American Sern FYFournal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY EDITORS IRA L. WIGGINS C. V. MORTON ROLLA M. TRYON JOHN H. THOMAS VOLUME 54 - 1964- MONUMENTAL PRINTING COMPANY, BALTIMORE, MARYLAND MISSOUR! BOTANICAL GARDEN LIBRARY Contents Votume 54, Numser 1, Paces 1-56, Issurp Aprin 15, 1964 The Genus Cyathea (sensu lato) in Malaysia _R. E. Holttum A New Species and Variety of Bolbitis from India B. K. Nayar and Prakash Chandra The Effect of Sucrose on Apogamy in Cyrtomium falcatum Presl Dean P. Whittier Ferns and es eae re Pine Hills Field Station and Environs (Illino: a: bert T. Mohlenbrock and Jane Hinners Engh Shorter Notes: Litobrochia in Florida Notes and News: 1963 Fern Foray to the Woodstock-Dorset Area of Vermont; Growing Berry Bladder Fern; An Appeal for Help_.. American Fern Society: Annual Meeting; Report of the President; Report of the Secretary; Report of the Treasurer; Report of the Judge of Elections; Report of the Librarian and Curator; New Members; Notice of the Colorado Fern Foray VoLtuME 54, Numper 2, Pages 57-104, Issurp Junp 25, 1964 Development of the Stelar Cylinder in the Rhizome of Bolbitis and Egenolfia Surgit Kaur A New Species of Pyrrosia from India Prakash Chandra Notes on the Hawaiian Fern Genus Adenophorus.__ Kenneth A, Wilson New Combinations in Lycopodium. C. V. Morton Some Hints for the Fern Culturist__ Irving W. Knobloch nese aae Uses of California Pteridophytes by Western American ._—Kobert M, Lloyd _ Phytogeography of Selaginella douglasii..........George Neville Jones — =) 44 Retention of Viability in Lyophilized Spores of the Fiddlehead he Matteuccia pensylvanica__________-_________- W. G. Baker and R. G. Whit Shorter ae Ferns and Fern Allies on Bonaventure Island, Quebec; A Note on Elaphoglossum erinitum; Observations on the Sensi- tive ro Supplement; Growing Mother Fer Recent Fern Literature pat NE Be he Notes and News JSD dee ener As Eh Nei AEA AxeOe Ye Me treane ve cemue mas." Peter ign Porn MOCleby 8 ea eg ee ee ae moblarce GN Votume 54, NumBer 3, PAcEs 105-176, Issuzep NoveMBER 12, 1964 A Preliminary ‘gacindiisl yim Study of Eastern Anericas DryOptonins. = ee Rainer W. Scora and Warren H. Wagner, Jr. Ferns Associated with Ultramafie Rocks in the Pacific Northwest Arthur R, Kruckeberg Seen tink of Polypodium virginianum forma brachypteron (Ridlon) Her eopold A. Charette Sporogenesis in Pteris eretica with Special Reference to the Cytoplasmic Nor PE VSN LCT ee ee ee a il are ae man P. Marengo Florida Strap Ferns and Their Culture.._____-__- Alex D. Hawkes An American Species of Stegnogramma__...___----------. Kunio Iwatsuki Some New Name-combinations for Southeastern Ferns Edgar T. Wherry Camptosorus rhizophyllus forma boycei C. L. Wilson Leopold A. Charette Shorter Notes: Asplenium X< ebenoides R. R. Scott in Kentucky Recent Fern Literature a Notes and News Serene American Fern Society: An Open Letter from the President; List of Member 1 toate aie oo “1 105 PR yer o 12 1 bo c=) 136 141 143 147 VotumE 54, NumsBer 4, Paces 177-216, IssueD JANUARY 15, 1965 Edwin Bingham Copeland (1873- -1964) and His igo that Pteridology______— W. H. Wagner, ys 177 Observations on Drought as es in Selaginella densa Rydb. y R. Webster and Taylor A, Steeves 189 Southwest Vacation... Thomas Darling, Jr. 197 Shorter Notes: Ferns in the Florists’ Trade in 1964.._________- 206 Notes and News: Boston Fern Collection Finds a New Home; Dr. C. Drummond; Edgar T. Wherry Honored______._-----__---------- 207 American Fern Society: Fern Foray; sp cctatelat: fork. 2. Bs: 8. Meetings; List of Honorary Member 2 B09 Index to Volume 54 VoL, 54 JANUARY-MarcH, 1964 No. 1 American Fern Yournal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY ts IRA L. WIGGINS Cc. V. MORTON ROLLA M. TRYON JOHN H. THOMAS ts CONTENTS i The Genus Cyathea (sensu lato) in Malaysia________. R. E. Hotrtum A New Species and Variety of ge from India YAR AND PRAKASH CHANDRA 9 The Effect of Sucrose on aur in Cyrtomium faleatum Presl Dean P. WuirtizeR 20 Ferns and Fern Allies of Pine Hills Field Station and Environs (Illinois) ____ Rosert H, MoHLENBROCK AND JANE HINNERS ENGH 25 Shorter Notes: Litobrochia in Florida 38 Notes and News: 1963 Fern Foray to the Woodstock-Dorset Area of Vermont; Growing Berry Bladder Fern; An Appeal for Help 39 American Fern Society: Annual Meeting; Report of the President; Report of the Secretary; Report of the Treasurer; Report of the — of Elections; Report of the Librarian and Curator; New Members; Notice of the Colorado Fern Foray__ Missour; Boramicay AY APR 21 p64 The American Fern Hociety Council for 1964 OFFICERS FOR THE YEAR Donovan §. CorgELL, Texas Research Foundation, Renner, Texas President Donatp G. Hurrteston, Longwood Gardens, Kennett Square, Sagas ania — LENETTE R. ATKINSON, 415 8, Pleasant Street, Amherst, anucnu ae Ricuarp L. Havxs, Department of Botany, University of Rhode Island, Kingston, Rhode Island reasurer Ina L. Wicatns, Dudley Herbarium, Stanford University, oe Calif. ditor-in-Chief OFFICIAL ORGAN American Fern Journal EDITORS Ira L. *Soccomiemirpmasicnes Herbarium, Stanford University, oe Calif. C. V. Morron_______———SsSmithsonian Institution, Washington 25, D. ©. Rotia M. eon Gray Herbarium, Harvard University, Cambridge 38, Mass. Joun H. THomas_Dudley Herbarium, Stanford University, Stanfo rd, Calif. An illustrated cognbvenk devoted to the aan study of ferns, owned b the American Fern Society, and published | 110 Elm Ave., Baltimore iL Mad. Sonietulam o som pai aid at Baltimor Matter for publication should be wtivtaasd to ne L. Wiggins, Dudley Herbarium, Stanford University, oe Califor Subscription $4.50, exclusive of agenc aaition: seas sent free to —_ bers of the American Fern Society (annual dues, $4. 00; sustaining mem bership, $8.00; life — $80.00). Extracted reprints, if ordered in advance, will be furni authors at cost, plus postage. Back volumes $3.00 aaks tants back numbers 75 eents each; Cumulative = to vols. 1-25, 25 cents, Ten per cent discount on orders of six volumes "Theanine of address, — for membership, — orders for back numbers, and other business communications should be addressed e Treasurer, Dr. Richard L. Hauke, Department of Botany, ” University of Bhode Island, Ringsten, Rhode Island. Members and subscribers should allow two months for changes in address to take effect. LIBRARIAN AND CURATOR OF THE HERBARIUM Dr. W. H. Wagner, University of Michigan, Ann Arbor, Mich. department is * pescamrag in connection with the library rrow i are is published at intervals, to assist those interested in ciahaian rom different localities. American Fern FYournal VoL. 54 JANUARY-Marcu, 1964 No. 1 The Genus Cyathea (sensu lato) in Malaysia R. E. Hotttum I have lately completed a study of the family Cyatheaceae for Flora Malesiana (Holttum, 1963), and have been led to formu- late a new arrangement of infra-generic groupings of species within a comprehensive genus Cyathea. The present paper pre- sents comments on this classification, and incidentally poses some questions which may be relevant to a consideration of the classi- fication of the species of tropical America. In the area covered by Flora Malesiana there are almost 200 species of Cyathea. In the mainland of Asia and the adjacent non-Malaysian islands I can distinguish only 24 or 25; in the Pacific and Australasia are 50 to 60 species (no full compara- tive survey of these has yet been made). There is no species in mainland Asia or the Pacific region which has no close rela- tive in Malaysia. Thus a discussion of the genus in Malaysia applies in essentials also to the species of these other regions. The traditional basis for a classification of this group of ferns long has been the character of the indusium, upon which Rob- ert Brown (1810, p. 158) established the genera Hemitelia and Alsophila as distinct from the earlier Cyathea Smith (1793, p. 416). Although there were some minor differences as be- tween different subsequent authors in the definition of these genera, the generic characters may be thus broadly summarized : in Cyathea the indusium is cup-shaped, in Hemitelia it is at- tached only on one side of the receptacle, and in Alsophila there is no indusium. Careful inspection with the use of modern opti- 2 AMERICAN FERN JOURNAL cal equipment shows this to be an over-simplification, and in- deed Hooker (1844, pp. 28, 34) had some understanding of this, but the use of the three genera persisted for over a century. Copeland (1909, p. 353) was the first to point out clearly that, in the Malayan region at least, the grouping of species provided by the generic definitions is unnatural. His solution was to in- clude all species in one genus, Cyathea; but other authors thought that though this might be right for Malaysia, it was not necessarily so for species of the American tropics, for which reason Christensen continued the use of the three genera in the third Supplement to his Index Filicwm (1934, pp. 20, 58, 110), though in his taxonomic papers on ferns of Malaysia he fol- lowed Copeland (1934, p. 218). Copeland (1947, pp. 98, 99) later atouinied to segregate two genera in Malaysia, famaly Gymnosphaera Bl. and Schizocaena J. Smith, but in my judgment he did not clearly define them and included in each case diverse elements not conforming to the characters of the type species; in particular, he overlooked the characters of the scales (Holttum 1957, pp. 41-45). I believe that it is possible to establish two natural groups of species around the type-species of Gymnosphaera and Schizocaena (in- eluding in each case some that Copeland retained in Cyathea) ; but both groups seem to me to be parts of larger groups, and I do not think the latter are sufficiently distinct to warrant ge- neric separation. This judgment is supported by cytological evidence; the haploid chromosome number in all species investi- gated (including some from each of my major groups), is 69. My two major groups within Cyathea are: subgenus Cyathea, having flabelloid stipe-seales, and subgenus Sphaeropteris (Bernh.) Holttum having setiferous scales. The distinction be- tween these groups is not only in the characters of the edges of the scales but also in their development (Holttum & Sen 1961, p. 410). It appears to me that, judged by other characters also, this division is a natural one, though it is difficult to de- scribe the differences in precise terms. The following is a con- spectus of the subgenera and their sections. CYATHEA IN Manaysia 3 SUBDIVISION OF THE GENUS CYATHEA IN MALAYSIA Stipe-seales flabelloid; hairs on lower surfaces, if present, crisped and appressed; pinnules in most eases deeply lobed, basal basiscopie vein rarely from costa; indusia in many eases hemitelioid. Subgen Paria Indusiate Rene in some eases very small) or if dtndeaatats hairy lower surface of p achis; axes not very dark; Peer: ‘morphin etween ane ty nd ape pinnules Seetion Cyathea Exindusiate; axes very dark, not hairy beneath; fertile att ate pin- les usually very dimorphous Section Gymnosphaera Stipe-scales setiferous; hairs on lower surfaces, if present, rather thick and spreading; where pinnules are shallowly lobed, basal basiseopie vein always from costa; indusia complete, or lacking, e of separate scales oie a fay cases imperfect, and then not hemitelioid). Subgenus SPHAEROPTERIS Costules not widely spaced (rarely over 4 mm. apart in pinnules 10 em long); pinnules usually 10 em. or more long, lobed almost or quite to costa throughout, or fully pinnate..._____ Section Koagire Free tertiary leaflets few; indusia present or absent; s ered with overlapping scales. _............ Subsection ‘Sieacitattenie ie See leaflets many; no indusia; sori covered with ove agli ction Fourn cone, Sande spaced (at least 4 mm. apart except Mie nnn are under 4 em. long); pinnules mostly less than 10 em. long, not lobe to within 1 mm. of costa ee near base; basal pacha vein lways from costa; indusiate WOWuee a Section Schizocaena Seales i stipe 1 cell thick eesnpeaie Cc Subsection Schizocaena Seales of stipe thick and fleshy at base, tapering and flat distally. Subsection Sarcopholis Each subgenus is divisible into two sections. Subgenus Cyathea comprises the sections Cyathea and Gymnosphaera (B1.) ; sub- venus Sphaeropteris comprises the sections Sphaeropteris and Schizocaena (J. Sm.). The sections Gymnosphaera and Schizo- caena have the same type-species as the genera of the same names recognized by Copeland, but a different assemblage of other species. Species which lack indusia, and have therefore in the past been included in the genus Alsophila, are to be found in section Gymnosphaera, section Sphaeropteris and section Schizocaena. In sections Sphaeropteris and Schizocaena there are both indusi- 4 AMERICAN FERN JOURNAL ate and exindusiate species; the exindusiate species in each section seems to me more nearly related to the indusiate ones within the section than to exindusiate species in other sections. In section Gymnosphaera there are no indusiate species, but there are exindusiate species which appear to be on the border- line between sections Gymnosphaera and Cyathea. Thus, in my view, Copeland’s statement that a genus Alsophila defined solely by absence of indusia is an unnatural one is amply justified. But another definition of a genus Alsophila R. Br., and by modern standards a more important one, is that it is a genus which includes the original species of Alsophila, namely A. aus- tralis R. Br. This certainly does not belong to section Gymno- sphaera, as it lacks the characteristic dark axes and constricted fertile lamina; it lacks also small scales with dark median band shown by Miss Tindale to be normally present in that group (1956, p. 331). I would place A. australis in subgenus Cyathea section Cyathea. In its scales it is near C. pruinosa Rosenst. and other indusiate species of New Guinea, but it has no indusium. It has, however, a variable number of small seales attached to the base of the receptacle, and in this resembles Alsophila aspera (.) R. Br. of the West Indies (the seales in A. aspera are much larger than in A. australis). This might seem to justify the inclusion of A. aspera in the genus Alsophila as typified by A. australis. But some other tropical American species which are included in Alsophila (as defined by lack of indusium) lack such seales and are in other ways very different. It is interesting to note that in the subgenus Sphaeropteris a much larger number of species have small scales attached to the base of the receptacle, and in the subsection Fourniera these scales are usually quite large, overlapping and covering the sorus almost to maturity; the subsection Fourniera is characterized also by almost fully tripinnate fronds. The Malaysian members of this subsection (C. celebica BL, @. tripinnata Copel. and others) have been described as indusiate, but are not so; the scales which appear to form an indusium are quite separate, and have the cell-pattern of scales, not of indusia. CyaTHEA IN Mauaysia 5 The section Schizocaena appears to me a very natural one. In it I include the species which Copeland placed in Gymnosphaera sect. 3, which have scales exactly like those of the type species of Schizocaena. I include also some indusiate species (C. im- tegra J. Sm. and allies) which Copeland placed in Cyathea. I exclude the Ceylon species Cyathea sinuata Hook. and C. hookert Thw., which have very different scales and appear to me most nearly related to some species in Madagascar, not to any others in Asia. I have rather tentatively subdivided Schizocaena into two subsections, based on the fact that in New Guinea and the Pacific are species which have stipe-scales arising from ascending fleshy bases; these I place in the subsection Sarcopholis. I am however not sure whether there is a sharp distinction between these species and those with large thin scales, and I have not seen good fresh material of any species with fleshy scale-bases, so that I do not understand their structure and development. Subgenus Cyathea section Cyathea is by far the largest sec- tion, and includes species with indusia of every kind. Sen and I have argued that the Hemitelia type of indusium is primitive in the genus, and homologous with the “inner indusium” of Dicksonia. I suggest that the cup-shaped Cyathea indusium has evolved, perhaps on several distinct lines, from the H emitelia type. There are species in Malaysia which show transitions be- tween a Hemitelia-type of indusium and a shallow cup-shaped one, sometimes on the same leaflet, notably C. javanica BI., and C. sumatrana Bak. In Ceylon, the species Cyathea walkerae Hook., as usually interpreted, comprises forms with shallow cup- sued indusia as well as the normal form with a conspicu- ous hemitelioid indusium, and I have also seen one with a very small indusium attached to one side of the receptacle and quite hidden by the sporangia. Very small indusia of this type are characteristic of several species in Malaysia (e.g., C. latebrosa (Wall.) Copel.) ; such species were placed in the gens Alsophila by Baker and other 19th century taxonomists because with their optical equipment they did not see the indusia. A very similar 6 AMERICAN FERN JOURNAL small indusium oceurs in Alsophila aquilina Chr. of tropical America. Maxon thought this so different from the indusium of species of Hemitelia in tropical America that he saw no obliga- tion to transfer A. aquilina to Hemitelia (1928, p. 317). But in Malaysia there is every gradation from very large hemitelioid indusia to very small ones, and nearly all are dark at the base, as that of A. aquilina. I wonder therefore whether Hemitelia in the American tropics can be clearly distinguished on indusial (or other) characters from Old World species which have been called Hemitelia. Among the latter is H. capensis (L. fil.) R. Br., which occurs also in South America, so that the distinction is not a geographical one. If a clear distinction can be established be- tween New World Hemitelia (in Maxon’s sense) and Old World ones, then the former should retain the name Hemitelia; but I suggest that it ought to be as a section of the subgenus Cyathea. However, a comprehensive monograph of all tropical American species of Cyatheaceae is necessary before the status of such a group can be fairly judged. Just as I think the cup-shaped (typical Cyathea) type of in- dusium developed on more than one evolutionary line, so also the exindusiate condition developed on various lines by loss of the indusium. In Malaysia one can see every stage of reduction down to complete loss, in the species I have placed in section Cyathea. I have not seen such transitions among tropical Amer- ican species, but I have not attempted a full survey of them. I would only call attention to the remarkably similar vegetative character and scales in the type-species of Cyathea, C. arborea (L.) Sm., and in the quite exindusiate Alsophila leucolepis Mart.; surely these two species must be rather closely related. The Malaysian species formerly called Cyathea in the strict sense have sori completely covered by indusia almost to ma- turity. In some cases the indusium finally opens at the top and forms a perfect cup with a smooth rim; in other cases the swell- ing sporangia break the indusium, the final form of which is a cup with torn edges. There are other species the indusia of which superficially resemble this latter condition at maturity. CYATHEA IN MauaysiIA 7 and have been deseribed as cup-shaped, but which have in fact large indusia of Hemitelia-form ; an example is C. oinops Hassk. of Java. The indusium here is attached only to the costular side of the receptacle but covers the top of the sorus like a hood. In C. spinulosa Wall., where the indusium is very thin and is partly lost on breaking, the sori look like a true Cyathea when they are young and like Hemitelia when they are old; specimens in the latter condition were called Hemitelia decipiens. One fact that puzzles me considerably is that the hemitelioid condition does not occur in subgenus Sphaeropteris; one finds only complete indusia which break at maturity (never truly cup-shaped with smooth rim) or none, except in the few cases where there appear to be hybrids, in which various forms of rudimentary indusia occur, but not the hemitelioid condition. This consistent absence of hemitelioid indusia is another indi- cation of the distinctness of subgenus Sphaeropteris. But if the Hemitelia form of indusium is the primitive form in Cyathea, it must have been present in Sphaeropteris ancestors, and presum- ably has died out. The case of the probable hybrid group called Cyathea alter- nans (Wall.) Pr. is of considerable interest. C. alternans is very variable, not only in the extent to which the pinnae are lobed or partially pinnate, but also in the development of the indusium, and there is no clear correlation between indusial form and ex- tent of lobing of the pinnae. Some specimens show quite com- plete indusia, others every gradation down to a small irregular disc round the base of the sorus (only detectable by very careful observation). The presumed parent species are C. moluccana R. Br. (which is simply pinnate and normally has a complete indusium) and C. squamulata (Bl.) Copel. (quite exindusiate and fully bipinnate). ough C. moluccana normally has complete indusia, many specimens have been found, especially in Borneo, which are vege- tatively like normal (. moluccana but have only small fragments of an indusium, much as in some specimens of C. alternans; Copeland gave the names C. pseudobrunonis and C. kinabaluen- 8 AMERICAN FERN JOURNAL sis to such specimens. It seems to me possible that this may be a case where suppression (or almost complete suppression ) of the indusium has passed from one species to another as the result of a long series of hybridization ; but I see no indication of the re- verse process of the development of an indusium in ferns like C. squamulata, The nearest indusiate bipinnate species is C. as- similis Hook., but this seems to be quite distinct. There are tropical American species which (as judged by seale characters) appear to belong to the subgenus Sphaerop- teris; those I have noted are C. erassipes Sod., C. insignis Eaton and (. princeps (Linden.) Meyer. These are all indusiate (in- dusia quite covering young sorus, breaking irregularly at ma- turity) whereas a majority of Malaysian species of subg. S phae- ropteris lack indusia. A careful comparison of these tropical American species with Malaysian ones having similar scales seems to me desirable. After long study of all the many Malaysian Cyathea species, I believe I have found natural groupings among them; but I do not see how those groupings can apply to tropical American species. It appears to me that a full monograph of New World Cyatheaceae is much overdue, and I hope someone will find the time and patience to attempt it. Only by such a comprehen- sive study can inter-relationships within the family be appre- hended. The study of a limited number of species may be help- ful as a preliminary, but conclusions based on a small sample may not be valid when considering the whole. LITERATURE CITED Brown, Roserr. 1810. Prodromus Florae Novae Hollandiae. London. CHRISTENSEN, C. 1934. The ferns of Mt. Kinabalu. Gard. Bull. Str. Settl. 7: 191-324. _ 1934. Index Filicum. Suppl. Tertium, Copenhagen. CopeLAND, E. B. 1909, New species of Cyathea. Philip. J. Sci. Bot. 3: 353-365. / ___-_—, 1947, Genera Filieum. Waltham, Mass. Hourrum, R. E. 1957. The seales of Cyatheaceae, Kew Bull. 1957: 41-45. _____, 1963. Flora Malesiana, Ser. II Pteridophyta, part 2. and U. Sen. 1961. Morphology and classification of the tree New Bouertis From InpIa 9 ferns. Phytomorphology 11: 406-420. Hooxer, W. J. 1844. Cyatheae, in Species Filicum 1: 14-55, MAxon, ee R. 1928. A new tree fern from Haiti. fe Wash, Acad. Sci. 18: 316-317. pow ane 3 < 1793. Paates Pataca ne de filicum generibus dorsiferarum, . Ae. Turin 5: pate M. D. 1956. ree prageeiore. of Australia. Contr. N.S.W. Nat. Herb. 2: 327-361. Kew GArpENs, RicHMOND, SURREY, ENGLAND. A New Species and Variety of Bolbitis from India B. K. NAYAR AND PRAKASH CHANDRA Recently a detailed morphological study of the genus Bolbitis was undertaken by the Pteridology Laboratory of the National Botanic Gardens (India), and in this connection several species uf the genus from different parts of India were collected and studied (Nayar, 1960; Kaur, 1962). During November and De- cember, 1962, two new types of Bolbitis were collected from the Castle Rock area in the Western Ghats of South India. They were transferred to the fernery of the National Botanic Gardens at Lucknow, along with specimens of Bolbitis presliana, B. semi- cordata, B. subcrenata, and B. virens. One of them matches B. semicordata (Moore) Ching, except that its rachis and stipe are narruwly winged and the margins of the pinnae are conspicu- ously lobed. It is described as a variety of B. semicordata. The other is a large fern forming extensive colonies on gravel- ly soil on the western slopes of the hills in deep shade. This ap- pears to be an unrecorded species and is deseribed below. Her- barium specimens of both new ferns are deposited in the Her- barium of the National Botanic Gardens, Lucknow, India. Botsrtis kanarensis Nayar & Chandra, sp. nov. Rhizoma repens ca. 2.5 em. diam., erassum dense ogee filis sclerenchymatis in pulpa centrali dissitis praeditum ; Pp atrofuscae lanceolatae, basi auriculatae, acuminatae, nadatios ciliatae; folia bifaria alterna in dorso ‘rhizomatis ; frondes ster- iles ea. 150 em. longae, pinnatae, apice elongato linzuliformi pen- 10 AMERICAN FERN JOURNAL duli apice bulbifero; stipes usque ad 60 cm. longus, tetragonus, latere adaxiali bicanaliculatus, canalibus in rhachi continuis, basi dense paleatus, aerophoris cariniformibus lateralibus praeditus ; pinnae laterales numerosae, inferiores breviter petiolulatae, su- periores sessiles, supremae coadunatae, lanceolatae, ca. 25 X 2.5 non gemmiferae, apice acuminatae serratae, basi late uneatae, margine irregulariter undulatae; lamina plus minusve coriacea, supra atroviridis lucens, subtus pilis atrofuscis parvis sparsis praedita; venae laterales 30-40-jugae (apice excluso ) fere usque ad marginem distinctae, venulis secundariis 3 vel 4 utrinque latere, venulis 2 basalibus (raro 3) anastomosantibus, areolis goniopteroideis venulis excurrentibus 1—3 liberis vel unitis et ita areolis primariis in areolis secundariis 2 vel 3 divisis; frondes fertiles usque ad 100 cm. longae (stipite 40 cm. longo incluso), pinnis 10 0.2-0.3 em., linearibus, apice obtusis, terminalibus coadunatis plus minusve prolongatis et saepe gem- miferis ; sporae bilaterales 32 X 42 X 32 p, perino rugoso psilato, exino psilato. Type in the herbarium of the National Botanic Gardens, Luck- now, India, collected at Gudkewadi, Castle Rock, North Kanara, Mysore, India, alt. 400 meters, Dee. 9, 1962, P. Chandra 95146. Bolbitis kanarensis (Fig. 1) is a large fern, restricted to deep- ly shaded, moist localities, growing on gravelly soil and forming large colonies on account of its “walking habit.” The rhizome is short-creeping, tenaciously attached to the substratum by strong, wiry roots; it is soft, ca. 2.5 em. thick, and densely paleaceous. The paleae are dark-brown, lanceolate (Fig. 6), acuminate, gland-tipped (Fig. 8) ca. 6 mm. long, basally attached by a broad flat stalk, with small but overlapping basal auricles, and bearing many elongated, multicellular, uniseriate, gland-tipped hairs (Fig. 7) along the margin. The rhizome is parenchymatous, but a few, slender, dark-brown, irregularly cylindrical sclerenchyma strands occur scattered in the pith; sclerenchyma strands gen- erally are absent in the cortex. The stelar cylinder of the rhizome is composed of a broad, slightly curved, ribbon-like, ventral vascular strand and a small, cylindrical dorsal one separated from each other by large, broad, closely placed leaf gaps which alternate on either side. The leaves are in two closely placed alternating rows on the dorsal surface of the rhizome, and a New Boupitris From Inp1ia 11 Figure 1. Puorograru or Type SPECIMEN OF BOLBITIS KANARENSIS NAYAR & CHANDRA, SP, NOV. 49 AMERICAN FERN JOURNAL prominent lateral bud is associated with each leaf base on the side away from the median line of the rhizome. The sterile fronds are large, pinnate, upwardly spreading, and with a long, ribbon-like, terminal pinna which often reaches the substratum and roots at the tip. The stipe is up to 60 cm. long and 5 to 6 mm. thick, densely paleate at the base, glossy green and sparsely paleaceous upward, tetragonal (with smoothly rounded corners), and with two closely placed, parallel, adaxial grooves separated by a prominent, slender median ridge. The ridge becomes in- conspicuous and the grooves merge into one broad, shallow de- pression toward the base of the stipe. A pair of prominent, dis- colored, thin, ridge-like aerating bands (more prominent at the base) occur on both lateral surfaces of the stipe. The lamina is lanceolate in outline. The rachis is up to 80 cm. long, green, sparsely paleaceous, and possesses two broad adaxial grooves separated by a prominent median ridge. The lateral pinnae are numerous, in alternate to subopposite pairs, more or less loosely placed and attached to the lateral margins of the adaxial grooves on either side. The basal pinnae are short-stalked (the stalk 5 mm. long and with a slightly dilated base) ; the upper are sessile and gradually reduced in size toward the apex of the leaf, the uppermost few pairs being coadunate with the terminal pinna. The larger lateral pinnae are 25 * 2.5 em., lanceolate (Fig. 2) and with attenuated, long-acuminate, serrate apices, irregularly wavy margins, and broadly cuneate bases. They are deep green in color, coriaceous in texture, and glabrous above, but bearing brown, glandular hairs all over the lamina and small paleae along the main veins on the lower surface. The midrib is strongly raised on the lower surface; it bears (excluding the acuminate apex of the pinna) 30 to 40 pairs of lateral veins more or less horizontally (Fig. 2). The lateral veins are distinct to the mar- gin of the lamina and bear three to four subopposite or alternate pairs of obliquely placed secondary veins (Figs. 3-5). The basal secondary veins unite in pairs forming two (rarely three) rows of goniopteroid areoles, with one, or usually more, excurrent veinlets in each. The secondary veins are free toward the mar- New Botpsitis From Inpra 13 yj ? f PEST PTS Rh il Figures 2-9, BOLBITIS KANARENSIS; 10-11, B. SEMICORDATA VAR. INCISA. Figure 2. LareraL Pryna. Fires. 3-5. Parts or Lamina, Fic. 3, From Near Apex; Fig. 4, Near Mippte; Fic. 5, Near Base or Pinna. Fig. 6. : RG PALEA (S, STALK). FI Marcin ofr YOUNG PALEA BE G LAR Harrs. Fig Apex oF Mature Para. Fia. 9 RE, EQUATORIAL VIEW PF 0. Part or Racuis SHowing ATTACHMENT OF PINNAE AND WINGS Or B. SEMICORDATA VAR. INCISA. gin and their apices are clavate. The excurrent veinlets are irregular, sometimes uniting with the upper secondary veins and dividing the primary areoles into two or three secondary areoles. In some eases some of the excurrent veinlets from the outermost row of areoles extend to the margin and may unite with an up- 14 AMERICAN FERN JOURNAL per secondary vein on that side (Fig. 4). The areoles are irregu- larly formed and there is often a slight variation in the vena- tion pattern even in the same pinna, but the typically goniop- teroid pattern found in B. subcrenata rarely occurs. The vena- tion is free in the attenuated apical region of the pinnae. Vegetative buds are absent on the lateral pinnae. The terminal pinna is much elongated, ribbon-like, 20 to 35 em. long, grad- ually narrowed toward the apex and bearing a vegetative bud laterally on the dorsal surface of the midrib a few centimeters below the apex. The venation of the terminal pinna is simpler than that of the lateral, with rather distantly placed main lateral veins. The margin is more or less undulate, except at the apex of the pinna, where it is serrate. The outer margins of the dorsal grooves of the rachis merge with the lamina of the terminal na. The fertile fronds are usually smaller than the sterile ones, being up to a meter long, of which the stipe is about 40 em. The laminae of the pinnae are highly reduced. The larger fertile pinnae are up to 10 em. long and 2 to 3 mm. broad, linear, with a blunt apex and parallel sides. The lower are short-stalked. The terminal pinna is coadunate, up to 10 em. long, and often bears a vegetative bud as on the sterile leaves. Sporangia occur over all of the lower surface except on the midrib and the main lateral veins. The annulus is 16 to 18 cells long. The spores (Fig. 9) are bilateral, measuring 32 * 42 &K 324% (PX Ei X FE, exclusive of the perine), with a prominent, smooth perine which is folded into short, thin, sinuous ridges. Bolbitis kanarensis resembles B. virens, but can be distin- guished by its larger size, distinctive venation of the sterile leaf, and by the size and morphology of the spores (Nayar & Kaur, 1963). BOLBITIS SEMICORDATA var. incisa Nayar & Chandra, var. nov. Rhizoma breviter repens dorsiventrale, ca. 1 cm. diam., filis sclerenchymatis in pulpa centrali paucis praeditum ; stipes ‘fron- dium sterilium 15-20 em. longus tetragonus, basi dense paleatus, sursum parce paleatus, sursum latere adaxiali utrinque anguste New Bousitis From InpIa 15 alatus; lamina sterilis lanceolata, 25-30 12 em., pinnis later- oidea, areolis saepe bifariis serie secundaria per ven tiariam venulis primariis parallelem in areolam superiorem ma- jorem (saepe cum venula libera inclusa) et in areolam inferi- lateralibus simili; sporae bilstévales perino granulato in lobis conicis rugosae. Type in the herbarium of the National Botanic Gardens, Luck- now, India, collected at Gudkewadi, Castle Rock, North Kanara, Mysore, India, alt. 400 meters, Dec. 9, 1962, P. ‘Chandra 95145. Bolbitis semicordata var. incisa (Fig. 12) is a small fern easily distinguished from B. semicordata (Moore) Ching var. semi- cordata (see Copeland, 1928) by its lobed pinnae and winged rachis. It grows in small clumps on moist, rocky substrata. The rhizome is short-creeping, dorsiventral, and tenaciously attached to the substratum by strong wiry roots. It is ca. 1 em. thick, more or less soft and brittle, and has a few, rather thick, dark- brown to blackish, sclerenchyma strands scattered in the ground tissue. The ventral vascular strand is broadly gutter-shaped. The paleae are small, lanceolate, basally attached, and pseudo- peltate owing to the prominent overlapping basal auricles. The leaves are more or less crowded in two alternating rows on the dorsal surface of the rhizome, and associated with each leaf is a vegetative bud posteriorly lateral to the leaf-base on the side away from the median line of the rhizome. The stipe of the sterile frond is 15 to 20 em. long, ca. 2 mm. thick, tetragonal, densely paleate at the base, and sparsely so upward. Aerating strands form prominent ridges on each side toward the base. The adaxial surface is nearly flat in the basal half, but grad- ually becomes strongly ridged upward, the ridge being contin- uous on the rachis. On either adaxial margin of the stipe is 16 AMERICAN FERN JOURNAL FIGURE 12. PHOTOGRAPH OF TYPE Or BOLBITIS SEMICORDATA VAR, INCISA. New Bousirts From Inpta 17 a prominent, narrow wing that tapers gradually downwards and becomes inconspicuous nearly halfway down the stipe. The wing is prominent and continuous on the rachis except for interrup- tions at the insertion of the lateral pinnae, and curved upward so as to form two dorsal grooves, one on either side of the me- dian ridge of the rachis (Fig. 10). The lamina is lanceolate in outline, broadest slightly above the base, 25 to 30 em. long, and ca. 12 em. wide. The rachis is rounded on the lower surface; the upper has a prominent slender median ridge and two nar- row grooves on each side of it. The lateral pinnae are alternate, sessile, or the lower ones short-stalked, oblong-lanceolate, ca. 5 em., acute or rarely more or less acuminate at the apex, and devoid of vegetative buds. The bases of the pinnae are un- equal. The upper base is more or less auricled and often over- lapping the rachis on its lower surface; the lower base is ob- liquely cuneate (Figs. 13-15). The pinnae are generally reduced towards the apex of the leaf, the uppermost often being coadu- nate with the terminal pinna. The terminal pinna is more or less elongated, but not ribbon-like, and often bears a dorsal veg- etative bud on the midrib a little below the apex. The margins of all the pinnae are prominently lobed, with the lobes nearly semicircular and the incisions between them extending nearly a quarter of the way to the midrib (Figs. 13, 14). Each mar- ginal lobe corresponds to a main lateral vein of the lamina. The ultimate margin is crenate, each tooth receiving one of the free ending veinlets. Usually there is a prominent tooth at the bot- tom of the incision between the marginal lobes. This is usually red-tinged and curved towards the upper surface of the leaf, recalling the seta in some species of Egenolfia (Copeland 1947; Kaur, 1960; Holttum, 1954). The midrib is strongly raised on the lower surface and forms a blunt ridge on the upper surface. This ridge interrupts the lateral wing of the rachis at the base, but is not continuous with the median ridge of the rachis. The lat- eral veins are in subopposite or alternate pairs, ca. 5 mm. apart, distinct to the margin, and slightly raised on the upper surface. The areoles are generally in two rows on each side of the mid- 18 AMERICAN FERN JOURNAL ‘ FIgurRES 13-15. LATERAL st INNAE OF BOLBITIS SEMICORDATA. Fias. 13, 14. B. SEMICORDATA VAR. IN Fig. 15. BoLsitis SEMICORDATA VAR, SEMI- CORDATA rib; the costal areoles are large and the second row often is di- vided longitudinally into two, with the anterior secondary areole sometimes having a small excurrent, included veinlet. Beyond the second row of areoles there are many excurrent veinlets that New Bousitis From Inpta 19 are nearly parallel, extend to the margin, and have clavate tips. Both surfaces of the lamina are glabrous. There are a few nar- row paleae on the ventral surface, where the pinnae are at- tached to the rachis. The fertile fronds are seasonal, and up to 40 em. long, in- cluding the stipe which is ca. 15 to 20 em. long. The fertile pin- nae are up to 4 em. long and 2 mm. broad, with bluntly rounded apices. The terminal pinna is small and resembles the lateral ones. The wing on the rachis of the fertile frond is not promi- nent. The spores are bilateral, 25 & 37 & 28u (P X E, X Es, exclusive of the perine), and with a prominent, faintly granu- late perine which is folded into a few obconical lobes. Bolbitis semicordata var. incisa differs from var. semicordata in the deeply lobed margins of its pinnae (see Figs. 14, 15), and in having a prominently winged rachis. Both varieties grow together in localities where they are restricted to deeply shaded forest beds, growing generally on rocky substrata or on gravelly soil, and forming small clumps We gratefully acknowledge the keen interest evinced in the work by Professor K. N. Kaul, and C. V. Morton’s help in pre- paring the Latin descriptions. LITERATURE CITED CopeLAND, E. B. 1928. pies and genera confused with it. Phil. Jour. 6 33-4 947. Genera Pilisien. Waltham, Mass. Hourrum, R. : 1954. Flora of Malaya. II. Ferns. Singapore. Kaur, S. 1962. Bolbitis, sn an pe ferns. Ph. D, Thesis. Agra rsl Nayar, B. K. 1960. ona gad of two Indian species of Bolbitis. Jour. Indian hh Soe. 39: 259-277. D Kaor, 8. a. Spore morphology of some Indian mem- bers of ne Lomariopsidaceae. Pollen et Spores. 5: 87-94. Prertmo.ocy Lasoratory, NaTionaL Botanic GARDENS, LucK- Now, INnpIA. 20 AMERICAN FERN JOURNAL The Effect of Sucrose on Apogamy in Cyrtomium falcatum Presl' DEAN P. WHITTIER The phenomenon of apogamy provides an excellent oppor- tunity to study the direct origin of a vascular plant from a non- vascular fern gametophyte. Besides the investigation of the change from the gametophytic to the sporophytic developmental pattern, apogamy which is a deviation in the normal life cycle provides a site to study the factors controlling the alternation of generations. Studies by Whittier and Steeves (1960, 1962) have demon- strated a new method for controlling induced apogamy, i.e. the formation of a sporophyte directly from the vegetative cells of a gametophyte which is able to form sporophytes by fertilization under other conditions. Several species of fern gametophytes grown in sterile culture on nutrient media containing suitable concentrations of sugar produce apogamous sporophytes. In the absence of sugar no apogamous plants are formed by the gameto- phytes. The availability of supplementary sugar to the gameto- phytes has a direct relationship to the formation of the apoga- mous plants Obligate apogamy, i.e. the formation of a sporophyte directly from the vegetative cells of a gametophyte which is unable to form sporophytes by fertilization, has been described in many ferns, but this type of apogamy has been the subject of few experimental investigations. In one such study with Pteris cretica I., Bell (1959) reported apogamy oceurred sooner on He sopelion with sugar than others without sugar. In iew of this report and the effect sugar has on induced apogamy, oe study was undertaken to determine if sugar influences obli- gate apogamy other than accelerating its appearance. 1This investigation was carried out with assistance from the Virginia Agricultural Experiment Station and the University of Virginia, Mountain Lake Biological Station. APOGAMY IN CYRTOMIUM 21 MATERIALS AND METHODS Cyrtomium falcatum Presl. was chosen for this study because the spores of this ferns were readily available and the develop- ment of the apogamous plant had been described (DeBary, 1878; Allen, 1914). The spores were wetted with a 0.1% solution of Tween 80 and exposed to a 15% Clorox solution for two minutes to sterilize them. The sterile spores were collected on filter paper, washed several times with sterile water and suspended in sterile water. The spores were innoculated into square one ounce bottles con- taining 15 ce of nutrient medium. The nutrient medium was composed of Knudson’s solution of mineral salts, minor elements and 0.5% agar. This medium differed from the one reported by Whittier and Steeves (1960) in that the iron was supplied as ferric sodium ethylenedianime tetra-acetate (Fe-EDTA) instead of ferric citrate. Ina preliminary experiment the fastest growth took place on a medium containing 2.5% sucrose, therefore this concentration along with 0, 0.5 and 6.0% were employed in this investigation. The cultures were maintained at a temperature of 24 + 1° C with 12 hours of illumination every 24 hours from Sylvania Gro-Lux lamps at an intensity of ca. 100 foot-candles. The remaining sterile culture techniques were those employed by Whittier and Steeves (1960). The area of the gametophytes was found with a microprojec- tor and a polar planimeter. Tracings of the projected prothal- lial images were measured with the polar planimeter. The mean area of twenty gametophytes was employed as a measure of pro- thallial growth. The size of the gametophytes undergoing initial apogamy was determined by calculating the mean area of twenty prothalli with early stages of apogamous development. The mean cell size of the gametophytes was found by dividing the cell number of twenty similar sized gametophytes into the area of these gametophytes. A T—test was employed to determine if the differences between the responses on the concentrations of sucrose were significant. 22 AMERICAN FERN JOURNAL RESULTS The spores on all concentrations of sucrose germinated the seventh day after innoculation. The least growth took place on the medium without sugar and the largest growth on the 2.5% sucrose medium (Table I). The differences between the growth on the various media were significant at the 5% level. TABLE I. THe Errect oF SuCROSE ON APOGAMY AND THE GAMETOPHYTE OF CYRTOMIUM FALCATUM. Sucrose Mean Prothallial anes: “ = mm Day First Mean pane Size 0 Day Ini X th Apogamy Apogamy 0.0 0.430. oo Hey 041 41st 0. 