in the original, the plates are on both sides of the pages; reproduction is good. The single and very serious fault is the lack of an index. Each volume of the Philippine Journal of Science proper has an index although not including the plates (and plates are not referenced back to the text). The inexcusable failure to make an index to the reprint will waste a great deal of time for users. Copeland’s work on Hymenophyllaceae deals hardly at all with the three Euro- pean species or the ten temperate North American species. But it is absolutely indispensable for those with a broader interest in this principally tropical and south temperate family. And as the first modern effort to make sense out of the filmy ferns, it provides an overall perspective that is the requisite foundation for further work.—M. G. Price, University of the Philippines at Los Banos, College, Laguna 3720, Philippines. **“ENUMERATIO PTERIDOPHYTARUM JAPONICARUM, FILICALES”’ by Toshiyuki Nakaike. xii + 375 pp. 1975. University of Tokyo Press, 7-3-1 Hongo, Bunkyo-ku, Tokyo, 113-91, Japan. Distributed in the U.S. and Canada by Inter- national Scholarly Book Services, P.O. Box 4347, Portland, OR 97208. $29.50/ £ 12.90/8,000 yen.—This is a disappointing book, mostly because of what is not included. It is basically what the title states: an enumeration of Japanese Filicales, some 800 taxa. There are no keys, no descriptions, no discussions, no illustrations, and, except of genera, no citation of types. We are given the ac- cepted name, homotypic and heterotypic synonyms, and chresonyms [a useful term for sensu names coined by H. M. and Rozella B. Smith (Syst. Zool. 21: 445. 1972)], all with exhaustive literature citations to floras, checklists, and other taxonomic notes that indicate how these taxa have been treated historically. In addition, there is a brief statement of distribution and the Japanese common name. Essentially, then, we have one man’s statement at a point in time with regard to the Classification of Japanese ferns—the author’s stated objective. The effort involved in compiling the synonymy and references was no doubt staggering. For me, though, a classification presented in this manner does nothing to provoke or promote additional research. It implies something immutable about taxonomy, REVIEWS 31 when in fact it is likely that this classification will be supplanted by a new one ina few years. There are over 75 ‘‘new combinations”’ in the book, many of them involving the impermissible taxonomic category ‘‘monstrosity.”” Nakaike elevates two in- frageneric names to generic rank, Lacosteopsis and Nothoperanema. However, the latter generic name, as well as two species combinations made by Nakaike (V. hendersonii and N. shikokiana), should be attributed to Ching (Acta Phytotax. Sin. 11: 25. 1966).—Alan R. Smith, Herbarium, Department of Botany, Univer- sity of California, Berkeley, CA 94720 SERVICE ae ae OWNERSHIP my 112, 1970 See 45, Title 39. United States Code | 1. TITLE OF PUBLICATION 2. DATE OF FIL | AMERICAN FERN JOURNAL | September 25, 1975. SUBSCRIPTION PAIGE wobacr. 3. FREQUENCY OF ISSUE [ ly pe mem. $8.50 4 | v8. mat'l, Herbariim, Suitheontan Inst cievelon, Washington, DC 20560 5. LOCATION OF THE HEADQUARTERS OA GENERAL BUSINESS OFFICES OF THE PUBL HERS (Not pri U. S. Nat'l. Barber Saithsonian Institution ton 20560 6 MES AND ADDRESSES OF PUBLISHER, EDITOR, AND MANAGING EDITOR PUBLISHER |N d addre hemmcmeprnah FERN socterr, INC., Smithsonian Inst., Washington, DC 20560 ss EDITOR (Na: md addr _ Dr. David B.. LelLinger, Smithsonian Institution, Washington, DC 20560 7. OWNER (1 e dividual must be given.) fae aes va niee NAME DORESS AMERICAN FERN SOCIETY, INC Smithsonian Inst., De __ 20560 8. Sas NAME. ADDRESS _ None t status of This Have not changed “. tee blisher atus tor Federal |) during preceding pr s oe 12 months G NO. COPIE: ACTUAL NUMBER OF COPIES Of EACH ISS BINGLE ISSUE Aaoenyy geM ponces ISSUE CURING. W — | CEDING 12 EST_TO FILING D A TOTAL NO COPIES PRINTED (Net Press Run | 4925 +. 2000 PAID CIREULATION | SALES THR! DEALERS AND CARRIERS, STREET 0 o as telah VENOORS ae COUNTS EAL es + 2. MAIL SUBSCRIPTION 1355 1567 ©. TOTAL PAID CIRCULATION 1355 3567 o REE DISTRIBUTION BY MAIL, CARRIER OR OTHER MEANS 1 SAMPLES COMPLIMENTARY, AND OTHER FREE COPIES 1 + e 1356 1568 ee F> COPIES NOT DISTRIBUTED 632 ¥. OFFICE USE. | igh | 369 } — 2. RETURNS FROM NEWS AGENTS | 0 0 } G. TOTAL (Sum of FE & 4 | 1725 l vertity th bh and complete 32 AMERICAN FERN JOURNAL: VOLUME 66 (1976) Suggestions to Contributors Manuscripts (original plus one copy for reviewers) should follow recent Journal style and should be prepared in accordance with the AIBS (1964) **Style Manual for Biological Journal’ or the AIBS (1972) **CBE Style Manual.”’ For major articles with more than one literature reference, use the me and year’’ system for bibliographic references and the American Standards Association list of bibliographic abbreviations (AIBS, 1964, pp. 82-87), which may be supplemented by the list of Schwarten and Rickett (1958), or by those in the comprehensive ‘‘Botanico-Periodicum-Huntianum”’ (Lawrence et al., 1968). Put a single reference in a footnote or in the text. For shorter notes and reviews, put all references in parentheses in the text. Abbreviations of the names of herbaria should follow the list of Holmgren and Keuken (1974). Group all footnotes, tables, and figure legends at the end of the manuscript. Art work must be ‘‘camera ready”’; paste-ups are not permitted in tone copy (photographs), but if carefully done are acceptable in line copy (drawings). Scales should be included on figures and plates, rather than by indicating magnification in legends Manuscripts should have ample margins and should be typed double-spaced throughout, including the title, bibliography, and footnotes. Footnotes and tabular matter should be kept to a minimum. Reports of chromosome numbers will not be published unless documents by voucher specimens deposited in some herbarium. Reprints should be ordered when galley proof is returned to the editor. An order blank will be included with galley proof. The payment or non-payment of page charges by authors’ institutions or grants will affect neither the acceptability of manuscripts for publication nor the date of publication. LITERATURE CITED AMERICAN INSTITUTE OF BIOLOGICAL SCIENCES, Committee on Form and Style of the onference of Biological Editors. 1964. Style Manual for Biological Journals, ed. 2. Washing- ton, D.C eat on Form and Style of the Conference of Biological Editors. 1972. CBE Style ual, ed. 3. Washington, DC. aol cee PATRICIA K. and W. KEUKEN. 1972. Index Herbariorum. I. The Herbaria of the World, ed. 6. Reg. Veg. 92: 1-397. LAWRENCE, G. H. M. et al. 1968. Botanico-Periodicum-Huntianum. Hunt Botanical Library, Pittsburgh. SCHWARTEN, LAZELLA, and H. W. RICKETT. 1958. Abbreviations of titles of serials cited by botanists. Bull. Torrey Bot. Club 85: 277-300. See also the supplement in Bull. Torrey Bot. Club 88: 1-10. 1961. Exotic and Hardy Ferns ae BOLDUC’S GREENHILL NURSERY 2131 Vallejo Street St. Helena, California 94574 Open Saturdays and Sundays from 10 A.M. to 4 P.M. and by appointment Send a self-addressed, stamped envelope for our list Phone 963-2998—Area Code 707 Mail orders accepted Rs ae Oe ENERGY-SAVING DOUBLE’ WALL GREENHOUSES FROM JUST ‘499 - Now you can grow economically year ‘round in the only greenhouse that solves the energy-cost problem. Lab- oratory tested in Vermont, rigid double- pane thermal walls are 244% more heat efficient, shatterproof, and main- tenance free. Free-standing and lean- to models, no foundations required. Factory direct only, 5 year total house warranty. Call or Write for Intormation P.O. Box 2235, Dept. AF Grand Central Station New York, N.Y. 10017 (212) 686-0173 GROW FERNS! with Barbara Joe Hoshizaki’s FERN GROWERS MANUAL John Canaday says: ‘The perfect guide for serious amateurs, and a combination of stimulant and chastener for the invincible windowsill horticulturist.” The Miami Herald calls it “the first solid guide to contemporary fern growing.” Indoors, outdoors— everything the amateur or expert needs to know. With 315 illustrations. $15 at bookstores. Or send check to Alfred A. Knopf, Dept. B, 201 East 50th Street, N.Y. 10022. (Include sales tax where applicable) rs. TRIARCH 1976 - Our 50th year of slide manufacture and service to botanists. Your purchases have made our 50 years of exis- tence possible. To satisfy your continued need for quality prepared slides, address your requests for catalog 17 or custom preparations to: TRIARCH INCORPORATED P.O. Box 98 Ripon, Wisconsin 54971 AMERICAN FERN JOURNAL Volume 66 Number 2 April-June, 1976 A TRIBUTE TO EDGAR THEODORE WHERRY Edgar Wherry in Pennsylvania Perry Creek, Washington, a Fern-watcher’s Eldorado Origin of the Pteridophyte Flora of the Bahamas, Caicos and Turks Islands Spore Retention and Release from Overwintering Fern Fronds The Distribution and Abundance of Dryopteris reey Cystopteris bulbifera in the Southwestern United States Variation in North American Asplenium platyneuro W. JOHN M. FOGG, JR. 33 A. R. KRUCKEBERG 39 DONOVAN S. CORRELL 46 DONALD R. FARRAR 49 JAMES D. MONTGOMERY 53 TIMOTHY REEVES 60 CARL TAYLOR, ROBERT H. MOHLENBROCK, and FREDDA J. BURTON 63 The Distribution of Dryopteris spinulosa and its Relatives in Eastern Canada Ferns: Potential In-situ Bioassay Systems Aquatic-borne DONALD M. BRITTON’ 69 ms for Mutagens EDWARD J. KLEKOWSKI, JR. and DAVID M. POPPEL 75 Review Missour! SorTanicab Jit 22 1976 GARDEN LIBRARY Council for 1976 DAVID W. BIERHORST, Dept. of Botany, University of Massachusetts, Amherst, Mass. 01002. President RICHARD L. HAUKE, Dept. of Botany, University of Rhode Island, Kingston, R.I. 02881 Vice-President TERRY R. WEBSTER, Dept. of Botany, University of Connecticut, Storrs, Conn. 06268. ecretary JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916. Treasurer DAVID B. LELLINGER, Smithsonian Institution, Washington, D.C. 20560. Editor-in-Chief JOHN T. MICKEL, New York Botanical Garden, Bronx, N.Y. 10458. Newsletter Editor American Fern Journal EDITOR-IN-CHIEF DAVID B. LELLINGER Smithsonian Institution, Washington, D. C. 20560 ASSOCIATE EDITORS DAVID W. BIERHORST ..Dept. of Botany, University of Massachusetts, Amherst, Mass. 01002 GERALD J. GASTONY ....Dept. of Plant eet Indiana University, Bloomington, Ind. 47401 JOHN T. MICKEI w York Botanical Garden, Bronx, New York 10458 The ** American Fern Journal”’ is an illustrated quarterly devoted to the general study of ferns. It is owned by the American Fern Society, and published at the Smithsonian Institution, Washington, DC 20560. Second-class postage paid at Washington. Matter for publication and claims for missing issues (made within six months of the date of issue) should be addressed to the Editor-in-Chief. Changes of address, dues, and applications for membership should be sent to Dr. Terry W. Lucansky, Dept. of Botany, University of Florida, Gainesville, Florida 32601. Orders for back issues should be addressed to the Treasurer. General inquiries concerning ferns should be addressed to the Secretary. Subscriptions $8.50 gross, $8.00 net (agency fee $0.50); sent free to members of the American Fern Society (annual dues, $5.00; sustaining membership, $10.00; life membership, $100.00). Extracted offprints, if ordered in advance, will be furnished to authors at cost, plus postage. Back volumes $5.00 to $6.25 each; single back numbers of 64 pages or igo $1.25; 65-80 pages, $2.00 each; over 80 pages, $2.50 each, plus shipping. Ten percent discount on orders of six volumes or more, postage additional. Library and Herbarium Dr. W. H. Wagner, Jr., Department of Botany, University of Michigan, Ann Arbor, Michigan 48104, is Librarian and Curator. Members may borrow books and specimens at any time, the borrower paying all shipping costs. Newsletter Dr. John T. Mickel, New York Botanical Garden, Bronx Park, Bronx, New York 10458, is editor of the mati’ ‘Fiddlehead Forum.”’ The editor welcomes contributions from members and non- ncluding miscellaneous notes, offers to exchange or purchase materials, personalia, hor- ticultural ae and reviews of non-technical books on fern Spore Exchange Mr. Neill D. Hall, 1230 Northeast 88th a Seattle, Washington 98115, is Director. Spores exchanged and collection lists sent on reque Cut At +h sits and paisa: Ses kone L is #bL ant ected in ferns. Herbarium specimens, ‘bownical books, back i issues of the Journal, and cash or other gifts are always welcomed, and are tax-deductible. Inquiries should be addressed to the Secretary. idigeds abiaaadni asada AMERICAN FERN JOURNAL: VOLUME 66 NUMBER 2 (1976) 33 Edgar Wherry in Pennsylvania JOHN M. FOGG, JR.* Edgar Theodore Wherry was born in Philadelphia on September 10, 1885, and received his education at Friends Central School and the University of Pennsyl- vania. After teaching at Lehigh University from 1908 to 1913 and working in Washington, D.C. for seventeen years, he returned in 1930 to the city of his birth and has resided there ever since Long before he returned to Philadelphia to live, local botanists became ac- quainted with Dr. Wherry. Since his family resided in this city, it was his habit to spend the Christmas holidays with them. This led to his being invited to speak at the December meeting of the Philadelphia Botanical Club, an organization which had been founded in 1891. The earliest of his lectures which I can remember was in 1921, when his topic was *‘Our Native Plants and their Soil Preferences.’’ From that time on for many years, with only a few exceptions, his December presentation was an annual ‘feature of the Club’s programs, and we were favored with lectures on ferns, orchids, pitcher plants, heaths and heathers, and of course, Phlox at the time when our speaker was preparing his monograph on that genus. All of his talks were illustrated with handcolored slides. In 1930 Dr. Wherry joined the faculty of the Department of Botany of the University of Pennsylvania, where he taught until his retirement in 1955. He had already made substantial contributions to our knowledge of soils, had perfected a colorimetric method for determining soil pH, had investigated the remarkable stand of box huckleberry (Gaylussacia brachycera) in Perry County, Pa., and had published an account of the ‘‘Wild Flowers of Mount Desert Island, Maine.”’ In 1918, while still employed by the Bureau of Chemistry in the U.S. Depart- ment of Agriculture, Wherry became a member of the American Fern Society. He was president of that organization from 1934 through 1938 and was the author of many articles in the ‘‘American Fern Journal.’’ He assigned the royalties from his popular ‘‘Fern Guide’”’ to the Society. This is entirely typical of the selflessness of the man. Today he is deservedly an Honorary Member of the Society. In 1932 the University of Pennsylvania inherited the property which became the Morris Arboretum. Dr. Rodney H. True, who was then Chairman of the Depart- ment of Botany, assigned four members of his staff on a part-time basis to ad- minister the project, and Wherry was appointed ecologist. One of his first tasks was to conduct a detailed soil survey of the grounds. This revealed that within some 175 acres there was a wide diversity of soil types. The ridge which traverses the property from east to west is composed of quartzite which, being a metamorphosed sandstone, weathers slowly to produce an acid soil. On the south slope this gives way to circum-neutral soils derived from schistose rocks, while northward the underlying formation is Cambro-Ordovician limestone, a distinctly alkaline soil. With this information in hand, it was possible to develop planting *Arboretum of the Barnes Foundation, Merion, PA 19066. olume 66, number 1, of the JOURNAL was issued April 1, 1976. 34 AMERICAN FERN JOURNAL: VOLUME 66 (1976) plans in a scientific manner, making certain that each family or genus was estab- lished on soils with which it was compatible. In 1934 the Bowman’s Hill State Wild Preserve was established with the aid of the Works Progress Administration. This preserve, which is located near New Hope in Bucks County, Pa., is dedicated to the growing and preservation of plants native to the Commonwealth. With Wherry’s deep-seated interest in wild flowers it was inevitable that he should be attracted to this project, and in 1935 he became one of the Founders. For a time he was Chairman of the Preserve and has been one of its botanical advisors for 41 years. The Wherry Fern Trail was one of the first trails to be developed at the Preserve and appropriately commemorates Dr. Wherry’s dedication to that group of plants. In the mid 1930’s, work was begun in earnest on a state Flora at the University of Pennsylvania. The Department of Botany had purchased a twenty-passenger bus to transport students between the campus and the arboretum, and each spring, as soon as Classes were over, we rounded up half a dozen or so of our abler students, loaded up the bus with presses, driers, and collecting papers, and set off on a week’s botanizing trip. During several summers we moved from east to west across the northern tier of Pennsylvania counties, combing every available habitat. We hoped eventually to cover the entire state. Frequently we returned to the area later in the season to collect fruiting material. From each of these trips thousands of herbarium specimens were brought back to be incorporated into our records and finally to appear as dots on our outline maps of the state. Edgar Wherry participated in many of these trips, and his knowledge of geology, geog- raphy, and the more interesting wild flowers made him a valuable member of the party. Enthusiastic field man that he was (and at the age of 90-plus still is), he always welcomed the prospect of a collecting expedition. I recall with much pleasure a foray on which I accompanied him across Pennsylvania in the fall of 1940. We drove through Cameron, Elk, Jefferson, and Armstrong counties, stopping fre- quently to collect. After visiting the extreme southwestern corner of the state, we came back through Fayette County, stopping near Elliottsville to collect Clethra acuminata at what was then its only known station in Pennsylvania. We coul almost look across the state line into Maryland, but there was no doubt about it—the Clethra was definitely in Pennsylvania, although it is not so recognized in the eighth edition of **Gray’s Manual.”’ Next we visited Ohiopyle, in the same county, where several genera such as Boykinia and Marshallia are at or near the northern limits of their range. Continu- ing eastward, we stopped at a locality southwest of Reels Corners to collect that unique sedge Cymophyllus fraseri at one of its few known stations in the state. Each of us brought back from this brief trip about 400 numbers (many of them in replicate), which added a significant number of records to our study. On another occasion a friend of mine offered us the use of his hunting lodge near Bradford in McKean County in the extreme northern portion of the state. My wife and I, accompanied by Wherry and Joseph W. Adams, who was then on the staff of the Morris Arboretum, spent five days in a lovely spot scouring the coun- J. M. FOGG, JR.: EDGAR WHERRY IN PENNSYLVANIA 35 tryside, with the result that we brought back nearly a thousand numbers from an area that was almost entirely unknown botanically. I mention these two trips merely as an indication of Edgar Wherry’s active interest in the Flora project and his willingness to drop anything he might be doing in order to participate in the field work. In 1941 I accepted an appointment as Dean of the College of Arts and Sciences at the University, having been assured that my duties would not seriously affect my work on the Flora. Of course, no one at that time knew that soon the country would be at war and that afterward there would be a lengthy period of readjust- ment affecting all educational institutions. The result was that for twelve years I was able to devote very little time to the project. It might have been necessary to abandon it altogether, had not Wherry stepped in and assumed the responsibility for continuing it. Fortunately, also, at this juncture I was able to enlist the cooperation of my friend and colleague, Dr. Herbert A. Wahl, Professor of Botany at Pennsylvania State University. On'three separate occasions during my involvement in adminis- trative duties, Herb was able to obtain a year’s leave of absence from Penn State and to come to Philadelphia to work with Wherry on the Flora. I joined them whenever possible, but my appearances were few and far between. The various techniques adopted for handling our multitudinous records have been described in detail elsewhere, and need be mentioned only briefly here. Most of them were devised by the late Dr. J. R. Schramm, formerly Chairman of the Department of Botany, to whom tremendous credit is due for his support of the Flora project. Once the thousands of specimens of Pennsylvania plants we collected had been mounted, it was necessary for their identifications to be authenticated before they were ready to be recorded and mapped. In this operation Dr. Wherry assumed responsibility for such groups as the pteridophytes, orchids, Ericaceae, Polemoniaceae, and a few others. Herb Wahl, a recognized authority on Carex (our state’s largest genus), worked with the entire Cyperaceae, as well as the very difficult genus Potamogeton and many of the apopetalous families. I took over Juncus and most of the Sympetalae, and in this manner there gradually evolved a division of labor, with Wherry taking more and more groups as time went by and my association with the project decreased. After the name on a given specimen was authenticated, it was entered on a master record card that listed the state’s 67 counties. The exact locality at which each specimen was collected was then entered, in carbon-base ink, under its appropriate county, together with name of the collector, his number or date of collection, and a symbol indicating the herbarium in which the sheet had been examined. The last was thought to be of value since, in addition to incorporating about 100,000 specimens in our own herbarium, we attempted to record all of the Pennsylvania material in the Philadelphia Academy of Natural Sciences, the Car- negie Museum in Pittsburgh, the State Museum in Harrisburg, and the herbarium at Pennsylvania State University. Probably no other state Flora has ever been based on such a tremendous number of specimens. 36 AMERICAN FERN JOURNAL: VOLUME 66 (1976) The final step in the procedure was to place on an outline map of the state for each species a dot representing each entry on the record cards. Accuracy is of great importance because the state includes diverse physiographic provinces, such as the coastal plain, the piedmont, the valley and ridge province, and the Appalachian plateau. The state is also traversed by the terminal moraines of two recent glaciations. From almost the very beginning of the undertaking this has been Edgar Wherry’s sole responsibility. There is probably not a town, village, or hamlet in the Commonwealth of Pennsylvania for which he does not know the exact location. Needless to say, we have disregarded all specimens with vague data and have recorded only those giving precise information. Of the more than 3000 species of higher plants which occur in Pennsylvania, about 800 are introduced, either from abroad or from other sections of the United States. Of those which are native, the majority are widespread, occurring in every or almost every county (Fig. /). Others have a more limited distribution, being either predominantly northern, or southern, or on the coastal plain, or compo- nents of the Ohio River vegetation. Still others might be found only on particular soils, such as limestone (Fig. 2), serpentine, or shale barrens. As Wherry’s care- fully placed dots began to fill in the map-cards, often several hundred for each species, these correlations became abundantly evident. Another fact which emerged was that in countless instances species were totally lacking from counties where they might be expected to be common. Wherry then began making ‘‘Wanted Lists’’ for each county. If he knew of an active collector in one of these counties, he would send him a list and urge him to collect. More often than not he would execute this commission himself, spending days in a given county to collect the plants which had been neglected by other botanists. Usually these were common plants, for all too frequently rare or spectacular species are collected at the expense of more familiar ones. Another by-product of the maps is species lists for given counties, which Wherry has compiled. Several lists have already been published in ‘‘Bartonia,”’ the official journal of the Philadelphia Botanical Club. They are the basic bricks of which state and regional Floras are constructed. It was the original intention that the Flora of Pennsylvania should be biologi- cally oriented and contain diagnostic keys, brief descriptions, and interpretive information concerning distribution, which would elevate it above the level of a mere checklist. To that end, Edgar Wherry, Herb Wahl, and I prepared hundreds of pages of manuscript. Unfortunately, Wahl’s increasing preoccupation in monographing Chenopodium, his recent untimely death, and my increasing in- volvement in administrative duties reduced this objective to a rather forlorn hope. Also, the cost of publishing such a work, plus our many hundreds of range maps, would have been almost prohibitive. It is a pleasure, however, to report that our manuscript, incomplete though it is, has been turned over to Dr. Carl Keener of the Department of Botany at Pennsy!- vania State University. Carl feels that is may be possible to put it in shape for publication, and if he does, Wherry’s treatments of the ferns, orchids, phlox, etc. will constitute a substantial contribution. J. M. FOGG, JR.: EDGAR WHERRY IN PENNSYLVANIA 37 As matters stand at present, only the maps, in the form of an ‘‘Atlas,”’ are scheduled for publication at an early date. It is unlikely that anyone examining these maps will gain any real insight into the forty years of field work and her- barium study which underly them, but I hope it has been made clear that to a large degree their publication is due to the indomitable energy of one very versatile and energetic individual. i eames tiie shy ae QOg JERSEYAN MOR . - VV ¥ APPALACHIAN Gaaaian FRONT F AAA MOUNTAIN FRONT nt FALL LINE ‘MULL WISCONSIN MORAINE 222 eNenee OR ADTE vv Vv APPALACHIAN PLATEAU FRONT i AAA eeptiechare Adiga | evevaseet FIG. 1. Pennsylvania range of Polystichum acrostichoides. FIG. 2 . Pennsylvania range of Asplenium cryptolepis. 38 AMERICAN FERN JOURNAL: VOLUME 66 (1976) In the meantime, from 1941 to 1950 and again from 1953 to 1957, Dr. Wherry served as a member of the faculty of the School of Botany, Horticulture and Landscape Architecture of the Arboretum of the Barnes Foundation. This school had been established in 1940 by Mrs. Laura L. Barnes, Director of the Ar- boretum, and the courses offered at her invitation by Wherry during these two intervals included geology, soils, ecology, and plant physiology. His emphasis at all times was on the out-of-doors, where the study of botany truly belongs. In 1972 the officers of the Delaware Valley Chapter of the American Rock Garden Society approached the Barnes Arboretum with a proposal to establish a small rock garden in honor of Dr. Wherry. He had been editor of their quarterly publication for several years and is today Editor Emeritus. The chapter proposed to construct this garden if the arboretum would provide space for it. Today there exists on a south-facing slope a plot 75 x 10 feet named the Edgar T. Wherry Memorial Garden. In this garden grow only plants which Wherry himself has selected. They are either species which have been named for him (such as Silene wherryi and Tiarella wherryi) or those with which he has had some intimate association, either as discoverer, introducer into cultivation, or author, as in several species of Phlox. Another interesting feature of this garden is its ecological character. The soil at the western end is derived from sandstone and is therefore acid. This is followed by a section of flaky shale, which produces a neutral reaction. The eastern end is rendered circum-neutral by the addition of limestone chips. Needless to say, this dramatic demonstration of the correlation between plant species and soil types is of considerable interest to both students and visitors to the arboretum. At least one day a week throughout the growing season, Edgar Wherry may be found in his garden, weeding, mapping, or planting specimens which have been sent to him by friends from many sections of the country. Mrs. Barnes had always been interested in hardy ferns, and in the two acres of native woodland which occupy the southwestern corner of the arboretum had assembled a collection of about a hundred species, including some rather rare and interesting exotics, such as Dryopteris erythrosora, Arachniodes standishii and Osmunda japonica. _ During the summer of 1975 Dr. Wherry suggested that a fern trail be established in these woods in honor of Mrs. Barnes. He personally checked the identifications of all of the species, new plastic labels giving the botanical and common names were made, directional arrows were installed, and the Laura L. Barnes Fern Dell became a reality. It has already been visited by the newly formed Delaware Valley Fern Society, of which Edgar Wherry is an honored member, and we are confident that it will prove a valuable teaching adjunct as well as a feature of interest to others who visit the arboretum. Botanists here and elsewhere—especially members of the American Fern Society—may well be grateful for the “‘return of the native’’ to the state and city of his birth. AMERICAN FERN JOURNAL: VOLUME 66 NUMBER 2 (1976) 39 Perry Creek, Washington, a Fern-watcher’s Eldorado A. R. KRUCKEBERG*! The vascular plant flora in the major drainage systems of western Washington is highly predictable from one river valley to the next. The typical coniferous forest plants reoccur throughout the region. This expectation holds as well for the ferns and fern-allies in the cismontane Pacific Northwest. The same species, Bracken (Pteridium aquilinum var. pubescens), Sword Fern (Polystichum munitum), Lady Fern (Athyrium filix-femina), Oak Fern (Gymnocarpium dryopteris), and Deer Fern (Blechnum spicant), can be expected in the usual range of forest com- munities. But the botanist-naturalist is in for a big surprise along a two-mile segment of the Perry Creek trail in Snohomish County, Washington, about 40 miles northwest of Seattle. No fewer than 26 species of ferns and fern-allies are known from this remarkable locality. It is a truly exceptional habitat well within the lowland coniferous forest biome. The first detailed collecting in Perry Creek was that of Mr. John W. Thompson ‘in the 1930’s. Thompson, then a high school biology teacher in Seattle, had a keen eye for the unusual. His collections of Pacific Northwest plants turned up many a novelty. A complete set of his collections was acquired by the University of Washington Herbarium, where Thompson served as assistant curator from 1943 to 1960 Following Thompson’s discovery of the diverse ferns of Perry Creek, the local- ity has been visited frequently by botanists and amateur naturalists. In 1963, Dr. Warren H. Wagner, Jr. visited Perry Creek in the company of the author. Wagner’s interest in the locality was whetted by the earlier Thompson finds— seven species of Botrychium and the sympatric occurrence of three Polystichum species. It was on this foray that Wagner found the hybrid between P. andersonii and P. munitum (Wagner, 1973). The richness of the Perry Creek fern flora was revealed to other pteridologists in 1969. A trip to the area, led by Drs. Wagner and T. M. C. Taylor, was made during the 12th International Botanical Congress, held that year in Seattle. Members of the foray were able to verify the occurrence of many pteridophytes that Thompson had first found. PERRY CREEK FERNS The following paragraphs list the pteridophytes that occur along the first two miles of the Perry Creek Trail. This listing is compiled from observations and herbarium records (WTU) made over the years. The most recent corroboration of this list was made on September 11, 1975 by members of the Fern Study Group of the Northwest Ornamental Horticultural Society and the author. Although most of species listed are frequent to common in western Washington, it is exceptional to find them so closely associated in a single, rather small natural area. The occurrence of Polystichum andersonii, its hybrid with P. munitum, P. *Department of Botany, University of ear aay Seattle, WA 98195. ‘Tl am indebted to W. H. Wagner, Jr. and T. C. Taylor for their nue counsel oe the years on the ferns of Perry Creek. Two si of ie Fern Study Group (Northwest Ornamental Horticul- tural Society), Mareen S. Kruckeberg and Sue Olsen, have vigor helpful in T thie field corey. Thanks are due David Wagner for data from this recent forays in the a 40 AMERICAN FERN JOURNAL: VOLUME 66 (1976) lonchitis, Asplenium trichomanes, Dryopteris filix-mas, Cryptogramma crispa, three Lycopodium species, and seven Botrychium species are of special interest. Adiantum pedatum L. is frequent and often gregarious on lush, shady talus under hardwoods. Asplenium trichomanes L., so common along the Perry Creek Trail on wet talus mostly under hardwoods, is infrequent and local elsewhere. At Perry Creek it is nearly as abundant as Cryptogramma crispa, a fern found everywhere on the more open but moist talus. Athyrium filix-femina (L.) Roth is very common throughout the area; in forb- rich talus habitats it is the dominant forb. Blechnum spicant (L.) J. Smith is restricted to regenerating, clear-cut Hemlock-Fir forest near the trail head and is rare on wet talus. Botrychium boreale Milde, B. dusenii (Christ) Alston, B. lanceolatum (Gmel.) Angst., B. lunaria var. onondagense (Underw.) House, B. multifidum (Gmel.) Trev., B. simplex E. Hitchc., and B. virginianum (L.) Swartz are all infrequent on wet, mossy talus in hardwood glades. None is at all common, although B. mul- tifidum is not infrequently encountered from the coast of Washington up to mid- montane altitudes. Some years it has not been possible to locate the other species, even after intensive searches above and below the trail. It was considered the high point of the 1969 pre-Congress foray for the party to have rediscovered all of the Grape Fern species located many years before by Thompson. Cryptogramma crispa (L.) R. Br. is common on wet talus, with or without hardwood cover. The plants are unusually large and lush, and well below their usual lower altitudinal limit. In the Cascade and Olympic Mountains, C. crispa normally occurs on subalpine talus and rock outcrops. Cystopteris fragilis (L.) Bernh. is rare under the cover of hardwoods on talus. Dryopteris filix-mas (L.) Schott is another fern of sporadic and infrequent oc- currence in the west (Reisender, 1974), and is not common at Perry Creek, having been collected by Mickel in 1969 and Denton in 1974. It was from a plant collected at Perry Creek (Snohomish, not Chelan County) that Reisender obtained the first chromosome count (2n=82, tetraploid) of any western member of the complex. Dryopteris assimilis S. Walker (=D. austriaca (Jacq.) Woynar) is infrequent, mostly in coniferous woods. Gymnocarpium dryopteris (L.) Newm. is occasional but gregarious in conifer- ous woods and under hardwoods on talus. Lycopodium clavatum L. is infrequent in regenerating, clear-cut forests, with Deer Fern. Lycopodium miyoshianum Makino is present at Perry Creek, accord- ing to Mr. Joseph Beitel (pers. comm.). It occurs on wet, moss-covered talus in the shade of Vine Maples. Lycopodium selago subsp. patens (Beauv.) Calder & Taylor, although frequent at Perry Creek in habitats like those of L. miyoshianum, seldom makes more than a sporadic appearance in the Pacific Silver Fir zone elsewhere in our area. Polypodium glycyrrhiza D. C. Eaton occurs mostly on trunks and limbs of Maples. Polypodium hesperium Maxon (or possibly P. montense Lang) grows on rocks in crevices, and is infrequent. A. R. KRUCKEBERG: FERNS OF PERRY CREEK, WASHINGTON 4] Polystichum andersonii Hopkins is frequent under hardwoods on moist talus slopes. It is sporadic and infrequent in the Pacific Northwest; its few localities are widely separated throughout its range from northwestern Oregon to southeastern Alaska and east to Montana. The hybrid of P. andersonii with P. munitum is sporadic and rare, and usually occurs with both parents. It has been found on Mt Hood, Oregon, and on Mt. Rainier, Washington, according to David Wagner (pers. comm.). Polystichum lonchitis (L.) Roth typically is a fern of high, wooded to open talus in the fringes of the subalpine forest, the Mountain Hemlock Zone of Franklin and Dyrness (1973). The Perry Creek station is certainly the lowest in elevation for this fern in Washington. It is intermittent mostly under hardwoods on talus, but also occurs on open talus slopes. Obvious hybrids involving P. lonchitis with P. munitum or P. andersonii have not been discovered, despite attempts to find them. Polystichum munitum (Kaulf.) Pres] is locally abundant, elsewhere frequent, mostly in the shade of hardwoods or conifers. Pteridium aquilinum var. latiusculum (Desv.) Underw. is locally abundant, mostly along the Perry Creek trail. Selaginella wallacei Hieron. is frequent on moss-covered talus boulders, mostly in the open. VEGETATION OF THE PERRY CREEK AREA The Perry Creek fern habitat is a most exceptional enclave within the broadly distributed lowland coniferous forest biome (Western Hemlock or Tsuga heterophylla Zone of Franklin and Dyrness, 1973) in western Washington. One needs only to traverse similar tributary canyons of the major river systems of the Puget Sound basin to discovery why. Whereas most such drainages are uniformly clothed with a mixed coniferous forest of Douglas Fir, Western Hemlock, and Western Red Cedar, with associated typical understory species, the fernery of Perry Creek is located along a succession of hardwood glades, patches of excep- tional assemblages of conifers of smaller than usual stature, and open talus, all on steep, moist slopes. The hardwoods are chiefly maples: Acer macrophyllum of low and sparse profile, along with thickets of the shrubby A. circinatum and A. glabrum. The Maple thickets frequently are replaced by similarly dense stands of deciduous, shrubby Cornus stolonifera or pure forb communities dominated by Athyrium filix-femina and Rubus parviflorus. Hardwood thickets are the dominant community on wet talus, and contain no conifers. The seed plants and associated ferns are: Acer circinatum, A. glabrum, *Actaea arguta, Alnus rubra, Aruncus sylvester, Athyrium filix-femina, Carex mertensii, Cornus stolonifera, *Elymus glaucus, Epilobium angustifolium, Geum macrophyllum, Oplopanax horridum, Osmorhiza_ occidentalis, Pteridium aquilinum var. latiusculum, Ribes petiolare, Rubus parviflorus, R. spectabilis, Sambucus racemosa, Sorbus sitchensis, Thalictrum occidentale, *Tolmiea men- ziesii, Valeriana sitchensis, and Veratrum viride. The species with asterisks ap- parently are restricted to trailside. It is in the understory of these hardwood glades that most of the Perry Creek ferns occur, although several species extend beyond onto the massive bouldery talus tracks between the hardwood glades. 