0021-0. ‘00011 0.5 1.45+0.09 1.85+0.04 35th 0.00210.0001 2.5 1.89+0.08 1.46+0.02 30th 0.00220.0001 6.0 0.800.05 1.050.03 36th 0.00230.0001 1Standard error of mean. The prothallial development proceeded normally from the filamentous to the cordate stage. Some time after the cordate shape was attained a pale green area, which later turned brown, appeared a short distance behind the sinus. This pale green region formed behind the sinus after the cordate prothallus had become more than one cell thick in that area. Although this light region was not initially apogamous, it was considered the first stage leading to apogamous development because the apog- amous sporophyte originated from the cells of this area. The initiation of the apogamous plant occurred once the pale green area had become a few cells thick. On the 30th day the apogamous development was initiated on the 2.5% sucrose medium (Table I). This was five or six days earlier than on the other sucrose media and eleven days before apogamy started on the medium lacking sugar. At the time the apogamous plants were being initiated on the 2.5% medium, none of the prothalli without exogenous sugar were more than one cell thick or beyond the early cordate stage. The decrease in the size of the prothalli undergoing initial apogamy with the increase in the sucrose concentration was sig- APOGAMY IN CyYRTOMIUM 23 nificant at the 5% level (Table I). The mean cell size of the prothalli on the various sugar concentrations did not vary sig- nificantly from each other (Table I). Since the prothallial size is determined by the cell number and cell size, the decrease in prothallial size was due to a reduction in the number of cells per gametophyte because the mean cell sizes were not signifi- cantly different. DISCUSSION AND CONCLUSIONS The growth rate of the prothalli was increased significantly above the rate on no sugar by the addition of sucrose to the medium, although at 6.0% sucrose the growth decreased signifi- cantly from that on 2.5% sucrose. This decrease in growth on the high concentration of sugar probably was due to the high osmotic potential of the medium interfering with the growth. This has been found to be true in other studies on the growth of gametophytes in sterile culture (Mitra & Allsopp, 1959; Whittier, 1962). Besides the change in growth rate the size of the prothalli undergoing initial apogamy decreased significantly as the con- centration of sugar increased. This observation does not support Bell (1959) who reported no change in gametophyte size under- going apogamy on different concentrations of sugar. This dif- ference in size was due to a reduced number of cells in the ga- metophytes because the mean cell sizes on the various concentra- tions of sugar were the same. Consequently, less growth and fewer cell divisions were required for the prothalli to initiate apogamy on the media containing sugar. In terms of the number of days for the occurrence of apog- amy, it was initiated on the 2.5% sucrose medium about 15% sooner than on the other sucrose media and 25% sooner than on the medium without sugar which is in agreement with Bell’s report on P. cretica (1959). This was due to the gametophytes on the 2.5% sucrose medium having the fastest growth and a small prothallial size which would produce apogamy. It can be concluded that the time necessary for obligate apogamy is de- 24 AMERICAN FERN JOURNAL termined by the growth rate and the prothallial size undergoing apogamy on any concentration of sucrose. The sugar brought about the requisite conditions for obligate apogamy in the gametophyte sooner with fewer cell divisions and less prothallial growth. One of the required conditions is the thickening of the gametophyte which always precedes the development of the light green area behind the sinus and the apogamous plant. In other species the thickening of the pro- thallus before the initiation of the obligate apogamous plant has been reported (Duncan, 1943). Also, a thickened prothallial i i 1962; Whittier & Steeves, 1960). Thus, the thickening of the gameto- phyte is a requisite condition for both types of apogamy. In sterile culture the thickening necessary for induced apogamy, which is many times the thickening required for obligate apog- amy, is controlled by the sugar in the medium. Without sugar the necessary thickening of the gametophyte for induced apog- amy is absent, and apogamy fails to occur. The supplementary sugar causes the thickening required for induced apogamy and accelerates the normal thickening preceding obligate apogamy probably by modifying the carbohydrate metabolism of the pro- thallus. LITERATURE CITED V ALLEN, R. F. 1914, ema in spermatogenesis and apogamy in ferns. Trans. Wis. Acad. 17: BELL, P. R. 1959. The po investigation of the pteridophyte life cycle. Jour. Linn, Soc. Bot. 56: DeBary, A. 1878. Uber apogame Farne und die Erscheinung der Apog- amie im allgemeinen. Bot. Zeit. 76: 449-487. v/ hanes: R. E. 1943. Origin and development of embryos in certain apog- us forms of Dryopteris. Bot. Gaz. 105: 202-211. a eas a C. & A. Auusoprp, 1959. IT, The effects of sugar concentration on the development of the protonema and bud formation in Pohlia nutans (Hedw.) Lindb. Phytomorphology 9: 6 . Wurrtter, D. P. 1962. The origin and development of apogamous struc- tures in the gametophyte of Pteridium in sterile eulture. Phytomor- phology 12: 10-20 Ferns or Pine Hits 25 Wuirtirr, D. P. & T. A. Srerves. 1960. The induction of apogamy in the bracken fern. Canad. Jour. Bot. 38: 925-930. _ 1962. Further studies on induced apogamy in ferns. Canad. Jour. Bot. 40: 1125-1131. DEPARTMENT OF BroLoay, VirGINtIA PoLyTECHNIC INSTITUTE, BLACKSBURG, VIRGINIA, Ferns and Fern Allies of Pine Hills Field Station and Environs (Illinois) Rosert H. MomLENBROCK AND JANE HINNERS ENGH This is one in a series of studies which have emanated from the Southern Illinois University Field Station. A check-list of vascular plants has been prepared (Mohlenbrock & Voigt, in press). The study of the ferns and their allies of the area was completed during the summer of 1961 when the junior author participated in a National Science Foundation Research Par- ticipation Program.'? The authors are grateful to the graduate council of Southern Illinois University for assistance in this study. THE Pine HILLS AND ENVIRONS Two representative areas were chosen for exhaustive study of the fern and fern ally groups native to the Pine Hills and its environs. These areas were the Pine Hills Recreation Area and the adjoining Union County Forest Preserve. Both areas are located in the Shawnee Hills of Southern Illinois. Pine Hills Recreation Area is approximately 37 miles from Carbondale, Illinois, off of Illinois Route 3. The territory cov- ered in the survey extended from the second set of railroad tracks on the Aldridge Levee Road to Hutchins Creek, and from the Pine Hills Recreation Area entrance to the edge of Otter Pond. All of the roads, trails, pienie areas, and the penetrable interior of the Recreation Area were covered. IN. S. F. Grant 16,180. 2Contribution from the Pine Hills Field Station and the Department of Botany, Southern Illinois University. 26 AMERICAN FERN JOURNAL A less intensive search of the Union County Forest Preserve was conducted since there was considerable uniformity in eco- logical habitats in this area. Collections were made along roads, fire trails, picnic areas and a portion of the lower interior region near the entrance. History oF FERN COLLECTING IN THE PINE Hitis AREA Prior to June, 1961, only nine species of ferns had been re- ported from the Pine Hills Recreation Area. These were: As- plenum pinnatifidum (Hatcher, Oct. 21, 1949); A. resiliens (Hatcher, Oct. 21, 1949); Athyrium pycnocarpon (Mohlen- brock, July 1, 1954); Dryopteris marginalis (Evers, Oct. 2, 1948) ; Onoclea sensibilis (Mohlenbrock, July 1, 1954); Pellaea atropurpurea (Hatcher, Oct. 21, 1949) ; Polypodium polypodio- ides (Hatcher, Oct. 8, 1949) ; Peiustken acrostichoides (Moh- lenbrock, July 12, 1954) ; ; and Woodsia obtusa (Hatcher, May 28, 1954). Six additional species had been found near the Pine Hills area. Five of these were collected in the Union County Forest Preserve. They were: Adiantum pedatum (Sanders, June 26, 1952; Asplenium platyneuron (Kaeiser, Spring of 1952) ; Dryopteris austriaca var. intermedia (Sanders, April 7, 1952; D. austriaca var. spinulosa (Hatcher, May 21, 1949) ; and Botrychium dissectum var. obliquum (Swayne and Bailey, Sept., 1949). Azolla mexicana (Bailey, Sept. 22, 1947), was found floating on the waters of Wolf Lake. During the present study, 12 species previously unreported from the Pine Hills-Union County Forest Preserve were found: Asplenium trichomanes, Athyrium filix-femina, A. thelypter- woides, Botrychiwm virginianum, Camptosorus rhizophyllus, Chetlanthes feet, C. lanosa, Cystopteris bulbifera, C. fragilis, Thelypteris hexagonoptera, Ophioglossum vulgatum, and Pter- idium aquilinum. Only one fern ally, Equisetum hyemale, was found. EcoLocy or SOUTHERN ILLINOIS FERNS AND FERN ALLIES The ferns and fern allies of Southern Illinois that occur in our areas are listed in Table 1, with check marks to show their Ferns or Pine His 4 § usual environmental habitats. Table 2 lists ferns known from Southern Illinois, but not yet found in the Pine Hills-Union County Forest Preserve areas. ABLE 1 FERNS AND FERN ALLIES OF SOUTHERN ILLINOIS KNoWN TO OCCUR IN THE Pine Hmus RECREATION AREA AND THE UNION County ForREST PRESERVE, IN RELATION TO HABITAT he n pa i n > 3 3S oF 8 - Set ee ee 8 OR tee tee ae eee, gm aa = a a ej = ee = 3 | we Go eee ck od SE he AO 36 a BS 6 BF Aa AE ak Adiantum pedatum x Asplenium pinnatifidum Asplenium platyneuron x Ka Se splenium resiliens Be Asplenium spare x A fil x x & Athyrium pyecnocarpon x = inca Dekkedclics x zolla mexicana - Botrychium dissectum x x x Botrychium virginianum x x Camptosorus rhizophyllus x x x Cheilanthes eei x Cheilanthes lanosa x4 Cystopteris Btateck x se Cystopteris fragilis x * Dryopteris austriaca coh Pia Dryopteris margina x - Equisetum hyemale . Onoclea sensi x . Ophioglossum vulgatum x Pellaea atropurpu x olypodium polypodioides x = Polystichum acrostichoides x * Pteridium aquili x Thelypteris saat z Woodsia obtu * 28 AMERICAN FERN JOURNAL The Pine Hills Field Station is a region of limestone bluffs, moist woodlands, and swamps. It provides a variety of ecological habitats for ferns and fern allies, limited primarily by the ex- treme temperature ranges (—22 to 106°F.) in southern Illinois. High temperatures during the summer cause a correspondingly high transpiration rate in the plants during the time when water supplies in the soil and creeks are diminishing. Plants that survive this seasonal drought must either be adapted to high temperatures or grow in cooler, moister habitats. The woodland and water ferns are examples of plants that occur in a more temperate environment. The woodland ferns grow in moist, shaded sites, protected from direct sunlight most of the day. They tend to be tall, dark green, and delicate in texture and cutting of leaves. Aquatic ferns have an even greater water requirement than do woodland ferns. They grow in or on the surface of still, shal- low ponds or at the swamp’s edge. Equisetum frequently grows near creeks and rivers, not so much because it needs a great quantity of water, but because it requires large amounts of silica found in the sand and gravel of creek banks. For this reason, Equisetum is also common on sandstone rocks and beside well- graveled railroad tracks. The rock ferns have become adapted to extremes in temper- ature. Ferns best able to live in xerophytic or desert-like con- ditions grow on limestone, They are protected from excessive transpiration by having thin, wiry stems and small leaves pro- tected either by a covering of hairs or by a thick, leathery cuticle —or both. The smaller size of the plants enables them to expose less surface from which water escapes, while a thick cuticle helps to keep moisture inside the plant. Ferns that can survive with little water but not able to exist on the desert-like limestone out- vapor between their bodies and the rock, thus reducing the desiccation of their leaves, They also have a cuticle and thin Ferns or Pine Huis 29 FERNS AND Fern ALLies or SouTHERN ILLINOIS Not Founp IN THE PINE Hitis Recreation AREA AND THE UNION County Forrst PRESERVE, IN RELATION TO Hasirat Swamps and swamp margins Moist woodlands Stream beds Dry woodlands Open fields Limestone rock Caleareous soil Wet rocks and ledges On tree trunks Ditches Ponds and stagnant water 4 | Sandstone rock Asplenium bradleyi Dennstaedtia punctilobula quisetum arvense wo richomanes boschianum x stipes, but these features are not as highly developed in the sandstone and acid soil ferns. Rock ferns are scattered throughout bluff-top areas of the upper Pine Hills road. They are abundant nowhere, but the greatest variety of them occurs on land of the Southern Illinois University Field Station. he woodland ferns grow tallest and are most prolific on damp, wooded hillsides and along ditches or creeks. The best 30 AMERICAN FERN JOURNAL area for these ferns is near the picnic grounds at the entrance to the Pine Hills. At the Union County Forest Preserve they are abundant in all the shaded areas. Aquatic ferns occur on the wet mud or on the surface of swamps and ponds of the Pine Hills. A colony of Equisetum hyemale, the only fern ally found in the Pine Hills, grows on the banks of Big Muddy River near the railroad tracks. Loca DistrisuTION oF SOUTHERN ILLINOIS FERNS OPHIOGLOSSUM—ADDER’S TONGUE FERN These are short, green ferns, often overlooked because of their small size. They have an erect, scaleless rhizome with a smooth, fleshy stipe and an entire leaf blade. Arising at a point about half way up the stem is a fertile stalk bearing sporangia distally. The gametophytes are tuberous and exist underground for long periods. In the spring, the young sporophytes push straight up from the soil, unlike most ferns which arise in a coiled position. OpHIoGLossuM vuLGatum L. Adder’s Tongue Fern.—Local in moist woods throughout southern Illinois. A colony of var. pycnostichum Fern. occurs in a frequently inundated woods near Winter Pond in the Pine Hills. Variety pseudopodum (Blake) Farwell, not known to occur in the Pine Hills, is known from only a single station in southern Illinois. OPHIOGLOSSIUM ENGELMANNU. Prantl. Limestone Adder’s Tongue Fern.—Unknown from the Pine Hills Area, although to be expected on limestone ledges. Known from Randolph, John- son, and Hardin counties. BotrycHIuM—GRAPE FERN Both species of Botrychium found in southern Illinois grow in the Union County Forest-Pine Hills area. In our region, they are usually about 8-12 inches tall, with cut or lobed sterile blades. In one species, the leaves turn bronze-colored in the fall. They have a soft, fleshy stipe and a short, erect rhizome as in Ophioglossum, but their leaf veins are open and forked, unlike the closed veins of Ophioglossum. Ferns or Pine Hits 31 BorrRyCHIUM VIRGINIANUM (L.) Sw. Rattlesnake Fern— Abundant in moist woods or dry shaded areas throughout south- ern Illinois. BorrRYCHIUM DIssecTUM Spreng. Grape Fern.—Two varieties occur in southern Illinois, the much-divided var. dissectum, and the less-divided and more common var. obliquum (Muhl.) Clute. Variety dissectum is restricted to moist woods, while var. ob- liquum occurs also in dry open fields and ditches. OSMUNDA—ROYAL FERN No representatives of this genus have been found in the Pine Hills area, although O. regalis, O. cinnamomea, and O. claytoni- ana are known from southern Illinois. Members of this group have scaleless rhizomes and fibrous roots. OsmuNnDA REGALIS L. Royal Fern.—Only var. spectabilis (Willd.) A. Gray is found in southern Illinois. It differs from var. regalis in the absence of black hairs from the stems. Native in swamps, bogs, and on very wet ledges, although in southern Illinois it inhabits moist sandstone ledges. OSMUNDA CLAYTONIANA L. Interrupted Fern.—Reported from Union County in 1955, where it is very rare. It usually grows in moist woods and along swamp margins, although it occurs on sandstone ledges in southern Illinois. It has not been collected in the Pine Hills. OsMUNDA CINNAMOMEA L. Cinnamon Fern.—Oceasionally on damp ledges in southern Illinois. It is rare in this part of the State and unknown from the Pine Hills. TRICHOMANES—FILMY FERN In Illinois this genus has been collected only in Pope and Johnson counties. Trichomanes boschianum Sturm has a scale- less, branching rhizome with black root hairs. Its leaves are very fragile, being only one cell thick. They are cut into about six overlapping divisions. A distinctive feature is the character of its sori, with their tubular indusium with its included hairlike bristle. The fern grows only in damp acid soil in caves or in pockets on sandstone cliffs. 32 AMERICAN FERN JOURNAL AZOLLA—MosquiTro FERN Azolla mexicana Schlecht. & Cham., the only species reported from southern Illinois, is a tiny, ribas ik plant floating on the swamp waters in the Pine Hills area. A cluster of this little fern is about as large as a thumbnail. It has branched rhi- zomes, tiny, twice-lobed leaves, and sporocarps at the leaf bases. The fronds often become reddish in late summer. MARSILEA—PEPPERWORT Marsilea quadrifolia Li. is also an aquatic. It has been found twice in southern Illinois, both times in ponds in Carbondale. It has a branched rhizome rooting at the nodes, a frond that resembles a four-leaf clover, twice-branched veins, and sporo- earps at the base of the leaf stalks, with each sporocarp con- taining several spore cases. ADIANTUM—MAIDENHAIR FERN ADIANTUM PEDATUM L.—Common in the Pine Hills area, and usually grows in wooded areas along streams and occurs com- monly throughout southern Illinois. ASPLENIUM—SPLEEN WORT Four of the five species of this genus that occur in southern Illinois grow in the Pine Hills area. Only A. bradleyi is absent. ASPLENIUM BRADLEYI Eaton. Bradley’s Spleenwort—On sand- stone in southern Illinois; known only from Piney Creek (Jack- son and Randolph Counties) and Panther’s Den (Union Coun- ty). ASPLENIUM PINNATIFIDUM Nutt. Pinnatifid Spleenwort.— This fern usually grows on non-caleareous rocks. It was re- ported from the Pine Hills in 1949, but was not found during our survey. It is rather infrequent even in sandstone areas. The degree of leaf-cutting is variable. ASPLENIUM PLATYNEURON (L.) Oakes. Ebony Spleenwort.— The slightly serrated forma platyneuron is the more common form in our area. Forma serratum (KE. 8. Miller) Hoffm. is a minor variation with deeply jagged-serrate pinnae. Both forms Ferns or Pine His 33 grow in moist woods, ditches, and on wet rocks throughout southern Illinois. ASPLENIUM RESILIENS Kunze. Black Spleenwort.—This spe- cies is distinguished from A. platyneuron, with which it is often confused, by its black rachis and opposite pinnae. It grows in calcareous soils and on limestone rock in the Pine Hills. It is unknown elsewhere in Illinois. ASPLENIUM TRICHOMANES L. Maidenhair Spleenwort. This fern is distinguished from A. resiliens by its smaller size, purple stipe, and non-auriculate rounded pinnae. It is found on lime- stone rock and in calcareous soils of the Pine Hills and else- where in Southern Illinois. It occurs occasionally on sandstone. ATHYRIUM—LADY FERN All three species of this genus known from southern Illinois occur in the Pine Hills area. ATHYRIUM FILIX-FEMINA (L.) Roth. Lady Fern.—Only var. michauxti Farwell has been found in southern Illinois. It has black basal seales on the stipe, stalked glands, sori that are long, harrow, and slightly hooked apically, and a frond with the fourth or fifth pair of pinnae the largest. It is found near the Pine Hills Research Station and in many other parts of south- ern Illinois. It has not been found in Union County Forest. It grows in moist woods, in fields, and along stream banks. ATHYRIUM PYCNOCARPON (Spreng.) Tidestr. Narrow-leaved Spleenwort——In moist woods of Pine Hills, Union County For- est, and elsewhere in southern Illinois. ATHYRIUM THELYPTERIOIDES (Michx.) Desv. Silvery Spleen- wort.—In moist woods of Pine Hills, Union County Forest, and other parts of southern Illinois. CHEILANTHES—Lip FERN Both species of Cheilanthes reported from southern Illinois oceur in the Pine Hills. They are small, rock-inhabiting ferns. CHEILANTHES FEEI Moore. Slender Lip-fern—In ealeareous soils and on limestone rocks of Pine Hills and various other parts of southern Illinois. Not abundant. 34 AMERICAN FERN JOURNAL CHEILANTHES LANOSA (Michx.) D. C. Eaton. Hairy Lip-fern. —QOn dry rocks and shale outcroppings in Pine Hills and else- where in southern Illinois. It is distinguished from C. feet by its larger size and less dense, whitish, jointed hairs. CAMPTOSORUS—W ALKING-FERN This genus produces plantlets where the attenuated tips of the leaves touch the ground. The genus is closely related to Asplenium; some pteridologists consider the two genera to be the same. The only North American species, C. rhizophyllus (L.) Link, occurs in both limestone and sandstone areas throughout southern Illinois. CYSTOPTERIS—FRAGILE FERN Two species of this genus grow in southern Illinois, and both occur in the Pine Hills Area. CYSTOPTERIS BULBIFERA (L.) Bernh. Bulblet Bladder Fern.— On limestone rocks and in calcareous soils of southern Illinois, including the Pine Hill area. It can be distinguished from C. fragilis by the bulb-like structures on the rachis and by having veins that run to the sinuses rather than to the teeth of the leaflets. CYSTOPTERIS FRAGILIS (L.) Bernh. Fragile Fern—Three varie- ties grow in southern Illinois. Cystopteris fragilis var. mackayi Lawson has a short rhizome and subleaflets without a petiolule and with a cleft tip; var. protrusa Weatherby has a long- creeping rhizome and basal subleaflets petiolulate. Cystopteris fragilis var. fragilis has rhizomes similar to those of var. mac- kayi, but the sori in var. fragilis are nearly twice as large as those of var. mackayi, these structures being only about 0.5 mm. wide in var. mackayi. All three varieties are found in the Pine Hills Recreation Area and in Union County Forest Preserve. DENNSTAEDTIA—HAY-ScENTED FERN This genus is not present in the Pine Hills area. These ferns have scaleless branching rhizomes, smooth stipes, and much di- Ferns or Pine His 35 vided fronds that have an odor like hay. The sori are protected by a white, cup-like indusium. Only D. punctilobula (Michx.) Moore occurs in southern Illinois. DRYOPTERIS—SHIELD FERN, WOOD FERN Two species, one represented by two varieties, occur in the Pine Hills Area. All are ferns of moist woodland habitats. DRYOPTERIS AUSTRIACA (Jacq.) Woynar. Spinulose Shield Fern.—The two varieties are briefly characterized by the fol- lowing key: aera thick, suberect; stipe medium brown; gee aera ntugeor ges lobed, the lowermost shorter than the next ones above; i part way tips of veins intermedia sate oar tac reg oki es rome Ne sighty Ted the lowermost longer than others; sori at tips of vein arl paral Both varieties occur in Union County Forest, although neither is plentiful. They grow best in moist woods or along stream banks and ditches. DRYOPTERIS MARGINALIS (L.) A. Gray. Marginal Shield Fern. —Considerable variation occurs in leaf morphology. This species grows in moist woods in southern Illinois. It was reported from the Pine Hills in 1948, but was not found during our survey. It is unknown in the Union County Forest Preserve. ONOCLEA—SENSITIVE FERN ONOCLEA SENSIBILIS L. Sensitive Fern.—This is a weedy fern in woodlands and along swamp margins of the Pine Hills area. It is the only species of the genus known from southern Illinois. PELLAEA—CLIFFBRAKE One of the two species native in southern Illinois is repre- sented in the Pine Hills area. Members of this genus have shiny, wiry, red or purple stipes, leathery, once- to thrice-cut fronds, and sporangia borne near the margins and covered by the in- curved edges of the leaves. Pellaea grows only on caleareous rocks in our area. 36 AMERICAN FERN JOURNAL PELLAEA ATROPURPUREA (L.) Link. Purple Cliffbrake—In calcareous soils and on limestone rock at Pine Hills and else- where in southern Illinois, PELLAEA GLABELLA Mett. Smooth Cliffbrake—This species, with glabrous stipes and rachises, is known in southern Illinois only from Pope County. PoLypopIuM—POLYPopDyY Two species occur in southern Illinois. PoLYPoODIUM POLYPODIOIDES (L.) Watt. Resurrection Fern.— This species frequently is epiphytic on trees, although it occurs on rocks also. It was reported in 1949 that a large bed of this fern grew on a limestone bluff at Pine Hills. It has not been collected there or at the Union County Forest Preserve since. Our plants belong to var. michauxianum Weatherby. PoLypopIuM vuLGARE L. Common Polypody.—Only var. vir- gimianum (L.) Eaton is known in southern Illinois. It has smaller rhizome scales and its leaves are less scaly than those of var. vulgare. Common Polypody grows on rocks, cliffs, and moist bluffs. Sometimes it inhabits dead trees or logs. It is fairly common in southern Illinois, including the Pine Hills area, POLYSTICHUM—CHRISTMAS FERN PoLYsTICcHUM AcROsTICHOIDES (Michx.) Schott, the only spe- cies of the genus found in southern Illinois, is abundant in the Pine Hills area. Several forms or minor variations occur oc- casionally in southern Illinois. The forms in the Pine Hills area can be put into four groups: (1) long, deeply serrated, promi- nently barbed leaflets; (2) membraneous textured, rounded- barbed, very shallowly serrated leaflets; (3) short, rounded leaf- lets with almost entire margins and inconspicuous barbs; and (4) long, slightly serrated leaflets with a prominent barb. PrerIDiIuM—BRACKEN Only one member of this genus grows naturally in southern Illinois. Ferns or Pine Hits 37 PTERIDIUM AQUILINUM var, LATIUSCULUM Underw.—This va- riety has been reported from southern Illinois, and can be dis- tinguished from var. aquilinum in having an almost totally hairless stipe and leaf surface. Our variety grows in dry fields or burned-over areas in the Pine Hills area and a few other places in southern Illinois. It is not common in this area, prob- ably because farmers have tried to eliminate it because it is poisonous when eaten by livestock. THELYPTERIS—BEECH FERN Members of this genus have black, slender, scaly, creeping rhizomes, twice-lobed to once-cut, triangular, membranous, hairy fronds: and small kidney-shaped sori. Many taxonomists include this genus within Dryopteris. THELYPTERIS HEXAGONOPTERA (Michx.) Weatherby. Broad Beech Fern.—Abundant in moist woods of Pine Hills, Union County Forest Preserve, and other localities in southern Illinois. THELYPTERIS NOVEBORACENSIS (Li.) Nieuwl. New York Fern.— Found only once in southern Illinois (Seymour, 1880): It was then growing in a woods near Giant City State Park. THELYPTERIS PALUSTRIS Schott. Marsh Fern.—Grows natu- rally in mashy land only. It is known from a few scattered sta- tions in southern Illinois, but has not yet been found in the Pine Hills area. Woopsta—Woon’s Fern, CLIFF FERN Woopsta optusa (Spreng.) Torr. Blunt-lobed Woodsia.—This is the only species of the genus listed from southern Illinois. It is found at Pine Hills and its environs. FERN ALLIES Although nine species of fern allies (Lycopodium, two spe- cies, one of them with two varieties; Selaginella, two species; Tsoetes, two species; Equisetum, three species) occur in southern Illinois, only one grows in the Pine Hills area. Equiserum HYEMALE L. Common Scouring Rush.—Several varieties of this species occur in the United States. In our area 38 AMERICAN FERN JOURNAL var. elatum (Engelm.) Morton is dominant. It is distiguished from other varieties by its pointed teeth which remain attached to the sheath throughout the winter. The plant occurs in the Pine Hills area and elsewhere in southern Illinois. It is com- mon in sandy areas, such as along river banks and railroad embankments. SUMMARY The twenty-nine ferns and fern allies from the Pine Hills Field Station Area and environs reported here constitute 62 per cent of the ferns and fern allies known to occur naturally in the southernmost 17 counties of Illinois. SourHERN ILuinois UNIVERSITY, CARBONDALE, ILLINOIS. Shorter Notes LiIToBRocHIA IN FLorma.—I read with interest the article, “More Florida Rarities,” by Thomas Darling, Jr. (TH1s JouRNAL 52: 137-148. 1962). Especially notable to me was the author’s mention of finding Litobrochia (Pteris) tripartita in a hammock near the Plant Introduction Station, south of Miami. Mr. Dar- ling’s comment that this was one of the “ferns which formerly I had considered nearly extinct” is noteworthy, for this Giant Bracken is poorly known in South Florida. I would like to point out, though, that Litobrochia tripartita is not as rare in our area as is often believed. For instance, the most impressive colony of it that I know is in the Fairchild Tropical Garden, near Miami, where several hundred handsome plants, exposed to full sun, form an almost solid mass along a coral wall at the Bailey Palm Glade. Since this is one of our showiest ferns, the colony should certainly be protected. The Giant Bracken was formerly frequent along the road- sides, growing on the margins of cypress swamps in deep, rich, constantly moist soil and in semishade west from Fort Lauder- dale. This area, now destroyed by the encroachments of housing subdivisions, supported dozens of plants that often towered well over my head—and I am just a shade under six feet tall! Notes AND NrEws 39 Within the past few months, I have also found scattered plants of Litobrochia in marginal growth of several cypress sloughs along the Tamiami Trail, about fifty miles west of Miami, and also on the edges of wet thickets beside the Sunshine State Parkway, between Fort Lauderdale and Pompano Beach. In these localities, the plants are always, insofar as my experience extends, few and far between. This stately fern has also made its way into a few of our southern Florida gardens. I have two friends in the Miami area who have brought specimens of Litobrochia tripartita to me for identification within the past year, the plants having “just come up” amidst shrubbery in their private gardens. The Giant Bracken is among our finest Florida ferns, and it is encouraging to find that it is spreading within our region, in spite of the tremendous destruction of our forests and fields through the processes of “civilization.”’—AuLex D. HAwKkEs, Coconut Grove 33, Florida. Notes and News 1963 Fern Foray To THE WoopsTocK-DorseT AREA OF VER- mMonT.—A group of Fern Society members gathered at the Wal- lineford Inn, Wallingford, Vermont, on Thursday evening, Au- gust 22, where Dr. R. L. Hauke presided over the indoctrination meeting prior to the foray. Dr. Benjamin R. Allison, Mr. Henry Potter and Dr. Ralph C. Benedict discussed the places we were to visit and some of the ferns we could be expected to see. There was a slight disagreement as to the hour of departure in the morning, but a compromise of 9 A.M. was agreed upon. Due to slow service in the dining room, we were delayed until 9:30 when a convoy of ten loaded cars started off. After a stop between Tyson and Reading, where a number of ferns were seen, the convoy drove to the remote but beautiful home of Mr. and Mrs. H. W. Newton, Reading, Vermont. Everyone enjoyed a tour of Mrs. Newton’s extensive garden which contained the fol- lowing ferns: maidenhair, maidenhair spleenwort, a eut-leaf 40 AMERICAN FERN JOURNAL form of ebony spleenwort, lady fern, slender rock-brake, bulbous and fragile bladder ferns, hay scented, marginal and spiny- toothed wood ferns, ostrich, interrupted, royal, purple clifi- brake, rock polypody, Braun’s holly fern, northern holly fern, New York, marsh and rusty woodsia. Many people also enjoyed a tour of the very old house which Mr. and Mrs. Newton had beautifully restored and furnished with antiques. Lunches were delivered by Mr. Gibson of the Wallingford Inn but, since he got lost and didn’t find us until 2 P. M., everyone was ravenous. After lunch, most of the group walked up the mountain to the “stone chimney” which is the site of a long-gone colonial house. This trip was made mainly to see a beautiful colony of Braun’s holly fern, though many other ferns were seen along the way. That evening Mr. F. Gordon Foster showed some excellent slides of ferns in various stages of development, including many ‘superb close-ups. He discussed photographic techniques and also stressed the satisfaction to be derived from close observation of ferns with a lens. Saturday morning dawned cloudy, windy and chilly, but this did not deter us “from our scheduled rounds.” During the day there were a few sprinkles, but none caused any appreciable discomfort. The group first stopped on a dirt road near West Woodstock to see a colony of the rare North American male fern, Dryopteris filiz-mas. Despite the fact that the colony had been threatened by road widening last year, a number of healthy plants were found along the road and up the bordering hillside. In the rich woods surrounding this colony, a number of other ferns and fern allies were found as well as hybrids between D. filiz-mas and D. marginalis. After this stop we had lunch at Ruth’s Restaurant and then went on to the Summit Railroad Cut on the abandoned Rutland- Bellows Falls branch of the Rutland Railroad. This is a long, narrow cut through rock, and the walls are spotted with a sur- prising variety of ferns and other plants. Here many of the group saw more Slender Rock-brake than they had ever seen be- Notes AND NEws 41 fore. It was here that Mildred Faust coined a new name for a well-known fern, “The Briddle Braddle Fern.” Everyone agreed that this was much more fascinating than the prosaic Brittle Bladder Fern From the Cut most of the cars returned to Wallingford, but some went on to Plymouth Lake to see the green spleenwort. After dinner that night, Mr. James E. Wilkinson, State Forester for the southern half of Vermont, gave a very interesting illus- trated talk on forestry in his state which prompted many ques- tions from the group. Sunday was partly cloudy but warmer than Saturday. With our numbers somewhat depleted, we first visited an open, rocky hillside at East Dorset and saw a number of ferns such as purple cliff-brake and slender rock-brake. Before lunch a brief stop was made at Clarendon Gorge so that. the group could see the three horsetails, Equisetum hyemale, E. scirpoides, and E. variegatum growing together. Following lunch we congregated at Wallingford Inn and pro- ceeded to Elfin Lake where we saw a beautiful quaking bog with an abundance of bog elubmoss. Here Dr. Correll had found quillwort growing in the shallows along the edge of the lake. Several people had never before seen pie insignificant, grass- like fern allies. The last stop of the foray was made at - farm of Mr. Henry Potter, one of the leaders. Here, the sight of several of the group crawling on hands and knees in a mowed field caused a good deal of hilarity among the upright members, as well as considerable fascination among people driving by. We were looking for the Adder’s-tongue Fern, even though it was very late in the season for it. And we found it, so there! Mr. Potter had also collected a fascinating group of hybrid ferns, and many members had not seen most of these growing before. Back at the Inn after dinner, three members showed slides. Dr. Mildred Faust showed several pictures taken on last year’s foray in Oregon; Dr. L. K. Henry projected a number of slides of ferns we had seen during this foray; and Mr. Robert Aborn 42 AMERICAN FERN JOURNAL showed very good slides of Hawaiian ferns. Miss Jeannette Klute displayed several large color prints of her magnificent woodland photographs. This brought to an end the 1963 Fern Foray. There had been sixty-four participants on all or part of the trips. Everyone ex- pressed great appreciation for the excellent organization and leadership shown by Dr. Allison, Mr. Potter and Dr. Benedict and for Dr. Hauke’s coordinating ability. FERNS AND FERN ALLIES SEEN ON THE FERN FORAY Bekanaen virginianum Campto sta tae ea Gymnoc um dryopteris Pteridium aquilin Fhekepeone rphacipheris Adiantu a veboracensis Polen chesus — = is Pellaea Onoclea sibilis Seucneatee punctaobala tines. @rathionlva Osmunda cinnamome Cystopteris bulbifera td nag Woodsia ilvensis Seen um vulga Botrychium ves letearlfittent sh deshitors braunii lanceolatum rostichoides vi um Deventer erista ta multifi lin dissectum goldia dissectum var. obliquum paanes Equisetum arvense marginalis sylvaticum campyloptera fluviatile spinulosa scirpoides intermedia ret btagg filix-mas * marginalis hye eh abe ~ aithimatis Tsoetes cane spor cristata * spinulosa Lycopodium catia ate Athyrium pure arpon lucidulum thel Tioides bse filix-fem flabelliforme Asplenium ruta-muraria tristachyum platyneuron elavat korean es annotinum ride The above list contains 57 species, one variety, and three hybrids. Forty-three species and the variety are ferns, 14 are fern allies, and all of the hybrids oceur in the genus Dryopteris. Norrs AND News 43 Members and friends participating in the Foray were as fol- lows: Mr. Robert Aborn, Mrs. Perey W. Adams, Dr. and Mrs. Benjamin R. Allison, Mrs. Geoffroy Atkinson, Miss Helen M. Attridge, Mr. and Mrs. William A. Barnes, Dr. Ralph C. Bene- dict, Mr. Henry 8. Blethen, Miss Frances W. Brewster, Mrs. Orvis K. Collins, Dr. Donovan S. Correll, Mrs. Howard Davis, Dr. and Mrs. Lewis Dickinson, Miss Ann L. Dunham, Mr. Fred- erick Dunlap, Mrs. Richard Dunlap, Mr. David L. Emory and his father, Dr. Mildred E. Faust, Dr. and Mrs. F. Gordon Fos- ter, Dr. Paul E. Grattan, Miss E. Irene Graves, Mrs. Ronald 8. Gray, Mrs. Alice N. Gobin, Miss Inez Hartsoe, Dr. and Mrs. Leroy K. Henry and son John, Dr. and Mrs. R. L. Hauke, daughters Katy and Nellie and Grandma Hauke, Dr. Donald G. Huttleston, Mrs. W. A. Kenzie, Miss Jeanette Klute, Mr. George S. Lee, Jr., Mr, Elwin F. Leysath, Dr. and Mrs. Robert 8. Lom- masson, daughter Carol and son Timothy, Dr. and Mrs. Ray Martin, Dr. Jewel Moore, Miss Mary Murphy, Mrs. H. W. New- ton, Mrs. Gladys H. Nourse, Miss Elsa Potter, Mr. L. Henry Potter, Miss Ruth A. Reed, Mr. and Mrs. Alex D. Reid, Mrs. A. E. Scherrer, Miss Anna E. Seudder, Miss Eva Sobol, Miss Olive Stacey, Mrs. Harold Stillwell, and Mr. and Mrs. W. W. Willis. —Donaup G. Hurrieston, Longwood Gardens, Kennett Square, Pennsylvania. GRrowine Berry BLADDER F’erN.—For some years we have had a bed of Berry Bladder Ferns, Cystopteris bulbifera, in a semi- shaded spot in a slight hollow. Here they have multiplied hap- pily. When we dig up the larger ones, we find many bulblets (black) with the outer shell just partially cracked. If these are planted in black humus barely an inch or maybe a half-inch deep, and the humus then mulched with old straw, little green fronds soon appear. It takes some years for these to grow to maturity—Mariz SperKa, Woodland Acres Nursery, Route 2, Crivitz, Wisconsin. 44 AMERICAN FERN JOURNAL AN APPEAL FoR Hetp.—During recent months it has become increasingly apparent that many papers published both in the United States and abroad that would be of interest to members of the American Fern Society have not been reviewed or listed in the Recent Fern Literature section of the Fern Journal. The task of checking through the numerous journals and publica- tions from various laboratories and institutes in an attempt to search out such titles is too great a one for a single individual to attempt. In order to provide a somewhat more adequate cov- erage, the Editor will appreciate a card or note from any mem- ber who sees a paper or book that he or she thinks would interest other pteridologists. Just note the name of the author, the title of the book or article, and if the latter, the name, volume, and pages covered in the article, followed by the date of its appear- ance. Such offerings will be collected, evaluated for the relation- ship between interest and the space available in the Journal, and a selected list published in an ensuing number of the American Fern Journal. Authors who publish such papers in other journals can further help by sending two copies of a reprint of such a paper to the Editor. If the paper is of sufficient length and interest, an at- tempt will be made to find an impartial reviewer, whose critique will be published when printing schedules and space permits.— American Fern Society ANNUAL MEETING IN 1964:—The annual meeting of the Amer- ican Fern Society will be held at the University of Colorado in Boulder, with other Adherent Societies of the American Insti- tute of Biological Sciences. The AIBS meetings begin August 23 and run through the 28th. Plans are being made for two ses- Sions, one in the forenoon and one after the annual luncheon, for the presentation of contributed papers. Our society will probably cooperate with one or more other societies in sponsor- ing symposia of mutual interest. AMERICAN FERN SOCIETY 45 Each member of the Fern Society wishing to present a paper at Boulder should send the following information to Mrs. Lenette Atkinson, 415 South Pleasant Street, Amherst, Massa- chusetts, not later than May 15th: title of paper, time required for its presentation (10 to 20 minutes), size and kind of projec- tion equipment needed, if any. Please send in your title prompt- ly, because late requests for a place on the program may have to be denied. No summer meeting is complete without a field foray prior to or following the formal sessions, so a Foray will be planned and details will be included in the April—June issue of the Journal. Report of the President for 1963 During August, just prior to the annual meeting in Amherst, the Society was represented at the Adanson Bicentennial Sym- posium at the Hunt Botanical Library of the Carnegie Institute of Technology in Pittsburgh, Pennsylvania, by the Society Li- brarian, Dr. Warren H. Wagner, Jr. His report appears in this issue of the Journal. The annual foray sponsored by the Society was held in Ver- mont. Wallingford served as our headquarters site, and daily trips were made from there on August 23, 24, and 25. Dr. Rich- ard L. Hauke made local arrangements and was largely respon- sible for handling many of the details of the largest foray we have had in recent years. Mr. Henry Potter of West Rutland, Vermont, deserves special recognition for leading the trips and providing direction to the very interesting fern habitats in the vicinity. Besides being new to the duties of Secretary of the American Fern Society, Dr. Lenette R. Atkinson also had the added task of being local representative for our Society in planning the arrangements for our annual meeting held in conjunction with the AIBS meetings in Amherst, Massachusetts. She was assisted in this role by Mrs. Marion Rhodes. As program chairman Dr. Atkinson assembled a fine program of contributed papers that was given Tuesday, August 27, utilizing both the morning and 46 AMERICAN FERN JOURNAL afternoon sessions. Thanks are especially due also to the speak- ers for their excellent presentations. Between the formal ses- sions, the American Fern Society Luncheon was attended by 48 members and provided a welcome change of pace and time for general conversation and making several announcements. The American Fern Society also co-sponsored a symposium, “System- aties in the American Tropics,’ presented on Wednesday, August 28th. In September I solicited a limited number of the members of the American Fern Society for contributions to provide and erect a display case for fern pictures in Fern Valley at the Na- tional Arboretum in Washington, D. C. The response was most gratifying to me, and generous on the part of contributors, for a fund of $93.00 was turned over to the group in charge at Fern Valley. At a meeting of the Council of the American Fern Society in Amherst, it was decided that the Society should become an ‘“‘Ad- herent Society” of the American Institute of Biological Sciences if our membership was willing to support this move by approv- ing an increase in dues. Since the Society responded affirma- tively in the annual election, our society, as of January 1, 1964, will have full rights and representation under the new consti- tution of the AIBS Special service to the Society has been rendered this year by John T. Mickel, who served as Judge of Elections, and by Pro- fessor Elmer A. Palmatier, who served as Auditor. I extend the Society’s thanks to both of them It has been a distinct honor to me to serve as President of the American Fern Society, and I am sure the Society will prosper under the energetic leadership of my successor. Respectfully submitted, Ropert C. Lomnasson, President of the Secretary for 1963 With support from the President and the Treasurer, and with considerable aid from Dr. Huttleston, the past Secretary, I have come through the first year in office without receiving com- plaints. Or is everyone being polite? AMERICAN FERN SOCIETY 47 Membership in the American Fern Society at the end of 1963 stands at 857. California still heads the list with the largest number of members, with New York a close second. There are 79 members outside the United States, distributed among 28 countries. We still have no members in Nevada, New Mexico, or North Dakota. I regret to report the death of seven members during 1963: Mr. Peter Borree (1953), Mr. Thomas 8. Constantine (1936), Mr. Lewis A. Dyer (1956), Mr. Andrew T. Skidmore (1950), the Reverend Jesse F. Smith (1949), Mrs. Ora M. Wilson (1957), and Dr. Walter Dépp (1951). Dr. Dopp was one of our distinguished overseas members. His extended researches on ferns produced two important contributions: an elucidation of the mechanics of apogamy and the discovery of a biologically active substance in thalli of the bracken fern that induces and hastens the formation of antheridia. The annual meeting of the American Fern Society, held with the AIBS on the campus of the University of Massachusetts at Amherst, on August 27, was attended by well over 50 persons. Major assistance in arranging the program was given by Mr. Conrad Morton and by Mrs. Marion B. Rhodes. Their aid is eratefully acknowledged. The President of the Society, Dr. Robert C. Lommasson, presided at the formal sessions and at the luncheon. Titles of papers read follow in the order of presenta- tion: “First century of Louisiana fern records,” by Joseph Ewan; “Ferns and their habitats in Texas,” by Donovan Cor- rell; “Preliminary studies on the fern flora of Oaxaca, Mexico,” by John T. Mickel; “Recent data on ferns of Georgia,” by Wil- bur Dunean (read by title) ; “County distribution of ferns and fern allies of Rhode Island,” by Dorothy Crandall; “Contribu- tions toward a morphological classification of fern spores,” by Gerhard Kremp; “Sporogenesis in Pteris eretica with special reference to the cytoplasmic inclusions,” by Norman Marengo; “On the rhizophore of Selaginella,” by T. R. Webster and T. A. Steeves (read by Mr. Webster) ; “The Polypodium virgimanum complex,” by A. Murray Evans; “Fern gametophytes and devel- 48 AMERICAN FERN JOURNAL opmental plant physiology,” by John Miller; “Antheridium for- mation in ferns—a model for the aie: of developmental change,” by Ulrich Naf; “Morphological effects of ultraviolet radiation on prothalli of Onoclea sensibilis,’” by Larry Estes (read by R. C. Lommasson); “Eastern American spreading woodferns, Dryopteris campyloptera and D. ‘dilatata, ” by War- ren H. Wagner, Jr. (a timely paper as those on the foray saw some of these ferns) ; “The genus Cyathea (sensu lato) in Ma- laysia,” by R. E. Holttum (read by W. H. Wagner, Jr.). Miss Clara Hires had a fine exhibit concerned with fern indusia and other structures in the exhibit hall. On Wednesday afternoon, August 28, the American Fern So- ciety, the American Society of Plant Taxonomists, and the Sys- tematic Section of the Botanical Society of America cooperated in sponsoring a symposium on “Plant Systematies in the Amer- ican Tropics,” with Mildred Mathias presiding. Three papers were presented during this symposium: ‘The present state of knowledge and current field explorations,” by Richard 8. Cowan; “Opportunities for evolutionary studies,” by Herbert G. Baker; and ‘Facilities for student training and research,” by Walter H. Hodge. All members of the Council were present at a meeting August 26. The Council decided to have an attractive membership card printed, to be sent to each new member, with a statement of the purpose and history of the Society. It was suggested that Mr. Conrad V. Morton be asked if he would serve as a Clearing Offi- cer for exchange requests among members of the Society, for both living and herbarium specimens. Happily, Mr. Morton has accepted this responsibility. In view of the increasing number of requests for our membership list for various uses, it was voted to deny use of the American Fern Society membership roster for solicitation purposes. One of the distressing duties of the Secretary is to deny re- quests from organizations or individuals who wish loans of fern specimens for exhibition purposes. I suggest that careful con- sideration be given to some method for complying with such AMERICAN FERN Society 49 requests. Such a move would promote one of the aims of the Society: to spread information about ferns. Respectfully submitted, Lenerre R. ArKinson, Secretary. Report of the Treasurer for 1963 I am happy to report that the Society remains solvent. Thanks to the constitutional amendments passed in 1962 we were able to remove many delinquent members from the rolls without hav- ing to carry them in arrears for an extended period. This saves postage and copies of the Journal. A major expenditure this year was for reprinting back num- bers of the Journal that were out of print. We are now in a position to again supply complete sets of the Fern Journal. Be- cause of this and the cost of reprinting, I have increased the book value of our holdings, thus increasing the Society’s assets. The cost of reprinting, much larger than our cash account could cover, was met with funds from the Una Weatherby Fund. This raised the question of the distinction between that fund and the Reserve Fund. It was decided in the Council meeting at Amherst that in the absence of a definite difference between them the monies from the Reserve Fund could be combined with the Una Weatherby Fund. This will be done as of January 1, 1964. The amendments passed in 1963 have not yet become effective, but the future looks good. The budget for the Journal will be increased in 1964, enabling greater coverage of topics of interest to members. Continued association with the American Institute of Biological Sciences will enable the American Fern Society to retain its prestige in professional botanical circles. I hope that the necessary increase in dues will not cause an appreciable drop in our membership, and urge that any member deeply interested in the Society who finds this increase an insurmountable finan- cial burden will let me know. The Society’s application for tax-exempt status under Internal Revenue laws, which was dependent upon passage of one of the 50 AMERICAN FERN JOURNAL amendments to our constitution, has been filed. When it is ap- proved you will be notified, and gifts to the Society will be de- ductible in preparing personal income tax returns. Hopefully, this will encourage more people to make gifts or bequests to the American Fern Society. The financial report for the year follows: Receipts Cash on hand, stack! 