42 AMERICAN FERN JOURNAL: VOLUME 66 (1976) On slightly less precipitous slopes, one finds open, coniferous woods, but not of the same composition as those of the climax coniferous forests on adjacent slopes. Rather, Alaska Cedar (Chamaecyparis nootkatensis), Subalpine Fir (Abies lasiocarpa), Douglas Fir (Pseudotsuga menziesii), and Western Hemlock (Tsuga heterophylla), all of smaller than expected stature, occur interspersed with the same hardwood dominants mentioned above. Open, bouldery talus has a rich cover of mosses and lichens. The frequent ferns are: Cryptogramma crispa, Asplenium trichomanes, Lycopodium selago subsp. patens, and Selaginella wallacei. Very few angiosperms occur, among them Montia flagellaris, Saxifraga bronchialis, and S. ferruginea. Wooded, bouldery talus replaces hardwood thickets or open talus on gentler slopes, and yet has the same rocky nature. This habitat is dominated by Alaska Cedar, although Acer macrophyllum also is common. Douglas Fir and Pacific Silver Fir (Abies amabilis) are occasional. Thick carpets of moss occur on the shaded talus, commonly with Asplenium trichomanes, Cryptogramma crispa, Goodyera oblongifolia, Montia flagellaris, and Polystichum munitum. Nearly everywhere along the two-mile sector of the trail the substrate is rocky, ranging from gravelly textured scree to massive, jagged boulders arranged in long, steep talus slopes. Although the soil is shallow to non-existent, the rock is mantled with a luxuriant coat of mosses and lichens. This non-vascular mat serves as a substrate for the ferns, fern-allies, grasses, forbs, and even for some of the woody plants. In sum, we find a microcosm of wet, rocky talus surrounded by the climatically dominant coniferous forest; the former must be an enclave caused by a unique combination of environmental factors. In the absence of carefully instrumented monitoring of the habitat, some of the operational factors may be inferred by observing the topography and the biologi- cal indicators of the area. Perry Creek is a tributary of the south fork of the Stillaguamish River in central Snohomish County, western Washington (Fig. /). It is well within the lower to mid-montane transition zone along the western flanks of the Cascade Range. The terrain along this sector of the Stillaguamish drainage is one of spectacularly sheer and high peaks rising abruptly out of the valley floor, which lies at ca. 1700 feet altitude. Perry Creek itself is in a narrow, steep-sided canyon bounded on the east and west by two dominating peaks, Mount Dickerman (5266 feet) and Stil- laguamish Peak (5863 feet). Downstream from Perry Creek, just beyond its junc- tion with the Stillaguamish River, is the massive, sheer wall of Big Four Mountain (6120 feet), which has on its north-facing slope permanent snowfields and at its talus base low-lying snowfields that are sculpted into the well known Ice Caves near Big Four. At the Ice Caves, the local environment and flora are truly subal- pine, despite the altitude of only 1800 feet. Given the frigid shelter of towering mountain walls on all sides, the occurrence of a truncated and telescoped sequence of ‘‘life-zones”’ is not unexpected. Under the usual conditions of gradual increase in elevation, the shift from the Western Hemlock through the Pacific Silver Fir to Mountain Hemlock Zone, with their attendant plant communities, would be gradual and not perceptible over short A. R. KRUCKEBERG: FERNS OF PERRY CREEK, WASHINGTON 43 distances. But at Perry Creek and at other sites along the Stillaguamish River, abrupt topographic and associated local climatic changes cause similarly abrupt changes in the floristic assemblages. But this inference from topography and climate alone does not wholly explain the hardwood scrub and talus communities within the foreshortened zonation of coniferous forest life-zones. I believe it is a matter of two integrated physical factors. First is the local climate, which provides a cool, moist habitat, shaded for much of the day. Second is the unstable, boul- derly talus. Both climate and substrate provide niches for ferns and other plants from diverse communities. Further, the talus of the hardwood glades that contain the bulk of the fern species is rich with mosses, which provide an optimally moist substrate for ferns throughout their life-cycles. FIG. 1. Looking down Perry Creek to the canyon of the south fork of the Stillaguamish River and Big Four Mountain on the horizon. Photo courtesy Mr. J. W. Thompson. The nearest weather station to Perry Creek is at Silverton (1500 feet), about 5 miles west and downstream in the same river valley. From weather records (Table /) it is easy to see that winter temperatures decrease and precipitation increases across the altitudinal transect from Puget Sound to the lower montane elevation of the Stillaguamish River. Undoubtedly, winter temperatures are still lower and precipitation greater along Perry Creek. The Washington State geological map (Huntting, 1961) portrays the rocks in the region of Perry Creek as pre-Jurassic sediments. It has not been possible to identify the particular sedimentary rocks along the crucial stretch of wet talus that 44 AMERICAN FERN JOURNAL: VOLUME 66 (1976) supports the ferns. The entire formation is complex, and according to Huntting (1961), consists of ‘‘graywacke, argillite and siltstone with some slate and phyllite; [it] includes graywacke breccia and ribbon chert with minor local limestone lenses and basalt flows.”’ The entire drainage of the south fork Stillaguamish River is exceedingly rich floristically and includes other fern species besides those that cluster along the Perry Creek talus. From herbarium records and observations by the author, the following ten pteridophytes can be added to the Perry Creek list. Three species characteristic of ultramafic (high magnesium) rocks occur in the nearby upper Coal Creek drainage: Aspidotis densa (Brack. in Wilkes) Lellinger, Adiantum pedatum subsp. aleuticum (Rupr.) Calder & Taylor, and Polystichum mohrioides var. lemmonii (Underw.) Fern. (Kruckeberg 1969, p. 84). David Wagner (pers. comm.) confirms the presence of Polystichum kruckebergii Wagner along with P. mohrioides var. lemmonii on Devil’s Thumb, also in the upper Coal Creek drain- TABLE 1. CLIMATIC DATA FOR PORTIONS OF SNOHOMISH COUNTY, WASHINGTON' Station Distance (mi) Temperatures (°F) Precipitation (in) an rom Jan. July Annual altitude Perry Creek average average max. min. average Everett, 100 ft 40 38.0 61.2 99 3 32.74 Granite, Falls 391 ft 20 ee ashy as pe 59.41 Silverton, 1500 ft 5 33.0 61.2 103 0 94.80 ‘Data from U.S. Dept. of Agriculture (1941, polis): age, as reported first by Slater (1967). D. Wagner also found Lycopodium sit- chense Rupr. in the same area. Athyrium distentifolium is a common talus species in the subalpine sections of the drainage; it is also at the Ice Caves. Other ferns and fern-allies known to occur in the Perry Creek drainage basin are Asplenium viride Huds. (Mt. Dickerman), Thelypteris limbosperma (All.) H. P. Fuchs (Monte Cristo), T. phegopteris (L.) Slossen (Big Four Mtn.), and Woodsia scopulina D. C. Eaton (Mt. Dickerman). Expected, but not yet found, in the Stillaguamish River drainage are Cheilanthes gracillima D. C. Eaton, Crypto- gramma Stelleri (Gmel.) Prantl, Dryopteris arguta (Kaulf.) Watt, Pityrogramma triangularis (Kaulf.) Maxon, Selaginella oregana D. C. Eaton in Wats., and Equisetum spp. he Perry Creek fernery is within the jurisdiction of the U.S. Forest Service (Monte Cristo Ranger District, Snoqualmie-Mt. Baker National Forest). Al- though there seems to be no immediate danger to the habitat, it is conceivable that mining or logging operations in other parts of the Perry Creek drainage could pose a threat. When I first visited the site in the early 1950s, our party hiked in from the bottom of the river valley through a virgin Hemlock-Cedar- Douglas Fir forest before reaching the fern-hardwood talus. The lower coniferous forest was logged in the late 1950’s to within % mile of the critical fern habitat. It is to be hoped that ae a ™ A. R. KRUCKEBERG: FERNS OF PERRY CREEK, WASHINGTON 45 the Forest Service will take steps to preserve the Perry Creek fern area in per- petuity, either as a Botanical Area or as a Research Natural Area. At present the trail through the site is a tolerable and handy intrusion, but no further modification of this priceless botanical and scenic habitat should occur. LITERATURE CITED U.S. DEPT. OF AGRICULTURE. 1941. Climate and Man. Yearbook of Agriculture. FRANKLIN, "i F, and C. T. DYRNESS. 1973. Natural Vegetation of Oregon and Washington. U.S.D.A. Forest Service Gen. Tech. Rept. PNW-8. oe Ree M. T., et al. 1961. Geological map of Washington. State Dept. of Mines and Geology, aa KRUCKEBERG A. R. 1969. Plant life on serpentinite and other ferro-magnesian rocks in north- rm North America. Syesis pick DER, E. A. 1974, AAG of the Male Fern from the western United States. Amer. m J. 64: 81-82. ren. 7 R. 1967. More on fern distribution in Washington state. Occ. Pap. Dept. Biology, Univ. Puget Sound 32: 293-310. WAGNER, W. H., Jr. 1973. Reticulation of holly ferns (Polystichum) in the western United States and adjacent Canada. Amer. Fern J. 63: 99-115 46 AMERICAN FERN JOURNAL: VOLUME 66 NUMBER 2 (1976) Origin of the Pteridophyte Flora of The Bahamas, Caicos and Turks Islands DONOVAN S. CORRELL* The region under consideration might be likened to the open mouth of a vast sack of tropical and subtropical plants of all categories. It is a region, excluding Bermuda, that is occupied by the farthest northeastward extension of floristic elements which have their optimum development to the south (in Cuba, His- paniola, and Puerto Rico) and west (in Florida). Subtropical southern Florida and its keys also have been recipients of the Greater Antilles flora. According to the best authorities, the geological processes of emergence and subsidence have affected the low-lying Bahamas more than once in the past. During periods of greatest emergence, the islands of the relatively shallow Little Bahama Bank (primarily Grand Bahama and Abaco) were connected. The same is true of the islands of the Great Bahama Bank (primarily New Providence, Eleu- thera, Cat, Exuma, Long and Andros). However, the great depth of water be- tween the two banks has precluded their ever having been connected. All of the islands scattered to the south of the Great Bahama Bank also have such depths of water between them that apparently they have never been connected; the main islands in this group are San Salvador, Crooked, Acklin, Mariguana, Inagua, Caicos and Turks. Finally, the great depth of water between all of the above islands and Florida, Cuba, and Hispaniola would appear to have prevented their ever having been connected, except for a possible connection of Great Bahama Bank with Cuba during the last Ice Age, about 25,000 years ago. That connection would account for some of the animal life now known to occur on some islands in this region. The dissemination of plants into the islands from the south and west has had to rely entirely upon agencies other than overland migration, since such routes apparently have never existed. Although there are several factors that affect the introduction and distribution of plants in this region, the two most important are the mode of dispersal and the conditions for establishment. Hurricanes and violent winds, water, birds, and the activities of man, such as those utilizing airplanes, boats, bulldozers, and au- tomobiles, are unquestionably primary sources of seeds, spores, and vegetative parts. But the establishment of a species depends primarily upon the availability of a suitable habitat. For instance, only those species that can grow at low elevations can become established in the Bahamas; the highest altitude is 208 feet on Cat Island. Many pteriodophytes need the protected, moist environment of solution pits or ‘banana holes’’ for their establishment, and these preferably should be in coppices. Six species occur only in the Bahamian archipelago and in Florida. It is not possible to determine with absolute certainty the direction of their migration. But it is likely that they originated in the United States. These are Preris vittata L., Selaginella armata Baker, Tectaria lobata (Poir.) Morton, Thelypteris augescens *Fairchild Tropical Garden, Miami, FL 33156. D. S. CORRELL: PTERIDOPHYTE FLORA OF THE BAHAMAS 47 (Link) Munz & I. M. Johnst., and 7. ovata R. St. John. Osmunda regalis var. spectabilis (Willd.) A. Gray, which also occurs rarely in Jamaica, should be placed in this category. Only Marsilea nashii Underw. is strictly endemic in the region, from Great Inagua to Grand Turks. Five other species are quite limited in their distribution and could be considered wide endemics. These include another sterile Marsilea, possible the Hispaniolan M. berteroi A. Br., from Acklin Island and South Caicos, Anemia wrightii Baker from Andros, also found in Cuba; A. cicutaria Kunze from Abaco, Andros and New Providence, also found in Cuba and Yucatan, Selaginella bracei Hieron. from Andros, also found in Cuba; and Thelypteris cordata (Fée) Proctor from Andros, also found in Jamaica. Apparently Cuba should be considered the point of origin for most of these species. The remaining 30 species in the Bahamas also are found in Cuba, Hispaniola, and Puerto Rico. Many of these extend to islands in the Lesser Antilles, Jamaica, or even to Mexico, Central America, or South America. Most are found in south- ern Florida as well, with the exception of such species as Polypodium squamatum L. on Andros and Schizaea poeppigiana Sturm in dense coppices on Abaco. In my opinion, all of these have migrated northward into the Bahamian archipelago. Those species that thrive best in association with the native pine Pinus caribaea Morelet (found on Abaco, Andros, Grand Bahama and New Providence, with a disjunct occurrence in the Caicos Islands) are Anemia adiantifolia (L.) Swartz, Pteris longifolia L., Thelypteris normalis (C. Chr.) Moxley, and Pteridium aquilinum var. caudatum (L.) Sadeb. The Pteridium frequently forms impene- trable masses in the ‘‘pineyards’’ on the northern islands. The northern Osmunda regalis var. spectabilis is found in fresh water marshes and ponds in the pinelands of Abaco. Trismeria trifoliata (L.) Diels is found in similar habitats on Grand Bahama and Abaco. Blechnum serrulatum L. C. Rich., Acrostichum danaeifolium Langsd. & Fisch., and A. aureum L. are found in less fresh water. All occasionally are found to some extent on islands devoid of pines, but they attain their optimum development when associated with pines. Those islands or sections of islands devoid of pines, but which have an exten- sive coverage of broadleaf coppices, support various ferns in company with or- chids and bromeliads. All occur on tree trunks, fallen logs, and on exposed rock ledges and walls. These are Polypodium polypodioides (L.) Watt, P. heterophyl- lum L., P. phyllitidis L. and its var. latum (Moore) Proctor, P. plumula Humb. & Bonpl. ex Willd., Paltonium lanceolatum (L.) Presl, and Psilotum nudum (L.) Pal. Beauv. By far the greatest number of species occur in the solution pits, sink-holes, or ‘‘banana holes’’ that are formed in the limestone stratum common to all of the islands. The holes that support ferns usually occur in more shaded and more moist habitats than are found elsewhere on the islands. Although one or more species may often be found in these sinks, no single species can be said to occur in all fern-inhabited sinks. The four most frequently encountered are Adiantum 48 AMERICAN FERN JOURNAL: VOLUME 66 (1976) tenerum Swartz, Tectaria lobata, Thelypteris normalis, and T. reptans (J. F. Gmel.) Morton. In fact, some species are of singular occurrence and most are sporadic. These are Pityrogramma calomelanos (L.) Link on Andros, Tectaria heracleifolia (Willd.) Underw. on Grand Bahama and New Providence, and Thelypteris dentata (Forssk.) E. St. John on Great Exuma. Other such species with slightly more widespread distributions are Adiantum melanoleucum Willd., Asplenium dentatum L., Psilotum nudum, and Sphenomeris clavata (L.) Maxon. The only genus of pteridophytes that might be considered a weed is Neph- rolepis. The ruderal N. exaltata (L.) Schott is the most widespread, whereas N. biserrata (Swartz) Schott, N. multiflora (Roxb.) Jarrett ex Morton, and N. rivularis (Vahl) Mett. ex Krug are limited to only one or two localities. A unique habitat, the leaf bases of old arborescent palmettos, is favored by two species: the rather widespread Polypodium aureum L. and the less common Vit- taria lineata (L.) J. E. Smith, which is found only on Andros and New Providence. Although I have examined literally hundreds of palmettos hoping to find the Shoestring Fern, I have not been successful, mainly because most of the palms have been burned by malpais farming practices. My search for Vittaria recalls an interesting and most profitable trip that I made in company with Dr. E. T. Wherry and Mr. J. E. Benedict, Jr. in June, 1939, to central Georgia for the express purpose of seeing this species growing on clay cliffs. On our return northward into South Carolina, we visited the station for Hymenophyllum tun- bridgense (L.) J. E. Smith in Oconee County, which I later reported (Amer. Fern J. 30: 21-27, 1940). I have many fond memories of my various field trips and associations with Dr. Wherry. I wish to acknowledge National Science Foundation support of my Bahama Flora research (Grant No. GB-41190X), and the assistance and cooperation of various individuals during the course of my exploratory work in the Bahamas, Caicos and Turks Islands. AMERICAN FERN JOURNAL: VOLUME 66 NUMBER 2 (1976) 49 Spore Retention and Release from Overwintering Fern Fronds DONALD R. FARRAR* The need for ecological data on ferns is becoming increasingly apparent as more effort is directed toward an understanding of the significance of their mor- phological and physiological diversity (Wagner, 1973). If detailed ecological studies of fern sporophytes have to date been too few, such studies on the gametophyte generation are almost non-existent. The reproductive cycle of ferns has been known for over a century, and more than a thousand articles on fern gametophytes have been published, half of these in the last quarter century (Miller, 1968; Naf, 1975; Nayar & Kaur, 1971). However, nearly all data on gametophyte growth and sexual reproduction have been based on laboratory ob- servations. Several factors have contributed to this paucity of information on gametophyte ecology, but perhaps the most important has been a widely held notion that gametophytes cannot be found in nature, or if found, cannot be iden- tified. Several recent studies have indicated to the contrary, that in situ gametophyte studies not only are feasible, but that they are essential for the integration of existing laboratory data into studies on the natural history of ferns (Cousens, 1973; Holbrook-Walker & Lloyd, 1973; Lloyd, 1974; Farrar & Gooch, 1975). To investigate further the feasibility of studying fern gametophytes in nature, we have begun a long-term observational study of fern reproduction in Woodman Hollow, a relatively isolated canyon in central Iowa, in which 13 species and 11 genera of ferns occur (see Table 1). This study is designed to answer the following questions. When are spores available for germination? When and where does reproduction occur and how is it influenced by micro- and macroclimates? When and by what breeding systems are sporophytes produced? Does sexual reproduc- tion occur in nature on a regular basis for all species? Results of the first year of study (Farrar & Gooch, 1975) indicate that the data needed to answer these and other questions will be forthcoming. Here we report some unexpected data rele- vant to the question of when spores are available for germination. Observations made on the time of spore maturation and first release during the growing season gave results which were similar to those of Hill and Wagner (1974) for pteridophytes in Michigan. Differences found in the two studies were no greater than might be expected due to differences in latitude, climate, habitat, and seasonal variation. Our observations also support their estimate that most spores of a given species are released during a period of about two weeks. However, a two week period of maximum release, if taken as a guide to the duration of spore release, may be very misleading. Our observations at Woodman Hollow indicate that for most species, significant quantities of spores are retained on the fronds after the initial release period and may be dispersed during a much longer period. Only in Botrychium virginianum and Osmunda claytoniana were essentially all *Department of Botany and Plant Pathology, Iowa State University, Ames, IA 50011. 50 AMERICAN FERN JOURNAL: VOLUME 66 (1976) of the spores shed in a period as short as two weeks after spore maturation. These species have large, smooth-walled sporangia which mature simultaneously. Fur- thermore, these species have dimorphic pinnae, and the fertile pinnae wither and often disappear soon after maturation. In the remainder of the species in our study, all sporangia in a sorus do not mature simultaneously. This differential maturation in itself lengthens the period of spore release, but an additional effect of this mixed sorus condition is that late-maturing sporangia are frequently covered by older, dehisced sporangia. Asa result, they may be physically unable to dehisce and shed their spores normally, and often are prevented from opening at all. Thus, the sorus may remain indefi- nitely with a mixture of opened sporangia containing no spores (those which opened first), opened sporangia which still contain some or all of their spores, and unopened sporangia containing their full complement of spores. Spore retention on the fern fronds varies among species and is dependent upon several factors. These include the number of sporangia per sorus, the presence of hairs or an indusium over the sorus, the type and degree of persistence of the indusium, and the time of maturation of the fertile frond. We observed a number of species which, after an initial flush of fertile fronds, continued to produce additional fertile fronds for the remainder of the growing season. Fronds maturing late in the season, especially of Adiantum pedatum, Asplenium rhizophyllum, and Polypodium virginianum, often had large numbers, occasionally approaching 100%, of unopened sporangia. To quantify as much as possible this extended period of spore retention and release, fertile fronds of species in the study area were collected in December and again in March. The fronds were examined under a dissecting microscope and an estimate was made of the percent of unopened sporangia and the total number of Spores remaining on the frond. Because of the generally deteriorated condition of the fronds, precise counts of sori, sporangia, and free spores could be made only with considerable difficulty. Thus, the spore estimates were limited to the orders of magnitude listed in Table 1. Ten or more sori were examined on each frond. The estimates of spores per frond were based on the number of unopened sporan- gia, the number of spores which could be removed from the dried frond, and the number of spores which could be seen remaining on the frond. Taking into ac- count the inherent problems with the methodology of the spore assay and assum- ing that the estimates are not in error by more than an order of magnitude, it was obvious that for most species, large numbers of spores were retained on the fronds throughout the winter. Spore viability was also tested for each collection by sowing the spores on mineral nutrient agar and measuring percent germination after three weeks. Ger- mination in December ranged from 49% to 96%. Differences in germination per- centages obtained in December and March probably represent variation between plants of the same species, since the overall range in March was similar (48-88%) to that in December and the number of species showing an increase was nearly as great as those showing a decrease in germination (Table 1). The change in Cysto- pteris bulbifera was most dramatic and may represent a real decrease in spore D. R. FARRAR: SPORES OF OVERWINTERING FERN FRONDS 51 viability for this species. To determine whether unopened sporangia of late- maturing fronds contained mature and viable spores, fronds of Asplenium rhizophyllum were divided into three lots on the basis of the number of unopened sporangia. These lots, when tested independently, showed similar germination percentages, indicating that indehiscence of sporangia, at least in this species, was not due to spore immaturity. From December to March, a definite decrease in the number of unopened sporangia occurred only in late-maturing fronds of Asplenium rhizophyllum, which in December had greater than 90% unopened sporangia, and in Matteuccia TABLE 1. SPORE PRESENCE AND VIABILITY ON FERN FRONDS COLLECTED IN ECEMBER AND MARCH IN WOODMAN HOLLOW, IOWA No. fronds % sporangia No. sig aes % spore germination ned ve analyzed unopen x (number counted Dec. Mar. Dec. Mar. Dec. pe ec. r; Adiantum pedatum 6 3 1-1 same 10-100 1-10 77 (416) ~=67 grin Asplenium rhizophyllum Bes 3 1-10 same 1-10 same 81 (237) ~—87 (378) Asplenium rhizophyllum 6 — 10-20 _ 10-100 — 84 (210) — “Asplenium rhizophyllum 9 2 >90 >75 100-1,000 same 81 (308) a Athyrium filix-femina 3 4 1-10 same _ 10-100 same 49 (645) 73 (541) Cryptogramma stelleri 0 2 — S15 —_ 10-100 — 63 (183) Cystopteris bulbifera 4 4 <1 same 1-10 0.1-1 62 (444) 6 (226) Cystopteris fragilis 5 4 <1 same 1-10 same 68 (190) 58 (248) Dryopteris goldiana 4 4 1-10 <1 10-100 1-10 50 (231) 48 (306) Dryopteris spinulosa 4 4 <1 1-10 10-100 same 52 (307) 84(703) Matteuccia struthiopteris 3 1 >90 >80 100-1,000 same 96 (S99) 95 (668) Polypodium virginianum 3 5 10-50 same 100-1,000 same 70 (224) ~=88 (258) Woodsia obtusa 6 4 1-10 same 10-100 1-10 82 (403) 59 (230) Botrychium virginianum no fertile fronds found Osmunda claytoniana no fertile fronds found struthiopteris, which was observed to be releasing spores both in December and in March. The total number of spores per frond appears generally to have re- mained relatively unchanged; however, our method of analysis may not have been sufficiently sensitive to detect the changes that did occur. That some spores were released from December to March is indicated by the decrease in unopened sporangia in Asplenium rhizophyllum and Matteuccia struthiopteris and by a measurable decrease in numbers of spores per frond in Adiantum pedatum, Cys- topteris bulbifera, Dryopteris goldiana, and Woodsia obtusa. The data thus indicate that for most of the fern species in Woodman Hollow, sporophyte fronds of the previous year retain large numbers of spores throughout the winter and into the growing season of the following spring. Furthermore, it appears that some spores continue to be shed from these fronds as the winter progresses. The fate of the spores that are shed, or of those that remain on the fronds, has yet to be determined. The old, spore-bearing fronds are generally flattened against the substratum by early spring, and further release of their spores into air currents must be greatly reduced. Nevertheless, some spores could cer- tainly germinate in the vicinity of the sporophyte fronds if a suitable habitat were available. 52 AMERICAN FERN JOURNAL: VOLUME 66 (1976) As yet, we have detected establishment of large numbers of gametophytes of only three species, Adiantum pedatum, Cystopteris fragilis, and Woodsia obtusa, and this has occurred in the fall. Our failure to observe gametophyte establish- ment of other species, or of these species at other times of the year, may be due to unfavorable weather conditions, or may reflect the inability of some species to reproduce regularly or extensively through the production of gametophytes. It most certainly reflects the rudimentary state of our knowledge of fern reproduc- tion in nature. It may well be true that significant gametophyte establishment results only from spores released during the sporophyte growing season. How- ever, until this is proven, workers studying gametophyte ecology in temperate areas must consider the possibility that significant reproduction may also result from spores shed from overwintering fronds. LITERATURE CITED COUSENS, M. I. 1973. Reproductive biology and autecology of Blechnum spicant. Ph.D. Thesis, Washington State aoe an FARRAR, D. R., and R. D. GOOCH. 1975. Fern reproduction at Woodman Hollow, central Lowa: pT We pbicvetons and a consideration - the feasibility of studying fern reproductive biology in nature. Proc. lowa Acad. Sci. 82: De HILL, R. H., and W. H. WAGNER, Jr. 1974. Sai nin and spore type of the pteridophytes of Michigan. Michigan Bot. 13: 40-44. HOLBROOK-WALKER, S. G., and R. M. LLOYD. 1973. Reproductive biology and gametophyte morphology of the Hawaiian fern genus Sadleria (Blechnaceae) relative to habitat diversity and propensity for colonization. Bot. J. Linn. Soc. London 67: 157-174. LLOYD, R. M. 1974. Reproductive biology and evolution in the pteridophyta. Ann. Missouri Bot. Gard. 61: 318-331. MILLER, J. H. 1968. Fern gametophytes as experimental material. Bot. Rev. -440. NAF, U., K. NAKANISHI, and M. ENDO. 1975. On the physiology and beahews of fern an- theridogens. Bot. Rev. 41: 315-359. NAYAR, B. K., and S. KAUR. 1971. Gametophytes of homosporous ferns. Bot. Rev. 37: 295-396. WAGNER, W. H., Jr. 1973. Some future challenges of fern systematics and phylogeny. In A. C. Jermy etal. (eds.). The Phylogeny and Classification of the Ferns. Bot. J. Linn. Soc. London 67: Suppl. 1: 245-256. AMERICAN FERN JOURNAL: VOLUME 66 NUMBER 2 (1976) 53 The Distribution and Abundance of Dryopteris in New Jersey JAMES D. MONTGOMERY*:! The eastern American Wood Ferns are now reasonably well understood as to relationships. There is, however, little quantitative information concerning the local distribution, habitat preference, and abundance of Dryopteris hybrids, espe- cially in the northeastern United States. The purpose of this investigation is to document the abundance, distribution, and habitat of the Wood Ferns found in New Jersey. Although Wood Ferns are found throughout New Jersey, they are particularly common in the more rugged northwestern part of the state (Sussex, Passaic, Warren, and Morris Counties). Hybrids have been known since Dowell (1908) described Dryopteris clintoniana x intermedia and D. goldiana x marginalis from New Jersey. In ‘‘The Ferns of New Jersey’’ Chrysler and Edwards (1947) listed 14 hybrids and gave one record for each except D. x boottii; but no informa- tion was given about abundance, distribution, or habitat. Little specific informa- tion has been given by recent northeastern fern guides, although Dryopteris hy- brids are listed and briefly discussed by Wherry (1961). Wagner (1963) discussed the relative abundance of species and hybrids of Wood Ferns in Virginia, and Britton (1965) tabulated their relative abundance in Ontario; no quantitative data were given. Although hybrids are usually described as being rare, some fern hybrids are relatively common. Wagner (1969), in defense of the inclusion of hybrids in floras, pointed out that several Appalachian Asplenium hybrids are more common than the non-hybrid Hart’s-tongue (Phyllitis scolopendrium), and that Dryopteris x triploidea and D. x boottii (both sterile hybrids) are commoner than either D. celsa or D. clintoniana (both fertile). Herbarium records for Dryopteris hybrids are deceptive. Some of the hybrids are large and conspicuously different, and therefore frequently collected, whereas others are difficult to distinguish and less conspicuous, and so are less often taken. For this reason the abundance of hybrids was measured in two ways in this investigation: (1) the overall abundance, that is the number of stations at which each occurs in New Jersey as determined from herbarium records and field recon- naissance; and (2) the relative abundance as a proportion of the frequency with which a hybrid occurs when the parents are found growing together. PROCEDURE Populations of Dryopteris where hybrids might occur were located in three ways: (1) examining herbarium labels where reasonably exact data were given, (2) scanning topographic maps for likely habitat areas, and (3) extensive driving and hiking in the area where likely habitats might occur. Once a population had been oe Associates, P.O. Box 2, Stamford, NY 12167. 