1, 1963 $1,070.25 Membership d Arrears a Renewals $1,337.35 Sustaining 430.00 New 244.50 Advance payments 94.57 Life 100.00 $2,206.42 Subscriptions urrent and Arrears 247.00 Advance payments 495.80 742.80 Sale of baek numbers 813.14 Sale of reprint 312.87 Gifts 452.76 Fern Valley Fund 93.00 Extra et Journal articles 188.00 Advert 14.00 Tr hay hen Weatherby Fund 2,362.54 Miscellaneous 8.05 7,193.58 $8,263.83 Disbursements American Fern Journal Vol. 52, No. 4 $ 773.59 Vol. 53, 1 992.73 Vol. 53, No. 2 658.34 Vol. 53, No. 3 738.93 $3,163.59 AMERICAN FERN Society prints Envelopes and printing stationery Treasurer’s Stiees, Seeretary’s exp Shipping and handing back numbers Editor’s expen Life Honbenieg Fund Reprinting back numbers A. I. B. 8. membershi ern Valley Fund Miscellaneous Cash on hand, January 1, 1964 Statement December 31, 1963 Assets Cash in Industrial National ages t Savings Ss ba a Life see aii Fund Reserve und Una Weatherby Fund Accounts Receiv Inventory, rile Fern Journal ibrary Liabilities Advance dues colle $ 94.57 cted Advance subscriptions collected 495.80 Fund Balances Bissell Herbarium Fund Life ee Fund und Reserve Una i Fund General Fund $1,597.82 $ 590.37 815.59 1,198.92 2,232.81 1,213.62 7,009.31 6,666.01 $1,597.82 $13,060.62 $13,060.62 52 AMERICAN FERN JOURNAL Respectfully submitted, RicHarp L. HauKe, Treasurer. Report of the Auditing Committee I hereby certify that I have seen the books and accounts of Dr. Richard L. Hauke, Treasurer of the American Fern Society, Ine., and have obtained confirmation of the correctness of the Society’s balances on hand as set forth in detail in the accom- - panying report of the Treasurer. Etmer A. PauMATiER, Auditor. Report of the Judge of Elections The results of balloting for officers of the American Fern So- ciety are as follows: For President Donovan §8. Correll : 330 Ue EVO ee ee X At SW aaner at sh i 1 Grae t, Where 8 a er 1 For Vice-President POUL, Gr SUE BE i er 328 SNUG AONOEY hoa ee ea ote re 1 Richard L. Hauke 1 MEE Sa OS EES GG Ram ks Wan ese an Sem er ch ee 1 Dale De Sith ee ENR a | Hola Mi. Tryon Ge echt eae For Treasurer Richard L. Hauke. 329 aOR OI iri a i Warren Bo Weoner I ee 1 I therefore declare the following candidates elected to office: Donovan 8S. Correll, President; Donald G. Huttleston, Vice-Pres- ident; Richard L. Hauke, Treasurer. e four amendments to the Constitution were voted on as follows: AMERICAN FERN Socrery 53 oC pe III, delete Section 2 dda the Constitution. 7 Aga Under ae III, change Sections 2, " and 4 to increase ad- mission fee and annual dues from $2.50 to $4.00; change Section 4 to increase the sustaining membership from $5.00 to $8.00; change Section 5 to increase life membership from $50.00 to $80.00 For 299 Against 39 Under Article IV, Section 3, add “Program Chairman for the Annual Meeting” to < Ss as of the hig President. For. Agains Uader ‘Reticle V, ee new sections 3 a 4 to fulfill legal re- quirements as to a non- organization. For 334 Against I therefore declare eas amendments to the Constitution of the American Fern Society approved by the membership and now in foree. Respectfully submitted, Joun T. Micke, Judge of Elections. Report of the Librarian and Curator for 1963 In the present report I shall deviate somewhat from custom and refer the reader to an earlier statement (THIS JoURNAL, 52: 52-54. 1962.) to express the general picture of activities in the Society’s Library and Herbarium. I wish to describe here instead an important botanical meeting. During August the Adanson Bicentennial Symposium was held at the Hunt Botanical Library of the Carnegie Institute of Technology in Pittsburgh. It was my pleasure to be invited to - represent the American Fern Society on this occasion. Although many members of the Society are probably familiar with the Hunt Library, a few words about it may be of interest. More than 60 years ago, when she was a girl in her teens, the late Mrs. Rachel MeMasters Miller Hunt began her now famous collection of botanical books, paintings, and prints, supplement- ed by letters and portraits of distinguished botanists. In 1961 54 AMERICAN FERN JOURNAL the Hunt Library was formally opened in an excellent new building on the campus of the Carnegie Institute, where not only the significant collections will be cared for, but biblio- graphic research in botany will be fostered through the Hunt Foundation. This work is under the administration of George H. M. Lawrence, the Director, plus the directors of the Hunt Foundation and an advisory committee of six botanists from several countries. In addition to the research carried out by the staff and visitors to the Library, various other projects are sup- ported, including the Hunt Faesimile Series, in which rare and often unavailable botanical works will be reproduced; the Hunt Monographie series, for the publication of new botanical inves- tigations emphasizing historical aspects of the field; and the Bibliographica Huntiana, described as a “new Pritzel for the period of 1735-1850.” It seems especially timely that this im- portant establishment has been so efficiently organized so recent- ly, now that the academic world and the educated public at large have become very strongly concerned with the history of science. In botany the historical aspects are of special operational sig- nificance in the day-by-day work because of the necessity to establish a stable nomenclature for plants based upon the prin- ciple of priority and in accordance with our International Code of Botanical Nomenclature. It is appropriate too that it was the Hunt Library which cele- brated the bicentennial of the publication of Michel Adanson’s Familles des Plantes, one of the landmarks in the literature of plant sciences. Michel Adanson (1727-1806) was a French bot- anist and naturalist, whose name is much in the scientific lime- light currently because of his philosophical conceptions of the nature of systematics. How do we evaluate taxonomic relation- ships? How are species grouped ? Genera, and so on? Adanson argued for the use of all characteristics, all the information available about plants. His ideal methodology would be based upon not just a few characteristics (in ferns, for example, just the sorus and sporangium) but on all that we can learn about the plants. Furthermore, the idea that we do not know the rela- AMERICAN FERN Society 55 tive “value” of characteristics and therefore should treat all data (e.g., roots, stems, leaves, spore-cases, epidermis—every- thing) as equal seems to come from Adanson, whose ideas must have seemed heretical to some of his colleagues. Currently, with the appearance of the electronic computer as a tool in taxonomy, a whole new school of botany—“Numer- ical Taxonomy” or “Quantitative Adansonianism’—is appar- ently arising, and much of the philosophical groundwork traces back to Adanson. I am sure that many of the delegates and registrants at the Symposium found the purely historical papers of great interest, as I did. Out of the eight major talks that were presented (some in French, some in English), I was per- sonally most inspired by the lecture on “Mathematics and classi- fication from Adanson to the present” by Peter H. A. Sneath of the Medical Research Council in London, in which the influ- ence of Adanson was analyzed from the perspective of the present world. The program of the Adanson Bicentennial involved an exhi- bition of Adansoniana, the above-mentioned Symposium papers (held on the two days, August 18 and 19), a visit to the Car- negie Museum, and a Banquet. Botanists representing many viewpoints and backgrounds came together for the occasion. Each of the participants, undoubtedly, found the event a most stimulating experience, and I am grateful to the American Fern Society for being chosen as one of the delegates of the 18 botani- cal societies represented, Respectfully submitted, WARREN H. Waaner, Jr., Librarian and Curator. NEw MEMBERS Dr. Harry C, Blair, Neskowin, Oregon Mr. William D. Davis, Dept. of Biological Sciences, Purdue University, Lafayette, Indiana Mr. Kenneth J, DeNault, Box 5929, Stanford, California Mr. Erie Drew, 75 Bank Street, Apt. 2A, New York, N. Y. 10014 Mr. John Love, 1501 St. Christopher, Columbia, Missouri 65201 56 AMERICAN FERN JOURNAL Dr. Thomas R. Pray, Dept. of Biological Sciences, University of Southern California, Los "eet 7, Oalit, Mr. Robert ri Russell, 435 7 apa Fitchburg, Mas Mrs. 4 sell, 1580 S. E. 4th Court, eae se Seats Blends 33441 Mrs. ete ur, 104 East i ie Street, , Llino Mr. ober S. Sutherland, Space 128 Circle ae ook isa San Pablo n Pablo, Calif Mr. bk ta Show albot, Res 492, Bates College, Lewiston, Maine Mrs. H. Edward Walker, 4306 Newport Drive, Richmond, Virginia 23227 Mr. Walter W. Willis, 357 Marion Avenue, Mansfield, Ohio CHANGES OF ADDRESS Mr. Roger F. Anderson, 647 Alvord Street, Flint 7, Michig Mr. Mat Bishop, 633 South Orchard Street, Madison _ When 53715 Miss Ruth A. Bowers, 650 Poplar Street, Sharon Hill, 9079 Mr. Bernard Bowker, 9821 W. 47th Street, Miami, ee 33165 Mrs. Jolly H. Galt, 6515 El Nido Lane, Goleta, Calif. Mr. Robert per aebars c/o International Labour Office, Manpower Division, Geneva, Switzerland Mr. Edward N Talinan 714 N. Woodland Blvd., Deland, ae 32720 eroy E. L 4 Lake Mendota Drive, Madison, Wise Miss Joyee Y. ing 100% Coyle, Houston, iat 77 ay Dr. Albert C. Smith, i ity of Hawaii, Honolulu 14, eee Mrs. E. H. Streissguth, 101 W. Main Street, sigs Wash. 9827 Mr. Merrill H. Sweet, oe of Biology, Texas A & M Payot College Station, Texas 77840 Copy for a number of the Fern Journal goes to the printers before the 15th of the third month prior to the date of issue; before the middle of January, April, July, and October. Requests for change of address can not be published in the Journal sath the Gisend number if they reach the Baitor after these deadlines. Novice or CoLorapO FrrRn Foray Plans are being made for an American Fern Society field foray to be held prior to our annual meeting at the University of Colorado, Boulder, August 23-28, 1964. It is planned to have the foray in some area of Colo- rado approximately August 19-22. Details will be given in the next issue of the Journal as well as in a future issue of the AIBS BioScience. SUGGESTIONS TO CONTRIBUTORS TO THE AMERICAN FERN JOURNAL 1, Authors of papers submitted to the American Fern Journal should con- sult the Style Manual for Biological Journals (1960) and be guided by its examples, 2. The “name-and-year” system for bibliographic references will, except in exceptional cases, be used. (See Literature Cited below for an example). 8. Authors are encouraged to use the journal abbreviations set forth by Schwarten and Rickett (1958, 1961). 4. Abbreviations of names of herbaria will be those listed by Lanjouw and Stafien (1959) 5. Reports of chromosome numbers will not be accepted unless documented. 6. The use of footnotes and tabular matter should be kept to a minimum. 7, All manuscripts ae tips publication should be typed, double- spaced, and have ample ma 8. Reprints should be ordered when galley proof is returned to the Editor. An order blank will be included with the galley proof LITERATURE CITED American Institute of Biological Sciences, Committee on Form and Style of the Conference of Biological Editors. 1960. Style Manual for at Journals. American Institute of Biological Sciences. W: D. CO. 100 p. Langouw, J., and F. A, STaFuzv. = The Herbaria of the World. Index Herbariorum, Regnum Veg. 15: 1 Scuwarren, L., and H. W. Ricker, 1958. Abbreviations of Titles of Serials Cited by Botanists. Bull. Torrey Club 85: 277-800; 1961. Supplement I. Ibid. 88: 1-10. Von. 54 “ Aprit-JUNE, 1964 No. 2 American Pern Fournal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY ts IRA L. WIGGINS C. V. MORTON ROLLA M, TRYON JOHN H. THOMAS CONTENTS Development of the Stelar Cylinder in the Rhizome of itis and Egenolfia______________Sursir Kaur 57 A New Species of Pyrrosia from ee ee CHANDRA 62 Notes on the Hawaiian Fern Genus Adenophorus KennetoH A, Witson’ 68 New Combinations in Lycopodium _______________ C. V. Morton 71 Some Hints for the Fern Culturist______Irnvive W. Knostocne 73 Ethnobotanical Uses of California Pucidosheive by Western Aaneetenn Teenie ___Ropert M. Luoyp = 76 Phytogeography of Selaginella douglasii...Gzorcz Nevin Jonrs 82 Retention of Viability in Lyophilined Spores of the Fiddlehead Fern W.G. BarKER aNDR.G.WuHITE 87 Shorter Notes: Heie aa Fern Allies on Bonaventure Island, Quebee; A Note on Elaphoglossum crinitum; Observations on the Sensitive Fern: Supplement; Growing Mother Ferns. 89 Recent Fern Literature — ee a. Oe Notes and News ee a American Fern Society a a f MissouR! BoTANICAL JUL 01 1964 The American Fern Hocietp Council for 1964 OFFICERS FOR THE YEAR Donovan 8. CorrRELL, Texas Research Foundation, Renner, Texas President DonaLp G. Huttieston, Longwood Gardens, Kennett oe Foonsylvaws ice-President LENETTE R. ATKINSON, 415 S, Pleasant Street, Amherst, Masnchosti cretary Bicnarp L. Hauke, Department of Botany, University of itiods: —— Kingston, Rhode Island Tre Ina L. Wiaerns, Dudley Herbarium, Stanford University, Stanfor 4, Cal Edit eos mee OFFICIAL ORGAN American Fern Journal EDITORS Ina L. Wiaeins___Dudley Herbarium, Stanford University, Stanford, Calif. “a V. Morton_._._...____Smithsonian Institution, Washington 25, D. C. oN Gray Herbarium, Harvard University, Cambridge 38, Mass. Joun H. THomas_Dudley Herbarium, Stanford University, Stanford, Calif. An illustrated quarterly devoted to the — study of ferns, owned by the American Fern Society, and published at 3110 Elm Ave., Baltimore re Md. Gadoadchehe 4 postage paid at Baltimore, Ma Matter for publication should be addressed to aes L. Wiggins, Dudley Herbarium, Sins ford University, Stanford, Califor Subscription $4.50, exclusive of agency handlin ing cred sent free to — bers of the American Fern Society (annual dues, $4. 00; sustaining bership, $8.00; life Sr sege $80.00). Extracted sepeaees, if pot so in advance, will be furni authors at cost, plus postag volumes $3.00 Se single back numbers 75 pion ek: Cumulative ope by vols, 1-25, 25 cents. Ten per cent discount on +; seas of six volumes Chdiaaiil of address, applications for SRE EAT subscriptions, orders for back numbers, and other business comm ns should be addressed to the r, Dr. Richard L. Hauke, alae of Botany, University of Rhode Island, Kingston, Rhode Island. Members and subscribers should Sree ee Seen ae eae address to take effect. LIBRARIAN AND CURATOR OF THE HERBARIUM Dr, W. H. Wagner, University of Michigan, Ann Arbor, Mich. A regular loan department is maintained in connection with the library books and specim ying all postal or express @ pa also are open to members who wish to arrange exch e8; a membership list ro dage-mmanig at oe to assist those Sotaveaed & obtaining American fern Journal Vou. 54 APRIL-JUNE, 1964 No. 2 Development of the Stelar Cylinder in the Rhizome of Bolbitis and Egenolfia Sursgit KAuR Bolbitis and Egenolfia are Lomariopsidoid ferns of doubtful systematic position, relegated to the Aspidiaceae by most pter- idologists (Ching, 1940; Copeland, 1947). They are small or medium sized ferns with dimorphic leaves, acrostichoid distribu- tion of sporangia and dorsiventral epigeal rhizomes which may either be short creeping or long and sometimes climbing ver- tically on supports. During the course of a morphological in- vestigation of some species of Bolbitis and Egenolfia it was ob- served that the characteristic dorsiventral type of solenostele of the rhizome (Holttum, 1954; Nayar, 1960) develops from the juvenile protostele in an unusual manner in these genera. The common type of development of solenostele from the solid cylin- drical protostele in ferns by the formation of a central shaft- like pith connected to the cortical tissue through leaf gaps, is described by many workers (Bower, 1935; Gwyne-Vaughan, 1901). A variation of this type of development is reported in Danaea (Brebner, 1902). The present observations are based upon ten species of Bolbitis viz., B. costata (Wall.) Ching, B. crispatula (Wall.) Ching, B. deltigera (Wall.) C. Chr., B. diversifolia (Bl.) Schott, B. heteroclita (Pr.) Ching, B. presli- ana (Fée) Ching, B. semicordata (Moore) Ching, B. suberenata (Hook, et Grev.) Ching, B. subsimpler (Fée) Ching, B. virens (Wall.) Schott and five species of Egenolfia viz., E. appendicu- lata (Willd.) J. Sm., E. asplenifolia (Bory) Fée, E. helferiana Volume 54, No. 1, of the JourNaL, pp. 1-56, was issued April 15, 1964. 58 AMERICAN FERN JOURNAL (Kze.) C. Chr., E. sinensis (Bak.) Maxon and E. vivipara (Hook.) C. Chr. The observations were made by cutting serial transverse sections of young and adult rhizomes and foliar buds, and making drawings with a camera lucida, from which the stelar pattern is reconstructed to scale. These are then com- pared with the stelar cylinders obtained after prolonged boiling in strong KOH solution. STELAR CYLINDER OF THE ADULT RHIZOME The stelar cylinder of the adult rhizome in Bolbitis and Egenolfia is a hollow cylinder pierced on the dorsal surface by two alternating, closely set rows of large, prominently overlap- ping leaf gaps (Fig. 4-1), which dissect the stelar cylinder into a broad, gutter-shaped, root-bearing, ventral vascular strand (Fig. 4-d). In some species more than two rows of leaves are developed at maturity and then the dorsal strand is pierced by one or two rows of leaf gaps. Vascular connections to each leaf (each leaf trace, Fig. 4-f) consist of many cylindrical vascular bundles originating in succession from the lateral margins to- wards the abaxial end of the leaf gap. Vascular connection to a vegetative bud (Fig. 4-b) is associated with each leaf trace on its abaxial side, and a root trace (Fig. 4-r) originates along with the bud traces (either fused with the bud trace or sometimes independently) in all the species. DEVELOPMENT OF STELAR CYLINDER oF RHIZOME The juvenile rhizome has a solid, cylindrical, centrally placed vascular strand (Fig. 1-A). Leaf traces are single cylindrical bundles originating from the dorsal surface as superficial branches, unaccompanied by a leaf gap. Vegetative buds are absent. As the plant grows, the rhizome increases in thickness and the successive juvenile leaves increase in size. The stelar cylinder expands and successive leaf traces become alternately placed on either side of the dorsal median line ( Fig. 1-B). Be- tween the two rows of leaf traces a longitudinal groove is formed, which becomes progressively deeper and wider as the RHIZOME OF BOLBITIS AND EGENOLFIA 59 rhizome elongates. Later, each juvenile leaf is supplied by two vascular bundles originating from the opposite margins of the median groove (Fig. 1-C). The stelar cylinder expands consid- erably and the dorsal groove becomes more and more prominent till the stele is gutter-shaped, and later the margins of the gut- ter approach each other and fuse, forming a hollow cylinder with the cortical parenchyma forming a central pith (Fig. 1-D). One or two elongated lacunae (Fig. 1-p) may occur laterally on the stelar cylinder. Soon the dorsal surface of the stelar eylin- der opens as a leaf gap (Fig. 1-1) and a set of vascular strands (usually two or three on either side of the gap) are given oft to a leaf, the leaf in question being markedly larger than the previously formed ones. Vascular connection to a vegetative bud and the accompanying root (Fig. 1-b & r) is usually associated with this leaf trace on its abaxial side as in the adult condition. Vascular connection to the next leaf in succession is established in a similar way by the formation of a gap close to the first gap on one side. Thereafter, leaves are formed alternately in closely placed rows, and their gaps are prominent and considerably overlapping so that the dorsal median region of the vascular cylinder appears like a zigzag narrow meristele (Fig. 1-d) con- necting the two margins of the gutter-shaped ventral region (Fig. 1-v) STELAR CYLINDER OF FoLiar Bups Foliar buds are characteristic of most species of Bolbitis and Egenolfia. These buds oceur on the dorsal surface towards the apex of the terminal pinna (in some species on the lateral pinnae also), replacing one of the main lateral veins of the lamina. In many cases these buds strike root and grow into young plants while still attached to the parent leaf. The vascu- lar bundle entering the base of the rhizome of these young plants is a simple, cylindrical, solid strand (Fig. 2-A) resembling the protostele of the juvenile rhizome. Soon it expands consider- ably, becomes flattened and later gutter-like (Fig. 2-B), with its coneavity facing the lamina of the parent leaf (ventral sur- 60 AMERICAN FERN JOURNAL Fig 1. STELAR CYLINDER OF YOUNG RHIZOME OF BOLBITIS SUBCRENATA. Fig. 2. STELAR CYLINDER OF FOLIAR BUD OF THE SAME. Fia. 3. STELAR CYLINDER OF FOLIAR BUD OF EGENFOLIA APPENDICULATA. Fig. 4. A POR- TION OF THE VASCULAR CYLINDER OF THE RHIZOME OF B. SEMICORDATA. A—CYLINDRICAL PROTOSTELE; B—DEVELOPMENT OF DORSAL GROOVE ON STELAR CYLINDER; C—FORMATION OF MULTIPLE TRACES TO LEAF; D—For- MATION OF HOLLOW CYLINDRICAL STELE; E—-ADULT CONDITION OF STELE. |= Lear gap, f = Lear TRACE, b = BRANCH TRACE, r = RooT TRACE, d = DORSAL VASCULAR STRAND, v = VENTRAL VASCULAR STRAND, p = LACUNA. RHIZOME OF BOLBITIS AND EGENOLFIA 61 face of the daughter rhizome). The margins of the gutter-like stele approach each other and fuse (Fig. 2-D) forming a hollow cylinder which usually develops one or two elongated lacunae generally on the opposite side (dorsal surface of the daughter rhizome). A large leaf gap (Fig. 2-1) is formed dorsally, open- ing up the stelar cylinder, and a pair of leaf trace bundles (Fig. 2-f) originates, one from either lateral margin of the gap. The leaf gap next in succession originates lateral to the first gap and successive leaf gaps are formed in alternate succession dis- secting the vascular cylinder into a broad, ventral strand and a narrow dorsal one as in the adult rhizome (Fig. 2-E). In Egenolfia a vascular connection to the first leaf of the daughter plant originates before the stelar cylinder becomes siphono- stelic. The protostelic vascular cylinder entering the bud be- comes gutter-shaped, with its concavity facing the dorsal side (Fig. 3-B) and one or two vascular bundles are given off from either margin to supply the first leaf. Soon afterwards the mar- gins of the gutter unite to form the hollow, cylindrical stelar cylinder (Fig. 3-D) but generally an elongated dorsal lacuna (Fig. 3-p) is formed close to the point of fusion of the margins. After the formation of a hollow cylindrical stelar cylinder the adult condition is reached by the development of large leaf gaps in two alternating closely placed rows on the dorsal surface, but bud traces do not occur associated with the leaves though a root trace (Fig. 3-r) often is attached abaxially to each leaf trace. Thus the stelar cylinder of the adult rhizome of Bolbitis and Egenolfia, whether developing from the protostele of a vegeta- tive bud or the juvenile sporeling, is formed by flattening of the solid cylindrical protostele, accompanied by upward curving of the margins of the flattened strand and their final merging to form a hollow eylinder. The pith is connected to the cortical tissue and is clearly cortical in origin. I am grateful to Prof. K. N. Kaul, who has taken keen in- terest in this work. I am obliged to Dr. B. K. Nayar under Whose guidance this work was carried out. Thanks are due to 62 AMERICAN FERN JOURNAL Shri P. C. Roy who has helped in the preparation of figures. LITERATURE CITED Bower, F. O. 1935. Primitive land plants. Maemillan & Co., London. BreEBNER, G. 1902. On the anatomy of Danaea and other Marattiaceae Ann. Bot. 16: 517-552. Cuine, R. C. 1940. On natural classification of the “Polypodi .” Sun- yatsenia, 5: 201-268. CopELAND, E. B. 1947. Genera Filicum. Waltham, Mass. GWYNNE-VAUGHAN, D. T. 1901. Observations on the anatomy of soleno- stelic ferns. I. Lorsoma. Ann. Bot. : 71-98 R ‘ E, 1954. Flora of Malaya, 2: Ferns. Singapore. Nayar, B. K. 1960. Morphology of two Indian species of Bolbitis. Jour. Indian Bot. Soe. : 259-277, PrertpoLogy Lasoratory, National Botanic GarpEns, LucK- NOw, INDIA. A New Species of Pyrrosia from India PRAKASH CHANDRA During a fern collecting tour of northeastern India, organized by the National Botanic Gardens (Lucknow) in May and June, 1961, I collected a narrow-leaved species of Pyrrosia in Manipur. The specimen did not tally with any of the species described pre- viously. It was sent to Professor R. C. Ching (Peking Academy of Science), who confirmed that it is new, and suggested the name Pyrrosia nayariana, in honor of Dr. B. K. Nayar in recog- nition of his contributions towards a better understanding of the phylogeny of ferns, especially of the Indian species of Pyrrosia. Pyrrosia nayariana Ching et Chandra, sp. nov. Rhizoma breviter repens 2-3 mm. erassum, radicibus fascicu- latis nigris firmis, dense paleatum, paleis peltatis lanceolatis 2-3 mm. longis dentatis; folia conferta lineari-oblanceolata 15-20 em. longa, vix. 1.0 em. lata, sessilia vel subsessilia, apice acuta, basi gradatim attenuata, supra glabra, hydathodis punctatis in apice venulis ultimis liberis, subtus dense tomentosa, pilis stellatis tri- formibus, inferioribus densis ramis elongatis gracilibus glomer- atis, ‘superioribus floccosis ramis brevibus latis eymbiformibus, alteris ramis gracilibus acicularibus; folia fertilia saepe paullo NEw SPECIES OF PYRROSIA 63 angustiora quam sterilia, soris subtus ubique praedita, soris in 2 vel 4 lineis costae utrinque latere; sporangii annulus ex 16-20 cellulis compositus ; sporae oneae extine verrucoso. Type in the herbarium of the National Botanic Gardens, Luck- now, India, sheet No. 47,967, collected at Imphal, Manipur, India, at about 795 meters elevation, May 12, 1961, by P. Chandra (no. 74310). Isotype in the Botanical Institute, Academia Sinica, Peking, China. Pyrrosia nayariana (Fig. 1) grows in small epiphytic clumps on trunks and branches of trees in the plains. The fleshy rhizome is 2-3 mm. in diameter, short, branched, and enveloped by much- branched clusters of fibrous roots which form spongy masses at- tached to the substratum. The paleae (Fig. 3) are elongate- lanceolate, densely covering the rhizome, light brown in color, and peltate with an ovate basal region and a short, cylindrical stalk. The apex of the palea is acuminate and crowned by a sim- ple or branched, deciduous glandular hair ; the margin is dentate. The developmnt of the palea is as described earlier (Chandra, 1962; Nayar, 1961). Structurally, the rhizome is similar to that of P. flocculosa and P. mollis (P. fissa), but slender sclerenchyma strands, up to 16 cells thick, are irregularly distributed in the pith and inner cortex (Fig. 4). These strands are composed of very thick-walled, dark brown cells, with occluded lumen and prominent pit-connections in the walls. The cortical sclerenchy- ma sheath is 4-6 cells thick and golden brown in color. The vas- cular cylinder of the rhizome (Fig. 5) is a loose reticulum com- posed of many cylindrical vascular bundles, as in other species of Pyrrosia (Nayar, 1961). Leaf-traces to successive leaves origi- nate alternately on either side of the dorsal, median vascular bundle. Branching of the vascular bundles and the fusion of nearby branches is very common. Three slender branches, origi- nating from the dorsal median vascular bundle and a bundle lateral to it, supply each leaf (Fig. 51). The first-formed branch (usually from the median vascular bundle of the rhizome) forks once and forms the pair of abaxial bundles of the stipe. Compli- New Species oF Pyrrosta 65 cated vascular anastomoses occur before the separation of the leaf traces. Branches of the rhizome are associated with the leaves, and many are dormant, Each branch is supplied by a solitary vascular trace (Fig. 5 b), which originates along with the vascular connections to the leaf, as in P. flocculosa (Nayar, 1961). The leaves are articulated to short phyllopodia, the articu- lation being composed of a saucer-shaped pad of small parenchy- ma cells, as in Drymoglossum (Nayar, 1957). Peltate paleae cover the phyllopodium but paleae are absent on the leaf. The leaves are crowded, subsessile, and linear-oblanceolate (Fig. 1). The lamina is thick, leathery, glabrous on the upper surface, and densely covered by stellate hairs beneath (Fig. 2). A dense felt is formed by nearly hyaline, thin hairs with very long, more or less thin-walled, coiled, slender, frizzly arms (Fig. 6), which are entangled to form the dense felt. Above this layer are two types of hairs intermingled; one with short, broad, dark brown, boat-like, crowded arms ( Fig. 7) and the other of a lighter color and bearing few, long, slender, needle-like arms (Fig. 8). Promi- nent, pitlike depressions representing foliar hydathodes oceur on the upper surface. The venation of the lamina (Fig. 9) is funda- mentally similar to that of many species of Pyrrosia described by Nayar (1961). The midrib is prominent, and bears alternately placed, loosely arranged, immersed, lateral veins. The latter are at an angle of 30-40° with the midrib and are connected by three (or four) secondary branches at right angles to them, resulting in a row of elongated, rhomboidal areoles. The secondary veins bear two or three, free-ending, short, tertiary branches on the side facing away from the midrib. Most of these tertiary branches subtend hydathodes on the upper surface of the lamina. Each stoma is encircled by a single epidermal cell. The fertile leaves are similar to the sterile ones in structure and venation. The sori are punctiform and distributed over all of the under side of Sohal ariee, Sean Figure 1, PYRROSIA NAYARIANA CHING & CHANDRA. FIGURE 2. STELLATE HAIRS ON UNDER SURFACE OF LEAF, AMERICAN FERN JOURNAL NEw SPECIES OF PyrRRosIA 67 the leaf, directly above the apices of the tertiary veinlets. The sporangia are long-stalked. The annulus is 16-20 cells long. The spores (Fig. 10) are monolete (bilateral), plano-convex in equa- torial view, ovate in polar view, and 61 < 86 X 62 w(PX Ey X Ez). The laesura is thickened. The exine is golden brown and densely verrucose, the protuberances being small and more or less uniformly distributed. Pyrrosia nayariana resembles small-leaved specimens of P. mollis (P. fissa). In the latter the annulus of the sporangium is 20-24 cells long and the spores are more rounded (54 86 X 94 w) and smooth-walled, whereas the annulus is 16-20 cells long in P. nayariana and the spores are verrucose and conspicu- ously larger. I am grateful to Professor K. N. Kaul, Director, National Botanie Gardens, for his keen interest in this work, and to Dr... - K. Nayar under whose guidance this work was done. Thanks are due to Dr. R. C. Ching for kindly confirming the identifica- tion and Suggesting the specific name, and to C. V. Morton for the Latin diagnosis. LITERATURE CITED CHANDRA, PRAKASH. 1962. Development of the peltate palea in Poly- podiaceae. Current Science 31: 477-479 1 “Nayar, B. K. 1957. Studies in Polypodiaceae—IV, Drymoglossum. Jour. Indian Bot. Soe. 36: 169-179 - 1961. Studies in Polypodiaceae—VIII, Pyrrosia, Jour, Indian Bot. Soc. 40: 164-183, NATIONAL BoTANic GARDENS, LuckKNow, INDIA. Figures 3—10, STRUCTURAL DETAILS: 3, PALEA ; 4. TRANSVERSE SECTION OF RHIZOME, SHOWING SCLERENCHYMA AND VASCULAR BUNDLES; 5. VASCULAR STRUCTURE OF A PORTION OF THE RHIZOME (ROOTS OMITTED) ; 8. FOLIAR HAIRS ; 9. VENATION PATTERN; 10. SPORE, EQUATORIAL VIEW. (A = STALK OF PALEA; B = BRANCH TRACE; C = OUTER CORTEX; D = DORSAL MEDIAN VASCU- LAR BUNDLE; L = VASCULAR CONNECTION TO LEAF; M = LEAF MARGIN; MR = MIDRIB; R = ROOT; S = SCLERENCHYMA STRANDS; SC = CORTICAL SCLERENCHY MA SHEATH; V = VASCULAR BUNDLE). 68 AMERICAN FERN JOURNAL Notes on the Hawaiian Fern Genus Adenophorus KENNETH A. WILSON The majority of the Hawaiian representatives of the fern family Grammitidaceae have recently been interpreted as rep- resenting a single endemic genus (Copeland, 1947). This con- cept is not entirely new. As early as 1824, in a short report describing some new plant genera collected during Freycinet’s voyage around the world, Charles Gaudichaud (1824) published a description of the genus Adenophorus' in which he included three species. Two of these species had been described earlier Kaulf. (Adenophorus minuta Gaud.). The third species, Adeno- phorus bipinnata Gaud.) and Polypodium hymenophylloides Kaulf. (Adenophorus minuta (Gaud.). The third species, Adeno- phorus tripinnatifidus Gaud. was described here for the first time. This genus as originally described included those Ha- waiian grammitids with pinnatifid or bipinnatifid pinnae, bear- ing glandular epidermal hairs and possessing glandular-stipitate paraphyses in the sori. Evidently as a result of nomenclatural complexities, Desvaux (1827) later proposed the new name Amphoradenium for the described species of Adenophorus, and also provided a new epi- thet for Adenophorus tripinnatifidus in honor of Gaudichaud (Amphoradenium gaudichaudii Desv.). Clearly, however, the correct name for this group of grammitids when treated as a genus is Adenophorus Gaud. Amphoradenium is a synonym of this name. Amphoradenium, however, is the name that was adopted for this genus by Copeland (1947), who also made numerous new combinations Investigation shows that, according to Copeland’s concept of the genus, Adenophorus includes two distinct groups of species. 1The name leioten pots had been published earlier by Desvaux (1808) in a list of names without any description. According to Article 12 of the ei - Noresestatare i Moutccals this earlier name has no status under the code. 7S Nores ON ADENOPHORUS 69 The first group is characterized by having elongate creeping rhizomes, remote fronds, pinnate-pinnatifid to pinnate-tripinna- tifid blades, with an irregular epidermal layer bearing numerous appressed, usually unicellar, reddish-brown glandular hairs which are oriented so that they point to the apex of the ultimate blade segment or lobe. The second group includes the species with short, erect or ascending rhizomes closely invested by the crowded stipes, simple, pinnatifid or pinnate blades, with a smooth epidermis bearing scattered uniseriate, simple or more frequently branched, erect hairs. In the first ai belong Adenophorus hymenophylloides Hook. and Grey., A. tamaris- cmus Hook. & Grev. (as tamarisct), A. Mueens Gaud., A, hillebrandii (Hook.) K. A. Wilson? and A. abietinus (Eaton in Mann) K. A. Wilson.3 The second group includes Adenophorus sarmentosus (Brack.) K. A. Wilson,t Adenophorus haalilioanus (Brack) K. A. Wil- son® and Adenophorus pinnatifidus Gaud. The presence of clavate glandular paraphyses in the sori is the single morphological character that unites these two groups of Hawaiian grammitid ferns. The distinctness of these two groups raises the problem of the generic validity of Adenophorus. Considering only the Hawaiian species of the Grammitidaceae, it might be suggested that each one of the two groups merits generic status. The great similar- ity of the species within each group argues strongly in favor of their close relationship. The degree of relationship between the two groups, however, is not clearly established. Convincing evi- 2ADENOPHORUS hillebrandii (Hook.) K. A. Wilson, comb. nov. Basionym: P olypodium hillebrandii 4 Sp. Fil. 4: 228, tab. 279. 1864. SADENOPHORUS abietin + (Raton in Mann) K. ilson, comb. no Basionym: Polypodium ea Eaton in Mann, Proe. Am. Aead. Arts and Sei. 75 219. 1867. ADENOPHORUS sarmentosus (Brack.) K. A. Wilson, comb. nov. Basio- nym; en sarmentosum Brack. U. 8S. Explor. Exped. Bot. 16: 8, 9. 1854; pl. 2, fig 1855. py netid a dil haalilioanus (Brack.) K. A. Wilson, comb. asio- nym: Polypodium rene Brack. U. 8. Explor. Exped. Bot. Ter 5, 6. 1854; pl. 1, fig 4. 70 AMERICAN FERN JOURNAL dence could be presented in favor of treating these ferns as two separate genera. This situation is not uncommon throughout the entire. family Grammitidaceae. There seems to be general agreement that the genus Xiphopteris is polyphyletic and, although easily recog- nizable, an artificial grouping. The assemblage of ferns placed in the genus “Ctenopteris” (the name is illegitimate) includes numerous groups of ferns. The interrelationships of these ferns is very poorly understood. Prosaptia is another grammitid genus which is probably polyphyletic. Problems similar to those in “Ctenopteris” are also evident in the large genus Grammitis. Clearly, a detailed study of the entire family is needed in order to establish clearer, more natural genera. Until such a study is undertaken I consider it best to retain these convenient, although most likely artificial genera. LITERATURE CITED BRACKENRIDGE, W. D. 1855. In C. Wilkes, U. 8. Expl. Exped. 1838-1842. Vol. 16 COPELAND, E. B. 1947. Genera Filicum. Waltham, Mass. Desvaux, N. A. 1808. Sur les genres Thorea et Lemanea, de M. Bory de Saint-Vincent. Jour. Botanique Appliquée 1(2): 121-125. Bea Ay fe tiie de la famille des fougéres. Mem. Soe. Linn. aris, 6: rn ; gaia RE, C. 1824. Descriptions de quelques nouveaux genres de plantes ea dans le voyage autour du monde. Ann, Sci. Nat Bot. Zool. 3: 507-510. Hooker, W. J. 1863-64. Species Filicum. Vol. 4. London, Kauruss, G. F. 1824. Enumeratio Filicum ipsi MANN, ee iis, Enumeration of Hawaiian ‘pate ibe. Am, Acad. 7: 14 ee Pao VALLEY STATE CoLLEGE, NORTHRIDGE, CALI- FORNIA, New ComBINATIONS IN LycopopiuM 71 New Combinations in Lycopodium C. V. Morton About ten years ago, the U. S. National Museum lent a large number of unnamed specimens of tropical American Lycopodiums to Dr. W. Herter, the noted authority on the Lycopodiaceae. Dr. Herter found a number of new species among these collections, which he published in two papers, under the generic name Urostachys Herter. The generic name Urostachys, dating from 1922, is antedated by two or three older names for the same concept.! However, it does not seem to me either necessary or desirable to split up the genus Lycopodium, which certainly seems to be natural, although with plants as old as these and which offer so few characters it is hard to be sure. But I agree with Boivin,? that if one begins to split up Lycopodium it will be necessary to recognize not just two genera but five or more, which is surely extreme, considering the inadequate present state of our knowledge of the gametophy- tes, life history, and anatomy of the species. Apparently, this group of Lycopodium is rich in species, for most of those described by Herter seem to be well founded. Many of the specific epithets chosen by Herter are fanciful, derived from the names of mythical personages (cassandrae, dianae, poseidonis) or stars (arcturi, capellae, stellae-polaris, erucis- australis) or both (castoris), but why not? There are too many humdrum hirsutums and acuminatums. In order to refer in identification lists and otherwise to the various new species described by Herter based on material in the U.S. National Herbarium, I propose the following new combina- tions ; See Hans Peter Fuchs, ae nomen genericum conservandum?”, ven Naturf. Ges. Basel 66: 33-48. 1955. “Bernar Boivin, “The Problem of Gencrie ee ae in the Form-Genus Lycopodium,” Amer. Fern Jour. 40: 32-41. {2 AMERICAN FERN JOURNAL returi Herter, Rev. Sudam ahs 10: 118, 1953. LycopopIuM achat (Herter) Morton, es. Urostachys arthuri Herter, op. cit. Lycopopium bon ae-voluntatis (Herter) Monten, comb, nov, Lycopopium arecturi (Herter) Morton, comb. U ys oa Be LycopopluM ecassandrae (Herter) veneee comb. nov. Urostachys cassandrae eee op. cit, 116, LYCOPODIUM eastoris (Herter) Morton, comb. nov. Urostachys castoria Herter, op. cit. 11 ‘Lycopoptum chamaeleon (Herter) Morton, comb. 2° Urostachys chamaeleon Herter, Amer. Fern Jour. ore 82, 1958. LyYcopopIUM costaricense (Herter) Settee seas, nov. Urostachys costaricensis Herter, op. ae! Lycopopium crucis-australis (Herter) ae comb. Urostachys crucis-australis Herter, Rev. Sudamer. eee 10: 119. 1953. LYCOPODIUM ecuatrecasasii | Herter) Morton, comb, Urostachys hystrix Herter, op. cit. 120. LycopopIuM innocentium (Herter) atten: comb. nov. Urostachys innocentium Herter, op. cit. 127. bis Secale killipii (Herter) oy gah nov. achys killipii Herter, op. 128. ire see leptodon (Herter) ve in sched., comb. nov. Urostachys leptodon Herter, op. cit. 120. Lycopopium maebridei (Herter) sso a comb, nov, Urostachys macbridei Herter, op. cit. 115. Lycopopium poseidonis (Herter) Morton, comb, nov. Cae Lycopoptum soeratis (Herter) Morton, aad. nov, Urostachys socratis Herter, op. cit. LycopopiuM stellae-polaris (Herter) iiton: comb. nov. /rostachys stellae-polaris Herter, op. cit. 121. HINTs ror FERN Cuururists 73 Lycopopium trachyloma (Herter) Maxon, in sched., comb. nov. it. 113. Urostachys ulixis Herter, op. cit, 115 Herter’s reasonably good work was not duplicated by that of his student Hermann Nessel, whose book “Die Barlappgewachse” added little to our knowledge of the group. However, some of the species described by Nessel are good, and one of them is rather common in the Andes of Colombia: Lycopopium wohlberedtii (N essel) Morton, comb. nov. Urostachys wohlberedtii Nessel, Repert. Sp. Nov. Fedde 39: 69. 1935. SMITHSONIAN INsTITUTION, WasHINGTON, D. C Some Hints for the Fern Culturist) Irvine W. Knosuocu The culturing of ferns in all of its phases is of intense interest to many of our members. One of the most challenging aspects of this hobby is raising ferns from spores and bringing the spore- lings to maturity. A few of the more important articles on this Subject are those by Hires (1940), Benedict (1955) Klein- schmidt (1952, 1957), Boydston (1958). Fliflet (1961) sum- marizes much of the former material on growing ferns from Spores, and there is very little new that can be added. Fern enthusiasts might, however, be interested in a new sub- Strate for spore culture. The name of this material is Turface. Itisa ground, calcined, clay product designed as a soil amend- ment to loosen soil, stimulate root growth and for similar gar- dening uses. It is manufactured by the Wyandotte Chemical Corporation, J. B. Ford Division, Wyandotte, Michigan. Cur- rently it is sold in 50-pound bags but it is my understanding that it is soon to come out in smaller quantities and sold through chain stores. Spores can be sown on any reasonable substrate, of entries 1Contribution No. 63-1 from the Department of Botany and Plant Path- ology, Michigan State University. Photograph by Phillip Coleman. 74 AMERICAN FERN JOURNAL course. For example, one can hardly surpass oak soil (Klein- schmidt, pers. comm.) for many ferns. The very rough nature of a bed of Turface, however, seems to provide varying microeco- FIGURE 1. CULTURE )F PROTHALLIA AND YOUNG SPOROPHYTES IN FLASK. FIGURE 2. YOUNG SPOROPHYTES IN POT COVERED WITH PLASTIC FREEZER BAG. Hints ror Fern Cuuturists i logical niches for spores and we have had very good luck with the material so far. As seen in figure 1, we have been placing the Turface in small flasks. We soak the medium in a modified Knutson’s nutrient medium (almost any other will do), and then sterilize the flask in an autoclave for about 30 minutes. The spores are then sown through the mouth of the flask. One should have spores of known origin (uncontaminated) to begin with and Kleinschmidt’s method (1957) of sterilizing the fronds be- fore all of the spores have been shed is an excellent one. This precaution will largely prevent contamination by unwanted fern spores and also reduce fungal contamination. The sexual phase or gametophytes will develop from the spores if conditions are right. Usually one is only desirous of obtaining the sporophytes and one small trick here usually or frequently overlooked is to water the mass of prothallia from above (distilled water will do) So as to ensure fertilization and the development of the sporo- phyte. It should be mentioned in this connection that growing ferns in a closed flask largely eliminates as much watering as found in the potted soil method. One can tell when the clay needs watering by noting its color, which changes with varying percentages of water, When the sporophytes are about an inch high, they are ready for transplanting. They can be spaced in pots or flats after re- moval (by shaking or by long forceps) from the flasks. This Stage has always proven to be a hazardous one and many spore- lings die. A way to ensure success with the transplanting and to force the plants along, is to place the pots of transplants (in moist soil) in a plastic freezer bag, label, using a wooden label, and close the top of the bag with a rubber band (fig. 2). The humid atmosphere in the bag offsets transplant loss and, each bag, acting as a miniature greenhouse, causes the plants to grow much faster than they ordinarily would. LITERATURE CITED BENEDICT, R. C. 1955. Aids to Spore Culture. Amer, Fern Jour. 45: 60-64. 76 AMERICAN FERN JOURNAL .Boypston, Karuryn 1958. An Amateur Plants Fern Spores. Amer. Fern our, 48: 1-18. . Furrtet, THORLEIF 1961. Growing Ferns from Spores. Amer. Fern Jour. 5 13-127. . Hires, Ciara 8. 1940. Growing Ferns ee abl on Sterile Nutrient Media. Jour, N. Y. Bot. Gard. 41: 25 KLEINSCHMIDT, W. F. 1952. Growing nek ee Spores. Amer. Fern Jo 42: 117-124. 957. A Method of Preparing Spores for Fern Cultures. Amer. Fern Jour. ye 5- MICHIGAN STATE UNIvErsIry, East Lanstna, MICHIGAN. Ethnobotanical Uses of California Pteridophytes by Western American Indians Ropert M. Luoyp Utilization of pteridophytes by Western American aborigines played a minor but very interesting role in their lives. Most o the pertinent information is scattered. This paper attempts to summarize much of this information. The species known to have been used by the western North American Indians north of Mexico are discussed below. The botanical nomenclature is according to Munz (1959). Lycoropium cLavatum L. Club-moss. Although there is no evidence for use of this species by western Indians, Frye (1934) reported its spores were collected for dusting on open raw wounds and chafed infants. The spores are very fine and light, which en- ables them to repel water and prevent stickiness. Lycopodium selago L., a related species in the northwest was used by Indians as an intoxicant. The stem was chewed and the juice swallowed. It is said that three plants produced a mild intoxication whereas eight plants stupified the user. EQUISETUM ARVENSE L. Common horsetail. The major use of the horsetails was as an abrasive in polishing bows and arrows (Murphey, 1959). However, it was sometimes dried and burned and the ashes used on sore mouths. The Lower Chinook Indians ETHNOBOTANY OF CALIFORNIA PTERIDOPHYTES fi of Washington gathered, peeled and ate raw the stems of early reproductive shoots (Gunther, 1945). EQUISETUM HYEMALE L. var. ROoBUSTUM (A. Br.) A. A. Eat. Medical uses of this plant included treatment of the prostate gland with infusions made from dried material taken orally (Romero, 1954). Washington Indians boiled the stems and washed their hair to get rid of vermin. Infusions of boiled stems and willow leaves were given to girls to regulate the menstrual period. The Makah ate the reproductive shoot heads for diarrhoea and used the root juice as an eye wash (Gunther, 1945). Karok Indians used the stalk as an abrasive to sharpen the edge of mussel shells used in cleaning fish and to polish ar- rows. Schenck & Gifford (1952) reported that infusions were used to cleanse priests in the Karok’s First Salmon Ceremony. EQuIseTUM KANSANUM Schaffn. Hot infusions were drunk for backaches. Cold infusions were used as a lotion. This species was also used in the Waterway Ceremony of the Kayenta Navajo. Equisetum LaEvicaAtum A. Br. Hopi Indians dried and ground stems and mixed this with corn meal to make a mush used for food and in preparing ceremonial bread (Castetter, 1935). EQUISETUM TELMATEIA Ehrh. Giant horsetail. Portions of the root stalk were cooked and eaten by the Cowlitz Indians of Washington. Stems were also used for polishing and scouring and as fodder for horses. ADIANTUM CAPILLUS-VENERIS L. Venus-hair fern. Kayenta Navajo used infusions as treatment for bumblebee stings and centipede bites, and for insanity. It was also smoked in the Lifeway Ceremony. Romero (1954) reported some California Indians used this species in treating menstrual irregularities. ADIANTUM JORDAN C. Muell. California maidenhair. The most interesting use for any fern is that which Chesnut (1902) reports for this species. The Indians of Mendocino County valued the smooth black midribs for keeping and enlarging ear- ring holes. ADIANTUM PEDATUM JL. var. ALEUTICUM Rupr. Five-finger 78 AMERICAN FERN JOURNAL fern. Despite its wide distribution this plant was only limitedly used. Its black shiny midrib coverings were a main component for making designs on baskets by many tribes (Balls, 1962; Merrill, 1923). Washington Indians soaked the fronds and used the infusions on the hair. The fronds were chewed for sore chest and stomach trouble and to stop bleeding from wounds. For the latter reason it was sometimes carried with war parties. Ashes of the fronds were rubbed on the hair to produce shiny black braids. Schenck & Gifford (1952) reported that the fern was used for decoration in the Jump Dance dress of the Karok Indians. PELLAEA MUCRONATA (D.C. Eat.) D. C. Eat. Bird’s-foot fern. Infusions of the frond were drunk as a tea for both medical and pleasurable reasons by the Luiseno Indians (Sparkman, 1908). The Minok used the tea to stop nose-bleeds and to purify the blood (Barrett & Gifford, 1933). PELLAEA sp. (P. ATROPURPUREA in Romero, 1954). When steeped this fern produced a delicious ephedra-like tea to flush the kidneys and tone the blood. In summer it was used to pre- vent sunstroke. PITYROGRAMMA TRIANGULARIS (Kaulf.) Maxon. Gold-back fern. This species was chewed by the Minoks for treating tooth- aches. It was also applied in mitigating the afterpains of child- birth by the Karok. Sometimes its midribs were substituted for those of the maidenhair fern in basketry. Chestnut (1902) re- ported that the fronds were slapped on clothing by children to make temporary gold colored prints. {This practice is still fol- lowed by white children !—Ed.] PTERIDIUM aQuiLinum (L.) Kuhn var. LaNuGINOsuM (Bong.) Fern. Brake or Bracken fern. The bracken fern was perhaps the most widely used fern in California. In basketry untreated roots provided the only brown coloring for designs (Merriam, 1955). Frequently these were blackened by heating in water, the degree of coloring depending on the length and temperature of the hot-water treatment. The Karok cleaned salmon on beds of the fern and used the fronds for wrapping tobacco. Some ETHNOBOTANY OF CALIFORNIA PTERIDOPHYTES 79 California tribes cut and cooked the young sprouts to obtain a rich flavored oil and starch. Wittrock & Wittrock (1942) re- ported that rhizomes were used by many tribes as an important food. The white heart was roasted until it resembled the dough of wheat. This was relished as a nutritious but pungent food- stuff. Young sprouting shoots were also used in preparing soup, or were eaten raw (Yanovsky, 1936). Baked rhizomes could be stored for later use. The Quinault Indians of Washington were said to have used the fibers of the rhizome for making string (Gunther, 1945). They also made a fern-paste bread from the pulp. The Indians of Mendocino County used the large fronds for beating down grass fires and lining berry baskets (Chestnut, 1902). It was also used as a diuretic for horses. ATHYRIUM FILIX-FEMINA (L.) Roth. var. sIrcHENsE Rupr. Lady-fern. Roasted rhizome centers, roots, and new shoots were eaten by Washington Indians. The fronds were also used to wipe fish after cleaning. Boiling rhizomes produced a tea which was drunk to ease body pains. Boiled stem infusions were taken to ease labor pains. Dryopreris pitarata (Hoffm.) Gray. Wood-fern. Pounded roots of this species were applied to cuts and frond infusions were used as a hair wash. The rhizomes were sometimes baked and eaten. Dryopreris FILIX-MAs (L.) Schott. Male-fern. The only re- corded use for this fern is as a vermifuge (Wittrock, 1942). Potysticoum muNiTuM (Kaulf.) Presl. Sword-fern. Rhi- zomes were peeled and baked in pits with salmon eggs and eaten. The fronds were sometimes used to line the baking pits or to serve as racks for drying berries. The Quilente and Cowlitz Indians made mattresses from them. Medicinally, the young curled fronds were chewed raw and swallowed for sore throats or tonsilitis or to facilitate childbirth. Infusions of the frond were sometimes placed on sores and boils. Boiled rhizome in- fusion was said to eure dandruff. The sporangia were put on burns. Sword fern fronds were used in a game to determine long 80 AMERICAN F'ERN JOURNAL wind. Leaflets were touched or torn off beginning at the bottom of the frond and certain words were spoken for each. The win- ner was the one who reached the highest point on the frond in one breath (Schenck & Gifford, 1852). BLECHNUM spIcant (L.) With. Deer-fern. Fronds were only used in case of emergency by lost children or to prevent thirst on long journeys. Infusions of the leaves were drunk for gen- eral ill health. Green fronds were eaten for lung trouble, stom- ach distress, and colic. OODWARDIA FIMBRIATA Sm. in Rees. Chain fern. Root fibers were used in basket designs either naturally colored or dyed red. The Luiseno Indians used infusions from the roots to relieve pain from injuries. In basketry the midrib of the frond con- tained two fibers which were stripped and handled like lengths of yarn. Sometimes they were dyed with alder bark or with moss to a burnt orange. POLYPODIUM CALIFORNICUM Kaulf. California Polypody. Ches- nut (1902) reported that juice of the rhizome was rubbed on sores for healing and on the body to treat rheumatism. Root extracts were sometimes used as an eyewash. PoLypopiuM GLycyrRHIza D. C. Eaton. Licorice fern. The rhizome of this plant was roasted and chewed as treatment for coughs. The Cowlitz crushed and boiled the rhizome and mixed it with fir needles to treat measles. DENNSTAEDTIA PUNCTILOBULA (Michx.) Moore. Romero (1954) erroneously reported this eastern fern from the mountains of California. Its only use was in treating tuberculosis and other lung diseases by drinking oil extracts from the root. The iden- tity of this plant is unknown but it is not the species above. Of the almost 85 species of ferns and fern allies oceurring in California less than one-fourth are known to have been utilized by western American Indians. Upon close examination of the species discussed in this paper, most are found to be restricted in range to the cismontane and other mesophytic regions of the coastal and southwestern states. However, the majority of ETHNOBOTANY OF CALIFORNIA PTERIDOPHYTES 81 Indian tribes covered in this study inhabit these regions. Uses of ferns by desert tribes has been restricted to a few of the many species occurring in their region. The Hopi Indians of north central Arizona, for example, used only Asplenium tri- chomanes and Equisetum laevigatum (Whiting, 1939). Elmore (1944) found no pteridophytes being used by the Navajo, al- though they did have a name for Cheilanthes feet. Wyman & Harris (1951) reported two ferns used by the Kayenta Navajo. By contrast, Indians of more mesophytic regions made more varied use of a greater number of ferns. Schenck & Gifford (1952) listed eight species used by the Karok of Humboldt County. Chesnut (1902) said six species were utilized by Mendocino County Indians and Gunther (1945) found ten species being used by Indians of western Washington. None of the species listed was a major constituent of a tribe’s sustenance except perhaps Pteridium aquilinum var. lanugino- sum which was valued as food and for less important uses. Al- though a few of the remaining species were important in a utili- tarian manner most of them were luxuries. LITERATURE CITED Baus, E. K. 1962. Early Use of California Plants. Univ. of Calif. Press, Berkeley Barrett, 8, . & E. W. Girrorp. 1933. Minok material culture. Bull. Publ. Museum City of Milwauke wo Wa CastTetrer, E. F. 1935. if ciui i abinaeioid studies in the American Southwes 1. Uncultivated native plants used as sources of food. Bull. Univ. Av Mexico Biol. Ser. 4: 1-62. CHESNUT, V. K. 1902, Plants used by the Indians of Mendocino County, California. Contr. U. 8. Nat. Herb. 7: 295-422 ee F. H. 1944, edishotans of the Mavaie. Monogr. School Am - Santa Fe, No. 8. Pave, ‘4 C. 1934. Ferns of the Northwest. Metropolitan Press, Portland, re. GUNTHER, ERNA, 1945. Ethnobotany of Western Washington. Univ. Wash. Pub rai be MERRIAM, 196 > phates of California Indians. Univ. of Calif. Press, Berkeley 82 AMERICAN FERN JOURNAL MerriLL, Ruta E. 1923. Plants used in basketry by ye California Indian, Univ. Calif. Publ. Am. Archaeol. and Ethnol. 20: 242, Munz, % ve 1959. A California Fiore. Univ. £ Calif. a. Berkeley. MurpuHey, EpirH. 1959. Indian Uses of fav Plants. Desert Printers, Dicer, Calif. Rowrno, J. - 1954. The Botanical Lore of the California Indians. Vantage Press, New York. PEt ‘hace & E. W. Girrorp. 1952. Karok ethnobotany. Anthropol. Ree. 13: hag Si SPARKMAN, P. §. 1908. The olagh of the Luiseno Indians. Univ. Calif. Publ. . Archaeol. and Ethnol. 8: 187-234. igre 63 x 1939. Ethnobotany of the Hopi. Bull. Museum Northern Ari Wien, geet & a L. he 1952. Food plants of the Indians. Jour. N. Y. Bot. Gard. 43: 57-71. ect MARION. oes Petey from plants. Jour. N. Y. Bot. Gard. 43: fas Wray, L. C. HA Ss Hangs. 1951. nee ethnobotany of the Kayenta Navajo. this. Haw M eae YANovsKY, E. 1936. Food plants of as ok American Indians, U. 8. Dept. Agr. Mise. Publ. No DEPARTMENT OF BorTany, ees OF CALIFORNIA, BERKE- LEY, CALIFORNIA, Phytogeography of Selaginella douglasii GEORGE NEVILLE JONES This western North American species has a restricted geo- graphical distribution, being known with certainty only in Wash- ington, Oregon, and Idaho. From time to time it has been at- tributed to both California and British Columbia, but there seems to be no available evidence that it has ever been found in either of those places. My attention was drawn to this matter by the critical monographie work of Alston (1955), who pointed to certain ambiguities in statements dealing with its geographical range. According to the classification of Alston & Walton (1938) Selaginella douglasui belongs in the subgenus Stachygynandrum, PHYTOGEOGRAPHY OF SELAGINELLA DOUGLASII 83 series Decumbentes. It was described first as Lycopodium ovali- folium by Hooker & Greville (1829), but this binomial was in- valid when published because Desvaux had applied it to a differ- ent species earlier. In 1831 Hooker & Greville renamed the plant Lycopodium douglasii, and it appeared under this name in Hooker’s Flora Borcali-Americana (1840). Spring (1843), the monographer of Lycopodium and Selaginella, included it with Selaginella as “24. S. douglasii (Lyc.) Hook. et Grev.,” the “Lye.” standing for Lycopodium. Originally Hooker ined the species on a collection by David Douglas. One hundred and forty-one years later this plant is known to occur only near the type locality along the Columbia River, and in one disjunct area in northern Idaho, some three hundred miles northeast- ward. The reports of Selaginella douglasii as a Californian species originated apparently with D. C. Eaton, who in the Botany of California (1880) wrote that it oceurs “Probably in Northern California.” Other later published reports of this nature include those by Clute (1905) who wrote, “It is reported to grow in northern California, Oregon, Washington, and British Colum- bia.” Piper (1906) gave the range as “Washington to Califor- nia.” Piper & Beattie (1915) correctly gave simply, “On wet rocks, anne abundant in the Cascade Gorge of the Columbia iver.’ Maxor in Abrams (1923) gave the range as British Columbia and northern Idaho to California, oh Jepson (1923) wrote, “Damp shades, n. Cal.; n. to B. C.” Peek (1941) attri- buted this plant to “Moist rocky slopes in fe ae abundant on the south side of the Columbia River Gorge; B. C. to Calif.. east to Idaho.” Munz (1959) merely mentioned 8. douglasii as “reported from n. Calif.” The continued ascriptions of 8. doug- lasi to California are remarkable because there is no other eae of Selaginella north of Mexico that resembles 8. one lasti. D. C. Eaton (1880, p. 350) commented that it is “more like some tropical forms than any of the northern species.” There is now strong presumptive evidence that S. douglasi oc- 84 AMERICAN FERN JOURNAL eurs not nearer than about 240 miles north of California. The reports of S. douglasit from British Columbia also are re- markably persistent. The first came from Baker (1887), who wrote, “Hab. British Columbia, Oregon, and Washington Terri- tory. A well marked species.” Alston (1955) noted that this seemed to have arisen through confusion with the Columbia River, and added, “Hooker wrote ‘Columbia, Douglas’ on the Kew sheet, and Baker has added ‘British.’” It may be observed that the town of Vancouver is on the Columbia River in the state of Washington, while the capital city of British Columbia bearing the same name is situated on the mainland of British Columbia. Hieronymus (1902) attributed S. douglasw to “nord- amerikanisch Columbien und in Oregon,” presumably refer- ring to the western Canadian province of British Columbia. Another possible cause for ascribing S. douglasii to British Co- lumbia may have resulted from confusing it with S. selaginoides (L.) Link, which Henry (1915) listed from “Selkirks and Roe- kies, Ucluelet [Vancouver Island, British Columbia], Ounalashka [Unalaska, Aleutian Islands].” There apparently is no evidence that S. douglas occurs naturally in British Columbia or else- where in Canada. Relevant references to Selaginella douglasii include the fol- lowing : Selaginella douglasti (Hook. & Grey.) Spring, Bull. Acad. Sci. Belg. 10: 138, 1843; Mém. Acad. Roy. Belg. 24: 92, 1850.— Eaton (1880, p. 350) ; Baker (1887, p. 47) ; Hieronymus (1902, p. 689) ; Clute 1905, p. 160) ; Piper 1906, p. 87) ; Piper & Beattie (1915, p. 14) ; Maxon (1923, p. 46); Jepson (1923, p. 42) ; Peck (1941, p. 57); Tryon (1949, p. 422); Alston (1955, p. 238) ; Munz (1959, p. 23). Lycopodium ovalifolium Hook. & Grev., Icon. Fil 2: t. 177. 1829, non Desv., 1827. “Hab. In oris occidentalibus Americae septentrionalis. Dom. Douglas.’ Lycopodium douglasii Hook. & Grey. in Hook., Bot. Mise. 2: 396. 1831.—“Hab. Near springs in woody places, N. W. Amer- PHYTOGEOGRAPHY OF SELAGINELLA DOUGLASII 85 ica, Douglas—An extremely distinct and well-marked species, which I have never received from any collector except Mr. Douglas.” (Hook, Fl. Bor. Am. 2: 268. 1940 (?) Lycopodium denticulatum sensu Wilks, Jour. David Doug- las 145, 1914, non Linn. Type: Abundant in moist places on the hills near Grand Rapids, Columbia River above Vancouver, Sept. 1825, Douglas 482 WASHINGTON Cowlitz County: Kalma, Se hill (K). SKAMANIA County: Cape rn, Piper 4965 (ILL, WS); Suksdorf 2647 S); 7. J. Howell, June ‘bat (MO); Castle Rock (Beacon Rock) Suks dorf 8475 (WS); east side of Beacon Rock, near summit, shady rocky situation, J. W. Thompson, July, 1956 (WTU) OREGON CLACKAMAS CouNTY: Shady banks of Eagle Creek, May 20, 1928, J. W. Thompson 4259 a OSC, WTU); moist fir woods, on logs, mossy ground, ete., Eagle Creek, M. W. Gorman 5115 (WTU); Milwaukie, May 1881, at Ses Howell (OSC, rome a Howell 695 (WS); above Estacada, Aug. 1914, M. FE. Peck (MO, W gos River Country: Herman Creek trail, Aug. 15, 1915, L. N. Goodding (OSC), Aug. 15, 1930, S. B. eee (OSC); Columbia Gorge, Apr. 10, 1943, elen M, brasil (OSC); Bridal Veil, Columbia River Gorge, G. N. Jones 8767 (IL Poa a es rocks, MeCord Creek, Columbia River Gorge, G. N. Jones 10091 dL MULTNOMAH a. Damp shaded rocks near Bonneville, Suksdorf 832 tenage 6 (Ones: 3; on vies on trail to fa ils, Wahteona Falls, Sept. 12, eae FS ches & F. Nicol (OSC); on moist sunny cliffs, Cornell Road, Barina d, Henderson 1217 (OSC); vicinity of Portland, William Palmer Fas akc, Portland, Godman 342 (BM); damp shaded bank, McClay Park, Portland, June 15, 1915, M. A. Flinn (OSC); rocky places by spring just beyond Corbett, J. W. Thompson 297 9 (WTU); Multnomah Falls, J. W. Thompson 4192 (WTU), 4978a (WTU), 11368 (MO, WS, WTU). 86 AMERICAN FERN JOURNAL County Unknown: Columbia River, near spring in wood, Douglas (K); abundant in moist places on the hills near Grand Rapids, Columbia see er above Vancouver, Sept. 1825, Douglas 482 (ex Wilks); banks of the lumbia, Lobb (BM); banks of the Columbia River, western Oregon, 7. J. Howell 245 (MO). IpaHo CLEARWATER CouNTY: Granite cliffs along the Clearwater River, 5 miles south of Orofino, W. H. Baker 13979 (ID). IpaHo County: Wet sandy soil, South Fork, Clearwater Riv ver, R. J. spa 8401 (IDS, WS); on moist me in deep shade, about 25 miles west Elk City on the South Fork of the Clearwater River, C. L. Hitchcock rea (WTU); faite Falls, H. J. Rust 2595e (ID); on moist cliffs, ps side of Selway River, 20 miles southeast of Lowell, J. H rt 2 (WS); jae rock cliff, Selway Falls, J. H. Christ 2595 (ID); Sel- way Peahe . C, Aase 1774 (BM, MO, WS); Ownbey & Ward 3131 (WS); Three Devils Camp, Daubenmire £526 (WS); Three Devils Camp Ground, four miles west of Lowell, W. H. Baker 14804, 14526 (ID); southeast of Harpster, Daubenmire 47144 (WS); on cliffs along Lochsa, one mile east of Lowell, J. H. Christ 12081 (ID); rocky cliffs near Deadman Creek on the Lochsa River, Young (BM). Nez Perce County: Moist wooded slopes along the Clearwater River, five miles east of Spalding, W. H. Baker 6459 (ID). LITERATURE CITED AtsTon, A. H. G. 1955. a heterophyllous pean of continental North America. Bull, Brit, Mus, (Nat. Hist.) Bot. 1: 219-274. ~~ & J. Wanrox, 1988 Lycopene, in F. Verdoorn, Manual of ees a 500-506. The Ha BAKER, J. G. 7. Handbook ef the fox. abba London. CLurr, W. "se The fern allies bd North America north of Mexico. New York. Eaton, Ze C, ae Pteridophyta, in S. Watson, Botany of California, 2: 329- HENRY, ae 8 ‘1915, Flora of Southern British Columbia. Toronto. HIERONYMUs, G. 1902, Selaginellaceae, in Engler & Prantl, Nat. Pflanzenf. 1(4): 621-715. HOOKER, oe J. 1840. Flora Boreali- -Americana, vol. 2, Lon sates a & EB. KC Garvnre, 1890-31 tonne acu vol. 2. London Jepson, W. L. 1923. A manual of the flowering plants of California: Berkeley, aia VIABILITY OF LYOPHILIZED SPORES 87 eee W. R., 1x L. R. Aprams, 1923. Selaginella, in Illustrated Flora of he Pacific States 1: 46-50. Stanfor wens P. A. 1959. A California flora. Bankicley. Peck, M. E. 1941. A manual of the higher plants of Oregon. soley: Pirer, ©. V. 1906. Flora of the state of Washington. Contr. U. 8. Herb. 7. 87. . K. Brearrie. 1915. Flora of the Northwest Coast. Pull- m Spring, A. 1843. Enumeratio Lycopodinearum, Bull. Acad. Sci. Belg. en 1800. Moncarspne de la famille des Lycopodiacées. Mém. Acad. Roy. ae TRYON, ALICE. 1949. Bier res of the genus Selaginella in North America rth of Mexico. Ann, Missouri Bot. Gard. 36: 413-432, f. 1-32 on pl aaa WILkKs, W. 1914. Journal of David Douglas. Lond DEPARTMENT OF BoTANY, UNIVERSITY OF tained URBANA, ILLINOIS. Retention of Viability in Lyophilized Spores of the Fiddlehead Fern, Matteuccia pensylvanica' W. G. BARKER AND R. G. WHITE For many years, residents of New Brunswick, Canada, have used young tender rolled fronds of Matteuccia pensylvanica as a table vegetable. In recent times, this use has assumed such eco- nomic importance that curled fronds are now gathered in the spring as a fresh crop, or are commercially processed either by freezing or canning. There has been no attempt at raising them commercially. Instead, they are harvested with difficulty from their wild habitat in shaded places along rivers and streams, mostly by Indian laborers. Because of the increasing impor- tance of this crop, one of us (R.G.W.) has undertaken to study the propagation and management of these plants on a regular- ized basis. 1Contribution No. 128 from the Research Station, Canada Department of Agriculture, Fredericton, ew Brunswick. 88 AMERICAN FERN JOURNAL There is a somewhat sparse literature on the handling of fern spores. However, it has been noted (Bold, 1957) that spores of Osmunda spp. must be sown shortly after they mature. Fur- ther, Campbell (1887) reported that Onoclea sp. spores begin ger- mination within a very few days and therefore grow rapidly to form the prothallus. On the other hand, Bailey (1915) reported that these organs, properly stored, can be kept alive for long periods. No statement could be found concerning the possibility of lyophilization, although similar techniques had been effec- tively employed in the preservation of ginkgo pollen (Tulecke, 1954) (but not the pollen of Pinus palustris Mill.) (Hesseltine & Snyder, 1958) and in various fungal diaspores (Mehrotra & Hesseltine, 1958). In consequence, it was decided to obtain this information with respect to this fern species. Spores were collected September 23, 1962, as they matured in the wild. They were shaken free of the parent frond and placed in flasks of a Virtis Macro freeze-dry apparatus and frozen quickly by partial submersion in a mixture of dry ice and etha- nol. Subsequently they were freeze-dried under vacuum imposed by a Duo-seal pump for 8 hours. They then were placed in bottles and stored at —20°C. in desiccators over Drierite crystals. Following one and three month storage, the lyophilized spores were sown in several media: well rotted leaf mold, a mixture of peat, sand and soil, and in sterilized sphagnum. In every in- Stance, the spores germinated readily and produced vigorous thalli. Both plantings have developed the sporophyte generation and are growing well. There has been no apparent drop in the viability of the lyophilized material. Spores of the fiddlehead fern, M. pensylvanica, can undergo lyophilization and_ retain viability after storage for three months. How extended a storage period can be realized by this technique will be determined. It is not known at present whether this procedure will have application in the commercial growing of the fiddlehead fern, However, it may be of enhanced interest SuHortTER Nores 89 should it be demonstrated that other fern spp., especially those difficult to store, will survive this process. LITERATURE CITED Baitey, L. H. 1915. Standard Cyclopedia of Horticulture, Vol. III. The Jon Boup, H. C. 1957. Mirphaloxs of Plants. Harper and Brothers. New York p CAMPBELL, D. H. 1887. II. The Development of the Ostrich ee Onoclea struthiopteris. Memoirs Boston Soc. Nat. Hist. I HESSELTINE, C. W. and F. B. Snyper. 1958. Attempts re eseaes dry Pine Pollen for Protnged heal Bull. Torrey iad Club. 85: 134-1 Menrora, B. S. and C. W. HessevtTiIne, 1958, Further Evaluation of he Lyophil Process eats th oun of oe and Penicillia, Appl. Microbiol. 6: -183. TULECKE, W. R. 1954. = servation and Germination of the Pollen of Ginkgo Under Sterile Conditions. Bull Torrey Bot. Club. 81: 509-512 RESEARCH STATION, CANADA DEPARTMENT OF AGRICULTURE, FREDERICTON, N. B., CANADA. Shorter Notes FERNS AND FERN ALLIES ON BONAVENTURE ISLAND, QUEBEC.— Bonaventure Island off the Gaspé Peninsula has long been a mecca for ornithologists, and it was to study sea birds that I visited that island from July 5-25, 1946. The richness of the flora and a spell of inclement weather were instrumental in my obtaining a reasonably complete collection of the pteridophytes of the island and a small collection of flowering plants, many of them not mentioned by Adams in his preliminary list of the plants of the island.1 The flowering plants and one set of the pteridophytes were deposited in the herbarium of the University of California at Berkeley and the remainder in the herbarium of the University of Michigan, Of the 29 pteridophyte species collected, only three (Pteridium aquilinum, Thelypteris phegopteris, and Equisetum Adams, J. 1940. A oo list of the plants of Bonaventure Is- tins fannie: Field Nat. 54: 90 AMERICAN FERN JOURNAL sylvaticum) were listed by Adams! and five others (Dryopteris filix-mas, Osmunda cinnamomea, Botrychium lunaria, B. matri- carnfolium, and Selaginella selaginoides) were reported from the island by Scoggan.? Because of the paucity of published records of pteridophytes from this island and the desirability of a guide to the habitats and relative abundance of these plants, I wish to put the results of this collection on record. I am en- debted to Dr. W. H. Wagner, Jr., for his encouragement and help with the manuscript. An annotated list of the species found follows. POLYPODIUM VIRGINIANUM L. Not uncommon; a thriving colony was found growing with Empetrum on a shaded boulder in mixed woodland. PTERIDIUM AQuILINUM (L.) Kuhn. Found sparingly in open meadows. CRYPTOGRAMMA STELLERI (Gmelin) Prantl. Fairly common on rock ledges. Some plants were found which suienod to be the hosts of a spe- cies of rust Sistas VIRIDE Hudson. A single colony of several plants of this species was found on a limestone ledge by a stream deep in eoniferous forest t GYMNOCARPIUM DRYOPTERIS (L.) Newm. Common under conifer THELYPTERIS PHEGOPTERIS (L.) Slosson. Abundant under coniers DRYOTERIS CAMPYLOPTERA Clarkson. Abundant in recent tly eut-over areas in the coniferous eater where it grows waist-high with stain disturbance indicators as Ribes and young birch. Its appearance in disturbed areas parallels that of Pianos puta in many areas to the south YOPTERIS INTERMEDIA (Willd.) Gray. Uncommon under comiecrt CYSTOPTERIS BULBIFERA (L.) Bernh. Not uncommon in moist parts of the coniferous forest and on shaded rock ledges. CYSTOPTERIS FRAGILIS (L.) Bernh. Fairly common in coniferous forest. DRYOPTERIS FILIX-MAs (L.) Seott. Uncommon in coniferous forest. A SapSecearhad large plant was found in the bottom of a small temporary tream near its head. POLYSTICHIUM BRAUNIT (Spenner) Fee. Uncomon in moist coniferous forest. MATTEUCCIA STRUTHIOPTERIS var. PENSYLVANICA (Willd.) Morton. Al- *Seoggan, H. J. 1950. The flora ae at Bie and the Gaspé Peninsula, Quebee. Nat. Mus. Canada Bull. 115, -399, SHORTER NOTES 91 though very common on the mainland, this species was only encountered on the island in a few wet places in thin coniferous forest OSMUNDA CINNAMOMEA L. var. CINNAMOMEA and OSMUNDA CLAYTONIANA L, both fairly common in wet openings in the forest. BorrRYcHIUM LUNA L., B. MATRICARHFOLIUM A. Braun, and B. SIMPLEX Hitcheock. ae scattered colonies of all three in open meadows, growing with moss, Iris, and Polygonum vivipar BOTRYCHIUM VIRGINIANUM L. Rare. A ¢ sida fo growing in partial shade in a small sak, in the coniferous for EQUISETUM ARVENSE L. Common in wet mea nt WSs EQUISETUM PRATENSE Ehrhart. One colony found growing with &. arvense in a damp meadoy EQuiseTumM Soave prea A colony found growing in tufts on stumps in eleared ground. EQUISETUM SYLVATICUM L. var. MULTIRAMOSUM oe Very common, replacing E. arvense in moist areas in coniferous fore LycopopIUM ANNOTINUM L. Uncommon; collected in man-made clearing in coniferous forest, Lycopopium cLavatum L. and L. tucipuLumM L, Uncommon in coniferous forest Lycopopium ogscuruM L. Rare. Two small plants were found in conif- erous forest. SELAGINELLA SELAGINOIDES (L.) Link. A good-sized eolony was found in an open bog, where the species grew on small stumps in company with Parnassia and Drose Rogert W. Sn Museum of Zoology, University of Michi- gan, Ann Arbor, Michigan. A Note on ELAPHOGLOSSUM cCRINITUM.—A number of years ago during one of my frequent botanizing trips to Cuba I Spent some time at the “rest house” of the Hermanos de la Salle high up on the peak of Loma del Gato. This verdant mountain, one of the tallest in Cuba, lies not far from Santiago. Because of the extensive botanical activities of the Brothers of the Cole- gio de la Salle, both in Santiago and in Havana, its flora is reasonably well known. Especial attention was paid, by several of the noted Cuban botanists, to its extraordinary fern flora, which ranges into the hundreds of species, and includes a re- markable number of endemies. 92 AMERICAN FERN JOURNAL The entire upper third of Loma del Gato—which means “Hill of the Cat,’—is covered by a dense forest. Constant very high humidity, coupled with frequent heavy rains throughout the year, create conditions which are ideal for the development of ferns. A variety of immense, graceful tree ferns is found here, virtually every tree-trunk is heavily clothed with dozens of dif- ferent kinds of epiphytic ferns, and the moist ground under the trees is generally a solid sheet of other ferns in almost over- whelming quantities and kinds. In sunny places where the in- digenous trees and shrubs have been disturbed, Gleichenia and Dicranopteris take over in almost impenetrable thickets. In such a pteridologist’s paradise, I felt uncomfortably aware of my failings and wished that I knew more about the intricacies of fern determination. But I did find, among others, one par- ticular species of fern which was known to me, since it is a famous one which has long been prized by connoisseur collectors in all parts of the world. This was the strange Elephant-Ear Fern, Elaphoglossum crinitum. Widely distributed in the West Indies, Mexico, and portions of Central Amrica, this remarkable fern is a common species in the summit forests of Loma del Gato, and has also been found on a few additional peaks in southeastern Cuba, such as Gran Piedra, Pico Turquino, ete. It has, in the past, been known as the type of a separate genus, Hymenodium, but Copeland and others consider this to be referable to the polymorphic group Elaphoglossum. The initial specimens of this Elephant-Ear Fern (its very heavy, russet-hairy fronds certainly do look somewhat like pachy- derms’ ears!) which I found were growing in a habitat which was characteristic of all those encountered. It was perched in regular array on fallen, partially rotting, moss-covered logs in the darkest shaded spots in the forest. Though I found occa- sional sporelings on the bases of mossy tree-trunks (even those of the lovely Palma justa, Euterpe globosa), no mature plants were found other than on the fallen and deteriorating logs. SHorter NOTES 93 Growing with the Elaphoglossum, in considerable profusion, were creeping colonies of the very delicate and delightful Rhipidopteris peltata, which has long been one of my personal favorites of all ferns, even though I cannot successfully main- tain it here in my Coconut Grove garden The hirsute covering of this Elaphoglossum varies from russet- brown to almost black, forms of the latter appearing rather as if they were dead, instead of living organisms. The fertile fronds are shorter-stalked than the sterile ones, smaller, and paddle- shaped; they bear a solid mass of sporangia on their undersides and are very distinctive, though seemingly rarely produced in the wild. According to expert fern-growers, this Elephant-Ear Fern does well in a friable rich compost under high humidity at all times, though over-watering should be avoided. I would assume, from having seen the plants in the wild, that enough water should be given them at the roots to keep them constantly moist, but the mossy logs on which they perch would also afford them adequate drainage even in nature. Though Elaphoglossum crinitum is a spectacular fern which has long been in cultivation, I am interested to find that in D. G. Huttleston’s recent valuable “Fern Sources in the United States” (Tuis JournaL 52: 97-109. July-Sept. 1962), it is not listed. If it is available in this country, I would be most anxious to hear of a source——ALEx D. Hawkes, Coconut Grove 33, Florida. OBSERVATIONS ON THE SENSITIVE FeRN: SuppLEMENT—Last year in my paper “Observations on the Sensitive Fern,” I asked the question, “What is the critical period during which destruc- tion of vegetative leaves will induce primordia of stig ag 7 change their course of development?” (THIs JOURNAL 53: 1963). A simple experimental test of plants in a ten foot square at Pilot Knob indicates that, at least in this case, twenty days is long enough. On June 16th the large vegetative leaves of the selected area 94 AMERICAN FERN JOURNAL were mowed off with a machete. Some small leaves, arising from tiny branches or young sporelings, were left untouched. Adjoin- ing this area there flourished more extensive stands of Onoclea growing under similar conditions, but left undisturbed. On July 6th, observations showed at least twenty specimens which could be classified as Onoclea sensibilis forma obtusilobata. Only normal types of sporophyls were observed in the adjoining, unmowed plot. Most of the intermediate leaves in the ex peri- mental plot were of the twice-pinnate type with which the name “obtusilobata’”’ is usually associated. A few were not two- pinnate ; their lateral divisions were narrowly linear and in this respect could be referred to as “lorinserioid.”—-RaLpu C. BENE- pict, Pilot Knob, New York. Growing Moruer Ferns.—I was interested in Mr. Morton’s recent article on the Mother Ferns (Asplenium bulbiferum and A. daucifolium). I was surprised to hear that the little plantlets of this fern are ever difficult to grow. I have a plant of A. bulbi- ferum which I bought in this country. Its older fronds are weighted down with very large number of plantlets with little fronds up to about six centimeters in length. These plants drop off, and they litter my greenhouse. If T remove them from the frond I find them very easy to grow, if they are not taken too young. As I am not much more than a beginner at growing ferns and have no special skill, I am sure that anyone could grow my plantlets. My fern has two types of fronds; some intermediate fertile fronds have narrow ultimate segments, mar- ginal sori, and bulbils on the upper surface. I am interested in Australian ferns, which I grow from spores, when kind friends send me any. I wish I could see some of your American ferns. You must have a bewildering variety.1—M. I. TerLey, Valley Howe, Cartmel, Grange-over-Sands, Lancashire, England. ‘Following the receipt of this communication, IT wrote to Miss Tetley asking if she would like to receive spores of American ferns, and she re- plied that she would be very happy to have them, especially spores of tem- perate ferns that would be likely to be hardy in Laneashire.—C.V.M. So) On RECENT FERN LITERATURE Recent Fern Literature Tue DEVELOPMENTAL ANATOMY OF IsoETES, by Dominick J. Paolillo, Jr., Illinois Biological Monographs, No. 31. University of Illinois Press, Urbana, 1963. $2.50 paper, $3.50 cloth. 130 pp., including 19 plates.—This recent book of interest to many pteridologists has come to my desk. It is a handsome book, well- printed and bound and of pleasing format. In addition to the general introductory sections there are three parts to the work headed “The Shoot,” “The Root-Producing Meristem,” and “The Apical Meristem of the Root.” Each section consists of a review of literature, observations, and discussion, plus a summary and conclusions. The literature reviews are quite detailed. The observations consist of careful descriptions of structure as seen in serial sections. Perhaps anatomical descriptions are by nature difficult to follow, but at times it does seem that the author could have found some way to make this less laborious. Considering the contents, the title of this book is misleading. It would be more aptly titled “Some aspects of the develop- mental anatomy of Isoetes,” since it deals with only the rela- tively mature sporophyte, and does not include the leaf. In fact, in this reviewer’s opinion, it would have been better published as a series of papers in a journal rather than as a book. Such a highly specialized and detailed book as this one sah’ ably will not be added to many private libraries. But it is contribution to pteridology and should certainly be part of any University library —Ricuarp L. HAUKE, Department of Botany, University of Rhode Island, Kingston, Rhode Island. Fora or Missourt,! by Dr. J. A. Steyermark, contains a com- plete account of the ferns (49 species, ineluding two introduced species and two admitted hybrids, in 24 genera) and fern- allies (13 species in four genera) of Missouri. The figures cited show 1Flora of aero by J. A. Steyermark, pp. i- — a 1725, pl. 1-390. 1963. Published by Ode State ‘University Press, / ee wa, $18.50. By a strange eeunehe the part of the printers, early co ae the book con- Pats no date of Shiba ion: According to Iowa State University Press the ate was Woweaities 12, 1963. 96 AMERICAN FERN JOURNAL that Missouri is not so rich in ferns as some states, but neverthe- less there are many interesting species. The treatment is modeled on Deam’s famous “Flora of Indiana,” and compares favorably with that monumental work. Like Deam, Steyermark has spent many years in the study of the flora of his state, and the results, in the keys and comments, indicate that the book is not a com- pilation but shows an intimate knowledge of the plants both in the field and in the herbarium. There are no descriptions, but the keys are unusually full and usable. Each species is accom- panied by a map showing its distribution by counties. An initial key to the species is followed by a key based wholly on sterile material, which should be useful. The nomenclature in general follows that in Gleason’s New Illustrated Flora. A few of the names are not in the form required or recommended by the Code of Nomenclature. According to Art. 73, note, the diaeresis should be used in the name Isoétes, genitives from personal names such as Engelmann should be corrected to engelmannii (even though originally spelled “engelmanni”) (Art. 73, Note 3), a terminal “y” is regarded as a vowel, and so Cystopteris fragilis var. mackayii should be corrected to “mackayi,” the double “i” being considered as an orthographic error, and finally Athyrium thelypteroides should be thelpyterioides, the original spelling, which cannot be considered an orthographic error since it is etymologically possible. The only misprint that I have noticed is an unfortunate one: On page 30 in the key to the species of Asplenium the species A. < ebenoides appears as “A. X asplenioides.” An observation of interest noted by Tryon in “Ferns and Fern Allies of Minnesota,” p. 138, is that Lycopo- dium selago var. patens has stomata on both surfaces of the leaves, whereas in L. lucidulum the stomata are on the lower surface only. These species and their varieties are perplexing and little known. Since Lycopodium possesses very few charac- ters, this one of the stomata may prove to be of importance. Of course, the major portion of the book is devoted to the flowering plants. Altogether, according to the summary on p. 1665, there - Recent FERN LITERATURE 97 are 799 genera and 2438 species treated, and in addition hun- dreds of subspecies, varieties, forms, and hybrids. Dr. Steyer- mark is certainly to be congratulated on a fine and scholarly piece of work, which is going to be permanently useful, not only to those working on Missouri plants but to systematic botanists in general.—C. V. Morron ARNOLD, CuEsTeR A. & Lyman E. Davauerty. 1963. The fern genus Acrostichum in the Eocene Clarno Formation of Oregon. Contr. Mus. Paleont. Univ. Mich. 18: 205-227 (6 pls.) —A brief resume of the ecological requirements of ferns in this genus sets the background for speculation about the conditions under which the fossil materials were deposited and preserved. The descrip- tions of the petrifactions, which occur in chert, are excellent and the photomicrographs of the thin sections are superb. The paper is technical, but will be of interest to many of the Fern Society’s members. Fapsri, F. 1963. Primo supplemento alle Tavole Chromo- somische Della Pteridophyta de Alberto Chiarugi. Caryologia 16: 237-335.—Continuing the work done by Chiarugi. The dis- cussion in this paper is in English. Heviy, Richarp H. 1963. Adaptations of cheilanthoid ferns to desert environments. Jour. Arizona Acad. Sci. 2: 164-175.— Contains some interesting and instructive ecological information. Nayar, B. K. 1963. Contributions to the morphology of Lep- tochilus and Paraleptochilus. Amer. Jour. Bot. 50: 301—308.— Technical description of morphology and anatomy. Illustrated with line drawings and half tones. Nayar, B. K. & Farrun Kazi. 1962. Ferns of India. IV. Plagiogyria. V. Hemionitis. VI. Cheilanthes. VII. Actiniopteris. Bull. Nat. Bot. Gard. (Lucknow) No. 64, 67, 68, and 75, respectively. — Each part is bound as a small booklet that contains long descriptions, considerable morphology, keys to the Species, and with halftone reproductions of varying quality Showing different parts of the plants. Line drawings used to illustrate the morphology and anatomy are good. There is an 98 AMERICAN FERN JOURNAL index to the species and references to pertinent literature. ScamMan, Epirn. 