'The ® thank W. H. Wagner, Jr. and F. S. Wa 5 a! for their suggestions and Anas a oe nt, an E. Fairbrothers for permitting the use of Botany Department facilities at Rutgers Chives Financial aid was received from the Upsala Cites Faculty Research Fun 54 AMERICAN FERN JOURNAL: VOLUME 66 (1976) located where two or more species of Dryopteris were growing intermixed, a careful search was made to locate all hybrids. Identifying a plant of Dryopteris as a hybrid is usually not difficult; hybrids have abortive spores (Whittier & Wagner, 1971) and frequently sporangia as well, and are generally intermediate in morphology between their parents (see Wagner, 1971). Some hybrids are relatively easy to identify by morphology alone: those involving as parents D. marginalis (marginal sori and a dense tuft of tawny scales at the base of the stipes), D. goldiana (abrupt taper at the tips of the large blades and very dark, shiny basal scales), D. intermedia (glandular blades and indusia and twice pinnatifid pinnae), and D. spinulosa (eglandular blades and indusia and twice pinnatifid pinnae). The hybrid between D. intermedia and D. spinulosa can be distinguished from its similar parents by abortive spores (Tryon & Britton, 1966). Although spore abortion can often be determined with a 10x lens in the field, in this investigation spores were examined with a light microscope at 100. The hybrids involving D. celsa, D. clintoniana, and D. cristata can be recognized as such, but determining parentage from morphology is often quite difficult. Chromosome counts and pairing behavior are very useful here. Slides were exam- ined from both transplanted and wild plants, following methods given in Montgomery (1975). Voucher specimens for field and cytological studies were deposited in the Chrysler Herbarium of Rutgers University (CHRB), with dupli- cates in the herbarium of Upsala College (EONJ); duplicates are fronds from the same plant. For ranges and habitats, as well as abundance, material was examined from the following herbaria: AFS, CHRB, EONJ, MICH, NY, PENN, PH, US, and Staten Island Museum (SIM). Habitat information was tabulated from her- barium sheets where sufficient information was given. RESULTS Six species and 13 hybrids were found in field studies. The relative frequency of the hybrids was tabulated as a percentage of the number of populations where both parents were present (Table /). The hybrid with the greatest frequency of occurrence when the parents were found together was D. goldiana x marginalis (100%), followed by D. intermedia x spinulosa (84%). It should be noted that the first combination of parents was found together only twice, whereas D. inter- media and D. spinulosa were found together 45 times. The overall abundance of the species and hybrids, as determined from her- barium records (including the author’s material) is given in Table 2. In this table a record refers to a distinct locality; duplicates from a locality at the same or differ- ent times are eliminated. Dryopteris marginalis was the most abundant species, from 176 localities. D. intermedia and D. spinulosa were nearly as abundant. The most abundant hybrids were D. intermedia x spinulosa and D. cristata X inter- media. Both of these, as well as D. cristata x marginalis were recorded from more stations than the least common species, D. celsa, D. clintoniana, and D. goldiana. Chromosome counts made to verify taxa are given in Table 3. Additional counts from New Jersey were given by Montgomery (1975) and Wagner (1971). J. D. MONTGOMERY: DRYOPTERIS IN NEW JERSEY 95 TABLE 1. FREQUENCY OF Dryopteris PARENTS AND HYBRIDS IN NEW JERSEY FIELD POP ULATIONS. x cristata X goldiana x intermedia n f(%) n f(%) n f(%) D. clintoniana 17 58.8 2 50.0 15 40.0 D. cristata 2 50.0 31 58.1 D. goldiana 1 0.0 D. marginalis n = number of populations with both parents present. f = percent of populations (n) where the hybrid was found. x marginalis n f(%) 2 25.0 15 18 55.6 35 2 100.0 1 27 37 45 x spinulosa n f(%) 26.7 TABLE 2. NUMBER OF LOCALITIES OF Dryopteris HYBRIDS IN NEW JERSEY DS. Number of OM HERBARIUM RECOR Number of localities for hybrids x * localities x x x for species clintoniana cristata goldiana_ intermedia marginalis spinulosa 2 0 4 3 2 3 | D. clintoniana 24 13 6 10 4 5 D. cristata 108 2 45 29 1] D. goldiana 19 1 1] 0 D. intermedia 138 2 46 D. marginalis 176 2 D. spinulosa 139 TABLE 3. CHROMOSOME COUNTS IN NEW JERSEY Dryopteris. ] I] Source of Material’ D. cristata 0 81+1 25466c, Macopin, Passaic Co. 0 82 25473y, Green Pond, Passaic Co. 0 81+1 8250p, Wawayanda, Sussex Co. D. goldiana 0 4] 8155b, Netcong, Morris Co. D. intermedia 0 41 ceeieh Macopin, Passaic Co. 0 423d, Wallpack, Sussex Co. * D. cristata X intermedia me x boottii) 123 0 25466b, Macopin, Passaic Co. [222 0-1 25466d, Macopin, Passaic Co. 123 10528g, Macopin, Passaic Co. D. cristata X marginalis (D. x slossonae) 12123 0 8250), Wawayanda, Sussex Co. 123+2 0 10528m, Macopin, Passaic Co. D. cristata X spinulosa (D. X uliginosa) 76 43 78 wartswood, Sussex Co. dD: Leben x spinulosa 0 x iWolbideay 40+ 1 40+ 010c, Greendell, Sussex Co. ‘Collected by the author; voucher at CHRB. 56 AMERICAN FERN JOURNAL: VOLUME 66 (1976) DISCUSSION In general hybrids were found with both parents nearby, usually in sight. Occa- sionally, however, a hybrid plant was found with only one parent very close by. Dryopteris x triploidea was found six times with only D. spinulosa present; D. x boottii was found once with D. cristata about 0.25 mile away and no D. inter- media in the vicinity, and one other time with only D. cristata present. Wagner (1971) referred to this phenomenon as “‘hybridization by remote control,”’ and it presumably happens when a spore from the missing parent is blown into the vicinity of the gametophyte of the other parent so that cross-fertilization can occur. Extinction of one species or the other, or both, at a locality after hybrid formation is another possible mechanism; the hybrid can persist and even spread slowly by rhizome growth. These conditions are unusual, however, and so parent species are usually found in the vicinity of the hybrids. The abundance of the hybrids would be expected to be related to the abundance of the parental taxa. The quantitative information presented here indicates that this is true in some cases; thus, hybrids involving D. goldiana are rare in New Jersey. Dryopteris intermedia x spinulosa (D. x triploidea) and D. cristata x intermedia (D. X boottii) were common, and the parent species were very com- mon. These were also the only taxa involved in ‘‘remote control’’ hybridization. Hybrids were also more common if the parents had similar habitat preferences. Dryopteris clintoniana x cristata was found relatively frequently, both in her- barium records (Table 2), although often identified as one or the other parent, and in relative abundance in field studies (Table /). Both parents occupied similar swampy habitats. On the other hand, D. clintoniana x marginalis was relatively uncommon, and these parents occupied rather different habitats. The most spec- tacular hybrid populations occurred where a wooded ridge occupied by D. inter- media and D. marginalis sloped into a wooded swamp occupied by D. clin- toniana, D. cristata, and D. spinulosa. In such locations the edge of the swamp may be virtually lined with hybrids! At Big Spring, where D. goldiana also oc- curred, six of 15 possible hybrids were found and the number of hybrid plants was unusually great. In two other areas, Wawayanda and Macopin, five species were present and six of ten possible hybrids were found. Certain hybrids were inexplicably rare. The most notable case was D. inter- media x marginalis. The parental species were abundant and occupied similar habitats, and so large intermixed populations were encountered frequently. This hybrid is also relatively easily seen if the population is searched carefully. The same general situation applies to D. cristata x spinulosa in New Jersey. NO explanation is known for the lack of hybridization between these species. Experi- mental studies with the gametophytes may be helpful. The relative abundance of hybrids found in New Jersey agrees in general with that found in Virginia by Wagner (1963) and in Ontario by Britton (1965). Both reported D. intermedia x spinulosa as the commonest hybrid combination, as in the present study. Both stated that D. intermedia x marginalis and D. marginalis x spinulosa were rare, as documented here. Britton listed D. cristata * mar- J. D. MONTGOMERY: DRYOPTERIS IN NEW JERSEY 57 ginalis as ‘‘rare to very rare’’ in Ontario; in New Jersey it was recorded from 29 stations (third highest) and found 55.6% of the time when the parents were found growing together. Dryopteris clintoniana x intermedia was also more frequent in New Jersey than in Ontario. The distribution, habitat preference, and abundance of each Dryopteris species and hybrid in New Jersey are summarized below. Dryopteris celsa (W. Palmer) Small. Very rare, perhaps extinct. Known from four localities in Bergen County (two by hybrids only); all from swamps that have been destroyed by urbanization. Dryopteris clintoniana (D. C. Eaton) Dowell. Uncommon. In low woods or wooded swamps; several locations in Sussex County, and a few each in Passaic, Warren, Morris, and Essex Counties. Dryopteris cristata (L.) Gray. Common. Usually in wooded swamps, growing on old stumps, logs, and hummocks; occasionally in wet meadows or damp woods; recorded from all counties except Hudson and Cumberland, but more common in the northern part of the state. Dryopteris goldiana (Hook.) Gray. Uncommon. In rich woods or ravines, especially on limestone in New Jersey; several records (mostly old) for Sussex and Warren Counties, and one or two each in Bergen, Morris, Essex, and Hun- terdon. Dryopteris intermedia (Muhl.) Gray. Abundant. Rocky, wooded slopes, espe- cially north- or east-facing, sometimes in wet woods or swamps especially in the southern half of the state; known from all 21 counties. Dryopteris marginalis (L.) Gray. Abundant. Rocky woods, often drier than D. intermedia, and only rarely in swamps; recorded from all counties except Cape ay. Dryopteris spinulosa (O. F. Muell.) Watt. Abundant. In woods, nearly always in moist areas (springs, etc.), or in wooded swamps as D. cristata. Recorded from all counties except Union. Dryopteris celsa hybrids. Five hybrids involving D. celsa are recorded from New Jersey: D. celsa x cristata, D. celsa x goldiana, D. celsa x intermedia (D. x separabilis (Palmer) Small), D. celsa x marginalis (D. x leedsii Wherry), D. celsa x spinulosa; all from Bergen County (Montgomery, 1975). Dryopteris clintoniana X cristata. Uncommon. In swamps, as the parents, but only a few plants; recorded from Sussex, Passaic, Warren, and Morris Counties. Dryopteris clintoniana x goldiana. Uncommon. Edges of swamps with D. goldiana above and D. clintoniana below; recorded from Sussex, Passaic, War- ren, and Essex Counties. Dryopteris clintoniana X intermedia (D. x dowellii (Farw.) Wherry). Uncom- mon; edges of swamps or wet woods; type from Macopin, Passaic County, plus several other records in Sussex and Morris Counties. Dryopteris clintoniana x marginalis. Rare. In swamps, three localities in Sus- sex County, one in Warren County. Dryopteris clintoniana x spinulosa (D. xX benedictii Wherry). Rare. In swamps; five records in Sussex, Passaic, and Morris Counties. 38 AMERICAN FERN JOURNAL: VOLUME 66 (1976) Dryopteris cristata x goldiana. Doubtfully present. One sterile collection from Lodi, Bergen County, which could be a D. celsa hybrid, and a possible plant from Morris County. Dryopteris cristata x intermedia (D. x boottii (Tuckerm.) Underw.). Common. Most commonly in swamps or wet woods, but also on wooded slopes, or even on rock walls; occasionally several plants, but more commonly only a few; most of the northern counties, plus Gloucester and Cape May. Dryopteris cristata X marginalis (D. x slossonae Wherry). Common. Edges of swamps, nearly always on hummocks or stumps; recorded from all northern coun- ties, plus Mercer, Middlesex, and Monmouth in central New Jersey. ryopteris cristata xX spinulosa (D. xX uliginosa Druce). Uncommon. In swamps, rarely more than one or two plants; four localities in Sussex, and one or two each in Passaic, Bergen, Morris, Middlesex, Monmouth, and Burlington Counties. Dryopteris goldiana X intermedia. Rare. One record in Bergen County from an area now destroye Dryopteris goldiana X marginalis (D. x neowherryi Wagner). Uncommon. At borders of swamps or in damp rich woods; several records in Sussex County, plus two each in Bergen and Morris Counties, and one in Hunterdon County. Dryopteris goldiana X spinulosa. Unknown in the state. Dryopteris intermedia < marginalis. Rare. Only two records: in 1914 from Sussex County, and by the author in 1973 in Union County; the latter with the parents on a steep, northwest-facing slope in hemlock woods. Dryopteris intermedia Xx spinulosa (D. x triploidea Wherry). Common. In woods, especially open areas such as pine plantations, or edges of swamps, banks of streams, etc.; the commonest hybrid, frequent with the parents and sometimes outnumbering either or both; recorded from nine counties, mostly in the northern part of the state. D. marginalis x spinulosa (D. x pittsfordensis Slosson). Rare. Only two rec- ords, one from a swamp in Middlesex County, the other, possibly from cultivated plants, in Monmouth County. LITERATURE CITED BRITTON, D. M. 1965. Hybrid wood ferns in Ontario. Michigan Bot. 4: 3-9. CHRYSLER, M. A. and J. L. EDWARDS. 1947. The Ferns of New Jersey. Rutgers Univ. Press, New Brunswick, N.J. DOWELL, P. 1908. New ferns described as hybrids in the genus Dryopteris. Bull. Torrey Bot. Club. MONTGOMERY, J. D. 1975. Dryopteris celsa and D. clintoniana in New Jersey. Amer. Fern J. 65: 65-69. TRYON, R. M. and D. M. BRITTON. 1966. A study of variation in the cytotypes of Dryopteris spinulosa. Rhodora 68: 59-92. WAGNER, W. H., Jr. 1963. Pteridophytes of the Mountain gg area, Giles Co., Virginia, including notes from Whitetop Mountain. Castanea 28: 113-1 . 1969. The role and taxonomic treatment of ty Bioscience 19: 785-789. J. D. MONTGOMERY: DRYOPTERIS IN NEW JERSEY 59 WAGNER, W. H. Jr. 1971. Evolution of Dryopteris in relation to the Appalachians. Jn P. C. Holt (ed.). The distributional history of the biota of the southern Appalachians. Virginia Polytech. Inst. Res. Div. Monogr. 2: 147-192 WHERRY, E. T. 1961. The Fern Guide. Doubleday & Co. Garden Citys N.Y: WHITTIER, D. P. and W. H. WAGNER, Jr. 1971. The variation in spore size and germination in Dryopteris taxa. Amer. Fern J. 61: 123-127. 60 AMERICAN FERN JOURNAL: VOLUME 66 NUMBER 2 (1976) Cystopteris bulbifera in the Southwestern United States TIMOTHY REEVES* The Bulblet Fern occurs primarily in eastern North America and reaches its southwestern distributional limit in Arizona (Fig. ]). The eastern distribution is fairly continuous as far west as the eastern edge of the Great Plains, where populations become sporadic. A gap of about 1000 km separates known localities in eastern Nebraska and Oklahoma from sites in New Mexico and Texas (Wagner, 1972). Anderson (1974) discussed the distribution of this species in the Great Plains but not in the Southwest. Cystopteris bulbifera (L.) Bernh. is known in Texas only in the Guadalupe Mountains, Culberson Co. (Correll & Johnston, 1970). New Mexican collections have been made in these same Guadalupe Mountains, Otero or Eddy Co. (Blasdell, 1963, p. 67); in the White Mountain wilderness, northern Otero Co. (C. R. Hutchins, pers. comm.); and north of Taos, Taos Co. (Dittmer, et al., 1954, p. 28). In Utah, Flowers (1944) reported the species from Zion National Park, Washington Co.; Elk Mountain, San Juan Co.; and Brighton and Little Cottonwood Canyon, Salt Lake Co. There are no reports of this species from Colorado, Nevada, or farther north. In Arizona, C. bulbifera has previously been known only from the West Fork of Oak Creek Canyon, Coconino Co. (Kearney, et al., 1960, p. 44). I have recently collected specimens of it in Walnut Canyon, Coconino Co., at a site about 40 km northeast of the Oak Creek locality. A recent floristic study of Walnut Canyon by Joyce (1974) does not mention the occurrence of C. bulbifera, but my examination of his collections has revealed one specimen of this species Joyce WC634, ASC) misidentified as C. fragilis (L.) Bernh. Blasdell (1963, p. 28) stated that C. bulbifera usually grows on neutral soils associated with limestone, whereas Anderson (1974) indicated that in Nebraska it occurs on sandstone. In Arizona, I have collected this species on both substrates. Cystopteris bulbifera is abundant in the lower portion of the West Fork of Oak Creek Canyon (1550 m), occurring in large colonies on the red sandstone talus and cliffs capped by Kaibab limestone. Associated fern species are Adiantum capillus-veneris, A. pedatum, and Polypodium hesperium. At higher elevations in the canyon (2000 m), where C ystopteris fragilis, Dryopteris filix-mas, and Poly- stichum lonchitis are found, the Bulblet Fern occurs on limestone outcrops and boulders. At Walnut Canyon, C. bulbifera occurs in a few small colonies at 1800 m in the narrow gorge to the southwest of the Walnut Canyon National Monu- ment Headquarters. Here, the plants grow in shaded seepage sites on vertical cliffs of Kaibab limestone. Associated species are Cheilanthes feei, Cystopteris fragilis , and Selaginella underwoodii. Putative hybrids between C. bulbifera and i fragilis from Walnut Canyon are currently under investigation. Since the only previously reported chromosome counts for C. bulbifera were based on specimens from eastern North America, an attempt was made to obtain “Department of Botany and Microbiology, Arizona State University, Tempe, AZ 85281. T. REEVES: CYSTOPTERIS BULBIFERA IN THE SOUTHWEST FIG. 1. Known distribution of Cystopteris bulbifera in the southwestern United States. an % a@ Beles a e Catv? ME oy a, mes 10 um FIG. 2. Camera lucida drawing of meiotic chromosomes of Cystopteris bulbifera at metaphase | showing 42 pairs of chromosomes (Walnut Canyon, Coconino County, Arizona, Reeves 3030A, ASU). 62 AMERICAN FERN JOURNAL: VOLUME 66 (1976) counts on Arizona material. Specimens of this species were collected in both Oak Creek and Walnut Canyons on 17 June 1975. Pinnae were fixed in modified Carnoy’s fixative (6 chloroform: 4 ethanol: | glacial acetic acid). After 24 hours the material was transferred to 70% ethanol and refrigerated until studied. De- veloping sporangia were stained in iron aceto-carmine and squashed in Hoyer’s medium. All cells studied had normal chromosome pairing with 21 =42 pairs (Fig. 2). This agrees with all previously reported counts for the species (Table 1). The counts reported here are the first reports for C. bulbifera in the western United States, indicating that this species apparently occurs as a diploid throughout its range TABLE 1. CHROMOSOME NUMBERS IN Cystopteris bulbifera. Number Voucher or reference Locality n= 42 Britton (1953) Canada: Ontario 42 Wagner (1955) Ohio: Ross : 42 Wagner (1955) Michigan: Oakland Co. 42 Wagner & Hagenah (1956) Michigan: Ionia Co. 42 Wagner & Hagenah (1956) Michigan: Eaton Co. 42 Reeves 3026A (ASU) Arizona: Walnut Canyon, Coconino Co. 42 Reeves 3030A (ASU) Arizona: Walnut Canyon, Coconino Co. 42 Reeves 3046D (ASU) Arizona: Oak Creek Canyon, Coconino Co. I am grateful to Dr. W. H. Wagner, Jr. for his suggestion that the genus Cysto- pteris be studied carefully in Arizona. I wish to thank the Superintendent of Walnut Canyon National Monument for granting permission to collect plants. | express my gratitude to the curators of Arizona herbaria (ARIZ, ASC, ASU, and MNA) for use of their facilities and the loan of specimens. Doctors D. J. Keil, D. J. Pinkava and W. H. Wagner, Jr. have reviewed this manuscript and their as- sistance is gratefully acknowledged. LITERATURE CITED ANDERSON, G. J. 1974. Cystopteris bulbifera new to Nebraska. Amer. Fern J. BLASDELL, R. F. 1963. A monographic study of the fern genus Cystopteris. ita fans Bot. Club 21(4): 1-102. BRITTON, D. M. 1953. Chromosome studies on ferns. Amer. J. Bot. 40: 575-583. CORRELL, D. S. and M. C. JOHNSTON. 1970. Manual of the Vascular Plants of Texas. Texas Research Foundation, Renner. DITTMER, H. J., F. CASTETTER, and O. M. CLARK. 1954. The Ferns and Fern Allies of exico. Univ. New Mexico Press, Albuquerque. FLOWERS, S. 1944, Ferns of Utah. Bull. Univ. Utah 35(7): 1-87. JOYCE, J. F. 1974. A Taxonomic and Ecological Analysis of the Flora of Walnut Canyon, Arizona. Unpublished M. S. Thesis, Northern Ariz. Univ.. Flagstaff. KEARNEY, T. H., R. H. PEEBLES, and collaborators. 1960. Arizona Flora. 2nd ed. Univ. Calif. Press, Berkeley & Los Angeles oe pel Jr. 1955. Cytotaxonomic observations on North American ferns. Rhodora 57: - 1972. Disjunctions in homosporous vascular plants. Ann. Missouri Bot. Gard. 59: 203-217. segshighe HAGENAH. 1956. A diploid variety in the Cystopteris fragilis complex. Rhodora AMERICAN FERN JOURNAL: VOLUME 66 NUMBER 2 (1976) 63 Variation in North American Asplenium platyneuron W. CARL TAYLOR, ROBERT H. MOHLENBROCK, and FREDDA J. BURTON* One of the most common and widespread of the eastern North American spleenworts is the Ebony Spleenwort, Asplenium platyneuron (L.) Oakes ex D. C. Eaton, which ranges from Quebec to Ontario, south to Colorado, Texas, Florida, and the West Indies. It is also known from South America and South Africa (Mohlenbrock, 1967, p. 157). Because it is a common, attractive, and variable species, a number of varieties and forms have been recognized by both professional pteridologists and amateur fern enthusiasts. We have found the litera- ture to contain nine infraspecific names accounting for variations in frond and pinna form or in stipe and rachis branching or proliferations. The purpose of this paper is to account historically for these taxa, to review their taxonomy and nomenclature, and to provide a key for their identification. The stimulus for this report comes from the discovery of the striking cut-leaf variant A. platyneuron f. hortonae, which is reported here for the first time from Illinois. Our studies have revealed that much herbarium material of A. platyneuron is incompletely or incor- rectly determined below the species level. KEY TO INFRASPECIFIC TAXA OF NORTH AMERICAN ASPLENIUM PLATYNEURON 1. Stipe and rachis unbranched and not proliferous. 2. Longest pinnae less than 3.5(4) cm long; pinnae subentire to nearly pinnatisect; erect fronds with or without sori. 3. Pinnae subentire to crenulate or serrulate 3. Pinnae doubly serrate to deeply incised or pinnatifid to pinnatisect. 4. Pinnae doubly serrate to deeply incised; all or nearly all of the pinnae cut less than 4/5 of the way to the midvein; erect fronds normally soriferous 1b. var. incisum 4. Pinnae pinnatifid to pinnatisect; all or nearly all of the pinnae cleft more than 4/5 of the way to the midvein; fronds without sori c. f. hortonae 2. Longest pinnae more than 3.5 cm long; pinnae often coarsely serrate-incised; erect fronds nor- mally soriferous Id. var. bacculum-rubrum 1. Stipe and/or rachis branched or proliferous. 5. Stipe and/or rachis branched le.-f. furcatum 5. Stipe and/or rachis proliferous If. f. proliferum la. var. platyneuron — la. Asplenium platyneuron (L.) Oakes ex D. C. Eaton var. platyneuron, Ferns No. er. 1: 24. 1 Acrostichum platyneuros L. Sp. Pl. 2: 1069. 1753. Asplenium ebeneum Ait. Hort. Kew. 3: 462. 1789. Chamaefilix platyneuros (L.) Farw. Amer. Midl. Nat. 12: 269. 1931. Most American botanists in the nineteenth century chose to use Aiton’s name A. ebeneum for this species. Fernald (1935, pp. 382-384) discusses the difficulties in typifying Acrostichum platyneuros. Typical A. platyneuron has erect fertile fronds up to 40 cm long with as many as 50 pairs of pinnae (Fernald, 1950) and a lustrous, dark brown, unbranched stipe *Department of Botany, Southern Illinois University, Carbondale, IL 62901. AMERICAN FERN JOURNAL: VOLUME 66 (1976) Pinna forms in Asplenium platyneuron. FIG. 1. var. platyneuron, Montgomery Co., Ark., Taylor 1073 (SIU). FIG. 2. var. platyneuron, Sharp Co., Ark., Taylor 1819 (SIU). FIG. 3. var. incisum, Williamsburg Co., So. Car., Godfery & Tryon 443 (GH). FIG. 4. var. incisum, Garland Co., Ark. Demaree 68587 (SIU). FIG. 5. var. incisum, Fleming Co., Ky., Braun 1750 (GH). FIG. 6. vat. incisum, Middlesex Co., Mass., Davenport s.n. (GH). FIGS. 7 and 8. var. bacculum-rubrum, Prin- cess Anne Co., Va., Fernald & Long 1222] (GH). FIG. 9. var. bacculum-rubrum, Polk Co., Ark., Taylor 1061 (SIU). FIG. 10. f. hortonae, Rensselaer Co., N.Y., Harrison s.n. (GH). FIG. 11. f. ao Jackson Co., Ill., Taylor 878 (SIU). FIG. 12. f. hortonae, Windham Co., Vt., Horton s.n. TAYLOR, MOHLENBROCK & BURTON: VARIATION IN ASPLENIUM 65 and rachis (Fig. 13). The blade is linear-oblong or linear-oblanceolate. Medial pinnae are oblong to oblong-lanceolate and auriculate at the base on their upper and also often on their lower margins. In addition, the pinnae are serrulate and rarely over 2 cm long (Figs. ]-3). Fernald (1950) described typical A. platyneuron as having ‘‘pinnae minutely crenulate, dentate or fine serrulate’’; Morton (1952) referred to the pinnae of sterile fronds as ‘‘remotely serrulate’’ and those of the fertile fronds as ‘‘serrate.”’ 1b. Asplenium platyneuron var. incisum (Howe ex Peck) B. L. Robins. Rhodora 10: 29,1 Asplenium ebeneum var. incisum Howe ex Peck, Ann. Rep. Regents Univ. N.Y. 22: 104. 1869. TYPE: Poestenkill, Rennselaer County, New York, E. C. Howe (NYS, photos GH!). Asplenium ebeneum var. serratum E. S. Miller, Bull. Torrey Bot. Club 4: 41. 1873. TYPE: Wading River, Suffolk Co., New York, E. S. Miller (not located). Asplenium platyneuron var. serratum (E. S. Miller) B. S. P. Prelim. Cat. Pl. New York 73. 1888. Asplenium ebeneum f. serratum (E. S. Miller) Clute, Our Ferns 314. 1901. Asplenium platyneuron f. serratum (E. S. Miller) R. Hoffm. Boston Soc. Nat. Hist. 36: 193. 1922. Chamaefilix platyneuros var. serrata (E. S. Miller) Farw. Amer. Midl. Nat. 12: 270. 1931. The first variation based on pinna characters was described in 1869 by Peck as var. incisum: ‘‘In this form the pinnae are about one inch long, and all except the extreme upper and lower ones are deeply incised-pinnatifid; the pinnules are rather strongly 3-5 crenate toothed. I have thought best to give it the name suggested by its discoverer [Howe].’’ Howe’s type material does posses several nearly pinnatifid pinnae; however, most of the pinnae are doubly serrate or serrate-incised. It appears that the doubly serrate or serrate-incised to nearly pinnatifid pinnae separate var. incisum (Figs. 4-6) from var. platyneuron, al- though numerous intermediates are readily found. Essentially the same taxon was described by Miller **. . . with the fronds wider than usual and the segments very sharply serrate, which he [Miller] proposes to ticket at Dr. Gray’s suggestion as var. serratum.”’ Although we have been unable to locate Miller’s specimens, an examination of authentic material, as determined by Asa Gray, indicates that both var. serratum and var. incisum are referable to the same variety. lc. Asplenium platyneuron f. hortonae (Davenp.) L. B. Smith, Rhodora 30: 14. 1 928. Asplenium ebeneum var. hortonae Davenp. Rhodora 3: 1. 1901. TYPE: Brattleboro, Windham County, Vermont, Sept. 1900, F. B. Horton s.n. (isotypes GH!, US). Asplenium ebeneum f. hortonae (Davenp.) Clute, Fern Bull. 14: 86. 1906. Asplenium platyneuron var. hortonae (Davenp.) Clute, Fern Bull. 17: 21. 1909. Asplenium platyneuron f. dissectum Bened. Amer. Fern J. 37: 11. 1947. TYPE: Lanedon, St. Mary’s County, Maryland, 21 Oct 1945, J. E. Benedict 5230 (US!). Mrs. Francis B. Horton first found the feathery-fronded form of A. platyneuron in September, 1900, in Brattleboro, Vermont. She sent material to Davenport, who described it as A. ebeneum var. hortonae. Davenport commented that the plant was so different in appearance that he first thought it to be a new species. Asplenium platyneuron f. hortonae has been found only sterile. Its pinnae are deeply pinnatifid to pinnatisect, with the ultimate segments ovate to oblong or 66 AMERICAN FERN JOURNAL: VOLUME 66 (1976) g 4 SE ~S TS —ft> <—4A- —f > ieee, es pe et | — > pals at = ; a os be, ae Sa as, = ey os) tee pee ae as” cf ee Lone e Ce 2239) — es —— ty Zo kA aa Ry Ree Est Te tere CH] kx AS 3 i, ow we Bae, ie Tae ‘a tet —i ead tS A Frond forms in Asplenium platyneuron. FIG. 13. var. platyneuron, Montgomery Co., Ark., Taylor 1073 (SIU). FIG. 14. f. furcatum, after Clute (1909). FIG. 15. f. ‘‘multifidum,’’ Upshur Co., W. Va- Tetrick s.n. (WVA). FIG. 16. f. proliferum, after Marshall (1923). TAYLOR, MOHLENBROCK & BURTON: VARIATION IN ASPLENIUM 67 obovate to spatulate and crenulate to serrulate-incised (Figs. 10-12). Due to its sporadic occurrence and distinctive appearance, Clute’s treatment of this taxon as a form is quite logical, although some specimens approach var. incisum or var. bacculum-rubrum. Asplenium platyneuron f. hortonae was found in Jackson County, Illinois, ap- proximately 4 miles SW of Ava, SW % of sect. 10, T8S, R4W, in July, 1971 (Taylor 878, SIU). A single plant occurs with several of var. platyneuron on a west-facing hillside woods dominated by Quercus stellata, Q. velutina, Q. alba, Ulmus alata, and Carya ovalis. Other species in the immediate vicinity of the plant include Fraxinus americana, Ostrya virginiana, Eupatorium rugosum, Sanicula canadensis, Acalypha rhomboidea, Woodsia obtusa, and Botrychium dissectum var. obliquum. The Illinois material of f. hortonae is sterile, with 22-28 pairs of deeply pinnatifid pinnae on fronds up to 35 cm long. The pinnae are up to 3.2 cm long, with the basal pair of lobes usually larger and often at right angles to the pinna midvein. Benedict’s f. dissectum is based on a large and much-divided specimen of A. platyneuron. Although three fronds of the Benedict specimen bear pinnae that are more divided than those of type material of f. hortonae, the other fronds are typical for f. hortonae. In addition, the Benedict specimen is sterile like those of all collections of f. hortonae. On the basis of pinna length alone, Benedict’s collection, with pinnae up to 4 cm long, would fit the dimensions of var. bacculum-rubrum. However, its sterility and pinna dissection place it clearly with f. hortonae. 1d. Asplenium platyneuron var. bacculum-rubrum (Featherm.) Fern. Rhodora 38: Asplenium ebeneum var. bacculum-rubrum Featherm. Rep. Bot. Surv. So. Centr. Louisiana 1870: 75. 1871. TYPE: Near Baton Rouge, Louisiana, Featherman (not located). Asplenium platyneuron var. euroaustrinum Fern. Rhodora 37: 382. 1935. TY PE: Munden, Princess Anne County, Virginia, 1 Aug 1934, Fernald & Long 3603 (GH; isotype US). Featherman described A. ebeneum var. bacculum-rubum from plants found near Baton Rouge, Louisiana as follows: ‘‘Stipe and rachis purplish brown, glossy, tall, one to two feet high. Fronds linear, lanceolate, accuminate, pinnate. Pinnae numerous, sessile, auricled on both sides of the base, coarsely serrate, the pinnae below the middle gradually decreasing in length. Fruit-dots elongated, from twenty to thirty on each pinna. Pinnae distinct.” A year after Fernald described var. euroaustrinum, Fernald corrected his error when he made the combination A. platyneuron var. bacculum-rubrum (Feath- erm.) Fern. This mainly southern variety has fertile fronds which are up to 70 cm long, with each frond bearing up to 70 pairs of pinnae. The pinnae are frequently coarsely serrate-incised and are typically longer than 3.5 cm (Figs. 7-9). The entire aspect of this variety is coarser than that of var. platyneuron and, at its extremes, is found to intergrade with var. incisum. le. Asplenium platyneuron f. furcatum Clute, Fern Bull. 17: 89. 1909. Asplenium platyneuron f. multifidum Tetrick, Amer. Fern J. 39: 92. 1949. TY PE: Upshur County, West Virginia, 21 June 1946, R. M. Tetrick IT (WVA)). 68 AMERICAN FERN JOURNAL: VOLUME 66 (1976) TYPE: Asheville, Buncombe County, North Carolina, Wright (not located). Clute’s f. furcatum is based on a collection sent to him from Asheville, North Carolina by Miss Frances M. Wright and about which he states, ‘‘The plant was normal in all respects with the exception of the fronds . . . which were much branched at the apex.”’ An illustration accompanies Clute’s description (Fig. 14). Apparently unaware of Clute’s description, Tetrick described f. multifidum stating, ‘fronds much branched, the ultimate divisions crested.’ Comparing the holotype of f. multifidum with Clute’s illustration of f. furcatum (Figs. 14 and 15), it appears that these two forms are essentially the same. 1f. Asplenium platyneuron f. proliferum (D. C. Eaton) Tanger, Amer. Fern J. 23: 16. 1933. Asplenium ebeneum var. proliferum D. C. Eaton, Bull. Torrey Bot. Club 6: 307. 1879. TY PE: Near Ocala, Marion County, Florida, Mar-Apr 1879, Capt. J. D. Smith. (isotype US). Asplenium ebeneum f. proliferum (D. C. Eaton) Clute, Fern Bull. 14: 86. 1906. D. C. Eaton described var. proliferum from specimens collected by Captain J. D. Smith near Ocala, Florida. Proliferous plantlets are normally quite small, inconspicuous, and located near the base of the frond (Fig. 16). They have been noted on taxa otherwise referable to var. platyneuron, var. incisum, or var. bacculum-rubrum. The authors are grateful to Dr. Delzie Demaree, Dr. Karl Schwaab, and Mr. John White for their aid in this study. LITERATURE CITED CLUTE, W. N. 1909. Rare form of ferns.-XI. A forked ebony fern. Fern Bull. 17: 88-89. FERNALD, M. L. 1935. Midsummer vascular plants of Virginia. Rhodora 37: 378-413. . 1950. Gray’s Manual of Botany, 8th ed. American Book Com mpany, New York. MARSHALL, M. A. 1923. Proliferous Ebony Spleenwort. Amer. Fern J. 13: 7-1 ee Cee H. 1967. The Illustrated Flora of Illinois: Ferns. Southern Illinois University ss, Carl MORTON, C. V. 1952. Pteridophyta. In H. A. Gleason, Illustrated Flora of the Northeastern United States and Adjacent Canada. New York Botanical Garden, New York. AMERICAN FERN JOURNAL: VOLUME 66 NUMBER 2 (1976) 69 The Distribution of Dryopteris spinulosa and its Relatives in Eastern Canada ONALD M. BRITTON*! There are some excellent distribution maps of vascular plants from Quebec and Labrador in Rousseau (1974). However, his map for Dryopteris spinulosa (O. F. Muell.) Watt actually groups data for all the species of Dryopteris which have been referred to as the D. spinulosa complex, much as Boivin (1966) did when considering D. austriaca (Jacq.) Woynar. This conceals the interesting geographic patterns found in eastern North America, where there are three northern limits in Quebec and Labrador and one southern limit. Biosystematists (Wagner, 1971; Wherry, 1961) recognize two diploid species and two tetraploid species in this complex. The diploids are D. intermedia (Muhl.) A. Gray and D. assimilis S. Walker (which also has been called diploid D. dilatata or diploid D. spinulosa var. americana). The tetraploids are D. spinulosa (O. F. Muell.) Watt (sometimes called D. carthusiana (Villar) H. P. Fuchs) and D. campyloptera Clarkson. After determining that the northern Clay Belt plants near Amos were in fact diploid and not tetraploid D. campyloptera as expected (Britton, 1967), I decided in the tradition of Wherry to attempt to delineate the ranges of the D. spinulosa complex species in Quebec, and more specifically to see if the ranges of the diploid D. assimilis and the tetraploid D. campyloptera overlapped. For studies of genomic analysis and chromatography, a concerted effort was made to find hy- brids between these four species of the complex, which were found growing together at Métis and Mt. Albert in 1968. Dryopteris assimilis was looked for unsuccessfully in Prince Edward Island, Nova Scotia, and New Brunswick in 1970. In 1971 D. campyloptera was studied in Vermont and north in the Lauren- tians to St. Donat de Montcalm and Mt. Tremblant. In 1972 I collected material north of Quebec City and along the north shore of the St. Lawrence River to Sept Iles and from there to Labrador City. The cytological vouchers, which represent over 200 different plants, have supplied material for chemotaxonomic studies (Widén & Britton, 1971; Britton & Widén, 1974; Widén et al., 1975), and are a sample from which extrapolations can be made to similar phenotypes for the purpose of mapping. The maps in this paper have been prepared from selected specimens seen in the following herbaria: GH, MTMG, MT, MTJB, QFA, QUE, QMP, SFS, ULF. Dryopteris intermedia.—There are scattered collections of this species (Fig. /) from the boreal forest, but it is much more abundant in the mixed forests (maple- beech) in southwestern Quebec. One station with cytological specimens is near Amos (ca. 48° 30'N), which is near the northern limit of its range (ca. 50° N). Only four stations are plotted north of this latitutde. Ferns with a similar distribution mapped by Rousseau (1974) with their respective map numbers are: Osmunda cinnamomea 28, Osmunda regalis 30, Dryopteris cristata 46, and Pteridium aquilinum 67. *Department of Botany and Genetics, University of Guelph, Guelph, Ontario NIG 2W1, Canada. 'The author thanks the National Research Council of Canada for financial support for this research. Si i bs ly diated el ci e oeaag es bee ee EE SCOTS Es on ay me es eA Seat Ee a gee rae ESD AP 1 Re ae tS ee eRe Dee coe SN Pui aman ae ray paws oa lees pte ‘POUIULIDIOP A[[BdTOJOIAD suondsa][O9 = W ‘syiwissy “Gq °7T OA ‘Vlpausaiui *q ‘| “‘O{A “Aopeiqge’] pue sagan ul suaidoduq jo uonnquysig = = SS aa pi. : "| => oS, ne 4 , eer \\3 Hoes Fe \ = | WAN Cites k ! : | ahs x Y, ‘ el aS 5 - | | < L577 " nN e 5 } is S Lee eS ame a Le at es ’ \ | ——— | Ae ee tb AR. . ey ar A By Au ad eur | a DON St aes a Grn fo é 5 i] ‘ / / Be T os. é x i | A a ets | Pei aay! ae i > t ve Re Aare ais Pe, (Hae |e 1 oS ™ fap) so wo wo lu = oe oO > = oO -_ = o Lad as od o lu = ie ay 4 z Gs ta 1 hy a I, a ce yr AA ie Fé ss" sa 2 a Se a eS SS D. M. BRITTON: DISTRIBUTION OF DRYOPTERIS IN CANADA 73 sis and the latter species had the same ancestral genomes as D. assimilis; and (4) The analysis of pairing in hybrids of D. assimilis x campyloptera showed more than 41 pairs (46-49) of homologous chromosomes. We now have the geographic evidence to consider. Dryopteris assimilis is a northern species and the derived D. campyloptera has a northern limit not unlike that of D. intermedia. There is very little overlap between D. assimilis and D. campyloptera. This evidence favors the view that D. campyloptera is indeed a derived allotetraploid of D. intermedia and D. assimilis. Wagner (pers. comm.) has pointed out that studies on spore sizes (Britton, 1968) also suggest that one parent of D. campyloptera should have small spores, e.g., D. intermedia, since Dp, assimilis and D. campyloptera have overlapping ranges of spore sizes. In the northern part of the range of D. campyloptera there are great difficulties in separating this species from D. assimilis unless one has cytological as well as morphological evidence. Also, the two species hybridize with each other more readily than with any other Dryopteris species. Their phenotypic similarity suggests that their genotypes must be very similar. In order to separate D. cam- pyloptera from D. assimilis, one must find the very small contribution in D. campyloptera from a parent such as D. intermedia: more finely cut leaf, some evidence of firmer leaf texture and subevergreen leaves, some darker stipe scales, some influence on the shape of the basal pinnae, a more upright form, etc. These are all characters which are difficult to quantify and equally difficult to reduce to a simple, reliable key. Dryopteris assimilis in Ontario.