1962. The genus Eriosorus in Costa Rica. Contr. Gray Herb. No. 191: 81-89.—A key to the five species, brief general descriptions, statement of range, synonymy, and citation of species are included. Line drawings are very good. Tryon, Auicre. 1962. A monograph of the fern genus Jame- sonia. Contr. Gray Herb. No. 191: 109-197, plus index.—In- cludes a key to the species, distribution maps, halftones and line drawings to illustrate various critical features. This is an indis- pensable book for anyone dealing with the ferns of the Andes, the mountains of Central America and southern Mexico. One who has tried to determine a species of Jamesonia without it can fully appreciate the service Mrs. Tryon has rendered all pteridologists ! Tryon, Rouua. 1962. Taxonomic Fern Notes. III. Contr. Gray Herb. No. 191: 91-107. Five short notes on systematics of vari- ous genera. Includes new a generic name, Blotiella, based on Lonchitis Kiimmerle (not Linn.), and seven new combinations in Blotiella. It includes also one new species, Dorypteris allenae; a new combination in Alsophila, some clarification of the genus Saccoloma Kaulf., and a note about the type species described by Kaulfuss, including a reproduction of Kaulfuss’ signature. WESSELS Borr, J. G. 1962. The New World species of Tricho- manes Sect. Didymoglossum and Microgonium. Acta Bot. Neer- landica 11: 277-330.—A technical monograph carefully done. SuaTer, JAMES R. 1964. Fern distribution in Washington State. Occ. Papers dept. Biol. Univ. Puget Sound No. 27: 242- 257 + 2 unnumbered, double page tables.—Fifty-two taxa based on specimens examined, with county occurrences given under each, are recorded, and county records for 23 added taxa taken from the literature are included in the tables only. Small type and extensive use of abbreviations make use of the booklet slow at first. It will be of considerable value to visitors wishing to find a particular fern within the state of Washington. NoTEes AND NEws 99 Notes and News THE British Prermo occa Socrery.—The objects of the Society are to study and encourage interest in the Ferns and Fern Allies. The British Fern Gazette, published annually by the Society, contains papers on the horticulture and taxonomy of ferns from all parts of the globe. Membership, including a subscription to the Gazette, is $2.00 per year. Further particu- lars may be obtained from the Hon. Secretary-Treasurer, J. W. Dyce, Esq., 46 Sedley Rise, Loughton, Essex, Great Britain.— Rotia Tryon, Gray Herbarium, Harvard University. DeatH Takes Notep Prermoiogist.—On March 16, 1964, Professor Edwin Bingham Copeland, prominent American au- thority on ferns, a world leader in the study of their classifica- tion, and an Honorary Member of the American Fern Society, died in Chico, California, a few months past his ninetieth birth- day. All who knew him will remember his dry wit and the caustic turn of his comments when he encountered careless work. We will miss him from among the workers on ferns and as a stimulating personality. Proressor R, C. CHine AND THE STupy oF FERNS IN CHINA.— During two weeks in October-November 1963 I was privileged to visit China as a guest of Academia Sinica. At Peking I met Prot Ro Ching, and found him in good health and still very actively concerned with fern taxonomy. He is 65, but still able to climb mountains and enjoy field work. Though there has been a delay in the intended program of publication of the new Flora of the People’s Republic of China (of which the first volume is- Sued, in 1959, contained the first part of Prof. Ching’s account of the ferns) two new volumes have recently been issued, and Prof. Ching told me he expected his second volume to appear in 1964, with two or three more volumes in later years. His pre- liminary studies of the genera allied to Thelypteris and to Athy- rium have recently appeared in Acta Phytotaxonomica Sinica (vol. 8: 289-335, 1963; vol. 9: 41-84, 1964). Prof. Ching is head of the division of Taxonomy and Plant 100 AMERICAN FERN JOURNAL Geography of the Botanical Institute of Academia Sinica. The herbarium, with a very good and up-to-date library, is housed in the main building of the Institute (formerly the Botanical Institute of the National Academy of Peking), with other divi- sions temporarily installed in neighboring smaller buildings. At Canton I visited the Botanical Institute of Academia Sinica for South China, of which Prof. W. Y. Chun is head; in this Institute is the herbarium started by Prof. Chun in 1929 as part of Sunyatsen University. Near the Institute (and part of it) is a Botanic Garden of 100 acres, founded in 1958 and still in an active state of development. The garden has a recently established collection of about 100 species of native fern plants; Mr. C. H. Wang, who assisted Prof. Ching in the preparation of the first volume of his fern flora, is actively interested in this collection. Mr. Wang also went with me to a National Park 90 miles from Canton. This National Park includes 500 acres of natural forest, in which are many species of ferns, and also an area of pine plantation with grass-covered hillsides beyond it, on which reforestation is planned. I took some young fern plants from the forest (among them Cyathea podophylla, abundant in shady gullies) and Brainea insignis from the pine plantation. These plants have now started growth in the tropical fern house at Kew. Many trees in the National Park, near the paths, carry labels painted with their botanical names in roman letters. I was impressed by the competence and the enthusiasm of the staff of the botanie garden and the national park—R. E. Hourrum, Royal Botanic Gardens, Kew. A Book Barcatn.—The remaining copies of the late Profes- sor Jesse M. Shaver’s fine book “The Ferns of Tennessee” are being offered for sale at the bargain rate of $5.00 each by his widow, Mrs, Daisy Shaver, 1706 Linden Avenue, Nashville, Tenn. AMERICAN FERN Soctrety 101 American Fern Society CoLoRADO FERN Foray AND ANNUAL MEETING Two full-day field trips are saa led for gaits of the American ] Fern Society. On Friday, August 21, Dr. William A. Weber will lead the foray from Boulder to Green Mt., ae Mt., “ee e Rocks and the Sates ” On Saturday, August 22, Mrs. Ruth Ashton "Neloon will lead e foray from Boulder to Rocky Maanists National Park. It will be pos- ait to observe many of the ferns of Colorado on these two trips. Partici- pants will not be permitted to collect botanical specimen mbers who register with AIBS and plan to alten ‘the foray will be housed in Kittredge Hall near the southeast corner of the campus of the University of Colorado (south of the Observatory). Members who do not plan to attend the meetings or do not expect to register with the American Institute of Biological Sciences are weleome on the foray, but they must arrange for their own housing. Those who do plan to register with the AIBS wil be sent forms for doing so if they will request them from Dr. Hauke. There will be two sessions for presentation of papers, one in the morning and one in the afternoon on en August 24. The annual meeting and luncheon will be held at noon the sa e day, at the University of Colorado. you plan to attend the foray aes eos: please fill out the fol- lowing and return as soon as possible to Dr. Richard L. Hawke, Department of Botany, University of Rhode Island, scicupisn, Rhode Island. Ido (_ ) or do not (_ ) plan to attend the luncheon. IT plan to attend the foray, August 21-22. 00 There will be ___. persons in my party. I need transportation for persons. I can provide transportation for ——......... persons. ; plan ( ) or do not plan ( ) to register with the AIBS and will () or will not ( ) need forms for doing so PuaLAKes pavkau Kiendinels Address a AMERICAN FERN Society Preregistration and Housing Application AIBS-Sponsored Meetings of Biological Societies University of Colorado, August 23-28, 1964 Please type or print 103 PHENO ie ei ohh a 2 CL Prot ike Mrs. Home address _..... (Street) (City) (State) Prof. or business address Roommate Request: “hs 1 Lina nic ea gee errs Are you a direct member of AIBS? (Yes or No) Are you a graduate student? 20 (Yes or No) Society of Primary interest. (Must accompany this form) Name(s) & Relationship I will be accompanied by: (Give age of children) (Continued on next page) Travel arrangements: Expected time of Arrival By plane at Denver (Date & hour: Flight No.) By train at Denver 2. Departure from Boulder Byuvar at Bowider: 2.03 ore (Date & hour) Housing: I have arranged my own housing at I request University housing assignment, American Plan — Modified American Plan _. Buropean Plan 2.00. Premeeting housing __...____ Special requirements — (i.e., Health, ete.) Note: (1) Please preregister prior to August 1, 1964. (2) Registration Fee: $10.00 for AIBS members, $15.00 for non- members, $3.00 for students. $2.00 extra after August 1. (3) Make check payable to “American Institute of Biological Sciences.” (4) Mail this form and check to: AIBS Registration, Room 508 2000 P Street, NW., Washington, D. C. 20036 Registration fee check in amount of $2). is enclosed. isle alin aah ieee ae Da re aE EY (feat eres Rae eral Suaa@EsTIONs TO CONTRIBUTORS TO THE AMERICAN FERN JOURNAL 1, Authors of papers submitted to the American Fern Journal should con- sult the Style Manual for Biological Journals (1960) and be guided by its examples, 2. The “name-and-year” system for bibliographic references will, ex except in exceptional cases, be used. (See Literature Cited below for an example). 3. Authors are encouraged to use the journal abbreviations set forth by Schwarten and Rickett (1958, 1961). 4. Abbreviations of names of herbaria will be those listed by Lanjouw and Staflen (1959). 5. Reports of chromosome numbers will not be accepted unless documented. 6. The use of footnotes and tabular matter should be kept to a minimum. 7. All manuscripts segnansd for publication should be typed, double- spaced, and have ample ma 8. Reprints should be ordered when galley proof is returned to the Editor. An order blank will be included with the galley proof. LITERATURE CITED American Institute of Biological Sciences, Committee on Form and Style of the Seseagered oe Biological Editors. 1960. Style Manual for Biologi- cal Journals, A can Institute of Biological Sciences, Washington, D. C. 100 p. Langouw, J., and FP. A. Starzzv. 1959. The Herbaria of the World. Index Herbariorum, Regnum Veg. 15: 1-2. Scuwarren, L., and H. W. Ricker. 1958, Abbreviations of Titles of Serials Cited by Botanists, Bull. Torrey Club 85: 277-300; 1961. Supplement I. Ibid. 88: 1-10, 4 Vou. 54 JULY-SEPTEMBER, 1964 No. 3 American Fern Journal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY ts IRA L. WIGGINS 0. V. MORTON ROLLA M. TRYON JOHN H. THOMAS CONTENTS A Preliminary Chromatographic Study of Eastern America Dryopteris _____ Ratner W. Scoza AND WaRREN H. al JR. 105 Ferns oe with Ultramafic Rocks in a: Pacific Northwes TauR R. KRUCKEBERG 113 Rediscovery — a virginianum forma mend ag (Ridlon LD A. Cnaxeren 126 OGRA in Pteris cretica with a ase reer to the Cytoplasmic Inclusions _____-___ NorMAn P. Mareneo 129 Florida Strap Ferns and Their Culture ____AtEx D. Hawxzs 136 An Amerian Species of Stegnogramma ____ Kun10 IwaTsuxt 141 Some New Combinations for Southeastern Ferns__Epgar T. WHERRY 143 Camptosorus rhizophyllus forma boycei C. L. ete teh A. CHARETTE 147 Shorter Notes: Asplenium ebenoides R. R. Scott in Kentucky. 149 Recent Fern Literature 150 Notes and News . 152 American Fern Society — 156 Missour; BoTanicar NOV 30 1964 Che American Fern Society Council for 1964 OFFICERS FOR THE YEAR Donovan 8. Corre, Texas Research Foundation, Renner, Texas eles: Donatp G. Hurtizeston, Longwood Gardens, Kennett Square, chapel ce-Pr bape LENETTE R. ATKINSON, 415 S. Pleasant Street, Amherst, Siassamuaad Secretary Ricnarp L, Havxe, Department of Botany, University of Rhode Island, Kingston, Rhode Island reasurer Ina L. Wiaams, Dudley Herbarium, Stanford University, Stanford, Calif. Editor-in-Chief OFFICIAL ORGAN American Fern Journal EDITORS Ika L, Wicamns___Dudley Herbarium, Stanford a went Stanford, Calif C. V. Morton______"_ Smithsonian Institution, Washington 25, D. 0. Botta M. Trron Gray Herbarium, Harvard University, Cambridge 38, Mass. Joun H. Tuomas Dudley Herbarium, Stanford University, Stanfo rd, Calif. ay illustrated quarterly devoted to the een study of ferns, owned by erican Fern Society, and published at 3110 Elm Ave., Baltimore 11, Ma. Second class postage ald a Balt teas Ma. er for publication should be addressed to Ira L. Wiggins, Dudley Herbarium, Sta nford University ; Stanford, Califor rnia. authors ts cost, plus postage. Back volumes $3.00 aie single back numbers 75 cents each; Cumulative apees to vols. 1. 25, 25 cents, Ten per cent discount on orders of six volumes more. P yao age Botany applications for membership, subscriptions, orders an business communications should be addressed . ard L. Hauke, D t: , Universi of Rhode —— Psp aveat T Me ost ne Sy of Botany, University American Fern Journal Vou. 54 JULY-SEPTEMBER, 1964 No. 3 A Preliminary Chromatographic Study of Eastern American Dryopteris Rainer W. Scora AND WARREN H. WaGner, JR.! The use of chemical methods—especially serology, electro- phoresis, and chromatography—is fairly new in plant taxonomy. In fact, our knowledge of chemotaxonomy is still so far in its embryonic stages that probably the significance of chemical data is not at all understood. The present investigation was made to determine what chromatographic results could be obtained from studying eastern American woodferns, Dryopteris. These ferns are among the best known in the United States cytogenetically, thanks to the work of Walker (1955, 1959, 1961, 1962) and are accordingly an ideal group to examine, especially because nu- merous hybrids are known. The work focused not only upon the Species, but the interspecific hybrids as well. The chromatographic method makes it possible to take ex- tracts from different kinds of plants and to compare the individ- ual compounds present. It has been found that species, varieties, and hybrids of plants and animals often show strikingly differ- ent chromatographic patterns (Alston and Turner, 1963). The application of chromatography to flowering plants has been fairly widespread during the past few years, but such study of ferns has been slight. Recently Smith and Levin (1963) investigated the Appalachian Aspleniums chromatographically and found striking evidence in support of a theory of reticulate nee Bias. vish to thank Dale J. Hagenah for supplying plants, Ara Paul fo of Mirtibetoe uate equipment, and Katherine Lim Chen, for technical sects The research was made possible in cast by NSF grant G-10846. LES se RI ee AO A Volume 54, No. 2, of the JouRNAL, pp. 57-104, was issued June 25, 1964. 106 AMERICAN FERN JOURNAL evolution for these plants. By the use of chromatography Fiken- scher and Gibson (1962) compared phloroglucinol derivatives among the woodferns Dryopteris intermedia, D. spinulosa, D. in- termedia X spinulosa, D. clintoniana, D. clintoniana X< goldi- ana, D. arguta, D. goldiana, and D. marginalis. They concluded that the hybrids showed additive phenomena of the compounds present in the parent species. This kind of result did not prove to be the case in the present study, as will be described below. MATERIALS AND METHODS Plants of various species and hybrids were grown under es- sentially uniform conditions in Birmingham, Michigan, by Hagenah of the Cranbrook Institute of Science. On July 30, 1963, comparable leaves, all with sori present, were collected from each plant. Vouchers for each plant are deposited in the University of Michigan Herbarium. Leaf material of D. goldiana was taken from a natural habitat near Ann Arbor on July 31 and that of D. clintoniana * spinulosa from the University of Michigan Botanical Gardens on the same date; all others were provided from the Hagenah garden. Because of their special interest, the D. spinulosa and the D. cristata complexes were studied in two dimensions. (These and other hybrids were ex- amined in one dimensional chromatograms also but will not be reported here.) Mature, healthy fronds were cut finely and extracted for 30 hours in 1 pereent methanolie IN HCl. These extracts were con- densed and 60 d spotted on 46 X 57 em. sheets of Whatman no. 1 filter paper. The two-dimensional descending method was em- ployed, using N-butanol: acetic acid: water (at volume/volume ratio of 4:1:2) for the first separation of 20 hours and 1 percent HCl for the second dimension of four hours, both at 21°-+1°C. The solvents were mixed six hours prior to use. Equilibration of the chromatographic chamber lasted five hours in both eases. All equipment and solvents were stored two days prior to use at the same temperature. The chromatograms were dried and CHROMATOGRAPHIC StupDy OF DRYOPTERIS 107 inspected for position of the solvent fronts and position and intensity of compounds under short and long wave ultra-violet light. Of several chromogenic sprays tested, 2N KOH gave the best results and was used throughout. The chromatographic patterns were copied before and after spraying with differently colored wax pencils on sheets of clear acetate (Seora, 1964). These acetate sheets were then superimposed to show pattern deviations. RESULTS The smallest number of spots in the chromatograms was found in Dryopteris campyloptera < intermedia, and the largest num- ber of chemical substances in D. dilatata < intermedia and in D. clintoniana * goldiana—a range from 6 to 16. The effects of hybridization were striking, and all of the fol- lowing situations were observed in our chromatograms: 1. Substance present in both parents and present in hybrid. 2. Substance present in both parents and absent in hybrid. 3. Substance present in one parent and present in hybrid. 4. Substance present in one parent and absent in hybrid. 5. Substance absent in both parents and present in hybrid. The chromatographic entities that illustrate the second and fifth of these situations are marked by the corresponding num- bers on the diagrams (Figs. 1-3). The most striking chromatogram obtained is the one involving the hybrid of Dryopteris dilatata and D. intermedia (fig. 1). This hybrid, first reported by Wagner and Hagenah (1962), was found in the Huron Mountains of Michigan, and subsequently grown from an offset in company with the parents. One-dimen- sional chromatograms only were made of D. dilatata * margi- nalis and D. intermedia X marginalis, but these were different in nature from similar one-dimensional chromatograms of D. dilatata < intermedia. Dryopteris dilatata X intermedia shows all of the five situations listed above. The most conspicuous result of this study is that wherever 4 108 AMERICAN FERN JOURNAL taxa of different polyploid levels are involved, the hybrid chro- matograms usually tend to have more substances in common with the parent of higher ploidal level than with the one of lower level. This is shown by Dryopteris campyloptera (4X) X intermedia (2X); D. intermedia (2X) X spinulosa (4X)—(fig. 2); and D. cristata (4X) & intermedia (2X )—(fig. 2). D. INT. ote) = 2.01. BOS S oe ro) “6 AO ro) © O | | or AB oS a O.DIL X INT D. CAMP. a 56 2 ae a Bae : eth 20% is OO S oe D.CAMP X INT D. SPIN. © = a are O US a2) Cx aa of CO fil oa ee CHROMATOGRAPHIC Stupy OF DRYOPTERIS 109 The one exception to this pattern is an unusual hybrid in- involving Dryopteris clintoniana, a hexaploid, and D. spinulosa, a tetraploid (fig. 3). In this plant the hybrid is unusual in show- ing more resemblance to the lower polyploid than to the higher. However, as has been found by Walker (1955, 1962) at least half of the genetic influence of D. spinulosa is probably already present as one of the three genomes of D. clintoniana (Walker, 1955, 1962). Thus the composition of D. clintoniana < spinulosa cannot be represented by the formula ABC plus DE, but rather ABC plus CD. Perhaps this genome homology between D. spinu- losa and D. clintoniana is responsible for this turnabout in the pattern previously indicated (namely that the higher polyploid tends to have more influence than the lower). DIscussION It is clear from the results obtained in this study of members of the Dryopteris spinulosa and D. cristata groups that (1) fairly clear-cut differences exist between the species—as has been suggested by the previous work of Fikenscher and Gibson ; and (2) interspecific hybrids by no means show clear-cut blend- ing of parental patterns. The hybrid_ patterns are neither pre- cisely intermediate, nor are they shown to be additive by the methods we used. If such a study as the present one had been made on only two parental species and a suspected hybrid, the worker might have concluded that the suspected hybrid was not a hybrid on the basis of the chromatic patterns even though it ne Figure 1. Two DIMENSIONAL CHROMATOGRAMS OF Dryopteris intermedia, dD, dilatata, D. dilatata < intermedia, D. campyloptera, D. campyloptera intermedia anp D. spinulosa, AS SEEN IN LONG WAVE U/V LIGHT AFTER K 3 i DENOTES “HYBRID CHARACTERISTIC SUBSTANCE” ABSENT IN BOTH PARENTS. HORIZONTAL ARROW INDICATES DIRECTION OF FIRST SEPARATION WITH N- BUTANOL: ACETIC ACID: WATER. eae ARROW INDICATES DIRECTION OF SECOND SEPARATION INVOLVING 1% HC 110 AMERICAN FERN JOURNAL D- INT X SPIN D-CRIST O @? OOo ® @) Ta Ps O Ce atone D.CRIST X INT 0.GOLD. 28 oe Q i, O O # O ©) . Upper Oreina pein S (dunite) POmol Steep talus of huge boulders Twin Sisters Mts., 3800 CHsi with sparse conifer-for) ADpal cover. . 14. rites Mine area, 8S. § (dunite CHsi Steep bouldery slope; seat: — ca pomsaek« Blver, ~ conglom.)-PTaq tered stunted DF & LPP; — ground layer either Rhaco- 4 15 and eriecaceous shrubs). E pas Eagle Lakes, S$ (per.) CHsi Steep rock ledges and faces — 3800’ (S-NS contact at POmol forbs and grasses in rock — V-shaped — steep aw ADpal ee above lakes) 16. (Same as 15) NS CRae ics yee eee saa walls; Alaska cedar, eT eads, forbs and grasses. +7, Southeast tip of Cypress ae S (dunite) CHsi Open balds and stony slopes FERNS on UttTrAMAFIcC Rocks 121. 28, Grasshopper Mountain, upper Tulameen River NS (shale) WOor Tsl., 500’ with seattered DF, LPP, and JUse cae: co oe grasses 18. Fidalgo Head, w. end S (per.) CHsi Beco: “opin pag eS a of Fidalgo Isl., 50’ heavy moss - lichen - grass - forb cover; occas. stunted DF. 19. W. slope of Sumas Mtn.,_ 5S (serp.) CHsi Open promontory on wooded 1000’ w. slope, dwarfed DF LPP, yew, and grass-forb cover on ledges and slopes. Grant and Josephine counties, Oregon 20. Buck Cabin Creek,Grant SS (serp.) CHsi Open steep stony outcrop Co surrounded by DF-PP for- sparse grass - forb cover. 21. Murderer’s Creek, Grant S(serp.) CHsi Open stony slope with scat- C tered DF, ; oc, an grass-forb co 22. Baldy Mtn., Grant Co. S$ (per.) CHsi Massive rn “ talus 7634! -facing slope of sum- mit; rich tgotgane cover 23. Strawberry Lake, Grant NS (vol- None! ck outcrops on shore Co. (ea. 5 mi. e. of #22) canics) of lake; herb layer luxu- riant but poor in spp 24. Road to Galice, Jose- § (serp.) CHsi Raw serp. cliffs above Rogue phine Co. River. Trinity county, California 25. Deer Lake, Trinity Alps SS (serp.) POmol Open rocky slopes with scattered MH and WWP British Columbia °6. Christina Lake S (serp.) CHsi Steep cliffs and talus bor- WOor dered by DF forest 27. Near Eholt S (perid.) CHsi Loeal barren outcrop bor- dered by DF-LPP-larch orest Outerop and talus’ with sparse shrub an orb- grass cover bo ni ot ary The high constancy and fidelity of Cheilanthes siliquosa and Polystichum mohrioides var. lemmonii for ultramafics in the Pacific Northwest suggests a close conformity of plant to sub- strate (Fig. 3). Only rarely does C. siliquosa oceur on non-ultra- mafic outcrops, and I have yet to find P. mohrioides var. lem- monv on other substrates than ultramafics. On the other hand, the rather characteristic ferns of nearby non-ultramafic rock out- crops — Cryptogramma acrostichoides, Cheilanthes gracillima, Polypodium vulgare var. hesperium, and Woodsia scopulina rare- ly, if at all, grow on soil of ultramafic origin. Of the two species commonly found on ultramafies, C. siliquosa is the most frequent, and through a wide altitudinal range. It is at sea level in the San Juan Islands and on up to 4000 feet in the Wenatchee Moun- tains and even higher in Oregon and northern California. P. mohrioides var. lemmonii, however, does not occur below 3000 feet in the areas I have visited. We may invoke an explanation to account for narrow restriction of these ferns to ultramafies that has been exploited in connection with angiospermous ser- pentine endemies (Kruckeberg, 1951, 1954, and Walker, 1954). Survival on soils high in ferromagnesian minerals but deficient in calcium requires a physiological eapability for efficient with- drawal of what little calcium is present and as well to accumulate other essential elements in low supply; failure to expand their range onto adjacent non-ferromagnesian soils may be due to the increased biotic (microbial and higher plant) competition en- 122 AMERICAN FERN JOURNAL Same area as 28 S (perid.) CHsi Outerop and talus with JUco and sparse grass- forb cover | het Mountain, upper S (perid.) CHsi Steep talus with grass-forb Tulameen Riv se and shrub co Pepa tis pci above S (dunite) CHsi Barren local outcrop sur Bralorne se ounded by dense brush CRae (rare!) and ( . B.C. Nickel Minesabove S$ (pyrox- CHsi Open treeless talus bordered — Choate enite) ADpal by MH and Alaska cedar — CRae FERNS ON ULTRAMAFIC Rocks 123 countered on more fertile soils. It should be possible to test the latter hypothesis in spore germination tests on the two soil types and in the presence of competition. Sporelings of Cheilanthes siliquosa frequently occur spontaneously on serpentine soils that I have used in the greenhouse for testing edaphic responses. Dispersal and establishment of ferns with such a disjunct dis- tribution and fastidious preference for substrate present a host of attendant problems. One is led to assume that spores of these “serpentinophytes” are widely dispersed or at least in a region- ally broad “chain-mail’” fashion, but only establish populations following germination on soils of ultramafic origin. The distribu- tion of C. siliquosa spans the North American continent. The easternmost point in its distribution—Mount Albert on the Gaspé Peninsula—is a world-famous alpine serpentine area. In the known localities intervening between Quebec and the Pacific Coast states, I cannot find accounts of the substrate. The same Species is common on serpentines of the Coast Ranges in Califor- nia (personal observation), but apparently it is not restricted to ultramafics. It has been collected on granite in the Sierra Nevada and from other areas unlikely to have ultramafic substrates. Polystichum mohrioides, as represented by variety lemmonii in the Pacific Northwest, appears to be exclusively on serpentine. Therefore I am suspicious of the granitic habitat ascribed to it by Maxon in Abrams (1923) and repeated by Munz (1959). The type of var. lemmonii—“near Mount Shasta’”—could easily be on ultramafic rock; serpentine is common in the lithology of north- western California. I can find no mention of substrate preference for the subantaretic and western South American congener, P. mohrioides,? though its unique bihemispheric distribution is fre- quently mentioned (Gams in Verdoorn, 1938, Christ, 1910). Ferns which are characteristic on ultramafic rocks elsewhere 2 Sr. José Diem, of Villa la pasate Neuquén, Argentina has wieder me that “This fern prefers open or semiopen sites at the base o n fis- Sures in granitic rocks and other Saran aan] but also is found in Y eer open woods pe at the edges of arroyos where it has developed other varie- ties and form 124 AMERICAN FERN JOURNAL in the world have intrigued botanists repeatedly through the years. The degree to which fern species are restricted to ser- pentine varies widely. Some are apparently true endemics at the species level, others are morphological and ecological variants of species possessing broader tolerance. Then some occurrences on serpentine are merely unusual range extensions wherein the fer- romagnesian substrate somehow extends the distribution of a spe- cies. Still other species are apparently indifferent to changes in substrate; these have been called serpentine-wandering ubiquists (“serpentinvagen ubiquisten,” Krause, 1958). I have compiled in the following table a list of those ferns which are known to inhabit soils of ultramafic origin. The list in- eludes species which show varying degrees of edaphic restriction, from endemies to ubiquists. This compilation undoubtedly will be incomplete; reports of additional instances of ferns inhabiting ultramafies will be welcomed by the author. OCCURRENCES OF FERNS ON ULTRAMAFIC RocKS ELSEWHERE IN THE WORLD Asplenium adiantum-nigrum L. Sweden (Rune, 1957), Balkans (inel. var. cuneifoliwm) (Krause & Ludwig, 1956), Italy (Messeri 1936, Pichi-Sermolli 1948 A, adulterinum Milde. Sweden (Rune 1957) A, onopteris L. var. davallioides Italy (Messeri 1936) Heufl. A, ruta-muraria L. var. brunsfelsii Italy (Pichi-Sermolli 1948) Heufl. A. trichomanes L, Japan (Kitamura 1950), Italy ere 1936, Pichi-Sermolli 948 A. viride Huds. nade (Rune 1953), but calei- cole in Quebee (Scoggan 1950), Finland aaciad 1956) Adiantum pedatum L, var. aleuticum Quebee (Seoggan 1950) Rupr. Cheilanthes siliquosa Maxon Page (Seoggan 1950) Ceterach officinarum Lam. & DC, Italy (Messeri 1936, Pichi-Ser- ’ donk 1948) FERNS oN UutTRAMAFIC Rocks 125 et te dichotoma (Thunb. ) Japan (Kitamura & Momotani Ber (=Gleichenia linearis 1952—probably of low fidelity) co Hints var Notholaena maran 2. Clay Italy (Pichi-Sermolli 1948) a See es aa Bs ) ne Quebec (Scoggan 1950, Rune 1953) Max Po than vulgare L, Italy (Messeri 1936) Gymnocarpium robertianum (Hoffm.) Japan (Yamanaka 1952) e Cryptogramma crispa R. Br. var. ja- Japan (Kitamura 1952) ponica Miyabe & Kuc Pteridium aquilinum eae. var. latius- culum (Des.V.) Unde Pennsylvania (Wherry 1932) LITERATURE CITED Curist, H. 1910. Die enw der Farne, G. Fischer Verlag. Gams, H. 1939. Okologie der istiist Ley Pteridophyten. (in “Manual . Nij of ga abet A ae i Fry) pineal 1950. ear Sekinomiya, Prov. Tazima. 6. pentine flora of a Hayachine, Prov. Rikuchu, Japan. Acta Phytotax, “Get 14: 177-180. Kiramura, 8S. A . Momor ar 1952. bse flora of aso Suga- shima, piov eh Japan. Acta Phytotax. Geobot. 14: 118- Krause, W. 1958. Andere Sa Fae ecinpeiih of eat ota ology 4: 755-806. Krause, W. an UDWIG. auf Sid Wendie des Balkans. 1. Halac: Dorfl. Ber. ener) Bot. Gesell. 69: 417-428. Ae Re 1951. iy 1956. Zur Kenntnis der Flora and Vegetation acsya sendtneri (Boiss. ) KRucKEBERG, A, 1. Intraspecific variability in the response of cer- ain native plant species to serpentine soil. Amer. Jour. Bot. 38: 408-419 1954. Plant species in relation to serpentine soils. Ecology 35 267-274. Lounamaa, J. 1956. Trace elements in plants growing wild on different rocks in Finland. Ann. Bot. Soc. ‘Vanamo’ 29: 1-1 Maxon, . R., in Abrams, L, 1923. Illustrated Flora of the Pacifie States. ol. k Gi diaviusevcea to Aristolochiaceae. Stanford University Press. ' Messer, A. 1936. La vegetazione delle roccie ofiolitiche me Monte Ferrato (Presso Prato.) Nuovo Giorn. Bot. Italiano, n.s. 43: 277-372 scopulinum is oceasionally found on or adjacent to ultramafies in the jo Northwest (Wagner & Kruckeberg, personal observation ). 126 AMERICAN FERN JOURNAL Muwnz, P. A. 1959. A California Flora. University of California Press PIcHI-SERMOLLI, R. 1948. ora e vegetazione delle serpentine e delle altre ofioliti dell’alta valle del Tevere (Toscana). Webbia 6: 80. Pratt, R. M. 1958. The geology of the Mount Stuart area, Washington. le Rung, O. . Plant life on serpentines and related rocks in the north of Sweden. Acta Phytogeographica Suecica 31: 1-139. ie areca bein ¢ 1957. De serpentinicola elementen i Fennoskandiens flora. Seouk: Bot. pion 51: 43-105. Scoaean, H. J. 1950, The flora of Bie and the Gaspé Peninsula, Quebec. Nat. Mus. of Pre Bulletin No. 115 (Biological Series No. 39) 1-399. WALKER, R. B 4, isin: affecting plant growth on serpentine soils. WuHerry, E. T. 1932 _ Belg studies of serpentine-barren plants—I. Ash composition, Pe . Acad Sci. 6: 32-38, WHuittTakEr, R. H. eas he ed of serpentine soils: a symposium. I. Introduction. Ecology 35:258-259, Sor oe, span response to serpentine soils, Ecology ao: 275-288: YAMANAKA, T. 1952. Studies in the vegetation and the flora on serpentine. es. Rep. Kochi Univ. 1: 1-8. DEPARTMENT OF Botany, UNIVERSITY OF WASHINGTON, SEAT- TLE, WASHINGTON, Rediscovery of Polypodium virginianum forma brachypteron (Ridlon) Fernald Leopotp A. CHARETTE In 1921 H. C. Ridlon, of Bennington, Vermont, described Polypodium vulgare 1. f. rotundatum, a peculiar, probably ab- normal form, in which the segments of the blade are reduced to semicircular or deltoid lobes, toothed at the apex. It was soon found that the name f. rotundatum had already been used, so it was changed to f. brachypteron (Weatherby, 1921). Ridlon did not give the origin of the specimen upon which the form was based, except to say that it came from Vermont. He neglected to record if a type specimen had been preserved. This has been a very rare form and nothing quite approaching it seemed to have been collected since its description until two POLYPODIUM VIRGINIANUM FORMA BRACHYPTERON 127 fern enthusiasts collected it in Colchester, Vermont, on Novem- ber 1, 1961, and on May 30, 1962. Their specimens came from a colony of 16 plants. Dole (1937) gave the only definite data as to the type station. He wrote, “Rocks, Jamaica—type sta. 1919 (Ridlon).” It is sur- mised that Dole’s information was based on his personal knowl- edge, of which he left no further written record. Other workers on our ferns, such as Weatherby (1921) and Fernald (1922) made no mention of the type station or of the type specimen. In an endeavor to procure more data, and hoping to locate the type specimen, I wrote to various individuals and institu- tions. The replies produced no clue to a type. One correspond- ent, Mr. G. L. Kirk, advised that Mrs. Polly Ridlon Wilson was one of Ridlon’s daughters and that she might have some infor- mation. A letter to her inquired about the final disposition of the Ridlon herbarium and requested any information she might have pertaining to her father’s collection of forma brachypteron. Mrs. Wilson replied as follows: “Unfortunately, I could not locate any correspondence nor even any notes regarding his study of this form of the Polypody. I do not know whether the station is still in existence, and I never had any knowledge of its exact location. However, tucked inside an old edition of The “My father’s herbarium is now at Bennington College, ign we ie hope that the type specimen of this fern is there. . Bloodsucker Pond is a local name for Adam Pond, which can be found on the Londonderry, Vt., Quadrangle of the U. 8. Topographic Survey map. Mr. Ridlon’ s daughter was unable to consult her father’s herbarium until March 9, 1963. Concerning her search for the type specimen Mrs. Wilson wrote: “At last I have been in touch with Mr. Woodworth at Bennington College, and regret to report that there is no good news. I spent ast Piaied carefully checking my father’s herbarium there and am kk fied that there is no specimen at all of the form of Polypody which we are sea 128 AMERICAN FERN JOURNAL “T am sorry to have to thus dash your hopes of finding it in Benning- ton. I had hoped and really expected to find at least something in con- where, but it is strange that you cannot locate it in any of the majo1 herbariums where it might logically have been deposited.” The only documentary evidence of the type specimen seems to be the photograph mentioned above. This photograph has been reproduced and together with data pertinent thereto mounted on sheets for deposit in the herbaria at VT, US, GH, NEBC Bennington College, and the herbarium of the American Fern Society. The discoverers of this form in Colchester, Vt., have offered to make available a few sheets from the new station for deposit in the aforementioned herbaria. It should be mentioned that the distribution of this form, as given by Fernald (1950) was incorrect when published. He wrote, “—, very rare in w. Vt.,; —.” It should have read, “— very rare in s. Vt.; —.” The known occurrence of the forma is as follows: WINDHAM County: “Rocks, Jamaica—type sta. 1919 (Ridlon).” Cuqrr- TENDEN County: on rocks in Colchester Township, 16 plants observed growing with the typical form, Mrs. Lawrence P. Howe and Mrs. Oliver R. Eastman, Nov. 1, 1961 (immature fronds), May 30, 1962, mature fronds). LITERATURE CITED Dote, E. J. 1937. The Flora of Vermont. 3rd Revised Ed. Free Press, Burlington, Vt. FERNALD, M. L. 1922, Polypodium virginianum and P, vulgare. Rhodora 24: 1 a 1950. Gray’s Manual of Botany. 8th Ed. American Book Co., Boston. Ripton, H. C. 1921. A new Polypodium from Vermont. Amer. Fern Jour. 11: 46-48. 7 pl. WEATHERBY, C. A. 1921. A Correction. Amer. Fern Jour, 11: 122, —————. 1937. A List of varieties and forms of the ferns of Eastern North America. Amer. Fern Jour, 27: 51-56. PRINGLE HERBARIUM, UNIVERSITY oF VERMONT, BURLINGTON, VERMONT. SPOROGENESIS IN PTERIS CRETICA 129 Sporogenesis in Pteris cretica with Special Reference to the ytoplasmic Inclusions NorMAN P. MARENGO INTRODUCTION.—Studies dealing with the cytoplasmic inclu- sions during sporogenesis in polypodiaceous ferns have indicated that in at least three species, Nephrodium molle (Senjaninova 1927), Onoclea sensibilis (Marengo 1949) and Polypodium vir- gintanum (Marengo 1959), cytokinesis of the spore mother cell results in the formation of a quartet of tetragonally arranged, and upon maturity, monolete, spores. Adiantum hispidulum, a polypodiaceous fern bearing trilete spores was similarly investigated (Marengo 1962). In this species the cytoplasmic inclusions preserved by mitochondrial fixatives were found to be arranged, prior to cytokinesis, in a pattern of six planes marking the sites of the internal walls of the four tetrahedrally arranged spores arising from the spherical spore mother cell. The cytokinetie basis of tetrahedral Symmetry in this species was thus found to be identical to that described in Osmunda regalis (Marengo 1954). n the basis of the cytokinetic similarity of the maturation divisions of these two unrelated species, both producing trilete, tetrahedrally symmetrical spores, it was suggested that further studies might show in other species a similar sequence of events in meiosis, and thus possibly reflect a pattern of cytokinesis common to the production of all trilete spores, regardless of the taxonomic position of the species considered. Through the courtesy of Dr. William Steere, material of Pteris eretica was made available at the New York Botanical Gar- den. This member of the Polypodiaceae produces typically and obviously trilete spores. It is the major object of this paper to describe the cytokinetics of the maturation divisions in this species and to compare them with previously described processes in Osmunda regalis and Adiantum hispidulum, and to clarify the suggestion that a common process and sequence hold for the production of all trilete spores. 130 AMERICAN FERN JOURNAL MATERIALS AND MetHops.—Fertile leaf margin pieces under 5 mm in length were excised from greenhouse plants and im- nae 3 2 Fies. 1-2, EARLY STAGES OF SPORANGIAL DEVELOPMENT. 1578. FIs. I, EARLY Pegpeasicii) M SHOWING INTERPHASE OF INITIAL SPOROGENOUS CELL PR TAL CELL FORMATION r 2: — oe ASE OF CENTRAL SPOROGE zou; : CELL SURROUNDED BY EARLY TAPE TUN SPOROGENESIS IN PTERIS CRETICA 131 mediately placed in fixative. To preserve the cytoplasmic inelu- sions, the material was fixed from twelve to twenty-four hours in a solution freshly prepared as follows (Huseby 1946): 10% Commercial formalin, 100 ml; Normal sodium hydroxide, 1 ml; Pyrogallol, 7 g. Pieces were rinsed briefly in running water, dehydrated with isopropyl aleohol and embedded in 56°-58° Fisher Tissuemat. Sections three microns in thickness were cut without difficulty. Sections were mordanted twenty-four hours in 2 percent ferric alum, stained twenty-four hours in 0.5 percent hema- toxylin and differentiated with microscopic observation in 2 per- cent ferric alum. Following a thirty-minute wash in running tap water, slides were dehydrated in isopropyl alcohol, counter- stained with Orange G in clove oil, cleared in xylene and mounted in gum damar. Material similarly fixed and sectioned was stained with acid fuchsin and methyl green. Observations and photomicrographs were made with a Bausch and Lomb Dynazoom microscope with a 97X achromatic objec- tive; 4” x 5” negatives were obtained using a Brinkmann Model “U” photomicrographic camera with Kodak Royal Pan Sheet film. Negatives were obtained also with Bausch and Lomb 35 mm photomicrographic camera, using Panatomic-X film. OBSERVATIONS.—The mature spore of Pteris eretica bears a characteristic tri-radiate sear which marks the internal edges of the triangular spherical pyramid which originated as one of the four tetrahedrally symmetrical spores in the young quartet arising by meiosis from the spherical spore mother cell (Fig. 12). Its original pyramidal shape is obscured by enlargement and spore coat formation. The early development of the sporangium is of the usual leptosporangiate type. A large sporogenous cell is separated at an early stage from the cells to become the sporangial wall (Fig. 1). This cell (Fig. 2) soon gives rise to tapetal cells and then to a central group of sporogenous cells. Prior to the maturation divisions, the sporogenous cells assume the rounded- 432 AMERICAN FERN JOURNAL 3-8, SPOREMOTHER CELLS IN VARIOUS STAGES OF MATURATION DIVI- SIONS» chang 3-4 X 1429. Figs. 5-8 & 1578. Fig. 3. Young sPpOREMOTHER CELLS IN INTERPHASE ae ip MEIOSIS, nn 4, PACHYTENE STAGE, FIRST MEIOTIC DIVISION. Fia, sighined PHASE, FIRST MEIOTIC DIVISIon. Fia. 6. INTERPHASE FOLLOWING - FIRS MEIOTIC Bbinabi Fig. 7. METAPHASE OF SECOND MEIOTIC DIVISION, Fic. 8. TELOPHASE OF SECOND MEIOTIC DIVISION. SPOROGENESIS IN PTERIS CRETICA gist up shape of the young spore mother cell (Fig 3). The mito- chondria of the spore mother cell in the interphase preceding maturation are small, granular or rod-like, and of uniform dis- tribution throughout the cytoplasm. At the pachytene stage of the first meiotic prophase, the chromosomes are polarized into a “bouquet,” with an accom- panying polarization of the cytoplasmic inclusions. A small concentration is at the base of the bouquet and a larger mass at the opposite pole of the spore mother cell (Fig. 4). At the metaphase of the first meiotic division, the cytoplasmic inclusions are almost completely localized in a ring or torus surrounding the metaphase plate. In a pole-to-pole section of this stage, this ring appears in cross section as a mass of granules at the edge of the metaphase plate (Fig. 5). As the first divi- sion proceeds, this ring of granules moves centrally into the spindle area. At the time of interphase following the first divi- Sion this ring has been converted into a compact, flat disc interposed between the nuclei (Fig. 6). There is no indication of cytokinesis up to this point. The first indication of cytokinesis appears at the metaphase of the second division. Favorable sections at this stage show the mitochondrial plate with a two-layered structure, suggesting that a cell plate has formed within it, although light microscopy has not as yet verified its presence (Fig. 7). By late telophase of the second division, a cell plate can be detected in favorable sections longitudinal to one of the second division spindles. This cell plate is formed completely independent of the mitochondrial localization which at this time assumes a folded structure, showing as a “V” in cross section (Fig. 8). As the second maturation division is completed, the eyto- plasmic inclusions are regularly arranged along all the internal walls of the newly-formed spores (Fig. 9). Prior to a cigtabe enlargement and quartet separation, these granules lose their localization and assume a uniform distribution in the cytoplasm of the young spore (Fig. 10). : It is difficult to properly fix and stain the young enlarging AMERICAN FERN JOURNAL a * : Bias. 9-12. SPORES neooregihac ty COMPLETION OF MEIOSIS. Fires. 9, 11, 12 x 1429. Fie. 10 & 15 Fic. 9. INTERPHASE IMMEDIATELY FOLLOWING y ME ISION. Fig. 10, SLIGHTLY OLDER QUARTET THAN Fig. 9. Fig. 11. ENLARGING SPORES. Id, 12, MATURE SPORE SRC: TMONED AND SHOWING INTERNAL VIEW OF TRILETE SCAR, SPOROGENESIS IN PTERIS CRETICA 135 spore. This difficulty may be associated with vacuole formation and spore coat deposition (Fig. 11). Stained sections through mature spores show the nucleus at one of the ends of the trilete scar, and numerous large granules along the scar itself (Fig. 12). The precise nature of these granules in the mature spore, and their genetic continuity with the mitochondria of the spore mother cell have yet to be established. Discussion.