—The only major additions to the distribution map for D. assimilis (as D. dilatata) given by Britton and Soper (1966) are Rez- nicek’s collection at Moosonee and Riley’s collection in the Cochrane District at the latitude of Amos in Quebec. These two collections are connecting links with the collections of D. assimilis on the eastern side of James Bay (Fig. 2), and remove this species from one found only in the Lake Superior basin of Ontario. In 1934, Dr. Wherry collected a specimen (TRT Acc. No. 118558) from Beaver Pond in Algonquin Park. He kindly gave me the exact location for this collection, which he said was unfortunately near the edge of the lake where there would be a good deal of ice movement in the spring. Several prolonged searches in the im- mediate area of this collection have failed to produce any living plants for cytol- ogy. Algonquin Park is noted for the presence of plants such as Saxifraga aizoon and Lycopodium selago with northern affinities, as well as plants such as Picea rubens with Appalachian affinities. Accordingly, this collection of Dr. Wherry could belong to either D. assimilis or D. campyloptera; however, its location in the more mesic hardwoods would suggest that it be referred to D. campyloptera, in which case it is our only specimen for this species in Ontario! Hybrids.—The only common hybrid in this group is Dryopteris x triploidea Wherry (D. intermedia x spinulosa). One can usually find this hybrid wherever D. intermedia and D. spinulosa grow together. A distribution map for this species should be very similar to that for D. intermedia (Fig. 1). The only other hybrid involving just these four species of the Dryopteris spinulosa complex that has been collected in Quebec is D. assimilis x campyloptera, from near Mt. Albert 74 AMERICAN FERN JOURNAL: VOLUME 66 (1976) (Widén & Britton, 1971). It should be stressed that the morphological variation that one finds in these four species can not be attributed to the occurrence of hybrids. A concerted effort was made to find hybrids of D. intermedia x cam- pyloptera, D. campyloptera x spinulosa, D. assimilis x intermedia and D. as- similis X spinulosa, and none were found. They must be of rare occurrence (Widén et al., 1975). LITERATURE CITED BOIVIN, B. 1966. Enumeration des plantes du Canada. ae Canad. 93: 253-274. BRITTON, D. M. 1967. Diploid Dryopteris dilatata from Quebec. Rhodora 69: 1-4. . 1968. The spores of four species of spinulose wood ae (Dryopteris) in eastern North Amnctics, Rhodora 70: 340-347. ———, and J. H. SOPER. 1966. The cytology and distribution of Dryopteris species in Ontario. Canad. J. Bot. 44: 63-79 , and C.-J. WIDEN. 1974. Chemotaxonomic studies on Dryopteris from Quebec and eastern North America. Canad. J. Bot. 52: 627-638. ROUSSEAU, C. 1974. Géographie Floristique du Québec-Labrador. Les Presses de 1’ Université Laval, Québec. WAGNER, W. H., Jr. 1963. Pteridophytes of the Mountain Lake area, Giles Co., Virginia including notes from Whitetop Mountain. Castanea 28: 113-150. - 1971. Evolution of Dryopteris in relation to the Appalachians. Jn P. C. Holt (ed.). The Distribiitional History of the Biota of the Southern Appalachians, Part II. Flora. Virginia Polytech. Inst. Res. Div. Monogr. 2: 147-192. WALKER, 2 — Cytogenetic Studies in the Dryopteris spinulosa complex. II. Amer. J. Bot. 48: 607 WHERRY, a z 1961. The Fern Guide. Doubleday, Garden City, New York. WIDEN, C.-J. and D. M. BRITTON. 1971. A chromatographic and cytological study of Dryopteris dilatata in North America and eastern Asia. Canad. J. Bot. 49: 247-258. , D. M. BRITTON, W. H. WAGNER, Jr., and F. S. WAGNER. 1975. Chemotaxonomic stagies on hybrids of Dryopteris in eastern North America. Canad. J. Bot. 53: 1554-1567. AMERICAN FERN JOURNAL: VOLUME 66 NUMBER 2 (1976) 75 Ferns: Potential In-situ Bioassay Systems for Aquatic-borne Mutagens EDWARD J. KLEKOWSKI, JR. and DAVID M. POPPEL*! It is now apparent that cancer constitutes one of the leading health problems in the United States. A major investment of the energies and resources of the scien- tific community is being made to understand something of the causes of cancer as well as its cures. Parallel, and in many cases included, with these studies are investigations into the mechanisms of mutation of the genetic material (DNA), which, given the heritable nature of the cancerous state in a given cell line, may shed light on the mechanism of cancer induction and growth (Freese, 1971). Equally apparent in recent years is the increasing introduction of man-made chemicals into our environment, either by their disposal in industrial wastes or via their use in pesticides, plastics, food additives, drugs, etc. Environmental groups raise a cry of warning against these pollutants, while biochemists show that an increasing number of the chemicals now present in our soil, air, water, food, and manufactured products are chemical carcinogens and mutagens. The debate has thus begun as to the benefits and risks of using such chemicals. But in order for the crucial debate to be meaningful, much precise information must be collected. Various methods are needed to assay the impact of technology on biology, both in the laboratory and in the field, using rapid microbial systems, critical mammalian systems, and on-site monitoring of native plant and animal populations. Plant geneticists and cytogeneticists can contribute their study of mutations to the problems of cancer and environmental carcinogenesis. The value to cancer research and cancer prevention of studying mutations and mutagens has been demonstrated in recent years by the significant correlation between the cancer- inducing properties of various chemicals and their capacity to induce mutations in DNA in a wide variety of organisms (Miller & Miller, 1971; Ames et al., 1973, 1975). To quote from the Millers’ survey: ‘In summary, it appears that many, perhaps all, chemical carcinogens are potential mutagens. Similarly, many, but possibly not all, mutagens are potential carcinogens.’’ Test systems used to estab- lish these correlations have utilized organisms ranging from viruses, bacteria, fungi, angiosperms, and fruit flies to mammals; even extracts from mammalian tissue are tested in the more practical and efficient bacterial systems. A wide survey of carcinogens, many of them known human carcinogens, in one such bacterial system found that 90% (156/174) of carcinogens tested are also mutagenic (McCann, et al., 1975). ‘ : The presence and activity of these and other suspect chemicals in our environ- ment must be detected and analyzed. Such screening and subsequent elimination from the environment may provide the major means of cancer prevention. We are allowing ourselves to be exposed increasingly to chemical carcinogens and muta- . iversi ‘ setts, Amherst, Massachusetts 01002. iGaue ael eal ne fy lal Sr ont (GB-8721 and GB-31990) and from the Office of Water Resources Research, Department of the Interior under the Water Resources Research Act of 1964, as amended, supported this work. 76 AMERICAN FERN JOURNAL: VOLUME 66 (1976) gens that are being more strongly implicated as factors initiating most human cancer (Cairns, 1975; Epstein, 1974; Higginson, 1969). Cairns (1975) said that ‘‘a substantial proportion of cancer deaths could be prevented by controlling appro- priate constituents of the environment.’’ The problem then, to a great extent, becomes one of characterizing mutagens/carcinogens and detecting their presence and activity in the environment. The screening of entire ecosystems for the presence of mutagens may become a tractable problem if the appropriate bioassay systems are available. One method of approaching this problem is to measure mutation rates in some component of the flora of a given ecosystem. For example, the detection of increased mutation rates in certain species of an aquatic ecosys- tem which is characterized by the presence of certain kinds of industrial pollution certainly would warrant further laboratory testing of the pollutants for mutagenic activity. The value of in-situ plant bioassay systems for mutagens in these situa- tions is that such bioassays serve as continuous and cumulative monitors, and so long periods of time are screened. Samples of water or undiluted effluent will vary in their mutagenicity since given mutagens resulting from industrial pollution vary as industrial production and methods change during the course of time. In con- trast, the mutations of a submersed or semi-submersed plant species in a riparian ecosystem should reflect the cumulative genetic damage that has occurred during a long period of time. This period of time may be a single growing season or many years, depending upon the plant. In some bioassays it may be possible even to date when mutation rates changed; this refinement requires a detailed knowledge of the organography and ontogeny of the plants. This paper discusses a fern bioassay system with which a riparian ecosystem was screened for mutagenic activity and an attempt made at dating the mutational events. For the past seven years, the senior author has been developing experimental techniques for the detection of genetic variability in fern populations (Klekowski, 1970, 1973). Recently these techniques have been applied to fern populations growing in environments that are polluted heavily with industrial wastes (Klekowski, 1975, 1976; Klekowski & Berger, 1976). These techniques, based on the Royal Fern, Osmunda regalis var. spectabilis (Willd.) Gray, involve the detection of both gene and chromosome mutations and whether such mutations have been induced post-zygotically in the parental sporophyte. The Royal Fern is admirably suitable for storing environmentally induced mutations. It has rhizome apices and leaf primordia based upon single apical cells that divide and give rise to all subsequent cells of a given organ. The induction of a mutation in these apical cells results in the mutation being passed to all subsequent cells of the organ. Therefore, as long as the mutations are not dominant cell lethals, they would be expected to accumulate in the genotypes of those organs as the organism grows and as mutations occur. Dominant deleterious mutations, involving those loci which control meiosis and aspects of early embryogeny, as well as recessive and dominant mutations at those loci involved in spore development, gametophyte maturation, gametangia development, zygotic development, and early em- bryogeny may be expected to accumulate in the apical cells of these organisms. Techniques are available to screen the genotype of a sporophyte for just such KLEKOWSKI & POPPEL: IN-SITU BIOASSAY SYSTEMS 77 mutations (Klekowski, 1971). Spore samples obtained from single sporophylls of a given sporophyte can be used to generate cultures of gametophytes in the labora- tory. The frequency of viable and inviable spores can be scored readily. The gametophytes can be scored for normal and abnormal morphology, and the former used in genetic experiments to determine whether their genotypes contain reces- sive or dominant zygotic or sporophytic lethals. Techniques to do this involve the establishment of cultures which have one gametophyte each. Such gametophytes become hermaphroditic, forming both antheridia and archegonia simultaneously, and self-fertilization results in formation of homozygous zygotes from which homozygous sporophytes develop. Such sporophytes can be scored for normal or abnormal development. Meiotic samples taken from the same sporophytes can be screened for chromo- some mutations. It is fortunate that in O. regalis var. spectabilis very large chromosomes are present at meiosis in spite of the fact that 21=44. Two types of chromosome aberrations can be detected readily, both of which require two chromosome breaks for their origin. Reciprocal translocations can be detected at meiosis by the presence of multivalents (associations of four chromosomes at metaphase I or trivalents and univalents). It is the latter configuration which is detected most easily as the univalents very often fail to align in the metaphase I plate and with subsequent divisions of meiosis are left in the cytoplasm as mi- cronuclei. Thus a given meiotic sample can be screened readily for the presence or absence of these micronuclei. Where micronuclei occur, further investigation of metaphase I and the pre-metaphase I stages of meiosis reveals the presence of multivalents. Another chromosome mutation that can be screened for in the meiocytes of this fern is the presence of paracentric inversions. Such chromosome mutations result in the formation of bridges and acentric fragments at anaphase I or anaphase II of meiosis. The occurrence of these configurations is based upon the position of cross-overs in the bivalent containing the inverted segment. The presence of bridges and acentric fragments in a sample of meiocytes can be taken as evidence of inversion heterozygosity (see Klekowski and Berger, 1976, for further discussion on the detection of chromosome aberrations in ferns). In New England the Royal Fern is found commonly in most moist habitats, swamps, and bogs, and very often occurs as a component of the riparian flora. In the latter cases, the fern grows at the edges of watercourses, and very often its rhizomes are submersed periodically. The fern occurs only in rivers where silting is not a normal situation. The Royal Fern population which we investigated grows along the Millers River below Erving, Massachusetts. The Millers River, a tributary of the Connecticut River, is approximately 50 miles long and drains 300 square miles of north central Massachusetts and 70 Square miles of southern New Hampshire. Along its length the quality of the mainstem and its tributary the Otter River varies and is classified from B (good) to D (poor) based on a scale of the Massachusetts Division of Water Pollution Control. The 1.5 mile portion in which the Osmunda population grows Is clas- sified currently as D. The population consists of approximately 100 plants along the south bank of the river one mile below the outfall of the Erving Paper Com- 78 AMERICAN FERN JOURNAL: VOLUME 66 (1976) pany, Inc. Many of the plants have their rhizomes and shoot apices submersed most of the year. Fronds are initiated below the surface and emerge from the water as the fiddleheads uncoil. Only submersed plants were studied genetically. Meiotic samples collected in the spring of 1973 from the Millers River popula- tion revealed that approximately 43% were heterozygous for chromosome muta- tions such as paracentric inversions and reciprocal translocations, whereas less than 1% of meiotic samples collected from nearby, non-polluted control popula- tions gave evidence of such mutational heterozygosity. These control populations were taken from areas within the Millers River watershed. Further analysis of the chromosome mutations present in the Millers River population was undertaken in the spring of 1974. Meiotic collections were made to determine the nature of the cytogenetic chimeras present within the sporophytes. The patterns of chromosome mutations were analyzed in an effort to date the time of induction. After extensive analysis, it was found that practically all the chromosome mutations detected in 1973 and 1974 represented mutations that had occurred since the sporophytes were growing in the Millers River, i.e. were post-zygotic mutations. It was found also that 64% of these chromosome muta- tions were induced since 1969. Thus, it was concluded that the waters of the Millers River were active mutagenically. The frequency of Royal Fern sporophytes which are chimeric for gametophytic and sporophytic lethals also was studied in both the Millers River and control populations. A hybridization program was designed to determine whether the several interconnected rhizome apices resulting from the continued growth of a single sporophyte were heterozygous for allelic lethals (for details see Klekowski, 1976). Where the genotypes of these apices differed with reference to these leth- als, post-zygotic mutations have occurred. Approximately 40% of the sporophytes studied from the Millers River population were chimeric for such deleterious mutants, whereas chimeras were absent from the control population in the non-polluted environment. This value (40%) is remarkably similar to the fre- quency of sporophytes exhibiting chromosome mutations (43%) based upon the cytological investigation previously discussed. The similarity of the genetic and cytological studies suggests that both methodologies give useful estimates of the amount of post-zygotic mutational damage present in the population. Because of the laborious and expensive nature of the genetic investigations (due to the pro- longed culture and maintenance of thousands of gametophyte cultures), it appears that cytological methods offer an easier method of detecting post-zygotic muta- tional damage in Royal Fern populations. These studies of the Royal Fern suggest that other ferns growing in riparian situations may be useful for in-situ bioassay systems of water-borne mutagens resulting from industrial pollution. Species such as Lorinseria areolata (L.) Presl, Matteuccia struthiopteris (L.) Tod., Onoclea sensibilis L., and members of the genus Acrostichum have the appropriate ecologies. But whether these species have the appropriate sensitivity, as well as suitable cytological and cultural characteristics, must be investigated before their usefulness for mutagen bioas- says can be determined. KLEKOWSKI & POPPEL: IN-SITU BIOASSAY SYSTEMS 79 LITERATURE CITED AMES, B. N., W. E. DURSTON, E. YAMASAKI, and F. D. LEE. 1973. Carcinogens are muta- gens: a simple test system combining liver Pegraricwrozay for activation and bacteria for detection. Proc. Nat. Acad. Sci. U.S.A. 70: 2281-2 ——., J. MCCANN, and E. ERP 1975. Methods for detecting carcinogens and mutagens with the Salmonella/mammalian-microsome mutagenicity test. Mutation Research 31; 347-364. CAIRNS, J. 1975. The cancer problem. Scientific American 233: 64-78. EPSTEIN, S. S. 1974. Environmental determinant of human cancer. Cancer Res. 34; 2425-2435. FREESE, E. 1971. Molecular mechanisms of mutations. Jn A. Hollaender (ed). Chemical Mutagens—Principles and Methods for their Detection. Plenum Press, New York. HIGGINSON, J. 1969. Present trends in cancer epidemiology. Canadian Cancer Conf. 8: 40-75. KLEKOWSKI, E. J., Jr. 1970. Populational and genetic studies of a homosporous fern-Osmunda regalis. Amer. J. Bot. 57: 1122-1138. ————. 1971. Ferns and genetics. BioScience 21: 317-321. pent Genetic load in Osmunda regalis populations. Amer. J. Bot. 60: 146-154. 75. Evidence for the presence of mutagenic pollutants in the Millers River: Results of a plant iossy system. Univ. Mass. Water Resources Res. Center Publ. 56. 6. Mutational load in a fern population growing in a polluted environment. Amer. J. Bot. _ (In press). F and B. fet BERGER. 1976. Chromosome mutations in a fern population powns ina polluted en. ent. A bioassay for mutagens in aquatic environments. Amer. J. Bot. 63: 239-246. MILLER, E. C. and J. A. MILLER. 