—The behavior of the cytoplasmic inclusions during the first meiotic division in Pteris cretica appears to be essentially like that observed previously in Onoclea sensibilis and Adiantum hispidulum. The arrangement of the late pro- phase chromosomes into a bouquet is accompanied by polariza- tion of cytoplasmic inclusions comparable to the same stage in the other two species mentioned. The localization of granules into a compact plate between the interphase nuclei resulting from the first division is observed in the same stage in Onoclea sensibilis and Adiantum hispidulum as well as in Osmunda regalis. In the two trilete species, Osmunda regalis and Adiantum hispidulum, there is no indication of cytokinesis or cell plate formation prior to the interphase after the second division, at which point the mitochondrial plates separating the nuclei are in two distinct layers. In Pteris cretica eytokinesis of the first meiotic division has started by the time the second division has reached metaphase. This is suggested by the two-layered struc- ture of the mitochondrial dise formed following the first divi- sion. The completion of cytokinesis in P. eretica is accomplished by cell plate formation independent of any mitochondrial local- ization, and would suggest that in this species at least, this process and its relation to the establishment of tetrahedral sym- metry of the spore quartet is not related to the distribution of the cytoplasmic inclusions during meiosis. In Osmunda regalis, prior to cytokinesis, the future internal walls of the spores appear to be delineated by mitochondrial plates. This Suggests an inherent tetrahedral symmetry of the spore mother cell. The findings in Pteris cretica appear to support this idea, 136 AMERICAN FERN JOURNAL except that this symmetry is established independent of and unrelated to the cytoplasmic inclusions preserved by mitochon- drial fixatives. LITERATURE CITED Husssy, R. A. 1946. Hydroxybenzene compounds as cytoplasmic fixatives. Proe. Soe. Exp. Biol. & Med. 61: 122-125. : Mi niohos, N. P. 1949. A study of the Lpkergog inclusions gr Gee sporo- genesis in Onoclea sensibilis, Amer. Jour. Bot. 1954. The relation of the i topiente recline . ihe estab- lishment of tetrahedral ageangns in the spore quartet of Osmunda regalis, Bull. Torrey Club 81: 501-5 —. 1959. The otokineti Bis of the ropead spores of Polypodium virginianum, Bull. y Club 86: >=. 1962. The .cytokinetic baie Ze tet pata pleas in the spore quartet of Adiantum hispidulum. Bull. Torrey Club 89: 42-48. SENJANINOVA, M. on eer uae, bei Nephrodium molle Desv. Zeitschr. Zellf. Mikr. Anat. 6: 493-508. C Post Criss Lone IsuAnp UNIVERSITY, GREENVALE, Florida Strap Ferns and Their Culture ALEX D. Hawkes The fern flora of the state of Florida is a remarkable one, the largest of that of any of the United States, insofar as I am aware. Of this assemblage, one of the most fascinating genera is that which contains the Strap Ferns, the genus Campylo- neurum. Of the fifty or so species known to science (this genus is often included in Polypodium), four have made their way into our area, where one is a reasonably common indigene, while the others are definite and restricted rarities, Campyloneurum phyllitidis is by far the most frequent and most widespread of the Strap Ferns here. Small, in Ferns of Florida (1931), notes it as occurring in “hammocks, lower two- thirds of the Florida peninsula and Florida keys.” It is, how- ever, not strictly an inhabitant of our marvelous hammocks, Fiorma Strap Ferns 137 for I have encountered it on many occasions in cypress forma- tions, and even in the strange areas bordering ou the great mangrove swamps and buttonwood forests in the Everglades National Park. This handsome fern is often called an epiphyte, but I have very infrequently found it—from many years’ casual observa- tion—growing under conditions which I would term precisely epiphytic. Rather, it in general prefers to form colonies (these often very extensive—and certainly very spectacular when one chances upon them unexpectedly) on fallen, partially decayed logs, these seldom far from perpetually standing water. It may also be found on the mossy trunks of old Pond Apples (Anona glabra) or Bald Cypress (Taxodium distichum) in spots which never dry up entirely, but I would consider this a definitely secondary habitat for the species here. In several of the beautiful hammocks characterized by heavy stands of Wild Tamarind (Lysiloma bahamensis) in the Pine- crest region to the west of Miami, this Campyloneurwm has rather well adapted itself to a semi-terrestrial habitat. Here it grows—with the largest colonies of our pretty Vine Fern (Phymatodes heterophyllum) which I have ever encountered in Florida—in the thick leafy humus under the varied trees. Glorious tree-snails (genus Liguus) are found here, too, as are far too many robust specimens of the vicious Catch-and-Hold- Back vine (Pisonia aculeata), so that botanizing in this region is always a challenge. This Strap Fern, C. phyllitidis, is our largest native species of the genus, sometimes even approaching the grandeur of our Scarce indigenous Bird’s-Nest Fern (Asplenium serratum) with its ereet or gracefully arching fronds to more than three feet in length. These are produced from a stout, creeping rhizome. They generally taper somewhat at each end, are glossy bright- green in color, and are frequently conspicuously irregular and wavy on the margins (though the sawteeth of the Asplenium are never normally present). Sori are typically produced in 138 AMERICAN FERN JOURNAL considerable quantities, in several irregular lines, on each side of the midrib. I have succeded with Campyloneurum phyllitidis in my per- sonal collection for many years, and consider it a very easy fern to grow. My specimens have come from a variety of re- gions in the wild, but all of them are cultivated in identical fashion—in proportionately large, very well-drained pots, in a compost made up of chopped osmunda fiber and fibrous loam in about equal parts. I find that they must have almost total shade to reach their full beauty, and the more moisture and humidity they receive the better they seem to like it. Any protracted period of drought sends them into a serious decline, from which they only very slowly recover. I have noted no serious pests with them, though on occasion a wayward slug will nip off a few of the juvenile sprouting fronds. Several years ago I gathered a couple of smallish specimens of this fern in a swampy area some fifty miles from Miami, growing in close harmony with a stunted plant of Phlebodiwm aureum, and our odd “fossil plant,’ Psilotum nudum, on a large cypress tree. With a machete I managed to pry out of the crevice in the tree-trunk the entire interwoven mass, and upon my return home, quickly mounted it on a sizable slab of porous Guatemalan tree-fern fiber. This soon proved to be unwise of me, and I lost the Psilotum (a very touchy thing to transplant, in any event) and the larger of the Campyloneurums before I moved the entire apparatus into a pot with the above-noted mix- ture of osmunda and loam. This did the trick, and the two re- maining ferns are still thriving at this date. From this, I would assume that our “epiphytic” Campylonewrum does not take well to the treatment normally afforded true epiphytes in our orchid and bromeliad collections. The three other Campyloneurums native in Florida are, as I have already mentioned, all classed as definite rarities. These are C. angustifolium, C. latum, and C. costatum. Campyloneurum angustifolium is noted by Small as having Fuoripa Strap Ferns 139 been “collected but once within the geographic limits of this work. . . It occurs in Timms Hammock, where it grows mostly on live-oak trees. It was discovered in Florida in 1903, and is widely distributed in the West Indies and in continental tropical America.” I have collected what I assume to be this fern both in Cuba and in Nicaragua, in the first-named island on the marvelous “fern paradise’ of Loma del Gato, in Oriente Province ; in Nicaragua on the slopes of El Picacho, near Santa Maria de Ostuma (Hawkes, 1962). In both areas, it was in- variably truly epiphytic, usually growing low down on very mossy trees, with a host of filmy-ferns (genera Trichomanes and Hymenophyllum) as close companions. I have not gathered it in Florida, though I have seen a very few specimens pur- ported to have been collected in the great Fahakahatchee Swamp of the Big Cypress area. These were growing happily in pots— well-drained it must be assumed—in a compost of loose chunks of osmunda fiber, under the closest shade of a lath-house. This Species has numerous fronds, rather closely arranged on the slender, fuzzy rhizome, whieh in large specimens arch to a length of almost 20 inches, with a width of only about 14 inch. They are notably glossy (especially on the upper surface), and their frequent sickle-shaped form makes them unique among all native Florida ferns. Campyloneurum latum (again according to Small) “has been found in Florida only in the Hattie Bauer Hammock in Dade County. It grows about lime sinks, both on the rock ledges and on the bases of trees around the sinks.” I have recently seen cultivated specimens of it, growing in a fibrous medium which Seemed to consist mostly of the “parts” of the fern itself, here in South Florida—these again purportedly from another lo- cality, not one in Dade County—but I personally have never collected it nor grown it in my own garden. Darling (1962) has recently noted an unsuccessful search for this species near Homestead. Small’s comments on the fourth member of the genus, C. cos- tatum, in Florida are, I believe, worthy of quotation: “This 140 AMERICAN FERN JOURNAL strap fern is very rare in our range, and, like C. latum, it has been found in only one locality. It is strongly characterized by the long slender tip of the leaf-blade and the obscure veins. The low hammock is the home of this fern. It is epiphytic and grows on trees, on logs, and on cypress knees. It was discovered in Florida in 1904. Outside of the United States it is known in the West Indies. “It is not evident just how this fern escaped from the West Indies. The spores may have been carried by the wind or on the plumage of migratory birds. Where it began its career in Florda is another mystery. It may have been carried directly to its present isolated habitat in the Big Cypress Swamp or it may have started on the Florida Keys or in the Cape Sable region, and as it worked its way northward, died out in the rear of its course.” I have seen specimens of this Strap Fern relatively recently, ostensibly collected in the same Big Cypress area (Darling men- tions it from the Fahakahatchee, near Copeland, in what is now the Collier County Park), growing happily in pots in osmunda fiber mixed with chopped tree-fern fiber. I have collected what I assume to be this species in eastern Cuba (near the summit of Gran Piedra), growing high in large, moss- and lichen-laden forest trees, in a welter of other epiphytic ferns, orchids, brome- liads, ete. It has not as yet made its way into my personal ° garden here in Coconut Grove, though I hope to acquire it eventually, These are the fascinating and fabulous Florida Strap Ferns, the quartet of species comprising our indigenous representation of the unusual genus Campyloneurum. LITERATURE CrrED DARLING, THoMaAS, Jr. 1962, More Florida Rarities. Amer, Fern Jour. 52: 137-148 Hawkes, A. D. 1962. Notes on the vegetation of Santa Maria de Ostuma, Niearagua. Tropics Mag. I: 8-14, . 6-8, Sma, J. K. 1931. Ferns of Florida. New York. Coconut Grove 33, Fiorma. AN AMERICAN STEGNOGRAMMA 141 An American Species of Stegnogramma Kunio IwatsuKI In my recent revision of the genus Stegnogramma s. lat., (Iwatsuki, 1963), I enumerated eleven species from the Old World and stated that the species of the New World were doubtfully included within my concept of that genus. After publication of that paper, C. V. Morton kindly offered me an opportunity to examine several American specimens having linear or oblong, exindusiate sori. Examining these specimens, I concluded that Gymnogramma pilosa Mart. & Gal. may better be considered an American representative of the genus Stegno- gramma, and a new combination is proposed, as follows: STEGNOGRAMMA Pilosa (Mart. . Gal.) Iwatsuki, nye nov. Gym mnogramma pilosa Mart. & Gal. Mém. Acad. Brux. iad 7, pl. 4, g. 1. 1842; pce ae Selsk. Skr. Vers TST. sf aaa Lege (Mart. & Gal.) C. Chr., Ind. Fil. 284. sens Monogr. Dryopteris I. 196. 1913 Phatypie pilosa (Mart. & Gal.) Crawford, Amer. Fern Jour. 41: 16. 951, Seykseroles pilosa. is most closely related to the wide- ranging 8. pozoi, from which it is distinguished by the slightly yellowish color and the gradually narrowing lower portion of the frond. Pinnules are closer together in S. pilosa than in S. pozoi, though the hairs are not so densely crowded in the former species as in the latter. Hinton 3467 (US) is peculiar in having glandular rather than setose hairs on the sporangia, and with very long hairs on the rachis sometimes exceeding 2 mm. Stegnogramma pilosa is classified into three distinct varieties, vars. pilosa, major, and alabamensis. The known range of the species is from Mexico to Guatemala, var. alabamensis extend- ing into Alabama. Among these three varieties, var. major Seems to be the closest ally of S. pozoi. Transfer of Gymno- gramma pilosa to Stegnogramma necessitates new combinations for the two varieties. STEGNOGRAMMA PILosa (Mart. & Gal.) Iwatsuki var. major (Fourn.) Iwatsuki, comb. nov 142 AMERICAN FERN JOURNAL Gymnogramme pilosa var, major Fourn., Mex. Pl. 1: 73. 1872. Thelypteris pilosa var. major Crawford, Amer. Fern Jour. 41: 19. 1951, STEGNOGRAMMA PILOSA (Mart. & Gal.) Iwatsuki var. alabamensis (Craw- ford) Iwatsuki, comb. now. Thelypteris pilosa var. alabamensis Crawford, Amer. Fern Jour. 41: It is rather difficult to give the differences between Stegno- gramma and Thelypteris in a few words. One of the most dis- tinctive features between them is the structure of the sori. In Stegnogramma, the sori are linear or oblong, exindusiate, some- times extending along the veins and reticulate, the sporangia being setiferous. Among the species of Thelypteris, there are a few which have linear sori, exindusiate sori, or setiferous sporangia. Compared with the vegetative characteristics, how- ever, the soral features of Stegnogramma are distinct. As known in many other fern phylons, such a large genus as Thelypteris presents a wide range of variation even in the soral character- istics. Another important feature of Stegnogramma is seen in the trichomes. All axes as well as laminar surfaces are covered with two kinds of hairs, the longer hairs and the shorter ones. In 8. pilosa, the soral characters and those of the trichomes are quite identical with the same characteristics of the Old World species of Stegnogramma. Above these, we can safely add sev- eral other features which clearly indicate the generic status of. this species, such as the constitution of fronds, especially the basal condition of lateral pinnae, the texture and color when dried. Christensen (1913) included in his Sect. Leptogramma two other species, Dryopteris dasyphylla and D. polypodioides. I have not examined the former; this is a species described from cultivated material, the native locality being doubtful. The latter species has the same soral construction as that of Stegnogramma, but is distinct from this genus by features found in the trichomes, constitution of fronds, venation, tex- ture, and so on. Except for the form of sori, D. polypodioides belongs to Thelypteris, In the frond form and texture, this New CoMBINATIONS FOR SOUTHEASTERN FERNS 1438 species resembles such Old World species as the group of T. crassifolia. There are a few other American species haying linear, exin- dusiate sori. Dryopteris ptarmica var. asplenioides has such soral structure, although the type variety is distinct in having round sori with reniform indusia. The sori of Thelypteris linkiana, T. gracilis, T. atrovirens and others are also oblong and naked. The elongation of sori in the thelypteroid series is discussed critically in my recent paper on the morphology of that series (Iwatsuki, in ed.). I wish to express my sincere thanks to C. V. Morton, whose kindness made possible this study. LITERATURE CITED CHRISTENSEN, C. 1913. A rage of the genus Dryopteris. I. Vid. Selsk. Skr. VII. 10: (196-198). IwatsuklI, K. 1963. Taxonomic ssi of Pteridophyta VII. Acta Phytotax. Geobot. 19: 112-126. . 1963a. Taxonomy of the Thelypteroid Ferns with Special Ref- erence to the Species of Japan and Adjacent Regions I. General Con- siderations. Mem. Coll. Sei. Univ. Kyoto B. 30: 21-51. EPARTMENT OF Borany, Facuuty or Science, Kyoro UNI- VERSITY, Kyoro, JAPAN. Some New Name-combinations for Southeastern Ferns Epaar T. WHERRY While several popular guides to the ferns of the northeastern states are available, the only broad treatment of those occurring farther south is Small’s Ferns of the Southeastern States, whicb was published in 1938 and has long. been out of print. The writer has accordingly undertaken to fill this need, and a book, Southern Fern Guide, is to be published soon. In its prepara- tion several new name-combinations proved desirable. These are tabulated on page 346 of the book, but to render them fully valid, they are published herewith, accompanied by more exten- Sive literature citations and discussion. — 144 AMERICAN FERN JOURNAL a ASPLENIUM PUMILUM Sw. var. anthriscifolium (Jacq.) Wherry, stat. nov 03 Asplenium anthriscifolium Jacq., Collectanea 2: 103. Pl. 2, fig. 2-%, 1788 Because A. pumilum Sw. and A. anthriscifolium J acq. inter- grade in tropical America it has been usual to reduce the latter to synonymy under A. pumilum Sw. In Florida colonies, how- ever, they are markedly distinct in habitat, cutting, and pubes- cence at all stages of development from first blade to fertile maturity. Their segregation at some taxonomic level is accord- ingly deemed desirable. Since their differences are not especially fundamental, that of variety is here proposed. GONIOPHLEBIUM triseriale (Sw.) Wherry, comb. nov. . Polypodium triseriale Sw., Jour. Bot. Schrad. 18002: 26, 1801. i .M GONIOPTERIS sclerophylla (Kunze) Wherry, comb. nov. Aspidium sclerophyllum Kunze in Spreng., Syst. Veg. 4: 98. 1827. Dryopteris sclerophylla (Kunze) C. Chr., Biol. Arb. Tilegn. E. War- i 1911. ming 84, Thelypteris sclerophylla (Kunze) Morton, Amer. Fern Jour. 41: 87. 1951 While the genus Goniopteris Pros] (1836) has been accepted by such authorities as Christensen, Copeland, and Holttum, cogent arguments for denying generic status to such segregates have been presented by Morton ( 1963), who reduces this to Thelypteris subgenus Cyclosorus section Goniopteris. Although admitting that the intergradations of thelypteroid ferns over the world tropies justifies such an arrangement, the writer still feels that in the southeastern United States it is preferable to follow Small in recognizing minor groups as genera. The pres- NEw COMBINATIONS FOR SOUTHEASTERN FERNS 145 ent taxon having been discovered in Florida since Small’s day, anew combination is needed for it. LEPTOGRAMMA PILOSA var. alabamensis (Crawf.) Wherry, comb. now. Thelypteris pilosa var. alabamensis Crawf., Amer, Fern Jour. 41: 16. 1951. The generic problems discussed in the preceding paragraph apply here as well. Leptogramma J. Smith (1842) is reduced by Morton (1963) to Thelypteris subgenus Cyclosorus section Leptogramma. Had the present taxon been known in the United States in Small’s day, he would surely have accepted Lepto- gramma as a “good” genus and the combination L. pilosa (M. & G.) Underw. (1902). In any case, the writer favors doing this to make possible a simple key for use in the southeastern United States, contrasting the reniform-indusiate sori of Thelypteris with the sori formed by a streak of sporangia without indusium characterizing Leptogramina. MicrograMMA heterophylla (L.) Wherry, comb. 1. 1008. Polypodium heterophyllum L., Sp. i" Polypodium exiguum Heward, Mag. Nat. Hist. II, 2: 458. 1838 Polypodium swartzii Baker in Hook. & Bak., Syn. Fil. 3 8 d. ign a 3: 18. Phymatodes heterophyllum (L.) Small, pau "Fiseida oe out Microsorium heterophyllum (L.) Hawkes, Amer. Fern Jour, 41: 52. 1951. ee heterophylla (L.) Diddell, Amer, Fern Jour. 43: 114. (As C. heterophyllum). This ee already having been renamed so many times, the proposal of still another combination seems regrettable. How- ever, the writer can not but agree with Copeland (1947) that the genus Miecrogramma of Presl (1836) should be expanded to include Craspedaria of Link (1841) in that in view of their correspondence in all other respects, their difference in sorus outline (elliptical versus circular) is of no fundamental im- portance, THELYPTERIS X lindheimeri (C. Chr.) Wherry, stat. nov. Dryopteris normalis var. lindheimeri C. Chr., Danske Vid. Selsk. Skr. VII, 10: 1892. 3. 146 AMERICAN FERN JOURNAL Aspidium lindheimeri A. Br. ex C. Chr., loc. cit., as a synonym Thelypteris augescens var. tnehanecs ae hr.) R teh fife Sclat in Small, Ferns S . 241. 1938, Dryopteris Sasksdane var. hin chelinovs (C. Chr.) Broun, Ind. N. A. Ferns 62. 1938. This taxon combines the characters of augescens and nor- malis in such a way as to suggest a hybrid origin, which is confirmed by its abortive spores. THELYPTERIS NORMALIS var. harperi (C. Chr.) Wherry, comb. wl ki normalis var. harperi C. Chr., Danske Vid. ne oi Vid, 1 2. 19 Pokea: ely ste eris qieia P. St. John, in Small, Ferns SE. U. S. 230. 1938. Thelypteris ovata var. harperi (C. Chr.) R. P. St. John, in Small, op. cit. 233. 1928. Dryopteris ovata (R. P. St. John) Broun, Ind. N. Amer. Ferns 76. 1938, Dryopteris ovata var. harperi (C. Chr.) Broun, Ind. N, Amer. Ferns 76. 1938. LITERATURE CITED CopELAND, E. B. 1947. Genera Paes. pp. 247. 10 pls. Walth 4 Link, H. F. 1841. Filieum species in horto regio botanico ahi cultae. pp. 179. Berlin. Martens, M. & H. Gatzorrr. 1842, Mémoire sur les fougéres du Mexique, et considérations sur la géographie botanique de cette contrée. pp. 99. 23 pls. 53: PRESL, - nh 1836. Tentamen pteridopgraphia, seu genera filicacearum praesertim juxta venarum decursum et distributionem exposita. pp- 290. Prague. Morton, aed 1963. The classification of Thelypteris. Amer. Fern Jour. 154. Situ, J. 1842, An arrangement and definition of the genera of ferns, with observations on the affinities of each genus. Hook. Jour. Bot. 4: 38-70. Swartz, O. 1788. Nova genera et species plantarum prodromus. pp. 152. U a. » Li. M. 1902. American Ferns, IV. The pass lew peeeg of the Synopsis Filicum. Bull. Torr ey Club 29: 617-6 EPARTMENT OF Borany, UNIVERSITY OF Preece PHILADELPHIA 4, Piwiwytivanis. CAMPTOSORUS RHIZOPHYLLUS FORMA BOYCEI 147 Camptosorus rhizophyllus forma boycei C. L. Wilson LrEopotp A, CHARETTE On 14 August 1934 Mr. Guy H. Boyce discovered a strikingly beautiful variation of Camptosorus rhizophyllus (li.) Link in the township of Highgate, Vermont, which was named forma Boycet by Wilson (1935) in honor of the discoverer. It differs from the species in having auricles deeply lobed, margins of fronds irregularly dissected, with obtuse teeth 1-3 mm long. _ A search through literature (Broun 1938, Dole 1937, Fernald 1950, and Weatherby 1937) failed to disclose any previous or sub- sequent collection of this form. In late 1962 the writer was at the Gray Herbarium and in April 1963 at Yale University at which times specimens of typical Camptosorus rhizophyllus (L.) Link were studied. Two specimens among the collections at Yale University, from Connecticut, were found to be of this form. Surprisingly enough these had been collected 17 and 30 years prior to the form having been described and named. The exact location of the type station had never been pre- cisely recorded. Desiring to ascertain the present day status of the form I wrote to Mr. Boyce for first hand information. In his reply Mr. Boyee (1962) invited the author to visit him at his camp and further offered to act as guide to the station which he described as follows: “The place where we found the fern is some three miles from camp and quite close to the Cana- dian line, and rather hard to describe. . . . Sentember 8, 1962 was the date agreed upon for visiting Mr. Boyce. On this date the author and Miss Marion L. Smith were personally guided to the colony from which the type collection had originated. The station is located in an open woodland which has only light underbrush and is liberally strewn with large limestone boulders and outerops. The woodlands are adjacent to a cow pasture and the cattle have a path which, unfortunately, passes very close to the colonies of this fern. Ae In the four colonies pointed out to us by their discoverer, 148 AMERICAN FERN JOURNAL the plants in all but one colony were growing at the base of boulders or shelving outcrops and in habit were characteristic- ally closely appressed to the rocks which here seems to afford some protection from the passing cattle. In only one colony were the plants growing on the face or top of boulders. All the plants were in luxuriantly healthy condition. The typical form of the Walking Fern is present, but in no case is it abundant. In a single colony only were the form and typical species growing intermixed, a factor which marks forma boycet as an artistocrat of its clan, displaying a preference for growing in pure stands. The colonies observed are rather small, with perhaps from 3 to 7 or 8 plants only in each. The wood- lands in the immediate vicinity of the station present a notable absence of other ferns as the only other species noted was Adiantum pedatum I. It is gratifying to report that this interesting form is thriv- ing at its original station. At the request of Mr. Boyee and in the interest of conservation the exact location of the station is not being divulged. LITERATURE CITED Boyce, G. H. 1962, 2-paged letter to L. A. Charette dated at Proctor, Vermont, 26 August 1962, Pringle eae document No. 962- 476-1. —————. 1963. 2-paged letter to L. A. Charette dated at Proctor, Vermont, 11 February 1963, Pringle Herbarium document No. 963- Ae: i, WeaTHeRBy, C. A. 1937. A list of varieties and forms of the ferns of Eastern North America. Amer. Fern Jour. 27: 20-24, 19 Witson, C. L, 35, "4 new form of the Walking Fern. Amer. Fern. a Jour. 2 8. PRINGLE Meio UNIVERSITY oF VERMONT, BURLINGTON, VERMONT. SHORTER NOTES 149 Shorter Notes ASPLENIUM X EBENOIDES R. R. Scorr in Kenrucky.—EKarly in the winter of 1961, Dan Schreiber asked me to examine sev- eral fern specimens collected by one of his students, Donald Willoughby. Among the specimens, all collected in Madison County, Kentucky, were two fronds of a fern unknown to me. I thought they might be hybrids since they were different from ferns commonly found in the area. The fronds were sent to C. V. Morton who identified them as Asplenium X ebenoides Scott, commonly called Scott’s spleen- wort or walking spleenwort. This is the rather rare hybrid be- tween Asplenium platyneuron Oakes and Camptosorus rhizo- phyllus Lin One frond hed the general appearance of A. platyneuron, the common ebony spleenwort. The second looked quite different. Some of the pinnae were extremely elongated and resembled closely individual fronds of C. rhizophyllus in miniature, even to the somewhat irregular arrangement of the sori. Morton said these were the most robust plants of this hybrid he had ever seen, Since only individual fronds of this fern had been collected it was hoped that the specimen could be relocated. Mr. Wil- loughby described the site of the original collection as a wooded Slope near College Hill, Madison County. However, subsequent visits to the area failed to relocate the plant. The original col- lection is deposited in the United States National Herbarium. It was first believed that this fern had not previously been reported from Kentucky since it was not included in the lists published by Reed (Castanea 23: 1-13. 1961; op. cit. 26: 94- 96. 1961), and Smith (Castanea 24: 48-50. 1959). However, a recent list of additions by Reed (Castanea 27: 83-87. 1962) provides a clue to a previous record. Williamson’s Fern Etch- ings. Ed. 2, illustrating all the species of ferns indigenous to the northeastern United States and Canada, published in Louisville, Kentucky, in 1879, contains several illustrations of ferns which 150 AMERICAN FERN JOURNAL he recorded from counties in Kentucky. Plate XVII shows Asplenium ebenoides. According to this source, “Professor Wild- berger” collected A. ebenoides in Franklin County in 1878. Ap- parently this is the earliest published record for this hybrid in the state. I wish to thank Conrad Morton for identifying the plant, and Donald Willoughby for giving the specimens to the National Herbarium.—THomas A. Hurro, Biology Department, School of the Ozarks, Point Lookout, Missouri. Recent Fern Literature FLorRA oF SOUTHEASTERN WASHINGTON AND oF ADJACENT Ipano. 3rd Ed., by Harold St. John. Outdoor Pictures, Escon- dido, Calif. xxix, 1-583. 1 map, 11 figs. October 1963. $6.95.— The second edition, which appeared in 1956, included about 30 pages of addenda which have now been incorporated into the 3rd edition. The present book is essentially a facsimile reprint of the former editions with some additions and corrections. The sequence of families of angiosperms follows Engler & Prantl. Within each family the genera are arranged alphabetically, as are the species in each genus. One of the notable features of the 3rd edition is a revised taxonomy of ferns, particularly in the delimitation of families along lines proposed by R. C, Ching, E. B. Copeland, R. E. Holttum, and others, although apparently these authors are not mentioned. There is no evidence of ordinal concept, but there are three classes, with Filicineae inserted between Lyeco- podineae and Equisetineae. The families of ferns begin with Ophioglossaceae and end with Marsileaceae, with the old “Poly- podiaceae” represented by eight families. For one of these, the name Cheilanthaceae would have been preferable to Sino- pteridaceae. The families are arranged in the key on p. xvill into two groups according to whether the rhizome is dictyostelic, protostelic or solenostelic, The book is clearly printed and well-bound and will continue to be useful for identifying plants growing in southeastern RECENT FERN LITERATURE 151 Washington and adjacent areas. The author is to be congratu- lated on having kept the book more or less up to date, and the publisher on having produced an attractive, handy volume.— EVILLE JONES, University of Illinois, Urbana, Illinois. Fern MATERIAL Usep IN RESEARCH ON MORPHOGENESIS AND PHOTOPERIODISM.—For thirty years and more ferns have pro- vided material for important research in fundamental prob- lems of differentiation of tissues and organs. Leading work has been carried on in England by Wardlaw and his students and by Wetmore and his aides at Harvard. In a recent paper’ DeMaggio and Wetmore reported successful excision and culture on nutrient media of young embryos of Todea barbara, an Australian member of the Osmunda family. When embryos that had become spherical, 5-7 days after fertilization of the egg, are used the embryo could be carried to independent growth, with a root-stem-leaf system, on the medium. ith one-celled embryos, however, multicellular structures would be produced, but they were without tissue organization and resembled prothallia. Another pair of experimenters? report the effects of exposure to red light on early stages of the prothallia of Onoclea sen- sibilis. Like most fern spores, those of Onoclea require light to germinate. When germinated spores are then placed in dark- ness, growth is in the form of a slender filament. Brief exposure to red light will increase the rate of growth (elongation) of these filaments to about the 12th day. Thereafter, the elongation rate is reduced by exposure to red light—Raupu C. BENEDICT, Pilot Knob, New York Jor, BarBARA. Species of Dryopteris Cultivated in California. Baileya 11(4): 117-130. Illus. 1963—Provides a key, brief descriptions and good photographs of each of eight taxa. 1De di dis, A. E. an Wetmore. Growth of Fern Embryos in Sterile Culture. Nature ‘191 Libay: 94-95. Miller, John H., . Wright. An Age e-Depen ndent Change in the Response ot of an inert a to Red Light. Science 134 (3490): 1629. OV.- 17) J 1a2 AMERICAN FERN JOURNAL Wacner, W. H., Jr. Pteridophytes of the Mountain Lake area, Giles County, Virginia, Including Notes from Whitetop Mountain. Castanea 28(4): 113-150. Illus. 1963.—A_ check list with copious notes on geographic distribution, ecology, and cytotaxonomy of the ferns and fern allies studied. Notes and News Harry W. TRuDELL was born in Richmond, Virginia, May 2, 1879, but came to Philadelphia as a youth. Becoming an accountant, he was employed for many years by a large leather firm, and rose to the important position of Secretary. A nature enthusiast, he spent all available time outdoors, collecting both botanical and mineralogical specimens. He also took an active part in such organizations as the Philadelphia Botanical Club, which he served as treasurer for many years, and the Phila- delphia and Pennsylvania Mineralogical Societies, Friendly in manner and having a fine sense of humor, he was an ideal companion on field trips, and joined the writer on many searches for ferns. He was a member of the American Fern Society for 45 years. A hybrid between the Lobed and Mountain Spleenworts, which we found on the cliffs of the Sus- quehanna River in Laneaster County, Pa., was named Asplenium trudellii Wherry in his honor (Amer. Fern Jour, 15: 49, PI. 4, figs. 4-5. 1925). He enjoyed superior health for 80 years, then was stricken with Parkinson’s disease, and after gradually failing, passed away on January 26, 1964. His collections of ferns and other plants have been placed in the herbarium of the Academy of Natural Sciences of Phila- delphia and other similar institutions—Epcar T. WHERRY, Umversity of Pennsylvania, Philadelphia. XPERIENCES IN RaAIsinG FERNS FROM Spores.—Two or three years ago I decided to try raising some ferns from spores, being stimulated by Mrs. Kay Boydston’s article in the Fern Journal! 'Boydston, Kathryn. An Amateur Plants F Spores. A Fern Jour. 48: 1-18. 1958, lel al cineny Notes AND NEws 153 and by the existence of the Spore Exchange of the Fern Society. Mrs. Boydston kindly sent me some spores, and I decided to treat them according to her system, which seems to me the most practical of those I have read about. The spores were all sown at the same time in transparent plastic jars, with enough mois- ture, and placed in a well-lighted room, away from the direct sunlight. After two or three weeks some prothallia started to develop. But just then I had to be absent from home for two weeks, and on my return I found to my regret all the jars as dry as a bone, the prothallia all dead, and no new ones apparent. Unwilling to give up I sogged the whole mixture, put the whole ruined collection on a window sill in full sun, and hopelessly waited. In less than two weeks the fattest prothallia developed in all the jars. This shows that not all spores in a sowing develop at the same time. I let them grow under the same con- ditions until I was able to find time to transplant the plantlets into individual baby pots. I placed the pots in a sort of Wa ian case, but since I do not have room for Wardian cases every- where I selected only the best and strongest plants, which are how doing pretty well. Some of my ferns are attacked by seale insects. When young these look like transparent oval-shaped drops about one or two mm long. Later they harden and become darker brown. They Seem to start from the base of the plant and work their way upward, covering at last the segments of the fronds, the stems, and all. When crushed the shell bursts and the inside is whitish and sticky. The plants attacked as a rule suffer considerably, Stop putting out new fronds, and lose all vitality. I have used fish soap and Blackleaf 40, and have rescued half of the plants treated, but on some this seems not to work. None of the nur- series I have asked can give me a satsifactory solution to this problem. It is a discouraging business. If we do not watch our Plants carefully the scales spread very rapidly.? *Malathion is ee to be the most successful insecticide against Seale insects. [C.V.M.] 154 AMERICAN FERN JOURNAL My wife has about 200 varieties of plants growing inside the ouse—cacti, philodendrons, orchids, anthuriums, and so on, including a Christmas cactus (Zygocactus truncatus) hanging in a window where it gets all the sun it wants. One day I noticed something else showing besides the cactus. Of course the cactus was kept on the dry side, but there it was—a fern! In a few days another showed up and a third. I started picking them out and transplanting them into pots. Most of them turned out to be Adiantum hispidulum, of which I have a nice specimen in my fern collection. One was Pellaea rotundifolia, which I also have in my collection, and another I have not been able to identify yet. I have already given away four or five nice looking plants, and more are coming along—NicHoLas BaRTHO, RFD, Center Conway, New Hampshire. A Correction.—In Wilson’s paper on Adenophorus (Amer. Fern Jour. 54: 68-70, 1964), line 9 on page 68 should read: (Kaulfuss, 1824), as Polypodium tamariscinum Kaulf. (Adeno- Remove the parenthesis just before, but not after “(Gaud.).” in line 11, to read: Gaud.). AMERICAN FERN SOCIETY An Open Letter from the President Few scientifie societies are so fortunate as ours is to have AMERICAN FERN Society 155 have forged a unified bond of common interest for the advance- ment of our knowledge of ferns and for the pleasure of work- ing with them. This has been a healthy situation, and one that should be constantly encouraged. With the increased amount of leisure that is becoming avail- able to many of us because of shorter work time and longer va- cation periods, the need for recreational interests is going to become more pressing. I do not know of any more pleasant way in which to spend one’s free time than in studying and working seriously with our native plants, especially ferns. Also, the pleasure of getting out into the field with people of like inter- ests has a strong fraternal appeal. Coupled with the need for developing more avenues of recre- ational endeavor is the urgent need to assure that organizations such as ours, providing recreational outlets, be placed on a sound and permanent footing, so that they and their activities can be perpeutated for the enjoyment of future generations. With the steadily mounting costs of publishing our Journal, a most important adjunct to our activities, and the future need to help support certain special publications in the field of ferns and fern allies, we should all give serious consideration to what we might be able to do personally for the Society. The Amer- ican Fern Society has been fortunate in the past to have had Some members who generously contributed materially to its progress and well-being. Such a one was Mr. C. A. Weatherby, who not only during his lifetime personally contributed sub- stantially to the advancement of our Society, but who also left a generous gift to the Society in his will. mong our members are doubtless some who now or in the future may wish to contribute generously to the support and perpetuation of our Society. I appeal to these individuals to keep in mind the future needs and welfare of the American Fern Society.—Donovan S. CORRELL. 156 AMERICAN FERN JOURNAL American Fern Society List or MEMBERS Aborn, Mr. Robert G., Box 323, Millington, New Jersey oe: Dr. Ralph D., 2805 - 26th Street, Moline, Illinois eres Eugene B. 2112 E. 60th St., Tulsa, Okla. 74105 cae Mrs esse C., 200 E. 9th St., eae Akutow agit Frank, 2007 Harvey Road, Wises 3, Del. Alava, Dr. ee Dept. of Botany, University of Turku, Turku, Fin Albro, si Mary Stephanie, 29 Mosswood Road, Berkeley 4, Calif. Alexander, Miss Eva, 900 Graymont Ave. W., Birmingham 4, Ala. Alexander, Mrs. Leonore, 2104 Grant Ave., Redondo Beach, Calif. Allen, Dr. & Mrs. Edward P., 505 Ridge View Rd., Orange, Conn. Allen, Mr. Walter 8., 2 Pool Road, New Haven, Conn. Allison, Dr. Benjamin R., 26 Ives Rd., Hewlett, L. I., New York Amason, Mr. Carl R., Route 3, Box 180, El Dorado, Arkansas Anderson, Prof. Tewie E., 2020 Sunset Avenue, Durham, Anderson, Mr. Roger F., Mich. Chr. Jr. College, 800 W. ANoik Rd., Rochester, oe Andreas, Mr. Lee ihe eon State College, ive ens Point, Wis. Andrews, Prof. me . Jr., Botany Dept., University of Artz, Miss Lena, Waterlick, Virgin Atkinson, las (Lenette), ps South Pleasant St., Ambher Attridge, ae “Helen M., 358 oes Ave., Boston 15, Mass. Atwoo Baker, Miss Harriet E., 1927 Buckingham Rd., Los Angeles 16, C alif. Baker, Dr. William H. , University of Idaho, Moscow, Idaho Baldwin, Prof. ee College of William & Mary, Williamsburg, Va. s. F. L., Jr., 149 Northwestern Ave., Philadelphia 18, Pa. Ballard, Mr. Peden Royal Botanie Gardens, Kew, Richmond, Surrey, England 19 Bange, Mr. M., 24 Rue Grenette, Lyon 2, France Barnes, Mrs. Albert C., Lateh’s Lane, Merion, Pa. Barnes, Mrs. William A., 2645 Edgewood Road, Utiea 3, N. Y. Bartho, Mr. Nicholas, R.F.D., Center Conway, New Hampshire Bar ioe Mr. Ralph R., 517 E. Alder St., Brea, Calif. AMERICAN FERN SOCIETY Bartsch, Mrs. Paul, Gunston Hall Road, Lorton, Virginia Bauer, Mr. Bill, ee Hill, Route 5, Arnold, Missouri Bayley, Mrs. T., Da bur Park, P.O. Box 21 MR, Marlborough, Salisbury, S. Rhodes Bean, Mr. Ralph C., 48 pire St., Wakefield, Mass, Beard, Mrs. Elva eet Route 3, Box 354, Salem, Virginia Beckner, Mr. John 6 Myrtle Way South, St. plage pn Fla. Beeman, Miss tone 7325 Willow Lane, Rayt 33, Missouri 0 Mrs. Dorothy 442 ceckane Ave., tthsib ithe California enediect, Mr. J. E., Jr., 9304 Warren St., Silver Spring, Maryland neta Dr. Ralph = "Pilot Knob, New Yor Benedict, Dr. Ralph 3106 oo St., Rockford, Illinois Benedict, Mr. ee he 83-09 122 St., Kew Gardens, New York Benson, Dr, Lyman, Pomona ee Claremont, Califor Benton, Mr. Brantford B., 134 Cedar Lake West, bea ge Sods Berko, Mr. Stephan J., P. O. Box 111, Export, boigane heroes Berry, Mrs. Robert, 817 Fairview Dr., Aberdee ea sh. 98520 on Miss elie 12 Boynton St., Worcester ukh Singh, Botany Dept., Panjab str Chandigarh 3, ce dia Bishop, Mr. Earl, 633 South Orehard St., Madison, Agere Bittinger, Mrs. Charles, 3403 O Street NW, Washington 7, D. C. Blackley, Mr. W. J., 16 Almay Grove, Pig ph mes Australia Blair, Dr. Harry C., Neskowin, Orego Blake, Mrs. Stephen M., Kittery Paint Maine Blasdell, Dr. Robert F., Dept. of Biology, Canisius College, Buffalo 8, 1 ew York Blaydes, Dr. David E., Dept. of Botany & Plant Path., Michigan State University, East ee Michiga Blaydes, Mr. Glenn W., 214 Westwood Road, “Colum 14, Ohio i Mrs. Henry W., P. O. se 51, Ravena, New York 12143 her ding, Mas Siswiolee Prof. H. L., Duke University, Durham, cre Carolina oom, Mr. William W. 755 Dove Ave., Valparaiso, Ind. 4638 Boeck, Nona Ford, 4702 Dietrich Rd., San Antonio 19, Texas Boivin, Dr. Bernard, Div. of Botany, Central Exp. Farm, Dept. of Agriculture, nets: Ontario, Canada Bold, Dr. Harold C., Dept. of Bot., Univ. of Texas, Austin 12, Texas Borghese, Lydia, Via S. Conea 12, Rome, Ital Boutin, Mr. Fred C., 2620 N. Willard Ave., So. San Gabriel, Calif. aie Mr. Bernard, 9821 S.W, 47th St., Miami, Floric Boydston, Mrs. Kathryn E., Fernwood, Rt. 3, Niles, oe 158 AMERICAN FERN JOURNAT: sees Mrs. Herbert A., 8 Highland Ave., St. Augustine, Florida 64 Brannen, ee Now. Irs 3430 Ada Ave., Fort Worth, Texas 76105 1964 Branscomb, Mr. Donald L., Route 1, Bix 45, Willits, Calif. 1954 Branum, Miss Florence, 727 Rutter Ave., Lancaster, Ohi 947 Brass, Mr, Leonard J., Archbold lag age Sta., Lake Placid, Fla. 1952 Brauss, Mrs. Albert, 7517 - gi: sie Re dn ond, Washington 1964 nS ona oe Okr cae ad right, Ontario, Cana ~ 1957 Brennan, Mr. Patrick ae a ait Ave., South Burnaby, B.C. Rea 1962 Brewster, Miss Frances W., 294 Montgomery St., Bloomfield, N. J. 1955 Bristow, Miss Alice A., 141 Silvermine Road, New Canaan, Conn 1952 Bristow, Miss Helen G., 141 Silvermine Road, New Canaan, Conn 1952 Guelph, Ontario, Canada Brooks, Dr. Karl L., 431 E. 3rd St., Brooklyn 18, New York 1951 Brooks, Mr. Maurice G., West Virginia Univ., Morgantown,, W. Va. 1926 M Maurice, Hawk Mt. Sanctuary, R.D. #2, Kempton, Pa. 1934 8 = ° 5 ~ Brown, Mr. Roger H., 321 West Mountain Road re Mass 1964 Bruce, Miss Rena, Box 50, RD 2, Williamstown, Verm 1962 Bruce, Mr. Robert G., 10019 Montauk Ave., Bethesda 34, Coa 1960 Bryan-Arana, Mr, Carls, Agr. Ext. Service, P.O. Box 607, Rio ras, Puerto Rie 64 Bryngelson, Mrs. Ed, Box 359, Princeton, Florida 1962 Bugbee, Mrs. Lloyd H, Bradford, Vermont 1954 Buker, Mr, W 3833 Oswego St., Pittsburgh 12, Pennsylvania 1958 Bulger, Mr. William, 108 N, Bacay Street, St. Clairsville, Ohi 961 Burati, Mrs 1, 9201 Over Lea Drive, Rockville, Mary 61 Burton, Dr. Daniel F., 512 Hickory Street, Mankato “Mindaenis 1949 Burton, Mrs. Verona, 512 Hickory Street Mankato, Aba cain 1949 “wid Mrs, Harold E., 2163 No. Starr Rd., Columbus 12, Ohio 53 19 » Mr. James R., E. State St., Apt. 14-F, Athens, eon 45701 1964 “aan Mrs. te H., 4982 Cherry Bottom Rd. , Gahanna 1961 Calder, Mr. J. A., Plant Research Institute, Geuteal © cent Farm, Dept. of ray Ottawa, Ontario, Canada 1952 , Miss Elsie D., 1023 Mencher Blvd., Johnstown 1935 cai, Dr. James E, Dept. of Botany, Arizona State gar cue Tempe, Arizona 1964 Caponetti, Mr, James, Dept. of Bot., Univ, of Tenn., Knoxville, Tenn. 1963 AMERICAN FERN SOCIETY Capurro, Dr. Roberto H., Chareas 1014-40, Buenos Aires, Argentina Carlson, Mr. Thomas O., 1627 Peachtree St., N.E., Atlanta 4a fa r cio, Mrs. Peter J., 2598 sagen ds Avenue, West. an ‘U7, Ak Cerimele, Miss Carol, 6208 S. 286th, Kent, W Chamberlain i Pi ssie is Hk: :.. Chapin, Dr. Hdward A., 505 Main St., West Medway, Mas Chillas, Mr. Richard B., Jr., 233 Winona Ave., Dib St 44, Pa. Cing-Mars, Mr. Lionel, 1289 Jean Dequen, Ste.-Foy, Quebec, Canada Clark, Mrs. J. M., 1724 Rockwood Trail, Waybtlevills: Arkans Clarke, Miss Gladys, 209 Lincoln Ave., Takoma Park 12, Clarke, Prof. Herbert M., Birge Hall, Univ. of Wis., Madison 6, Wis. Clarke, Mrs. Kenneth L., 1420 Orangegrove, Orange, California Clausen, Dr. Robert T., Dept. of Botany, Cornell Univ., pL Bre a obb, Mr. Boughton, 116 E. 66th Street, New York 21, N Cody, Dr. W. J., Plant Research Institute, Central ns EA Farm, Ottawa, Ontario, Canada Coleman, Mrs John R., 3699 Lake Avenue, Rochester 12, N. Y. Coleman, Mrs. Stewart P., Barrett Road, Cedar ee leg Deas = Ye. Baton Soe Counter, Mrs. Vernon A., Box 38, Petersburg, Covert, Miss Ione, 1105 Elizabeth Street, South pemers 6, Va. Cozzens, Mr. James Gould, Shadowbrook, Williamstown, Mas Crandall, Miss ana 1 ies ee -Macon Woman’s pa Box 8, Lynchburg, V Crane, Mrs. Charles W., Che Summit Ave., Summit, New Jersey ahi Marguerite, 6 Coronation Rd., Auckland, 8.E. 3, New Zealand tT, Mr. Orvil «aoe a Pihinghan, Mr, Edward, 199 Washington St., Boston 8, Mass. Cunningham, Miss Ruth, 1034 Castle Rock Rd., Walnut Creek, Calif. Curry, Mrs. R. Granville, River Road, Rockville, Maryland Cutler, Mr. Alvin B., 7675 8.W. 128 St., Miami, Fla. 160 AMERICAN FERN JOURNAL alin Dr. Pierre, New York Botanical Garden, Bronx Park, ork 58, N. Y. 1948 Darling, Mr. Thomas, Jr., 3809 Larno Drive, sneered pecans 1952 Davidson, Mr. Donald William, ee of Biology, Univ. of Alabama, er Box: 1927, es iversity, Alabama 1962 Davis, Mr. Bill D., Dept. Biological ene Purdue University, afayette, Indian 1963 Dean, Mrs nche E., 232 Rosewood Street, Crestline Gardens, irmingham ries Alabam 1960 Deaver, Mr. Chester F., Adela State College, Science Dept., Flagstaff, pa 1960 Deitrick, Mr. Ralph, 19 bs Rd., Dexter, Michigan 1962 e Joncheere, Mr. G. Antwerpen N.V., St. Paulusstraat 42, Ant Bel 1952 Dell, Mr have L., 137 Piermont Rd., Hewlett, L. I., N. Y. 1960 ced ota, Dr. Elias R., Facultad Ciencias Naturales y Midesn: a Plata, mibaiieg: 1963 sinha Mr: C: FE. 0 Toni Drive, Miami 57, Florida 1960 Demaree, Dr. Delzie, ipa ysis Avenue, Hot Springs, Arakansas 1938 DeNault, Mr. Kenneth J., Box oe Stanford, California 1963 Derr, Mrs. Blanche A., 232 Summit Rd., Springfield, Del. ais 1959 Desmond, Hon. Thomas C., 94 Soden. ‘Seater urgh, N. 1943 Ce so, rat E., 65 Wan Shen St., Chingmei, bien Taiwan, e Chin. 938 Heated Mr. and ay Spates 401 Hasbrouch Blvd., bean: N. J. 1956 Dickinson, Dr. Lewis, 321 W. Washington, Glasgow, Hen 1959 Dickson, Mr. Warrick Biology Dept., MeNeese State seca ake Charles, La. 1960 Dillman, Miss Paula, P. O. Box 364, — Florida 1955 Divoky, Mr. Robert Eugene, 1168 E. 78th St., Kansas o 31, Mo. 1952 Doray, Mr. Robert A, 2 Orchard cee Greenfield, Mas 1941 Doubles, Dr. James A., Dept. of Botany, Birmingham asters College, Birmingham 4, Alaa ama ec Dr. tee ude E., Feura Bush, Albany County, N. Y., 12067 1951 Dowling, Mr. C. F., Jr., 11545 SW 107th Court, Miami, Fla. 1950 Drew, Mr. Eric, 6 ton St., Apt. 2A, New York, N. Y. 1963 Dunean, Dr. Wilbur H, Deck of Botany, Univ. of Ga., Athens, Ga. 1949 Dunham, Miss Ann L., 331 Abbey Court, Ridgewood, New Jersey 1953 Dunlap, Mr. Frederick 1410 University Ave., Columbia, Missouri 1960 sagt Pr ouglas W. Badd of Wis-Milw, 3203 N. Downer , Milwaukee 11, Wisco sin 1941 hee ee Edmond, 2106 taut Ave., Wilmington 6, Delaware 1962 Durfee, Mr. Horton H., The Darrow Scheel: New Lebanon, N. Y. 1952 AMERICAN FERN SOCIETY Dyee, Mr. J. W., 46 Sedley ge Loughton, Essex, England Earle, Dr. T. T., Dinwiddle Hall, Tulane Univ., New Orleans 18, La. East, Miss tana A., 167 Seymour St., Hartlord Conn 0 u bia, g Vermont ards, Mr. James L., 27 Stanford Place Wontenie Nod. Pathe. Mrs. James C., 2889 SW 35th ‘Aiea. cap 33, Florida Ellis, Mr. Erl H., 1955 Albion St., Denver 20, Color Emery, Mr. Dara E., 517 W. Junipero St., Apt. 2, ae Barbara, Califo Emmons, ue ‘hie, T., 177 Lewis St.,. Geneva, N. Y. Emory, Mr. David L., Hie 188, Mercersburg Acad., Mercersburg, Pa. Engle, Mr. J. M., 1421 8th St., North, St. Petersburg 4, Florida Epstein, Mr. Harold, 5 Forest Court, Larchmont, N. Y. dy Angie, 153 Navarre Drive, Miami Springs, Florida , Mr. A. Murray, Dept. of Botany, Univ. of Tennessee, as xville, Tenn Evans, Mr. Ralph M., ees Kodak Co., Kodak Park Bldg., 65, Evert, Mrs. ease 430 ioe s Ave., Riverton, Naw Jersey Ewan, Prof. Joseph, Dept. of er Tulane Univ., New Orleans, La. 1 ulberr an , Dr. Mildred E., 304 Euclid Ave., Syracuse 10, New York Felix, Mr. Charles J., Sun Oil Lab., 503 N. Central Expressway, Richardson, Texa Ferguson, RE ms 33 Central Ay a Staten Island 1, N. Y. Fisher, Mrs. C, Page, bax 5152, Ralei Fisher, = C. ee pie miinon irhorabory, Bromyard, erefordshire, Kyr , En Fliflet, Mr. Thorle ie, 12 Keni orth a Mountain Lakes, N. J. Flo, Mrs. Spence y Club Road, Greenfield, Mass. ‘GC. 772: Cou Flowers, Dr. Seville, Raine ae Utah, Salt Lake City 1, Utah Ford, Prof. Ernest S. , Dept. of Botany, Univ. of Florida, oe a. Fosberg, Dr. F. R., 212 Holmes Run Road, Falls Church, Va. Foster, Mr, Kastih T., Derby Lane, Vermont Poster, Dr. F. Gordo on, ‘10 North sha Maplewood, N.. J. Foster, Mr. Lincoln, Falls Village, Con Fowler, Mrs. Priscilla G., 2845 Carlisle Sona. Birmingham 13, Ala. Fox, Mr, David S., 6623 W. Markham St., Little Rock, Arkansas 162 AMERICAN FERN JOURNAL Francis, Mrs. Carl G., 8739 Satyr Hill Road, Baltimore 34, Maryland Frank, Mrs. Irving H., 247 Summit Ave., Mill Valley, nenk Gey Franzen, Miss Tina, 1801 Niles Ave., St. Joseph, Mich Frederick, Mrs. V. R., 145 Tanglewood Drive , Urban Nie rehse, Mrs. Robert M., 506 West t Maplehurt, Fenn 20, Michigan 1 Fried, Dr. Alan R., Livitig ston » New York Fuchs-Eckert, Dr. Satis Boks acces te Wassenaar (ZH), The Neth Gaede, Mr. Robert H., 27 Werimum Brook Road, Saddle River, N. J. Gaetzi, Dr. Walter, Duforstrase = St. rarer ieee Garbe le er, Mr. Miles T., Jr., R.D , Carlisle, Gardiner, Mrs. George N. Pine oe Ra. tas, Vermont Gardner, Mrs. George H., 1724 Asbury ies Evanston, Il. Garretson, Mr. J. D., Apartado Aereo 342, het, Colombia, 8. “A; Gaston, Mrs. John Zell, Box 147, Webster, T onda oe Joseph, 25 Ronald Road, West ‘Hollywood, Fla., 33023 Giau s. M. F. Ashley, 2643 Benvenue Ave., Berkeley 4, Calif. Cisse, ‘habs A., 14781 Farwell Ave., Sa — alif. Gier, Dr. L. J., Wil Bens Jewell oe. Liberty, M Gilbert, Mr. Alfred H., Dorset, Verm Gilbert, Dr. Neal W., Dept. phase Univ. of Calif., Davis, California Gilkey, Dr. J. V., 2170 E. 28th Street, Oakland 6, Cal Gillespie, Mr. ‘Tainhie P., Science Dept., Marshall rants ree oe West Virginia Glasgow, Mrs. Robert D., 1013 Washington Ave., Albany 6, N. Y. Gliddon, Mrs. Gordon H., 14 Rope Ferry Road, Hanover, N. H Glynn, Mrs. Henry, Frankfort, Kansas Pht Mrs. Alice N., P. O. Box 483, Paradise, Calif. y, Mr. Erie V., P.O. Box 1881, Bradenton, Pini Fase Mrs. Martin: 24 Willow St., Brooklyn, N. Goodman, Dr. George J., Dept. of Plant iiaace Univ. ie Oklahoma, Norman, Oklahom Boot pnstive Mrs. Ske 3407 Hopkins St., gto, Tenn. Goslin, Mr. Charles R., 726 E, Kin ng St., egteastoc. hio Graf, Mr. Alfred Byrd, Julius Roehrs Oa East es N. J, Graham, Mrs. Rose H., (Mrs. J. L.) 404 West Queens Rd., North Vancouver, B. C., Cana Grajek, Mr. A. C., 2928 So. 15th St., Milwaukee, Wis abas ig Mrs. Christopher, 6677 Mesdvthey Blvd.,, Washington 16, C. Grant, Mrs, Hannabell Horton, 5521 Amestoy Avenue, Encino, Calif. 1960 19 AMERICAN FERN SOCIETY ibang Paul N., O.D., 60 Third St., Waterford, New Yor Graves, Miss E. Irene, 122 Mt. Prospect Street, aie gi sa Mass. 0232 Graves, Mr. R. A., 4606 Ortego Blvd., Jacksonville 10, Fla. Gray, Mrs. Ronald S., 64 North Street, North Reading, Mass. Greene, Mr. Frank C., 2838 Forest Ave., Kansas City 9 Grether, Mr. David F., i Teachers College, St. Cloud, Minn. Groot, Jan E., 1427 E m Ave., Orange, California Grunden, Mrs. Watee;: 5 aoe Road, Mountain Lakes, Gulick, Mr. Edward V., Dept. of History, Wellesley ce Wellesley 81, Mass. Gunnison, Mrs. R. M., The White Sands, 7450 Olivetas, La Jolla, California Gwinn, Mrs. L. K., 4008 Urbana Avenue, Dallas 27, Texas a 2, Apopka, Fla = 12 Dale J., 164 Westchester Way, Binkinatan, Mich. etd Me. Robert, 1963 Glynn Court, Detroit 6, Michi ig Haldimand, Lois D., 56 ayes Saye , White Plains, N Hall, Mr. Neill D., 1225 7 i Bireck, oo 15, sabe taal Hand, Mr. Louis E., Box ga Lisbon beara Handley, Mrs. Jack, at ate “Rowi: ee alan Hansen, Mrs. Don W., 101 Forestdale Drive, Eten NEG: Hardy, Miss Ruth W., a, Cedar Wild Fir, Nurs., §. Canaan Rd., Rt. 7, Falls Village, t Harley, Mrs. ange i fe Merry Point, R.R. 1, Laneaster, Virginia Harrison, Mr. H. h, Dept. of Botany & Bacteriology, Montana State College, DEL Montana, 59715 Harsh, Mrs, George, 383 nee! Rd., ee 17, Tenn. Hart, Mr. Cecil, 132 N. 3rd St., Montebello, Cali canes oa Richard L., Dept. of Botany, Univ. of ¢ Rhode Island, ton, Rhode Island Haupt, ‘Mrs, pa J., 117 W. bri St., Tampa 3, ast Hawkes, Mr. Alex D., P. 0. B , Coconut Grove 3 Healy, Mr. John E., meth 18.W. pears Hill Ra., sane ae: 97219 1 Heath, Mr. Russell, 808 5th Street, Juneau, Alaska Heath, Mrs. Victoria, 630 Highland Ave., Piedmont 11, Calif. Hedden, Mr. Orve K., 758 Secrest Road, Wooster, Ohio Hegwood, Miss Muriel P., Box 904, be. pore Carolina Calif. Heinlein, Mrs, Herman, 19395 §.W. 248th St., Homestead, Florida Heinzelman, Mr. oslo G., 267 Southern Ave., Rt. 2, Chillicothe, O. ennig, Mr. R. W., R.D. 1, Middle Grove, N. Y Henry, Dr. es ee Curator of Botany, Carnegie Museum, Pittsburgh 13, Pideavivaula 164 AMERICAN FERN JOURNAL ag Mr. Teofilo, Instituto de Biologia, Ciudad solar rtado postal 29817, Mexico 18, D. F., Mex 961 fecni: Mrs. Howard M., Westchester Geuntry ps Abe ak: 1966 Hevly, Mr. Richard, Rataty Dept., Univ. of Arizona, Tueson, Ariz. 1959 Hiatt, Mrs. Julian E., 1776 Ohio Avenue, Long h Cali va 1963 icks, Mrs. Myers H., 540 Azalea Lane, Florence, South Carolina 1964 Higley, Mrs. Clifford W., 8 Cherry St., Hudson Falls, ed ae 1963 Hill, Mrs. A. G., 26 Dean Rd., Apt. 2, Brookline, Mass. 1964 Hill, Mr. Julian ie og Greenhill Avenue, Wilmington 5, nie 1960 Hiltunen, Mr. Jarl K., P. O. Box 640, U. 8. Fish & Wildlife Service, Ann Arbor, era 956 Hindel, Rev. Richard, O.S.B., pacleth St. Meinrad, Indiana 1962 Hires, Miss Clara S., 152 Glen , Millburn, N 1941 Hodge, i Walter H., ae aad Peananton Washington 25, D, 1956 Hol nee Dr. L. R., P. O. Box 2908, San Jose, Costa Rica 1949 Holliger, Dr. Herbert H., 1949 Holttum, Mr. R. E., 80 Fei hock Kew Gardens, Richmond, Surrey, England 1957 Hood, Mrs. Vanes 1508 34th Street N. W., Washington 7, D. C. 1951 Hoshizaki, Mrs. Barbara Joe, 565 N. Westmoreland. Los Angeles 4, California 1953 iy aie ig Frances (Albert), 625 So. Baywood Ave., San Jose 38, Califo 1958 Howard, Me Benjamn C., Amberley, R.F.D. 2, Annapolis, Maryland 1961 Howard, Mrs, Freda C., 357 Lake George Ave., Ticonderoga, N. Y. 947 Howell, Mr. John Thomas, Calif, Academy of Sciences, Golden Gate , San Francisco 18, Calif 952 Hubert, ae Douglas E., 10415 Little John Ct. ean bcm 1964 Hudson, Mr. Ral x S., Route 2, Box 12 205, Nokounk Flor 1961 Huisking, Rev. P. J., St. boosie Mission House, Grover Ohio 1964 Hull, Mrs. Lewis M. (Helen S.), R.F.D. No. 1, Boonton, N. 1960 Hunnewell, Mr. Francis W. ae St., Wellesley a Mas 1915 Hutchings, Mr. David W., Botany Department, Cazenovia Serie Cazenovia, New York 961 Hutchison, Mr. Paul, Univ. of California Bot. Gardens, Berkeley 4, alifornia : 1956 Huttleston, Dr. Donald G. , Longwood Gardens, Kennett Square, Pa. 1956 Hutto, Mr. Thomas a: , Biology Dept., School of the Ozarks, Point Lookout, Missou 1958 Hynds, Mrs. Claude : R.D. 2, 1000 Acre Road, Delanson, N. Y. 1961 Isenberg, Mrs, Gerda, 19500 Skyline Blvd., Redwood City, California 1962 AMERICAN FERN SOCIETY 165 Ito, Dr. Hirosi, % pay oe be sliapaei Faculty of Science, Tokyo Uni ?Bdaent tion, Otsuka-Kubomachi, Bunkyo-ku, Tokyo, Japan 1961 hen. Kunio, ee of Botany, Kyoto Univ., Kyoto Japan ; 1958 Jacob, Mr. Henry J., 906 8S. Court St., Cleveland, Miss. 1962 James, Mrs. Phil, a 67th Ave., Miami, Florida 1956 Jarrett, Dr. Frane e Herbarium, Royal Botanie Gardens, ‘Vitenats ceca England 962 oti Dr. Otto Emery, Carnegie Museum, Pittsburgh, Pa. 1911 Jermy, Mr. A. Clive, Dept. of oe British Museum (Natural History) Cromwell Road, London, S.W. ms England 1960 Johnson, eee Harold M., Via Hood a al, Brinnon, Wash., 98320 1964 Johnston mae W., Jr., Georgetown University, 3900 Reservoir Rd. pitts ciel 7, UC, 1952 Salateis Will m S., 65 pee Lane, agen New York 1950 Jones, Prof. G. Sate — of Botany, Univ. of IL, Mapa Ul. 1945 Joshi, Miss Sumati A., 530 Budhwar Peth, Poona % India 1960 Juilly, Mr. Marcel E., a as Blithedale, Mill Valley, Cait 1957 Kaur, Dr. Surjit, National Botanic Gardens, Lucknow, India 1961 Kaye, Mr. Fred W., 1961 W. Verdugo Blvd., iontesee: Calif. 1941 Kenzie, Mrs. W. A., 495 Deloraine Ave., Toronto 12, Ont., Canada 1960 Kiefer, Mr P.O. Box 32 , Gardena, Calif. 1962 Kikendall, Mr. Robert, Rte. 2, Eaton Rapids, egrnmee 1963 Kilbride, Mrs. J. B., 2255 Woodward Way N.W., Atlanta , Georgia 1958 Killip, Mr. Ellsworth P., % H. 8. Dilts, 4306 Glenridge, pa alifor 16 Kirk, Mr. Gers eorge L., 8 E. Washington St., Rutland, Vermont 1961 Klebingat, Capt. Fred K., 873 22nd St., San Pedro, Calif. 1957 Kleinschmidt, Mr. alae F., 1800 N. Dixboro Rd., Ann Arbor, Mich. 1952 Klute, Miss Jeannette, 59 Farragut St., Rochester, New York 1963 Knable, Mr. John P., II, P.O. Box 476, Village Station, Warren, Mich, 48092 1941 oe Dr. Irving W. a Botany & Plant Path., Michigan State Univ., E. Lansing, Michig 1933 Knox, Mie William C., 649 Tale a Palms, Ft. Lauderdale, Fla. 1951 Koster, Mr. Hollis, 1508 Sansom St., Philadelphia 2, Pa. 1940 Krajina, Dr. Vladimir J., Dept. of Biology, Univ. of British Columbia, Vancouver 8, B. , Can ada. 1958 Kramer, Dr, K. Li Bot sch Museum en Herbarium, Lange Nieuwstraat 106, Picci. Netherlands 1955 Sages Mrs. Helen, eng 1961 » Mr. Gerhard O. eae Laboratory, University ae Arizona, — Kihaciis " 166 AMERICAN FERN JOURNAL Kruger, Mr. George A., 377 ayo Blvd., Harvey, Illinois Kujawski, Dr. Walter F., 407 Barclay Rd., Grosse Pointe 36, Mic Kunkel, Mr. Gunther, hateiedee” St. Mary Platt-Seven Oaks, meet E d gl Kunneke, Mr. J. W., 2307 Village Drive, Louisville, Ky. Kurata, Dr. Satoru, % Inst. of Forest Botany, Faculty of Agr., University of Tokyo, Hongo, Bunkyo-ku, Tokyo, Japan Laitsch, Mr. John T., Me 21, East Liverpool, Ohio Lakela, Dr. Olga, og of South Florida, Tampa, Florida Lampton, Dr. Rober one Dept. of Biology, West Georgia Bulees: jaaeed ie iso Langfelder, Mr. Richard, iy nase Road, Chappaqua, N. Y. Lassiter, Mrs. James A., 1703 Ferris Ave., Tampa 3, Florida Lawalrée, Dr. André, Av. Van Elderen 3, ‘Gradnels 16, Belgium Lawton, Dr. Elva, Botany Dept., Univ. of Wash, Seattle 5, Wash. Learned, Mrs. W. S., 404 Riverside Drive, New York 25, N. Y. Leatherman, Mrs. Sylvia B., 2637 North Lee Ave., South El Monte, Calif. 91733 Lee, Mr. Ed, 6427 Pinehaven Rd., Oakland 11, California Lee, Mr. George S., Jr., 17 Chicester Rd., New Canaan, Conn. k ‘ailinodi: Mr. D. B., U. S. Natl. Museum, Sesttlidonian Thatitution, Washin P xe Liggette, Mr. W. E., 700 Swarthmore Lane, Vaiveraty oe 24, Mo. Little, Dr. Elbert i. Jr., Forest eh U.S.D.A., Wash. 25, D. C. Lloyd, Mr. Cyrus, Tiles Mills, I need Mrs. Edith M., 47 Bond eae North, Hamilton, Ontario, nada Loyd, Mr. Seb M., Dept. of Botany, Univ. of Calif., Berkeley 4, aliforn Lockwood, ie Marjorie P., 44 Sutherland iF Pittsford, N. Y. Logan, Mr. Harry, 9 Binnitias e Lane, Westport, Conn - Robert C., Dept. of aoa Div of Nebraska, n 8, Nebr raska ware og Bayard 250 Ashbourne Road, Elkins Park, Philadelphia fice Dr. ‘Bosltarié, Casilla 5542, Santiago 6, Chile, S. A. gton, 0 Lengel, Patricia Ass Sotenies Dept., State Univ. College, 1300 Elmwood Buffalo 22, New York 1 AMERICAN FERN SOCIETY Lord, Mrs. Marie L., 101 Clay St., Burlington, Towa Love, Mr. John, 1501 St. Christopher, Columbia, Missouri Lovejoy, Mrs. Gordon W., 1415 East 19th St., Tulsa, Oklahoma Lovis, Dr. John Donald, Dept. of Botany, Univ. of Leeds, Leeds, 2, Yo pha United Kingdom, England hi Mrs. Frank E., Box 65, Harrison, Mai Lownes, Mr. ene E., 16 Barberry Hill, rae fea sae ean. ak thts s., ‘149 Buena Vista Terrace, San Francisco 17, Califor Luhr, Mrs. pees Box 371, Murphys, California Lynch, Mr. David, 3679 Highwood Drive, S.E., Washington 20, D. C. Lyon, Mrs. A. S., 11815 Juniette Street, Culver City, Calif MeAvoy, Dr, Blanche, 5476 Foley Road, Cincinnati 38, Ohi Maebride, Mrs. Philip D., Hollywood Farm, os rasan McCain, Miss Bea, Box vio Derry, N. H., McClure, Claire, 29 Ardmore Rd., es eh o California MeCool, Mrs. Wi McCoy, Mrs. Juanita S., dete: Dept., Salem College, Salem, W. Va. McCoy, Supt. Thomas N., Catlettsburg City Schools, Catlettsburs, Ky. McDowell, Mr. Gladstone W., 6 Oak Ridge Road, Ashevi wh, McGilliard, Miss Eleanor, Dept. of Biology, Univ. of te iiacties Chatta _ Tenn. McGregor, Dr. Ronald L., Dept., of Héteny: Univ. of Kansas, Lawre se an MacInnes, Mr. ae Rockefeller Institute, New York 21, N. Y. ch ennon hes Marie Walsh, Box 986, Morro Bay, Calif. Mandel, Mr. William H., 100 Bank Street, New York 1 Mangino, — pee W., RFD No. 4, Landview “ery ‘rentinatan; jay oe Manly, Mr. ne Jr., 7396 S. Willow Ave., Fresno 25, Cal Manly, Mrs. A. M., 1394 Shoreline Dr., Santa Barbara, ctor Mansfield, Prof. Wm., 371 Kenwood Ave., Delmar, New Manton, Dr. Irene, Botany Dept., The University, Leeds 2, "Yorkshire, ngla Marengo, paid Norman P., 123 Leverich Street, Hempstead, N. Y. Marengo, Mrs. Norman P., 123 Leverich Street, Hempstead, 1, FU Ohio Marqua, Zoe Rentz, 5602 Clifton Ave., Jac Kbonvilis, ent 32211 Martin Mrs. Grinnell, King George Farm, Sutton, ont Martiz Jesse T., 2622 Liberty St., North Bend, Ars 97459 168 AMERICAN FERN JOURNAL Martin, a Ray, 5612 Apache Road, Louisville, Ky. 1963 Masek, Mr. John, Apopka, Florida 1935 Mason, vie Esther E., 2523 geet Street, Louisville % Ky. 1955 Mason, Mrs. James H., Box 251A, Rt. 1, West Terre Haute, Indiana 19690 Mehra, Dr. P. N.., Head, Botany Siene Pasiah sais Ghasdirack 3, Indi a nggne Mr. William B., 4963 Western Hills Ave., Cincinnati, Ohio 5238 oe Mr. Thomas, 12070 S.W. 47th Ae aaa prin 33165 1961 or N. errell, Miss Elinor, 18 East 69th St., Mickel, Dr. John T., Dept. of Botany & obi Rae Towa se Univ., Ames, Iowa 1957 Middendov?, Miss Mary H., 3154 Winlow St., San Diego 5, Calif. 1958 Milan, Mrs. Doris, 19349 Mills Tower Road, vere 16, Rancho Cordova, ‘alifornia 1954 Miller, Betty Jo, 868 S. Sunkist Ave., West Covina, Calif. 1964 itehell, Mrs, Feit S., 1150 Yew St., San Mateo, Calif. 1961 Modler, Mr. Arthur R., 6666 Garber Road, Dayton 15, Ohio 1958 Mohlenbrock, Dr, on ie H., Dept. of Botany, Southern Illinois Univ., Carbondale, Illinois 958 Monahan, Mrs. Alfred H., 1315 Tower Ave., Raymond, Wash. 1957 Montgomery, _ James D. , Biology Dept., Upants College, East Orange, N. J. 1962 Moon, Mrs. Neil S. Pee = Rochester, N. Y. 14617 1955 Moore, Mr. Charles F., P. O, 8, Brevard, North ete 1957 Moore, Prof. Dwight M,, 506 oe. Ave., Fayetteville, Ark. 1935 Moore, Dr. Jewel, Box 915, Arkansas State College, Conway, Ark. 1947 Moore, Dr. John W. , Dept. of Botany, Univ. of Minnesota, Minneapolis 14, Minnesota Moran, Dr. Reid, P. O. Box 1390, San Diego 12, Calif. 1962 Morgan, Mrs. Weld, 10 Roxbury St., Woreester, 1943 Morgenroth, Mrs. Vera, Beech Tree Rd., Brookfield Gas a 1961 Morton, Mr. C. V. Agee cio Tnatitation Washington, D. C. 20560 1940 Mosely, Mr. Hal M., 6664 Av alon, Dallas 4, Texas 1960 Mosier. Mrs. Martha oe W. Washington St., Martinsville, Ind. 1961 Motley, Mrs. Jesse W., Rt. 2, Box 325, Albany, Oregon 1962 Moul, Dr. Edwin T. ee oi Botany, Rutgers Univ., New Brunswick, New Je 1945 Mulloy, Mr. ae Mrs. M. S., 90 Pierpont Rd. , Waterbury, Conn. 1968 Murillo, Miss Maria Teresa, Apartado Nacional 2848, Bogota, ‘olombia 1960 Murphy, Miss Mary, 50 City Hall, Boston, Mas 1963 Murray, Mr, Philip W., 162 Mar Monte Court, Saleh Salt. 1961 AMERICAN FERN SOCIETY Mutehler, oy reshian 71 W. 8th St., Bayonne, New Jersey Myers, Dr. R. Maur "Westen Hi St. College, Macomb, IIl. Nagel, Mrs. iin a bie 0 Evergreen Ave., Juneau, Alaska Namegata, Mr. Tomitaro, a Narita, Narita-shi, Chiba-ken, Japan Naylor, Mr. Richard M., 2835 Ash Street, Denver 7, Colorado Neidorf, Mr. Charles, 1580 a 22nd St., Brooklyn 10, N. Y. Nejdl, Mr. Frank E., 310 East 17th St., Lombard, Il. Senet Mr. Larry, 204 Hayden, E.Q., Ann Arbor, Michi shida, Mr. Makoto, Biol. Inst., Chiba eat: es yar eg Ja apar Norcross, me Arthur D., 244 Madison Ave., New York 16, N. Y. Norton, Mr. Leroy F., Box 123, Presque Tse, Maine An's Ts. 16,:C., Ohba, Mr. Hideaki, 157-6 chome, i Pd ss -ku, Tokyo, Japan Olivarez, Mrs. R., 3024 Castro Street, ig Francisco 12, Calif. Oliver, Mr. Charles, 74 Charter Road, t Acton, Mass. O Otto, Mrs. Virginia D., 145 Ridge Park Ave., Stamford, Conn. Ownbey, Dr. Gerald B., Dept. of Botany, Univ. of Minnesota, Minneapolis 14, Minn, Paine, Mr, George H., Jr., 446 Ludlow Highway, Ludlow, Kentucky C Patnaik, Mr. 8. N., Lecturer in Botany, G. M. College, Sambalpur, Orissa, India Patterson, Sister Mary Thomasine, Mount Mary College, Milwaukee, Wiscons Payne, Mr. Aailoos H., 3511 Henderson Rd., aa N. C. Peasley, Mr. Clifford L., 7324 N. Washburne, Portland, Oregon Peck, Mrs, Arthur N. 5 215 Hollywood Crossing, panaheaing Long Island, New York Penfield, Mr. Paul, Jr., 17 Bradford Road, Weston 93, Mass. Perry, Mr. David H., 276 Shorewood Drive, Webster, New York Peterson, Mr. Douglas E., 2306 E. Andrews, Fresno 3, Calif. Phelps, Dr. Orr a A., Wilton, N. Y. Phillips, Mrs. Anatin S., 440 Weymouth Drive, Wyckoff, New Jersey Saute a Walter S | Beat of Botany, Univ. of Arizona, Tucson, bake Rcensill Dr. Rodolfo, Ist. Bot. Hanbury dell’ Universita, Corso Dogali 1/C, Genova, Italy 170 AMERICAN FERN JOURNAL Piternick, Mrs. George, 9803 - 19th Ave., N.E., Seattle 15, Wash. 1958 Plaster, Mrs. J. N., 1875 Meadowbrook Dr aes. Salem 5, N. C. 1962 Plate, Miss Emma, 344 Highwood Ave., ae 1955 Poe, Mr: Alfred: Iy., Jr.; P.O. Box 253, aha ‘ac 1962 Poelt, Dr. J., Gotanivehe Staatssammlung, Menzingerstrasse 67, unchen 38, Germany 1956 Pahanin’ Si Nicholas, University of Ife, Ibadan Branch, Ibadan, Nig 950 Postma, Me George, 847 Courtney St., N.W., Grand sans 4, Mich. 1961 Potter, Miss bos 33 Charter Road, gcse Con 1963 Potter, Mr. L. Henry, West Rutland, mon 1957 Pray, Dr. Thomas R. Hci Biological ipo Univ. Southern Calif., Los Angeles 7, Cal 1963 Pringle, Mr. James ail Royal Botanie Gardens, Box 399, Hamilton Ontario, Canada 5 Pritchard, Mr. Arthur Herbert, Jr., 42 Quarry Rid., Waterford, Conn. 1964 roctor, Mr. George R., Science sitet. The Institute of Jamaica, i Kingston, Sitiaies, B.W. 1938 Proctor, Mr. Russell, 111 E. Main St., Louisville 2, Kentucky 1964 Radcliffe, Mrs. W. F., 636 S.E. 5th Place; Hialeah, Florida 1964 Rapp, Prof. William F., Jr., 430 Ivy Ave., Crete, Nebraska 1943 Ray, Dr. R. P., Head, Dept. of Botany, Pink University, Patna, tS ai oy 1957 Raymond, Mr. Mareel, imate: Botanical Garden, 4101 E. Sherbrooke St., Montreal, Can 1952 Reed, Dr. Clyde F., ne ‘Harford Rd., Baltimore 34, Md. 1950 Reed, Mrs Wisncis C., Llewell Pack West Orange, N. J 1958 Reed, Mr. Louis J., 78 Mayfair Drive, Pittsburgh 28, Pa, 1964 Reed, Ruth ea 3 Gorham, Canandaigua, New York 1962 eve, Mr. Wi , Dept. of B sa Carleton Univ., Colonel By Drive, ee 1 Ontario, Canad 1963 Reeves, Mr. Alfred E., 960 Kenwood pou Turlock, Calif. 1959 Reichstein, Prof. hs Institut fiir Organ. Chemie, 19 St. Johanns-Ring, Basel, Switzerland 1962 Reid, Mrs. Alex 260 Boulevard, Mountain Lakes, N. J. 1948 enton, Mrs. I. M., Route 1, Box 111, Snoqualmie, W halons on 1959 Reynolds, Mr. John T., 12 E. Scott St , Chicago 10, Ilin 1959 Rhodes, Mrs. Marion B., hor ast Valley, Conn 1954 Rickson, Mr. Fred R., Dept. Botan y, Univ. of Calif. re 4, Calif. 1961 Ries, Mr. Victor H, Pers Lineoln na , Columbus 1964 , Ohi Rock, Dr. Howard F. L., Dept. of ology: Vanderbilt Univ., Nashville 5, vei 1954 AMERICAN FERN SOCIETY Rodin, Dr. Robert J., Biological Sciences as Calif. State Polytechnic College, San Luis Obispo, Cal Rogers, Dr. Donald Philip, Dept. of poet an of Illinois, Urbana, Illinois Rogers, Mrs. J. R., 3107 Union St., Eureka, Calif. ei Dr; an ie i eat York Botanical Garden, Bronx Park, w York 58, ee a Lieut. sb oy M.S8.C., Demonstration Co. MFSS, Fort Sam Houston, Texa Root, Mr. Winthrop H., the Main St., paarowcy Mass. Ross, Mrs. C. Ervin, otis Gardens, Camden, Main Ross, Mr. Harry H., 1426 Milvia Street, rsp Calif. Js , Mas Salsman, Orpha L., 14016 - 8th Ave. South, Seattle, Washington Sanborn, Mr, Gierge K., Hidden ria vipat Mass. rh Saunders, Mrs. P. Blake, 58 Highla fe ve a. 22) ONG UX. Scamman, Miss see 474 Portland Rd., Saco, Main Schaefer, Mr. Robert L., Jr., Dept. of Biology, aA College, Allentown, Hie Scherer, Mr. Milton E., 830 N. Tech Rd., Sault Ste Marie, Mich. Schevker, Miss Julia, 3627 ae Hill Rd., Baltimore, og 21207 Schmidt, Miss Claudia, 39 Ely Ave., W. Springfield, Mas: Schrock, Mr. - uld F., Dept. of Science, Kutztown State College, Kutztow Schultz, Mr. ye B., Jr., Penryn sie Port Hope, Ontario, Canada Schultz, Mrs, Clara, B.D. 1, Stillwater, N. Schumacher, Dr. H. C. Albert, ease 8, (22e) Waldbroel, Rhid., ermany Schurr, Mrs. Ruth, R.R. No. 1, Portage, Ohio Scott, Mrs. J. Lewis, 208 Camberwell Dr., Bitebach, Pa. 15238 Scovell, Mr. Ronald, 3 Sims Crescent, Rexdale, Ontario, Canada Seudder, Miss Anna E., St. Margaret’s School, 565 Chase Parking, Waterbury 20, Con Seudder, Mr. James H., eis 52nd Ave., N., St. Petersburg, Florida 33703 Seully, Dr. Francis J., 904 Medical Arts Bldg., Hot Springs, Ark. ith net Helen, “pte Broadway, poor Texas ; Mr. Jen , 5922 Prospect Rd., Peoria 4 seg 5 Ko, % paces ‘Miaidipal Secadidas 2-chome S ditatee Nishi-ku Osaka, Japan tis AMERICAN FERN JOURNAL Seymour, Mr. George W., Keuka Park, Yates County, N. Y. harp, Dr. Aaron J., Dept. of Botany, Univ. of Tenn., Knoxville 16, Tennessee Sherman, Dr. Hope, Pleasant Valley, Conn. eae Mr. rib M., 1249 South Basin Lane, Sarasota, Florida hur L., Devt: of Biology, Southwestern State Coll., Pa ad, Okla. Sichler, Mrs. Mary C., 1118 Lakeview Blvd., E., Seattle “ Wash. tigre Mr. David, 8120 S. Jeffrey, Nae Til, 6 on, Mr. Andrew, Bluemount Nurseries, Box 219, sida Md. ye me Miss Mary L., Kissimmee, Florida eons Mrs. Maxine B., P. O. Box 562, Sweet Home, Oregon , Mr. James R., Dept. of Biology, Univ. of Puget Sound, Presi: Slater, Mr. William A., P.O. Box 448, Kerrville, Texas 78028 Smith, Dr. Albert C., University of Hawaii, Honolulu 14, Hawaii i OO; , Smith, Prof. Dale M., Dept. of Botany, Univ. o 1, Urbana, Ill. Smith, Miss Marion L., 429 South Willard te; age thts Vt Smysor, Miss Bettie, 3384 Erie Ave., Cincinnati, Ohio Snyder, Mr. Franklin D., 198 Valley Road, aoa ited a Sobol, Miss Eva, 1080 ei hneie Ave., Bronx 52, N. Sodomka, Mr. R. R., 7824 Normal Ave., ee aoa oe Solmitz, Mrs. Hentia. 2128 Veteran Ave., Los Angeles 25, Calif. Soper, Prof. James H., Dept. of Botany, Univ. of Toronto, Toronto 5, Ontario, Canada Spongberg, Mr. Stephen, 1514 E. State St., eet Til. Stauffer, Dr. R. Eliot, Kodak Park Works B 5 » Room 470, Stedman, Mr. Carling H., 8401 SW. 68th St., ‘Miami, Florida 33143 Steere, Dr. William C. Director, Nick. Botanical Garden, Bronx Park, New York 5 peak oe Steeves, Dr. Taylor A., “ieee Biology, Univ. of Saskatchewan, Saskatoon, Sask., Canada Steil, Dr. William N., 6624 Revere Ave., Milwaukee, Wis. 53213 Steinhauer, Mr, ithoodore, 1750 - 18th fet San Francisco, Calif. Stevens, Mr. Charles E., 615 Preston Place, Charlottesville, Va. Stevens, Mrs. Leslie B., 30 Marboy Dr., Plantsville, Conn. Stewart, Mr. Fred A., 8606 East Las Tu unas Dr., San Gabriel, Calif. euler a Robert W., 100 Pitt St., Apt. 12C, New York, N. Y. AMERICAN FERN SoOcIeTy Stewart, Dr. William me Director, Los sion State & County Arboretum, Box 688, Areadia, Califor Steyermark, Dr. a A., Apartado sia ee. Venezuela Stitt, Dr. Lynn B., 6414 Lupton, Dallas mye Pie Stockman, Mr. Charles C., II, 153 High Str dept tin vic Mass. E. P., P. O. Box 955, Angle on Tex. Stokes, Mrs. Francis J., The Mill, mite ia Stokey, Prof. Alma G., Dickinson House, Dept. of Plant Sciences, Mt fass. Stolz, Mr. Robert G., 8320 Coral Drive, Norridge, Ill. Storer, Dr. Robert W., Museum of Zoology, Univ. of Michigan, Ann Arbor, Michigan Stoudt, Dr. Harry N., Science Dept., Glassboro State College, lassboro, New eee Stoutamire, Dr. Warren - Se. Institute of Science, Bloomfield ills, Michigan Streissguth, Mrs. E. H., 101 W. Main, Monroe, Wash. 98272 Swan, Mrs. Joseph R., be Cohen te onn, Swearingen, Mr. Joh , R.D. 3, Box 573, Oklahoma City 27, Okla. Sweet, Mr. Merrill K es of Biology, Texas A. and M, Univ., College Station, Texas Swihart, Mr. Arthur W., 4837 Kendall, Corpus Christi, Texas 78415 Sybertz, Mrs. Carl, South Miller Rd., Rt. 1, Box 298, Valrico, Fla. Taback, Mrs. Mary Ellen, 151 Shoe Lane, Newport News, Virginia Tagawa, Dr. Motozi, Dept. of Botany, Univ. of Rehe Kyoto, Japan Talbot, Mrs. G. A., 7114 W. 35th St., Berwyn, Illinoi Talbot, Mr. Stephen Snow, 35 Shaw Road, Bridgeport, iy ard 02324 Tardieu-Blot, Madame, Museum d’Histoire Naturelle, 16 Rue Buffon, Paris V, France Taylor, Prof. T. M. C., Univ. of British Columbia, Vancouver, Canada Taylor, Mrs. Thomas §., P. 0. Box 1812, Bridgeport, Conn. Taylor, Mr. Wm. Carl, 8914 Francis Court, St. Louis 23, Missouri Teresita, Sister M. , Holy Family College, RFD 1, Manitowoc, Wisconsin d. 19 homas, Mrs. Edward O., 209 W. Federal Street, Snow Hill, M Thomas, Dr. John H., Dudley Herbarium, Stanford Univ., Stanford, alifornia 94305 Thurston, Mr, Edward D., Jr., Sharon, Conn. ierney, Mrs. George, 593 Buchanan St., Hillside 5, N. J. Timpson, Miss M. C., 15 Albro Lane, Cedarhurst, L. a N.Y: Tindale, Miss M., Natl, Herbarium, Botanie Gardens, New South Wales, Syildes, Australia t 174 AMERICAN FERN JOURNAL Tracy, Mr. H. Harwood, Apt. O, Claremont Gdn. Apts., 848 W. Bonita Ave., Claremont, California 91712 Tripp, Mr. Henry, 92-06 Jamaica Ave., Jamaica 21, N. Y. Tryon, Mrs. Alice F., Gray Herbarium, Harvard Univ., 22 Divinity Ave., Cambridge 38, Mass. Tryon, Dr. Rolla Milton, oe Gray Herbarium, Harvard University, 2 Divi vini — Ave., Cambridge 38, Mass. Turnau, Mr. und A., Dent. of Biology, Laurentian Univ., Sudbury, Gaus Canad Utter, Mrs. Howard H., R.R. + Stanley, New Yor Van de Water, Mrs. Blinore S., 264 Hillside Ave., Chatham, Ny; Vogdes, Mrs. Jesse T., Lemon St. No. of 5th St., ray 2d: Von Sneidern, Mrs. Karl, 23 Long Ave., Belmont ai Mas Voth, Dr. Paul D., Dept. of Botany, Univ. of Chicago, aes oF; aah Wagner, Dr. Warren H., Jr., Dept. of Botany, Univ. of Michigan, Ann Arbor, Michigan Walker, Mr. Albert N., 2310 Edgewater Rd., North Sacramento 15, California Walker, eg H. aie (Gladys D.), 4306 Newport Drive, Richmond, Virgin Walker, ties. 805 pesaie Square, Lynchburg, Virginia Walker, Dr. Stanley, Dept. of Genetics, Univ. of Liverpool, Liverpool 3, England Walker, Dr. Trevor G., Dept. of Botany, The University, Neweastle upon Tyne 1, England Warrick, Mr. Herbert H., 6543 - 26th Ave., NE. Beeston bademws 98115 Was, Margaret prabe 164 Prebele Dr., Tustin, Califor Weishaupt, Dr. Clara G., Botany Dept., Ohio State Dee: in Neil Avenue, aca 10, Ohio Weld, Mrs. John S., Box 197, Maine, New York (State), 13802 poet — David (Frieda L.), 16632 Eagle Ridge Dr., Hopkins, Mini West, Dr. gran Florida Agr. Exp. Sta., Univ. of Florida, Gainesville, Florida Wherry, Prof. sl T., Botany Dept., Univ. of Pennsylvania, Philadelphia 4, White, Mrs. D. P., sede W. 14th Ave., Vancouver 8, B. C., Canada Whitehead, Mr, Tick. P. 0. Box 123, Kingman, Arizona Whitehouse, Dr. Eula, 3033 Fondren Drive, Dallas 5, Texas Whitlock, Mrs. J. C., 834 N. Norman Lane, Wheeling, Illinois Whitney, Mrs. Elsie G., 104 Adams Place, Delmar 953 AMERICAN FERN SOCIETY Whittier, Dr. Dean P., Biol. Laboratories, Harvard University, Cambridg e a Mas ss. Wiggins, _ Ira L., Dudley Herbarium, Stanford University, Stanford, Calif. Wiley, Miss 3 A., American Museum i Natural History, 79th St. 19 and Central Park West, New York , 20 Pa. Willi, Mr. Edward i 125 E. 91st St., New York 28, N. Y. Williams, Mrs. E. O., 2601 Acacia Court, Fort Roe e ee gre Willier, Miss Lillian E., 320 Sheridan St., N.W., Washi C Willis, Mrs. W. 2g 130 Sherman Ave., Mansfield, Ohi Wilson, Nei, J. H., P. O. Box 88, South Fort Smith, A Wilson, Dr. Ses ae A., San Fernando Mare State ae 18111 Nordhoff Street, Northridge, Califor Wilesi, Mr. Robert I., 1249 E, Millbrae ree oe Calif. Windham, Mrs. L. B., Rt. 5, Box 110, Tyler Winnie, Dr. William T., 2517 Troy Road, Scents 9, New York Wiper, E., 6507 Maple Street, Vancouve nada Womer, Miss Irene, 6310 Somerset a verte aa 20840 Woolsey, Mr. David T., Route 1, Box 140, oe California Yopp, Mr. John, 3443 Afton Ave., ogee h, Ken h b to Zachau, Mrs. Marie, 603 BE. Iso St., Inglewood, California Zelley, Miss Grace S., 1014 ania Road, Burlington, N. J. 176 AMERICAN FERN JOURNAL SuaGEsTIONs TO CONTRIBUTORS TO THE AMERICAN FERN JOURNAL 1, Authors of papers submitted to the American Fern Journal should con- sult the Style Manual for Biological Journals (1960) and be guided by its examples, 2. The “name-and-year” system for bibliographic references will, except in exceptional eases, be used. (See Literature Cited below for an example). 3. Authors are encouraged to use the journal abbreviations set forth by Schwarten and Rickett (1958, 1961). 4. Abbreviations of names of herbaria will be those listed by Lanjouw and Stafleu (1964). 5. Reports of chromosome numbers will not be accepted unless documented. 6. The use of footnotes and tabular matter should be kept to a minimum. 7. All manuscripts submitted for publication should be typed, double- spaced, and have ample margins. 8. Reprints should be ordered when galley proof is returned to the Editor. An order blank will be included with the galley proof. LITERATURE CITED American Institute of Biological Sciences, Committee on Form and Style of the Conference of Biological Editors. 1960. Style Manual for Biologi- cal Jou oe rnals. American Institute of Biological Sciences. Washington, [ 100 p. Lanjouw, J., and F. A. Stariev. scion The Herbaria of the World. Index Herbartoram. Regnum Veg. 31: 1—251., ScHwarrTen, L., and H. W. Rickert, 1958. nga of Titles of Serials Cited by Botanists. Bull, Torrey Club 85: 277-300; 1961. Supplement I. Ibid. 88: 1-10, Vou. 54 OctTosER-DECEMBER, 1964 No. 4 American Fern Fournal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY ts IRA L. WIGGINS 0. V. MORTON ROLLA M. TRYON JOHN H. THOMAS ts CONTENTS Edwin Bingham Copeland (1873-1964) and we! ospaaeg to Pteridology H. Wacner, Jz. 177 Observations on Drought oe a in Selaginella densa Rydb. . WEBSTER AND TayLor A. STeEvES 189 Southwest Vacation Tomas DARLING, JE. 197 | Shorter Notes: Ferns in the Florists’ Trade in 1964__________-_ 206 Notes and News; Boston Fern Collection Finds a New Home; Dr. W. C. Drummond; Edgar T. Wherry Honored_________—_ 207 American Fern Society... ____—————_________ 08 Index to Volume 54 ee ae Missour, BOTANICAL FEB 8 4965 mak. Nin The American Fern Hocietp Council for 1964 OFFICERS FOR THE YEAR Donovan 8. CorrELL, Texas Research Foundation, Renner, Texas President Donatp G. Hurrteston, Longwood Gardens, Kennett Square, a Vice-President Lenerre B. Arxinson, 415 S. Pleasant Street, Amherst, sikanaabasitiis Secretary Bicuarp L, Havxs, — of Botany, University of _ hee. Bhode Islan Tea L. Wiganis, Dudley a Stanford University, Stanfor 1. Calif, ditor-in-Chief OFFICIAL ORGAN American Fern Journal EDITORS Ina L. Wicerrs___Dudley Herbarium, Stanford University, Stanford, © Calif. G, ¥. —_ Institution, Washington 25, D. 0. TrYon Gray Herbarium, Harvard University, Cambridge 38, Mass. Jou» H. THomas Dudley Herbarium, Stanford University, Stanford, Calif. An illustrated quarterly devoted to the general study of ferns, owned by Fern Society, and published at 3110 Elm Ave., Baltimore 11, t timore, Md. 0s Matter for publication should be addressed to Ira L. Wiggins, Dudley Herbarium, Stanford Univer sity, Stanford, California. Subscription $4.50, exclusive of agency eng cd fees; sent free to mem- bers of the American Fern Society (ann es, $4. sustaining mem- bership, $8.00; life membership, $80.00). gg reer reprints if ordered in advance, e furnish uthors at cost, plus for back pant and other -soorst communications should be to the Treasurer, Dr. Richard L. Hauke, Naaamet 4 of Botany, Universi of Rhode Island, Kingston, Rhode Island. Members and subscribers should allow two months for changes in address to take effect. LIBRARIAN AND CURATOR OF THE HERBARIUM . W. H. Wagner, University of Michigan, Ann Arbor, Mich. reed or express charges. The pages of the J ournal to members who wish to arrange ae Bog a membership list is published at intervals, to assist those interested in 0 ns American fern Fournal Vou. 54 OcTOBER-DECEMBER, 1964 No. 4 Edwin Bingham Copeland (1873-1964) and His Contributions to Pteridology W. H. Waener, JR.! The death of Professor Copeland in Chico, California, on March 16, 1964, marked the end of a distinguished botanical career not only of a leading pteridologist but an administrator, a teacher, a physiologist, and a tropical agriculturist. Of all o Dr. Copeland’s contributions, his work on the ferns is best known, and his last publication was fittingly titled simply “Fern” (1964). He was recognized by the American Fern Society by election as an Honorary Member in 1948. He was born on September 30, 1873, in Monroe, Wisconsin, and acquired his deep interest in botany from his mother, Alice Bingham Copeland. His father, a graduate of Cornell University, died when Edwin was only three. He grew up in Monroe, and when he reached college age he went to the University of Wiscon- sin in Madison, some thirty miles away. After his third year there he transferred to Stanford University where one of his purposes was to study under the fern authority, D. H. Campbell. When he graduated from Stanford in 1895, he left for Europe where he attended the University of Leipzig and the University of Halle, specializing in plant physiology. He completed his Doetor’s degree at Halle in 1896 with a thesis on the influence of light and temperature on turgor. After his return to the United States he held several positions in different American universities. In 1903, he left with his wife and young son for the Philippine Islands, where he remained for fourteen years. His first appointment in the Islands was as Sys- bert en Preparing this biographical obituary I acknowledge the aid of ee rt F. Copeland, Conrad V. Morton, and Florence S. Wagner especially. dees notes on the life of Copeland and a complete bibliography are pub- shed in Taxon oo 54, No. 3, of the JourNAL, pp. 105-176, was issued November 12, Ms gs 54, No. 4, of the JouRNAL, pp. 177-216, was issued January 15, 65, AMERICAN FERN JOURNAL Fig. 1. Epwin BincHam CopELAND AT THE AGE OF TWENTY-FIVE Epwin BInGHAM COPELAND 179 tematic Botanist in the Bureau of Science. In 1909 he founded the College of Agriculture of the University of the Philippines (for which he received a Doctor of Laws, honoris causa 40 years later). As Dean of the College he was deeply involved in aca- demic administration and one would judge that when he re- turned in 1917 to the United States he was relieved to be his “own man.” In fact, for nearly a decade after his return he lived as a leading rice grower in the area of Chico, Butte County, California, and effectively cut his connections with the academic life. By this time his family had grown to five children. Not until 1927 did Copeland return to botanical research and administration, first as Associate Curator of the University of California (Berkeley) for four years, and then as Technical Advisor and Director of the National Economic Garden in the Department of Agriculture of the Philippine Government for four years. In 1935 he retired and became a permanent Research Associate of the Department of Botany of the University of Cali- fornia, thus entering a period of pure research on ferns that lasted nearly three decades. It was during this period that most American botanists remember him. His appearance was striking. Although his height was average and his build slender, his eyes were sharp and penetrating and he always wore a mustache and goatee. The photograph (fig. 1) shows him as a young man of 25 when his hair was bright red. When I studied with him as a graduate student at Berkeley in the years 1946-50, his hair was pure white and he was bald on top. His bearing was dignified, alert, and authoritative. Dr. Copeland was a heavy smoker of cigarettes, and one of our constant worries was that he might set fire to the building and destroy it, herbarium and all. Actually the waste basket into which he casually tossed his cigarette butts was metal; and I have often wondered whether he did not start the billowing smoke just to keep things lively. He had a wry sense of humor, and it was a delight to hear him tell a story. He was also abso- lutely forthright and he let the chips fall where they might. I 180 AMERICAN FERN JOURNAL recall with blushing the time I asked him to criticize my first attempt at a botanical Latin description. He read it over rapidly, seratched his head, and said “Are you fooling?” No subject was too small or too large for his analysis and criti- eism. His lively, critical mind was always in action. Airplanes, new styles, politics, English grammar—all were objects of care- ful attention. Once Dr. and Mrs. Copeland decided to drive down to visit Dr. Campbell at Stanford University and invited me along. Dr. Copeland, 75 years old at the time, was driving along the freeway south of San Francisco at 80 miles an hour, simultaneously watching a helicopter soaring along to the east. Fascinated by the “whirlybird” he got so interested in it that he missed his turn ates and we ended up having to back- track about five m He always ae us by his ability to sit down at his desk and write up descriptions of species, give historical references and so on, almost entirely from memory. He had a remarkably reten- tive mind and put down his thoughts in a laconic style that was anything but wordy. His letters, likewise, were crisp and he wrote many of them to his colleagues across the world including, over many years, such authorities as A. H. G. Alston (England), Carl Christensen (Denmark), R. E. Holttum (Singapore and England), W. R. Maxon (U. S.), Conrad V. Morton (U. S8.), M. Tagawa (Japan), and Mme. Tardieu-Blot (France). It would be impossible to review thoroughly all of the pterido- logical contributions of Dr. Copeland. During the course of his long career he described 35 new genera and some 600 new species of ferns. His personal herbarium (now at the University of Michigan, Ann Arbor) totals approximately 25,000 specimens. The published results of his work on ferns appeared over years (1902-1964). In his bibliography the titles pertaining to notes, articles, reviews, monographs, and books on the ferns number nearly 120. The following comments will emphasize sev- eral of the high spots of his pteridological investigations, and the titles chosen for the literature cited will be only those which are pertinent. Epwin BingHAM COPELAND 181 One of Dr. Copeland’s most interesting (and most often over- looked) papers was one of his earliest on the ferns—The com- parative ecology of San Ramon Polypodiaceae (1907). His in- troductory statement in this article could have been written today, nearly six decades later: Contemporary biological science is working toward two ends, namely: (a) The reduction of the processes taking place in living things to understand reactions of chemistry and changes of phys- ics; this is Physiology. (b) Assuming the existence of living things, the causal interpretation of the forms they assume and the elucidation of their relationships; this is Bionomies, sometimes called “Evolution.” The application of principles established in the study of physiology to the problem of bionomies is called Ecology. At this point in his research career, Copeland’s interests were beginning to shift from “(a)” to “(b).” In the ten years prior to his San Ramon paper, practically all of his works were physio- logical, dealing with problems of tropisms, turgor, transpiration, and the mechanism of stomates. After 1907 nearly all of his writings were taxonomic, evolutionary, and ecological. In the San Ramon paper, 80 pages long, he investigated the origin and geographical affinities of the ferns of the area, the local environ- ment and adaptations to it, and gave a systematic application of the results. This paper is of importance not only to students of the ecology of ferns, but it represents Copeland’s first venture into the problem of fern evolution. We find here the germs of many of his ideas, some later amplified, others modified. His concepts at that time are summarized in an evolutionary diagram (pl. IV), which anticipated many of the ideas later expressed by F. O. Bower, Copeland himself, and a number of other workers. The purely floristic-taxonomic investigations of E. B. Cope- land dealt mainly with the Old World tropics — Philippines, Southern China, Borneo, Sumatra, New Caledonia, New Guinea, Fiji, Raratonga, Society Islands, Hawaii, and so on; and they comprised the bulk of his floristie publications. He did, however, 182 AMERICAN FERN JOURNAL concern himself with New World ferns to some extent, and worked up, for example, the collections of Ynes Mexia from Bra- zil and Mexico. Most important herbarium materials from the New World tropics during the period of Copeland’s research career were sent to the U. 8. National Museum for identification. Copeland was the authority for the area he knew best. He re- ceived many specimens for study from such botanists and mis- sionaries as Rev. Copland King (the materials collected in Papua); C. J. Brookes (Borneo); J. C. Moulton (northern Sarawak) ; M. Abbé U. Faurie and J. F. Rock (Hawaii) ; R. 8S. Williams, J. B. Steere, and D. L. Topping (Philippines) ; H. E. and 8. T. Parks (Polynesia) ; S. F. Kajewski (Santa Cruz Is- lands); L. J. Brass (New Guinea); A. C. Smith (Fiji) and many others. He always felt a special indebtedness to Mrs. Mary Strong Clemens, who sent him extensive collections from both Borneo and New Guinea. The major results of his floristic researches are embodied in several publications that appeared mostly during his later years. Among the most important are the fern floras of Fiji (1929c), Society Islands (1932b), New Guinea (1940, 1941a, 1947a, 1949a, 1949b, 1949¢, 1952b) and the Philippines (in three volumes— 1958, 1960a, 1960b). In bringing together his floristic knowledge he concluded that the ferns evolved in the Antarctic region and migrated from there into more northerly regions (1939a, 1939b). In view of the present evolutionary activity of the higher ferns he decided that “the period since the Miocene may be judged sufficient for the evolution of most of the existing species and a large part of the genera, and for their spread over any expanse of suitable land area the globe affords.” He also came to the opinion that fern evolution leading to the forms of today took place chiefly in the tropics and that ferns migrated therefrom into the temperate zones. He was impressed by the abundance and diversity of ferns in the warm regions of the earth and cited a single mountain (Kinabalu) in Borneo where the fern flora comprises 437 species. Epwin BInGHAM COPELAND 183 Much of Copeland’s revisional work was woven into his floristic publications, but he did produce several significant monographic studies, of which that on the Hymenophyllaceae is the best known (1933, 1987, 1938, 1947b). His procedure in presenting this monograph was unusual in that he treated first all of the species of the traditional genus Trichomanes, following this by a similar treatment of Hymenophyllum, and then revised all of the species generically and divided the family up into 34 genera. For thus splitting the two classical genera of the filmy-fern family, Cope- land received much criticism. In his defense he had already argued that those of his predecessors who had examined the family in detail equivalent to his own had come to the same con- clusion about the need for many genera. Furthermore, as he pointed out, many species do not readily fall into either of the two traditional genera. For this reason, and the fact that there are approximately six hundred species of filmy-ferns to be dealt with in an over-all taxonomy, he argued that so long as the genera were natural it would be more convenient to recognize as many as he did. Twelve of his genera are more or less isolated from the others and are monotypic; of the others, the number of species range from several to 100 each, averaging ca. 20 species each. Pteridologists are today divided on whether or not to ac- cept Copeland’s generic treatment. Even in so large a fern flora as that of Malaya at least one worker has found that it has not proved convenient to recognize so many genera. The general re- action seems to be to uphold the two traditional categories Trichomanes and Hymenophyllum but to recognize Copeland’s genera as subgenera. The other large generic monographs of E. B. Copeland include Leptochilus (1928), Plagiogyria (1929), Grammitis (1951), Xiphopteris (1952a), and Ctenopteris (1956). It is upon matters of broad evolution and classification of the Filicineae that Copeland’s influence has been and will be felt the most. His tutelage under Douglas Houghton Campbell, combined with his own intimate knowledge and experience with perhaps as 184 AMERICAN FERN JOURNAL many species of ferns as any one who has ever lived, placed him in a unique position to evaluate the classification of ferns. It must be remembered that the taxonomy of ferns was fairly “settled” by the thorough work of Diels and others in Die natiirlichen Pflanzenfamilien, and the valuable bibliographic source work of C. Christensen in Index Filicum. During the years approximately 1910 to 1930, the popular works of the Glasgow morphologist, F. O. Bower, called attention to the abundance of parallel and convergent evolution in the ferns. Nevertheless Bower himself still continued in the old tradition of emphasizing the sorus as the most fundamental criterion of fern relationships. (Copeland always felt that Bower’s work suffered from lack of knowledge of broad fern diversity.) Although we find Copeland as early as 1907 discussing the evolutionary relationships of ferns (as noted earlier), it was not really until 1929 in his Oriental Genera of Polypodiaceae that his basic ideas became well established. He expressed his operational procedure in this way: “Systematic Botany today is dominated by two principles: 1. Its groups of plants, of generic or of whatever rank, must be nat- ural in the evolutionary sense. 