1971. The mutagenicity of chemical carcinogens: correlations; problems and interpretations. Jn A. Hollaender (ed). Chemical Mutagens—Principles and Methods for their Detection. Plenum Press, New York. McCANN, J., E. CHOI, E. YAMASAKI, and B. N. AMES. 1975. Detection of carcinogens as mutagens in the Salmonella/microsome test: assay of 300 chemicals. Proc. Nat. Acad. Sci. U.S.A 72: 5135-5139. Exotic and Hardy Ferns Begonias BOLDUC’S GREENHILL NURSERY 2131 Vallejo Street St. Helena, California 94574 Open Saturdays and Sundays from 10 A.M. to 4 P.M. and by appointment Send a self-addressed, stamped envelope for our list Phone 963-2998—Area Code 707 Mail orders accepted eran ene ee ee 80 AMERICAN FERN JOURNAL: VOLUME 66 (1976) Jf REVIEW -~-— “FERN GROWERS MANUAL,”’ by Barbara Joe Hoshizaki. xi + 256 + xiii pp. 1976. Published by Alfred A. Knopf, Inc. $15.00.—This manual is the best book on fern horticulture on the market at present. It is an instant classic, and is bound to be the reference on fern horticulture for the next decade. Barbara Joe Hoshizaki, in words, but best of all in pictures and diagrams, tells realistically how to grow and propagate many diverse true ferns, as well as the lower vascular plants like Equisetum, Lycopodium and Selaginella. This book goes beyond gen- eral statements (like not burying the crown when transplanting) to explain exactly how to repot ferns with long creeping rhizomes, those with erect, stocky crowns, or those sometimes difficult Staghorn ferns, which grow on plaques like the picture-frame plants that they are. This book should adequately answer most growers’ questions on fern horticulture, with chapters on soils and fertilizers, year around needs, landscaping with ferns, and growing special ferns like the Maidenhair and desert ferns. A long chapter named ‘‘Troubles”’ is probably the best treatment in print of fern pests and the appropriate remedies. The “Manual” is a technical book about a specific subject, yet it is written in non-technical terms. There is a glossary to guide the amateur reader, and three useful appendices are included to help the grower. These tell how to measure light intensity, how to find information and supplies from lists of commercial suppliers, and how to locate the world’s prominent fern societies. _ The simple and direct style makes this book a pleasure to read. It is a little shy in literature citations, especially taxonomic citations: to specific names. This makes it difficult for the reader to find the original taxonomic sources. A number of nomenclatural problems are straightened out, like the disparity between botani- cal names of Tree Ferns and the names used in the commercial trade. The names presented in this volume give us a reliable list of the ferns currently in cultivation in the United States. Very few good, reliable sources of information on fern horticulture are avail- able. This book provides extremely knowledgeable directions on how to grow ferns in the home, the greenhouse, or outdoors. The author has overcome the regional bias shown by many horticultural writers; her book has universal rele- vance. It would have been helpful if all the species and varieties mentioned in the list of 480 plants were pictured because many cultivars are hard to characterize in writ- ten descriptions. But pictures of everything are impossible in a book of this scope. However, an especially useful system of code-words gives a concise and quick key to the structure and cultural requirements for every species listed. The inclu- sion of common and uncommon, easy and difficult varieties in the trade and in cultivation Bives the fern gardener a list of possible plants to add to his collection. For everyone interested in growing ferns this book is an indispensable aid. The informative text, 240 photographs (15 in color), and 57 diagrams make this book a ie rr uce W. McAlpin, New York Botanical Garden, Bronx, NY ENERGY-SAVING DOUBLE-WALL GREENHOUSES FROM ‘h- 499 Now you can grow economically year ‘round in the only greenhouse that solves the energy-cost problem. Lab- oratory tested in Vermont, rigid double- pane thermal walls are 244% more heat efficient, shatterproof, and main- tenance free. Free-standing and lean- to models, no foundations required. eee direct only, 5 year total house warran >” INSULATED WALLS Call or Write for Information P.O. Box 2235, Dept. AF se Central Station w York, N.Y. 10017 ow) 686-0173 A Classic Reprint... ...Now Available The Plants of Southern New Jersey by Witmer Stone After 63 years, this monumental work of Witmer Stone is again avail- able. Encompassing 944 pages bound in green cloth over heavy binders board, this book attractively provides the most complete reference available of the flora of the New Jersey Pine Barrens and adjacent areas. It describes 1,401 individual plant species in infinite detail and includes 129 botanical plates taken from the original 1910 edition. Trees, wildflowers, shrubs, ferns, grasses, rushes, sedges, and quillworts are covered completely and scientifically. A new foreword by Pine Barrens naturalist Elizabeth M. Woodford describes the charm and dedication of conservationist Stone and his love for this ecologically unique land of scrub pine, sand, and brown cedar water. As well as scientifically examining and classifying the flora of the region, this book spins exciting narratives about the things Dr. Stone actually found in this area at the turn of the century. Read about the Curly Grass Fern, local folklore, cranberry picking, and the arduous treks on foot in search of the rare and not-so-rare plants of this special botanical kingdom. The Plants of Southern New Jersey stands as the Single great work on this subject. $30.00 Quarterman Publications, Inc. 5 South Union Street, Lawrence, Massachusetts 01821 TRIARCH 1976 - Our 50th year | of slide manufacture and service to botanists. Your purchases have made our 50 years of exis- tence possible. To satisfy your continued need for quality prepared slides, address your requests for Catalog 17 or custom preparations to: TRIARCH INCORPORATED P.O. Box 98 Ripon, Wisconsin 54971 a Serietrane tt a oan eran ae eee i | i { 2 AMERICAN FERN ges J O U N A [ July-September, 1976 QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY Vegetative Propagation in Asplenium exiguum JOHN T. MICKEL 8! Cold hesaeteaae of Several Ferns n Southeastern Michigan ROYCE H. HILL 83 Variation in Costa Rican Ophioglossum palmatum and Nomenclature of the Species LUIS D. GOMEZ P. 89 Anatomical ‘Studies of the bigatecaes Cyatheaceae. I. Alsophila and Nephele TERRY W. LUCANSKY 93 Prothallus Morphology in some Tectarioid Ferns SURJIT KAUR and SANTHA DEVI 102 The Occurrence of Thelypterin in Ferns G. H. DAVIDONIS 107 Cystopteris fragilis in the Western Himalayas S. S. BIR and CHANDER K. TRIKHA 109 Shorter Notes: Two New Sites for Ceratopteris thalictroides in Texas; Adiantum Capillus- veneris in the Bahama Islands; Vascular Cryptogams at a Site Deglaciated in 1880 is Review Missouri Boranicah OCT 26 1978 GARDEN LISAARY The American Fern Society Council for 1976 DAVID W. BIERHORST, Dept. of Botany, University of Massachusetts, Amherst, Mass. 01002. President RICHARD L. HAUKE, Dept. of Botany, University of Rhode Island, Kingston, R.I. 02881. Vice-President TERRY R. WEBSTER, Dept. of Botany, University of Connecticut, Storrs, Conn. 06268 Secretary JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, Tenn. 37916. Treasurer DAVID B. LELLINGER, Smithsonian Institution, Washington, D.C. 20560. Editor-in-Chief JOHN T. MICKEL, New York Botanical Garden, Bronx, N.Y. 10458. Newsletter Editor American Fern Journal EDIT EF DAVID B. LELLINGER Smithsonian Institution, Washington, D. C. 20560 ASSOCIATE EDITORS DAVID W. BIERHORST ..Dept. of Botany, University of Massachusetts, Amherst, Mass. 01002 GERALD J. GASTONY ....Dept. of Plant ee Indiana University, Bloomington, Ind. 47401 JOHN T. MICKEI w York Botanical Garden, Bronx, New York 10458 The ‘‘American Fern Journal” is an illustrated quarterly devoted to the general study of ferns. It is owned by the American Fern Society, and published at the Smithsonian Institution, Washington, DC 20560. Second-class postage paid at Washington Matter for publication and claims for missing | issues (made within six months of the date of issue) should be addressed to the Editor-in-Chief. Changes of address, dues, and applications for membership should be sent to Dr. Terry W: Lucansky, Dept. of Botany, University of Florida, oe Florida 32601. Orders for back issues should be addressed to the Tre. General inquiries concerning ferns should be oa y to the Secretary. Subscriptions $8.50 gross, $8.00 net (agency fee $0.50); sent free to members of the American Fern Society (annual dues, $5.00; sustaining membership, $10.00; life membership, $100.00). Extracted offprints, if ordered in advance, will be furnished to authors at cost, plus postage. Back volumes $5.00 to $6.25 each; single back numbers of 64 pages or less, $1.25; 65-80 pages, $2. 00 each; over 80 pages, $2.50 each, plus shipping. Ten percent discount on orders of six volumes or more, postage additional. Library and Herbarium Dr. W. H. Wagner, Jr., Department of Botany, University of Michigan, Ann Arbor, Michigan 48104, is teal and Curator. Members may borrow books and specimens at any time, the borrower paying all shipping costs. Newsletter Dr. John T. Mickel, New York Botanical Garden, Bronx Park, Bronx, New York 10458, is er of the newsletter “*Fiddlehead Forum.” The editor welcomes contributions from members and n members, including miscellaneous notes, offers to exchange or purchase materials, personalia, hor ticultural notes, and reviews of non-technical books on ferns. Spore Exchange Mr. Neill D. Hall, 1230 Northeast 88th Street, Seattle, Washington 98115, is Director. Spores exchanged and collection lists sent on request. ‘Gifts and Bequests fie ah ok. : + torested in ferns. Herbarium seecinens. ‘hotanical books, back i issues of the < ira and cash or other gifts are always welcomed, and are tax-deductible. Inquiries should be addressed to the Secretary. AMERICAN FERN JOURNAL: VOLUME 66 NUMBER 3 (1976) 81 Vegetative Propagation in Asplenium exiguum JOHN T. MICKEL*! Asplenium is one of the largest, most diverse, and widespread genera of modern ferns. Many of its nearly 800 species reproduce vegetatively by buds. In different species or species groups, buds have been reported on the roots (4. auritum, A. cuspidatum), in the axils of basal pinnae (4. monanthes var.), in the axils of several pinnae (4A. commutatum), on the rachis near the apex (4. sessilifolium, A. polyphyllum), on an attenuated rachis tip (A. radicans, A. rutaceum, A. rhizophyllum), and plantlets have been observed on the upper surface of the pinnae (4. bulbiferum, A. viviparum), and on stolonoid fronds (A. mannii, A. bipinnatifidum complex). See Faden (1973) for a more detailed treatment of buds and plantlets in Asplenium. The various means of vegetative reproduction in A. exiguum are reported here and its horticultural possibilities are discussed. Asplenium exiguum is a small spleenwort (fronds 3-20 cm tall) with an interest- ing disjunct geographic distribution. It is found in the Himalaya Mountains, the Philippine Islands, Mexico, and the southwestern United States. In 1971 the author discovered it in the Mexican state of Oaxaca near the village of Ixtlan de Juarez, growing on dry, exposed rocky cliffs and shaded ravines. Plants growing on the more exposed sites occasionally rooted at the tip of an extended rachis. This mode of reproduction has been noted by Bir and Shukla (1968), Copeland (1960, p. 442) and Knobloch and Correll (1962, p. 151) Living plants transplanted to the New York Botanical Garden greenhouses have survived well. One plant was placed in our cloud forest chamber (the type described by Farrar, 1968), where it had nearly 100% humidity. After a few months, three methods of vegetative reproduction expressed themselves. A few fronds had prolonged apices with rooting tips (Fig. Je); occasionally a frond forked at the tip (Fig. Jc), resulting in two proliferous apices. Some fronds de- veloped buds at the base of pinnae near the tip of the rachis (Fig. /e); this type has not been noted for this species previously. The most unusual development, how- ever, was that a plantlet developed at the apex of many pinnae, forming a necklace of young plants around the entire frond (Figs. 1b, d). This is most pronounced on the older fronds as they lie close to the ground. Careful examination shows that all pinnae, whether they are on living plants in the cloud forest chamber or the greenhouse or are on herbarium specimens, bear a bud in the notch at the apex (Fig. la). Apparently wild plants only rarely get sufficient humidity to allow the buds to produce plantlets, for only rarely on herbarium specimens was it possible to detect with magnification the slightest germination of these buds, and plantlets were never seen. These pinna proliferations have been noted only once before (Bir & Shukla, 1968), but their illustration (their fig. 27) shows the bud arising on the lower *The New York Botanical Garden, Bronx, New York 10458. 'This work was supported by grant BMS 75-08358 leh the National Science Foundation. I am grateful to Mr. Edgar Paulton for cone the drawin 82 AMERICAN FERN JOURNAL: VOLUME 66 (1976) surface of the lamina. This is incorrect; in all specimens buds arise in the terminal notch of the pinna. I know of no other species of Asplenium or of other ferns that bear buds at the pinna apices. With sufficiently high humidity, apparently this is a very efficient means of propagation in this species. It is curious, though, that the species is found in habitats that do not permit development of these buds. The fact that it produces buds in three different positions is equally remarkable. Many species of Asplenium have been used in cultivation, and Asplenium exiguum may well be added to the list. It seems to be ideally suited for horticul- tural use in terraria or bottle gardens. It can withstand the very high humidity, it grows rapidly, it remains relatively small, it is an attractive plant in its natural habitat, and it is even more striking with the small plantlets along the frond outline. It is propagated readily by placing the plantlets in soil or on wet peat, and can also be grown well from spores. Hopefully, the number of these plants in cultivation will be increased so this species can be more widely distributed. ci 1. Asplenium exiguum. FIG. la. Dorsal surface of pinnae with terminal buds. FIG. 1b. Dorsal = oS of pinnae with plantlets. FIG. Ic. Forking frond tip. FIG. 1d. Ventral surface of pinnae with plantlets. FIG. le. Frond apex with terminal plantlet and rachis bud. LITERATURE CITED BIR, S. S., and P. SHUKLA. 1968. Pteridophytic flora of Simla Hills (North Western Himalayas)-II. Ova Hedwigia 16: 469-482, 1. 79 -182, COPELAND, E. B. 1960. Fern Flora of the Philippines. Monogr. Nat. Inst. Sci. Tech., Manila. 6: 77-557. FADEN, R. B. 1973. Some notes on i i i i i , : the | East Africa. Amer. Peon |. Gs 04.40, gemmiferous species of Asplenium in Tropical cane te 3 R. 1968. A cultural chamber for tropical rain forest plants. Amer. Fern J. 58: 97-102. » |. W., and D. S. CORRELL. 1962. Ferns and Fern Allies of Chihuahua, Mexico. Texas Research Foundation, Renner, Texas. AMERICAN FERN JOURNAL: VOLUME 66 NUMBER 3 (1976) 83 Cold Requirements of Several Ferns in Southeastern Michigan ROYCE H. HILL”*:! Dormancy mechanisms constitute a major adaptation of plants in the colder temperate zones. Dormant organs are especially resistant to winter cold, and it is known that an annual cold period is not only tolerated but is actually required for the resumption of growth in some temperate zone species. A cold requirement for completion of the annual cycle of growth and development in flowering plants native to the temperate zones is well known (Salisbury & Ross, 1969). However, little or no information is available on the role of low winter temperatures in the life cycle of temperate zone ferns. Preliminary experiments suggested that six local fern species differed widely in their dependence on a cold treatment for renewed growth. Three species (Os- munda claytoniana L., Athyrium filix-femina (L.) Roth, and Matteuccia struthiopteris (L.) Todaro) required at least 30 days of outdoors cold treatment, during November-February, to produce appreciable bud-break indoors. The other species (Cystopteris fragilis (L.) Bernh., Adiantum pedatum L. and Thely- pteris palustris Schott) broke bud after a cold period of only 12 days. PROCEDURE Plants of seven species of ferns were collected from their natural habitats during September 23-30, 1970. Fronds of these were cut back to ground level and the rhizomes, including buds, were potted in a soil mixture containing equal parts of sand, loam, and leaf mold. The plants were divided into six groups of ten plants each. The untreated group was placed in the greenhouse immediately after collec- tion. The 12-week cold treatment group was placed in a cold room (a refrigerated storage room kept at 5°C and darkness) immediately after collection for 12 weeks. The other four groups were placed in a holding room at 15°C and 8-hour photo- period for varying periods of time before being placed in the cold room, resulting in different periods of cold treatment. For example, the 8-week treatment group was placed in the holding room for four weeks and then the cold room for eight weeks. With this experimental design, it was possible to remove all groups receiv- ing cold treatment to the greenhouse at the same time. None of the rhizomes broke bud in the holding room, and the cool temperature (15°C) probably was not sufficiently low to satisfy cold requirements of the plants; generally, temperatures effective in breaking dormancy in buds are those below 10°C (Salisbury & Ross, 1969). The groups receiving cold treatment were brought into the greenhouse on De- cember 28. Greenhouse temperatures were 20-30°C during the day and 15-25°C at night. Photoperiods were effectively 24 hours, as incandescent lights illuminated the plants continuously. *Science Department, Huron High School, Ann Arbor, MI 48105. 1] wish to acknowledge the guidance of Dr. Warren H. Wagner, Jr. and Dr. Edward L. McWilliams and the use of the facilities of the Matthaei Botanical Gardens, University of Michigan. 119 © 1014 824 os O. claytoniana a ° J = 64 ° > J c « € za = be i °o 27 “ a . 7 : 3 i : : ? ° oo ° $ 1 2 4 8 2 Chill Period in weeks (logyg scale) 944 ° 80+ . ° A. pedatum 654 8 e . > os z 3 ° > 514 ° a c ° i] E 6 o 374 3 = + z= . BS é ° 224 ; ; i ° eo 4 . . e ee es Oe Chill Period in week lige . 104 } : C. fragilis 9 1 . a > a = 6 ° cd . 8 o c J 5 as ee, + ~ = = 2 & 6 + = . : . 2 4 Chill Period in weeks Number of Fronds Fronds (g) Biomass of Length of Frond (cm) AMERICAN FERN JOURNAL: VOLUME 66 (1976) 7 8 : 64 3 8 t 5 . ¢