2. These groups should be con- venient—easy to recognize and to define.” He felt that “It is by its distinctive effort to interpret [evolutionary relationships] that systematic botany maintains the dignity of a science... ,” and he was concerned not only with mere classification but ales un- derstanding and insight. The “Oriental Genera” is of interest in other respects too. For example, he used here a neat numerical way of expressing divergent relationships in a linear series (an idea based upon methods used in family genealogies and library catalogues) with unit numbers, 1, 2, 3, ete., the first figures the main branches, the second figures the secondary branches, and so on—for example: 125 Athyrium, 12521 Blechnum, 12522 DOORS PSG eae a 1253 Asplenium, 125311 Ce- terach, 125312 Pleurosorus. With the encouragement of Ethel Faulkner Copeland, his wife, Dr. Copeland decided to summarize his conclusions regard- EpwiIn BINGHAM COPELAND 185 ing the taxonomic relationships of the ferns, an effort which culminated in the Genera Filicum (1947b). There, he put into practice his conclusion that the old ““Polypodiaceae” was a catch- all for many different lines of evolution. He divided it into a number of families, so that the total of filicinean families was 19 in all. He later added a twentieth, the Grammitidaceae (1951). This division of the Polypodiaceae sensu lato resulted in the recognition of such segregates as Pteridaceae, Davalliaceae, Blechnaceae, Aspleniaceae, and Vittariaceae. The idea of break- ing up “Polypodiaceae” had been anticipated seven years earlier by R. C. Ching (1940), who had recognized 33 segregates from what had been considered a single family (Copeland himself criticized this treatment in 1941). Since Copeland’s Genera Filicum appeared, evidence is accumulating that he was prob- ably too conservative if anything. Many workers today believe that his Pteridaceae and Polypodiaceae are too large and hetero- geneous to represent natural assemblages of ferns. Perhaps the major contribution of Genera Filicum was the active recognition that many soral types have arisen repeatedly in separate lines of fern evolution, and that the sorus is not so reliable an expression of relationships as had been thought, for example, by Diels, Christensen, and Bower. Such forms of sori as the “aerostichoid,’ ‘‘asplenioid,” “ovymnogrammeoid,” “polypodioid,” and “nteroid,” have appeared convergently in in- dependent groups. Such fern genera as Elaphoglossum (which he placed in Aspidiaceae), Dietyoxiphiwm (Aspidiaceae), Diellia (Aspleniaceae), and Prosaptia (Grammitidaceae ) show soral structures that are strikingly different from their relatives as he interpreted them in his revision. It was in keeping with his basic philosophy, however, that relationships should be deter- mined upon the totality of characters, whether or not a single feature or complex of features should be strongly divergent. The influence of Genera Filicum may be measured in part by the number of floras which adopted the same or a modified form of his classification (over a dozen since 1947), and in part by 186 AMERICAN FERN JOURNAL the extent to which it has stimulated and augmented research. In many respects Copeland’s treatment has been supported by new lines of evidence, as in the cytological findings of Irene Manton (1954, ef. especially Table 1, p. 176) and her followers. At about the same time that Gaora Filicum appeared, another system of classification was published, namely that of R. E. Holttum (1946, 1949) and their differing viewpoints made re- search on fern classification especially timely and stimulating. In the fifties several other systems have been proposed, of which those of Alston (1956) and Pichi-Sermolli (1958) have been especially important. Current thinking suggests some significant changes in the taxonomic concepts of Genera Filicwm. For example, there is evi- dence now that Copeland’s “Cyatheaceae” should include several of the genera that he included in his “Pteridaceae” (Holttum, 1963). The remainder of the family “Pteridaceae” should prob- ably be divided into two separate families (cf. Manton, op. cit.). There is a strong question whether Dipteris and Cheiropleuria should be included in the Polypodiaceae sensu stricto (cf. Wag- ner, 1952; Wilson, 1959). On the other hand, it is possible that the ieseuat of Copeland’s families, the “Aspidiaceae” (compris- ing nearly 70 genera and 3,000 species) may have to be enlarged even more to include “Davalliaceae,” “Blechnaceae,” and “As- pleniaceae” if it is to be natural. When one considers the vast labor involved in Copeland’s extensive work on the ferns, it seems hard to believe that he also had time to be a teacher, a tropical agriculturist, and an adminis- trator. Many botanists are not aware that he also succeeded in writing several books on non-pteridological subjects—‘Elements of Philippine Agriculture” (1908), “The Coconut” (three edi- tions, 1914, 1921, and 1931) and “Rice” (1924), and even a book on the principles of practical ethics entitled “Natural Conduct” (1928). Although he will probably be remembered most for his contributions to the knowledge of ferns, he was obviously a man of broad interests. Alert and in possession of his mental faculties Epwin BiInGHAM COPELAND 187 until the end of his long life, as his age approached 90 years he became blind and could no longer work. In what was surely one of the last letters Dr. Copeland received in those final months, his good friend C. V. Morton wrote to him as follows: “You have the consolation of knowing that your name is in constant use by fern students the world over.” LITERATURE CITED ALSTON, . = G. 1956. The subdivision of the Polypodiaceae. Taxon 5: Bower, F. a 1923- 28. The ferns, vols. I-III. University Press. Cambridge Cuine, R. C. 1940. On natural classification of the family “cpidiyppaianone. nyatsenia 5: 201-268. eae, 3 amie BineHam, 1907. The comparative a Meeen “8 San Ramon aka ee Philip. Your. Sci. 2 (C): 1-76, pls. 1 8. Leptochilus and genera confused with it. eg Jour. Sci. o 333-416, pls 1929a. The fue genus cs aiid Philip. Jour. Sei. 38: 07% 417, pls Thc leeeb - The oriental genera of Polypodiaceae. Univ. Calif. Publ. Bot. 16: 128. 1929¢. Pe rns of Fiji. Bishop Mus. Bull. 59: 1-105, pls. 1-5. 1932a. ee ferns collected by Ynes Mexia. Univ. Calif. Publ. Bot. 17: 1-8. 1932b. Perey a ‘of the Society Islands. Bishop Mus. Bull. 93: 1-86, pls. — 1933. beg Se Philip. Jour. Sei. 51: 119-280, pls. 1-61 pea es Hymenophyllum. Philip. Jour. Sei. 64: 1-188, pls. 1-89. 1938, Genera Hymenophyllacearum, Philip. Jour. Sci. 67: 1 110, pls. 1-11. ————— 1939a. Fern evolution in Antarctica. Philip. Jour. Sci. 70: am 39b. Antarctica as the source of existing ferns. Proc. Sixth ea Sci. Congr. 4: 625-627 1940. Oleandrid ferns ictal of New Guinea. Philip. Jour. Sei. 73: 345-357, pls. 1-1 1941a, ayer ayia of oie Guinea. Philip. Jour. Sci. 75: 347-361, mg SB tom ermer us oun gna American ferns. Univ, Calif. Publ. Bot. 19: 287-339, pls. 36-66. 188 AMERICAN FERN JOURNAL 1941e. Gah on natural classification of the family Poly- 3 - podiaceae by R. C. Ching. Sunyatsenia 6: 159-177 ———— 1947a. Cyathea in New Guinea. Philip. Jour. Sei. 77: 95-125, pls. 1-165, —1947b. Genera filicum, the genera of ferns, Chronica Bot., Wal- tham, Mass. 247 pp., pls. 1-10 9a. Pteridaceae of New Guinea. Philip. Jour. Sci. 78: 5-41, pls. 1-6, —————— 1949b. ml ae sige Blechnaceae of New Guinea. Philip. Jour. Sei. 78: pls. -———_——— 1949¢. (ones, of Se Guinea. Philip. Jour. Sei. 78: 389- 475, pls. 1-44 -———— 1951. Geaninitin Philip. Jour. Sci. 80: 93-271, pls. 1-6. 1952a. The oe species of Xiphopteris. Am. Fern Jour. 42: 41-52; 93-1 — 1952b. Grammitiaea of New Guinea. Philip. Jour. Sei. 81: 81-119, pls. 1956. ni Ale ed in America. Philip. Jour. Sci. 84: 381-473, pls. 1-16 1958. Fern Flora of the Philippines. Vol. 1. Pp. 1-191. Bur. of Printing, Mani 1960a. Ski. oe of the Philippines. Vol. 2. Pp. 193-376. Bur. of Printing, Manila. 1960b. Fern Flora of the Philippines. Vol. 3. Pp. 377-557. ~~ of Printing, Manila ————— 1964. Fern. “Feageas Britannica 9: 191-195, pls. 1, Holttum, R. E. 1946. A revised classification of the Iepasportniie ferns. aes one Soe. (Bot.) 53 (no. 350): . The FOR of ferns. Biol. a 24: 267-296. — AND . SEN. 1961. gis and classification of the tree ferns. Phytomorphology 11 (4): 406-420. MANTON, IRENE AND W. A. SLEDGE. 1954. Observations on the eytology and taxonomy of the pteridophyte flora of Ce nh hee Trans. Roy. Soc. London, ser. B (Biol.) 238: (no. 654): -185. PIcHI-SERMOLLI, Ropotro E. G. 1958. The higher taxa a the Pteridophytes and their classification. Acta Univ. Upsaliensis 1958 (6): 70-90. Waener, W. H., Jr. 1952. Types of foliar dichotomy in living ferns. Am. Jou Bot! 39: 578-592. ON, itclenasia A. 1959. Sporangia of the fern genera allied with Poly- podium and Vittaria. Contr. Gray Herb. 187: 97-127 DEPARTMENT or Botany AND HERBARIUM, UNIversItY OF Micuigan, ANN Arpor, Micu. WILS DrRouGHT RESISTANCE 189 Observations on Drought Resistance in Selaginella densa Rydb. Terry R. WepstTeR and TayLor A. STEEVES InTRopucTION.—During a recent study of Selaginella densa Rydb., the Prairie Club Moss, it was noted that this plant has a remarkable ability to survive conditions of extreme drought. The purpose of this paper is to record some observations made on a small mat of S. densa in the laboratory, and to make some remarks about drought resistance in this species. According to Maximov (1935), drought resistance can be defined as “the capacity of plants to endure drought and to recover readily after permanent wilting, with the minimum of damage to the plant itself.” In the xerophytic species of Selaginella there is no outward appearance of permanent wilting. Probably the the best evidence of reaching the permanent wilting point in these species is the tight folding of the leaves next to the stem (Daubenmire, 1959). He further stated that it is important to distinguish plants which can really endure drought (that is, plants which are drought resistant) from those which merely escape or evade drought (e.g. desert ephemerals). Drought resistance has been described for several mosses and liverworts as well as for Selaginella. Campbell (1930) noted that among the California species of the Marchantiaceae, many dry up during the summer and are revived with the coming of rains in autumn. In these Species, Fimbriaria californica, Targionia hypophylla, and Cryp- tomitrium tenerum, to cite a few examples, the growing point and neighboring tissues of the thallus survive. Richards (1932) described examples of drought resistance among mosses. Leaves of Tortella inclinata were air-dried for eighty weeks at room temperature and survived, and those of Grimmia pulvinata survived sixty weeks in a desiccator at 20 degrees centigrade. Stems and buds of these species were found to be even more resistant to drying. Shuster (1955) reported a case in which material of the liverwort Riccia atromarginata from Texas had been revived after one year and three months of drying. Growth 190 AMERICAN FERN JOURNAL was evident five days after the material had been moistened, and the plants continued to grow in a greenhouse for nearly a year. Certain species of Selaginella also exhibit drought resistance, the most familiar example being S. lepidophylla, commonly ealled the Resurrection Plant. Leclerc du Sablon (1889) noted the capacity of this species to withstand prolonged periods of desiccation, the plant remaining in a tight rolled-up ball in the dry state. After the application of water, it unrolls, the branches take on a green color, and growth resumes. Uphof (1920) con- eluded that about six per cent of the then-known 580 species of Selaginella are xerophytes. He discussed the anatomical features of this small group of species as well as the possible mechanism by which the cells of these plants protect their protoplasm during drought. Finally, Tyron (1955) reported that part of a mat of S. densa var. densa that had been stored for six months as an herbarium specimen grew after being planted and watered. Hagit anp Econoay.—S. densa is considered to be an ally of S. rupestris, resembling this species very closely. However, 8. densa is usually more western in its range, occurring from southwestern Manitoba to southern Alaska, and south to Texas, Arizona, and northern California (Tryon, 1955). It grows in the form of dense mats, with short, decumbent branches. The shoot possesses a helical phyllotaxy. The branching pattern is terminal and unequal, resulting in one shank being slightly smaller than the other. Four-angled cones or strobili arise at the ends of the branches. These grow in an upright position and may reach a length exceeding one centimeter. The cones contain both microsporangia and megasporangia, the sporangial arrangement fitting Pattern I in the scheme set forth by Horner and Arnott (1963). In this pattern, there is a basal zone of megasporangia above which is a zone of microsporangia. On the Saskatchewan prairie, S. densa must be considered an important part of the vegetation, covering up to twenty-five DrouGHt RESISTANCE 191 per cent of the soil surface in some areas (Coupland, 1950). This species grows in relatively open places where it is sub- jected to the extremes of a severe continental climate. During much of the year the Saskatchewan prairie gets little precipita- tion (Table 1). During prolonged dry periods the mats are composed of dormant shoots having a dull grayish-green color and non-living shoots which exhibit an ash-gray appearance. In the dormant state the leaves, each with a long white awn on the tip, are closely appressed to the stem (fig. 1). It has been suggested by Uphof (1920) that these awns serve to reflect sunlight away from the plant as well as to protect the growing point of the stem. . In the field it was observed that the dormant aspect described above changes literally overnight if any amount of moisture becomes available to the plant. In the field after a rainfall, the grayish-green color gives way to a more luxuriant dark green, giving the previously dormant mats the appearance of active growth. On the other hand, the ash-gray color of the dead stems noted above does not change. In the active shoots, the leaves are no longer closely appressed to the stem, but lie at an angle to the axis. The appearance of a healthy and actively growing mat of S. densa is seen in fig. 2. Table 1, Precipitation in Saskatoon, Saskatchewan, Canada (60 year average). Data courtesy of Physics Department, University 0 Saskatchewan Inches Inehes Jan 0.56 July 2.36 Feb 0.53 Aug 1.73 Mar 0.59 Sept 1.38 Apr 0.74 Oct 0.78 May 1.37 Nov 0.55 CARE ee PE Dee 0.51 192 AMERICAN FERN JOURNAL EXPERIMENTAL OBSERVATIONS.—In September, 1960, a sod containing a mat of S. densa was brought into the laboratory for observation and study. A piece of this clump that was not { is, S 7 $ 4 4g UU HOLEEAERDAGOADUANDUORAGUUNL eo os FIGURE 1. DorMANT MAT OF 8. DEN :\. SCALE IN CENTIMETERS. FIGURE 2. THE SAME M AT 2 MONTHS AND 27 DAYS AFTER INITIAL WATERING. ARROWS INDICATE CONES. SCALE IN ( uN TIMETERKS. DrouGHT RESISTANCE 193 | used was placed on a shelf where it lay without moisture for two years. Several times during this period small pieces of the clump were removed and watered. In a few hours the dormant shoots took on the aspect exhibited by actively growing shoots. Having observed this interesting phenomenon, it was ecided to test the ability of the remaining mat of dormant S. densa to survive a more prolonged period of dessication. On June 17, 1963, about two years and nine months from the date of original collection in 1960, the remaining clump of 8. densa was photographed in its dry state (fig. 1). It was then placed in a shallow dish of tap water. After two hours a change was noticeable. In some of the shoots the leaves no longer clasped the stem, but had spread and turned green, exhibiting the revived condition. The next observations were made nineteen hours after the application of water, and it are currently shedding their spores, while others exhibit spo- rangia at different states of maturation. At certain times during this project the clump has become dry, and a return to the dormant condition has resulted. However, with the addition of water, the mat of 8S. densa has always revived quickly with no apparent damage to the plant—further verification of the drought resistance of this species. Discussion.—Tryon (1955) pointed out that, in general, the allies of S. rupestris are restricted in their distribution to open habitats, which are frequently xeric. He further stated that they are able to persist in such areas because of their 194 AMERICAN FERN JOURNAL ability to survive desiccation. In his opinion, this ability lies in the physical and chemical properties of the contents of the cells. Such an explanation is supported by field observations on S. densa. In one particular site near Saskatoon, Saskatche- wan, abundant sporelings at all stages of development were encountered. Over fifty locations of sporelings (or in some cases small groups of sporelings growing together) were marked with stakes, and periodic checks have been made over a period of one year and three months. It is clear from the observations made so _ far, that these sporelings, no matter how small (fig. 3), Can survive the long periods of droght. Their fragile con- struction suggests that the ability to resist drought lies in some physiological mechanism of the cells themselves. Such a mechanism has not as yet been explained for xero- phytic Selaginellas, but certain information suggesting possible mechanisms should be mentioned. Leclere du Sablon (1889) early suggested that there is a reserve substance in the cells of FIGURE 3, SPORELINGS or § DENSA, X 4.5. DrouGHT RESISTANCE 195 S. lepidophylla which he thought might be responsible for the resistance of this plant to desiccation. He compared the appear- ance of the dense opaque protoplasm of the cells to what one sees in cells of cotyledons or in the endosperm cells of certain seeds, In a paper dealing mainly with anatomical features of xerophytie species of Selaginella, Uphof (1920) concluded that the main question arising from his study is that of the protec- tion and behavior of the protoplast of the cells under condi- tions of extreme drought. In cutting sections for microscopic examination, Uphof noticed many large oval-shaped droplets, which were found to be oil, coming from the open cells. During plasmolysis of the cell, the oil droplets fuse to produce larger ones. Uphof suggested that when the cells lose water, a film of oil is formed around the vital parts of the cell, preparing the eells for dormancy during periods of drought. He noted that neither mesophytic species (e.g. S. galeottw) nor xerophytie Selaginellas grown under conditions of abundant moisture produce oil. Evans (1958, 1959) found that in certain fila- mentous algae which are able to survive drought, the vegetative cells become modified with accumulations of oil. Other changes that he noted for these cells are thickened walls and mucilagi- nous sheaths. In a recent work by Ijin (1957), possible mechanisms by which cells in general resist desiccation were noted. Iljin re- ported that, according to numerous workers, species of plants living in dry habitats have smaller cells than plants of moist habitats. Mosses, lichens, algae and other lower plants adapted to dry locations were cited as having cells of small volume. Uphof (1920) noted this feature for the xylem elements and cortical cells of xerophytie species of Selaginella. Iljin sug- gested that smaller cells suffer less damage under conditions of desiccation. When a cell loses water from the vacuole, the opposite walls are caused to approach one another so that the desiccated cells may become separated. In larger cells, there 196 AMERICAN FERN JOURNAL would be a greater readjustment of the walls to each other and more possibility of damage than in smaller cells. The authors wish to express their appreciation to Dr. M. V.S8. Raju and Mr. Norman Ferrier for aid in photography. They also wish to express their indebtedness to Mr. Ted Cooper for assistance in laboratory and field work and to Dr. William C. Steere for his helpful suggestions. This investigation was sup- ported by a grant from the National Research Council of Canada. LITERATURE CITED CAMPBELL, D. H. 1930. The slap and Development of Mosses and Ferns. MacMillan Co. New Y CouPLaND, R. T. 1950. Ecology of ca Prairie in Canada. Ecol. Monog. 20: 271-315. DAUBENMIRE, R. F. 1959. Plants and Environment. (2nd Edition). John Wiley and Sons, Inc. New York. Evans, BS i 1958. The Survival of Freshwater Algae during Dry Periods. I. An investigation of the Algae of Five Small Ponds. Jour. Eco 46: 149-167. ———_—., 1959 - The Survival of Freshwater Algae during Dry Periods. II. Drying ee pcciicate! Ibid. 47: 55-71, Horner, H. T., Jz. and H. J. Arn 1963. Sporangial geting in North Paros Wes Species of Selaginella, Bot. Gaz. 124: Ingin, W. S. 1957. D ught Resi stance in Seay ris ‘Hou . The aes in Relation to Water. (2nd Edition). nglish Translation. Allen and Unwin Ltd. London fetus, P. W. 1932. Ecology. In Verdoorn, Fr. _ (Editor) Manual of Bryology. Martinus Nijhoff. The Hague. pp. —395. Scuuster, R. M 55. Boreal Hepaticae, a AA fey < the api ore of Minnesota sind Adjacent Regions. II, Eco ology. Amer. Midl. : 03-299, TRYON, R. M. a Sela Gard. 42: UpnHor, J. G. ash 1990, hare ee ey of eens Selagin- ellas. New Phytol, 19: 101~ ginella rupestris and Its Allies. Ann. Mo. Bot. DEPARTMENT oF Biowoey, UNIVERSITY or SAsKATCHEW AN, SASKATOON, Sask ATCHEWAN, SouTHWEST VACATION 197 Southwest Vacation THOMAS DaRLING, JR. In November 1961, after six consecutive years of fern explora- tion in Florida, I decided to visit the Southwest, concentrating on Arizona, with a brief visit to New Mexico Preliminary investigation of the possibilities of finding in- teresting ferns at this season of the year proved discouraging. A letter from Dr. Walter S. Phillips, Head of the Department of shea University of Arizona, had this to say: fk e your se P igyaatets: sans hunting in free This is a prob- lem theat is very difficult to answer. I am sure that if I went out with you for two weeks we mie: be able to locate some of plants in which you are rested, but it would be a long and rather arduous trip vers entails considerable walking through rough re and some cliff elim “Ferns in Arizona are not nearly as easy to find as they are in your eastern area. There are several difficult prc about this time of year for a trip like you are planning “Tf I were to advise you on coming out here to see ferns, I certainly would not recommend this time of year. After a wet summer, the last of August or the first of September is a good time, or better still after a wet spring. Even our desert ferns are hard to find most of the time, as they curl up and dry, “One of the best guides that I know of is Leslie N. Goodding who lives in St. David, Arizona. He is a retired botanist and an excellent fern man. He loves to go on trips and I think would be glad to guide you. When he was in my office the other day, however, he said, ‘This is not the time of year to see ferns!’ ” On November 4th I flew to Phoenix, Arizona, and the next day telephoned Mr. Goodding that I would go to southern Ari- zona fern hunting if he considered it worth while. He said that if I wished to make a trip to Sycamore Canyon near the Mexican border, he would act as guide. Furthermore, he believed that the ferns would be in good condition following recent rains. I boarded a south-bound bus, spent Sunday night in Tucson, and early Monday met Mr. Goodding at the University of Ari- zona. We used his ear, and reached our destination before noon. Sy camore Canyon lies only a few miles north of the border, its 198 AMERICAN FERN JOURNAL southern reaches extending into Mexico. The Arizona section of the canyon is within the Coronado National Forest. I was impressed by the wild beauty of Sycamore Canyon with its color- ful pinnacles of eroded rock. Mr. Goodding, a man in his early eighties, led me a merry chase up the steep gorges of side can- yons and scrambled along rock walls wherever it was possible to obtain a foothold. An ardent naturalist and conservationist, my companion had written two articles about this area (1946, 1961). During my visit he pointed out many rare plants, shrubs and trees, some of which grew almost exclusively in this canyon, such as the Alamo Lotus and the Goodding Ash. As for the main object of my search, ferns of the Southwest, in a few brief hours we came upon more than a dozen species. With the single exception of the beautiful Venus’-hair Fern, Adiantum capillus-veneris, all were new to me. The Adiantum was seen to best advantage in a cave-like overhang along the trail, carpeting the entire roof and sides with its delicate growth. Probably the most conspicuous and striking of all was the rela- tively rare Phanerophlebia auriculata, sometimes called the Trail- ing Fern, locally plentiful on cool damp cliffs. The genus Cheilanthes (Lip Fern) was represented by C. wootoni, C. len- digera, C. lindheimeri, and C. eatonii forma castanea. Pellaca longimucronata, the sharp-pointed cliff brake so common in the Southwest, was abundant. Less common was Bommeria hispida, with its generally triangular outline and pubescent lower sur- face. Only young sterile fronds were in evidence. I was especially interested in trying to locate two rare and in- teresting species of Asplenium, A. exiguum and A. palmeri, both known from Sycamore Canyon. The former is one of the rarest ferns in the United States, found in but one other locality in Arizona and in no other state. It was described from specimens collected in the Himalaya Mountains, and also has been found in China and Mexico. Its curious geographical distribution remains unexplained, Although Mr. Goodding had originally discovered SouTHWEST VACATION 199 A, exiguum in this locality, he was unable to relocate it during our visit. Exploring a side canyon, however, I was delighted to find A. palmeri in the rock crevices high up the cliffs. At first I thought this little plant was only the common A. resiliens, but closer inspection revealed the recurved fronds and proliferous tips that distinguish A. palmeri. At the time of my visit Mr. Goodding was greatly concerned about the future of Syeamore Canyon. Overgrazing has resulted in heavy runoff during rainy periods, accompanied by soil ero- sion and damage to some of the rarest plants. Due to the efforts of Mr. Goodding and others, the Forest Service has recently de- clared Sycamore Canyon a scenic area. The objective of this classification is to maintain the canyon as nearly as possible in an undisturbed condition, but allowing use of the area. Approach roads, trails, picnic grounds and parking areas will be located so as not to disturb the natural environment, but at the same time provide for some public use. It is hoped that in the future Syca- more Canyon will be treated in many respects like one of our National Parks. Collection of plants, animals and minerals will be strictly prohibited. Back in Tucson later that week I arranged to go on a fern ex- pedition in the Santa Catalina Mountains with Dr. Walter S. Phillips. In a single day, comprising two separate trips, he showed me a surprising number of ferns typical of the South- west. Snow already covered the upper slopes of Mt. Lemmon, closing the trail to the area where Asplenium septentrionale had been found, making search for this curiosity out of the question. In the morning we concentrated our attention on a small can- yon just north of Tucson in the foothills of the Santa Catalinas. Here the rather common eray-green Cheilanthes lindheimert was found in the same neighborhood with the rarer and delicately graceful C. wrightii. It was especially interesting to see Notho- laena standleyi and Pityrogramma triangularis growing side by Side in this locality. Although to a layman these ferns may super- ficially resemble one another (the fronds ef each showing a 200 AMERICAN FERN JOURNAL white or yellow powdery effect on the under surface), the Gold Fern is more triangular in appearance and the Notholaena pen- tagonal or star-shaped. Here also two fern allies were seen grow- ing together, carpeting the nearby rocks, Selaginella rupincola and S. arizonica. The entire area was filled with various species of cacti, and as I backed away from an interesting specimen to take a photograph, thereby making posterior contact with an- other prickly plant, Dr. Phillips gave me one of the primary rules of the desert: “Never back up without looking around £1" After lunch we set forth on the second phase of our fern hunt- ing expedition, a visit to the Molino Basin Area in the Santa Catalina Mountains. En route we skirted the Saguaro National Monument, famous for its forest of giant cactus Carnegiea gigantea, limited in distribution to southern Arizona, in north- ern Mexico, and a few plants in California. At Molino Basin, in addition to ferns which I had seen pre- viously elsewhere, Notholaena was well represented. It was an experience to find NV. aurea (dull green above, with golden under- surface), NV. grayi, and N. sinuata, frequently intertwined with a species of sharp-spined yucca. Cheilanthes covillei and mature fronds of Bommeria hispida were locally plentiful. In Phoenix I rented a Comet “compact” which I drove some 1600 miles on an eleven-day trip via the Painted Desert, Petri- fied Forest, Grand Canyon, Bryce and Zion National Parks in southern Utah, to Las Vegas, Nevada. On this part of the trip my main objective was sightseeing, but some unusual botanical adventures still remained. Having missed the opportunity to see Asplenium septentrio- nale near Tucson, I decided to search for this oddity along the Mogollon Rim southeast of White Oak Canyon where the rock spleenwort was reported to be locally plentiful. Early on No- vember 10th, with not a cloud in the sky and relatively warm weather for that time of year, I left Cottonwood, Arizona, for the Rim Road. Beyond Camp Verde the going became difficult. With SouTHWEST VACATION 201 a steep winding mountain road looming ahead, it was mid-after- noon before I reached my destination. At Long Valley I noticed the first evidence of snow, but at the approach to the Mogollon Rim Road I realized that the trip was probably a wild goose chase. Motorists were returning from the area with chains and pessimistic reports. “The road is impassable,” they told me. “The snow is several feet deep!” Determined to botanize at least the nearest likely area, I drove eastward a short distance, skid- ding frequently en route, to Baker Butte at an elevation of over 8,000 feet. Leaving my car just off the highway, | climbed up a snow-covered road to the fire tower from which I obtained a fabulous view over the Tonto Basin and the surrounding low- lands. On the rocks nearby was the cliff brake, Pellaca ternifolia, var. wrightiana. Search for Aspleniwm septentrionale proving unsuccessful, I returned to the car and slithered back to Long Valley. After a visit to the ghost town of Jerome, where copper for- tunes were once made and lost, then on to Sedona with its color- ful buttes and movie-like atmosphere, I drove northward up Oak Creek Canyon to Flagstaff. Immediately on arrival, Veterans Day, I called Chester Deaver of the Botany Department at Ari- zona State College, hoping to enlist his aid in discovering the extremely rare Asplenium adiantum-nigrum, a European species found in the United States only from northeastern Arizona, northeastern Colorado and southern Utah. It has been found in very limited occurrence on the south face of Elden Mountain, four miles northeast of Flagstaff. Mr. Deaver told me that he had never seen this rare fern on Mt. Elden, but would be glad to join me in a search for it the following afternoon. Sunday morning dawned clear and cold. Planning an advance expedition on my own, I arose early and journeyed to the base of Mt. Elden, feeling confident of finding A. adiantum-mgrum after reading Dr. Wherry’s account of its occurrence in this lo- cality (1941). Having arrived at the general area, I realized that it covered a 202 AMERICAN FERN JOURNAL far greater area than I had bargained for. It was like looking for a needle in a haystack. Selecting the easternmost approach, I picked my way precariously up the mountainside. Halfway up the steep cliff, with the sun at my back, I detected a cave-like formation with dark shadowy floor. Since ferns frequently grow in such a locality, I ventured close to the cave’s entrance, reach- ing up to touch the roof with the tips of my fingers. Suddenly a sixth sense made me freeze in my tracks. Immediately ahead I detected a sheer drop-off! Lying flat and peering over the edge of the abyss, I looked into an apparently bottomless chasm. My enthusiasm for an independent search ended abruptly ! That afternoon, with Mr. Deaver as guide, I again ventured up Mt. Elden. This time we chose the western side of the south face, threading our way laboriously up the steep rocky wall. Cheilanthes feet was in evidence on exposed ledges along the route. Near the summit we reached a moist shady cul-de-sac where high vertical cliffs prevented further progress. Here the presence of the Male Fern (Dryopteris filix-mas) and the Maid- enhair Spleenwort (Asplenium trichomanes) indicated (accord- ing to Dr. Wherry’s notes) that Asplenium adiantum-nigrum might be in the near vicinity. Since none was to be seen, how- ever, and since we had reached an impasse, we reluctantly had to give up the search. Our visit to Elden Mountain was none too soon, as the region was blanketed with heavy snow the fol- lowing day. Journeying eastward, I visited the Petrified Forest and Paint- ed Desert in a young blizzard, making the loop circuit and spending two nights at Winslow, Arizona, en route. Tuesday evening I reached the Grand Canyon and planned to make the descent to the bottom the following day. Bright and early the next morning I was driven to Yaki Point, the beginning of the extremely steep but spectacular South Kaibab Trail. I was anxious to make this trip on foot, rather than by mule train, so that I would be free to botanize and take pictures independently en route. Some six hours, eight miles and SouTHWEST VACATION 203 about one hundred color photos later, after crossing the suspen- sion bridge, I reached Phantom Ranch on the north side of the Colorado River. Although the desert plants had been most in- triguing, there had been little of pteridological interest along the trail. After spending the night at the ranch, I left early the next morning on the steep eleven-mile climb back to the South Rim via the Bright Angel Trail. Near the intersection of the latter with the South Kaibab Trail I spotted a small patch of Cheilanthes parryi on a rocky ledge, one of the few ferns seen during my expedition into the Grand Canyon. Shortly before dark I reached the South Rim just after a severe snowstorm set in which lasted all night and into the next morning. Several days later, after visiting Glen Canyon and Bryce Canyon National Park, I spent a few hours at Zion National Park in southern Utah. Although I realized that this was one of the places where Asplenium adiantum-migrum had been found many years ago, I realized that my chances of locating it during a quick visit were practically nil. Dr. Wherry wrote (1941) :— “Then it [A. Adiantum-nigrum] was found in Zion Canyon, Utah; but on visiting that National Park in 1940 I was unable to find anyone who knew on which of the myriad cliffs it oceurs.” I made the usual photographic tour of Zion Park, then fol- lowed the self-guiding nature trail to Canyon Overlook. At one point I left the main trial to explore an interesting side canyon. Looking up at the high cliffs above, I detected an evergreen fern that looked from a distance suspiciously like Asplenium adian- tum-nigrum, but it was too far out of reach and an attempt to venture closer from above ended in frustration. Unable to erase the possibility from my mind, however, I stopped at headquar- ters before leaving and gave details of the location to the young naturalist on duty. Although just an off-chance that this fern might actually prove to be the extremely rare A. adiantum- nigrum, I suggested that a more thorough search might well be justified, and that in any event I would appreciate receiving @ 204 AMERICAN FERN JOURNAL specimen for identification. Early in February 1962, after I had nearly forgotten the inci- dent, I received a letter from Carl J epson, Chief Park Naturalist. He explained that bad weather and numerous duties had pre- vented earlier search, but went on to say that he believed they had located the side canyon I mentioned. He continued, “En- closed are some pieces of a frond knocked from a fern growing about twenty feet above the floor of the canyon at its head. -Ac- cording to the keys in Kearney and Peebles’ Arizona Flora, ‘this fern appears to be the one in question, i.e., Asplenium adiantum- nigrum. A brief search was made for more of this fern, but none was found. We hope that later investigations will establish its habitats and occurrence more accurately. We certainly do ap- preciate your bringing this fern to our attention, for it was one that we had not seen before. We would like very much for you to confirm the identification.” I lost no time showing the fragments, fully mature specimens, to C. V. Morton at the National Herbarium, Smithsonian Insti- tution, who immediately agreed that the plant was indeed Asplenium adiantum-nigrum. He further said that judging by the size of the fragments the original plant must be unusually large for the United States, although it frequently attains luxuri- ant stature in Europe. was anxious to see, but which so far had eluded me—Asplenium septentrionale, the curious little Forked Spleenwort which I had missed at Mt. Lemmon and Baker Butte in Arizona. A study of herbarium specimens at the University of New Mexico at Albuquerque indicated that A. septentrionale had been found in the vicinity of Las Vegas, New Mexico. Informed that Dr. R. G. Lindeborg of New Mexico Highlands University in Las Vegas knew the location of this rarity, I called him long dis- tance and arranged a field trip. The next morning I drove north SouTHWEST VACATION 205 to Santa Fe and reached Las Vegas about noon. Dr. Lindeborg and I set forth in the mid-afternoon, heading for a box canyon nine miles southwest of Las Vegas. The trip involved rather a long hike from the point where we left the car to our ultimate destination, an extensive area of flat rock out- crops containing occasional fissures. In one particular crevice, my guide informed me, grew a small patch of Aspleniwm septen- trionale, but it could easily be overlooked since the other crevices were barren. We spent several hours searching to no avail, and as darkness approached Dr. Lindeborg had some doubts whether our quest would be successful. Soon afterwards, however, he called out that he had discovered the elusive ledge, and there in a erevice below the flat rock’surface was a small but luxuriant growth of A. septentrionale. Not far distant Cheilanthes eatonti was fairly abundant. On the return trip to the car we en- countered a driving sleet storm which stingingly pelted our faces. That evening I drove back to Albuquerque over treacherous roads, After spending Thanksgiving with relatives on a ranch in Quemado, New Mexico, I returned to Albuquerque, turned in my rented car and flew back to Washington after a most enjoyable vacation in the Southwest, which included some memorable fern discoveries, LITERATURE CITED Gooppine, Lestie N. 1946, A hidden botanical garden. Jour. N. Y. Bot. Gard. 47: 86-96. . 1961. Why Sycamore Canyon in Santa Cruz County should be preserved as a nature sanctuary or natural area. Jour. Ariz. Acad. Sci. 6 3-115. Wuerry, Epgar T. 1941. Asplenium adiantum-nigrum in Arizona. Amer. Fern Jour. 31: 97-100. 3809 Larno Drive, ALEXANDRIA, VA. 206 AMERICAN FERN JOURNAL Shorter Notes FERNS IN THE FLorists’ TRADE IN 1964.—Some months of getting my exercise by hiking among the canyons of Manhattan had indicated that ferns were much scarcer than formerly as florists’ merchandise. Most commonly seen were the ersatz types made from plastic which ranged from small fronds having some characteristics of polypody and Christmas fern eutting to a larger, more divided pattern. Two lines of enquiry recently took me to the wholesale florists area: What kinds of living fern plants were available? What was the status of the cut fern leaf trade? Of living plants, I found only five forms of V ephrolepis. None of these was accurately named. Two were of special interest. One consisted of thrifty stiff-leaved plants of the wild sword fern, N. exaltata. The dealer spoke disparagingly of these as “Florida plants, imported for the chain store trade.” The other was the cultivar, “Scottii,” forty years ago one of the most widely grown of the Boston Fern series but missing from the collection recently sent to the University of Michigan. In retail shops I have seen at least one other cultivar in the Boston Fern series and have learned that retailers may receive plants directly from growers and wholesalers in the metropolitan area. My enquiries led me to the Kervan Company on West 28th Street. There I had the good fortune to find as general manager, Mr. Harry Hyatt, whom I had last seen as a biology student of mine in Stuyvesant High School over 42 years previously. He had begun work in the Kervan store as errand boy after gradu- ation from high school. Four kinds of fern are represented in the cutleaf trade today. The most abundant is the “fancy fern” Dryopteris intermedia which is collected in Vermont. While question has been raised from time to time whether the extensive trade in this fern might be endangering its existence, the fear has never seemed justified and the present status of collection and demand remove even the slightest risk. In recent years, Mr. Hyatt assured me that Notes AND NEWS 207 the use of this fern had declined to no more than one tenth of the former volume. Demand has also decreased for leaves of the Pacific coast “sword fern,” Polystichum munitum, which, like the faney fern is a cold storage item. The other two kinds are picked and used fresh. The most ex- pensive is the splendid “California Woodwardia” or “Giant Chain Fern,” Woodwardia fimbriata, with leaves over four feet long and one and one-half feet wide. It can be ordered on a given day and received in New York the next by air freight. The fourth is the leather fern, Polystichum (Rumohra) adiantiforme, native in the West Indies and in southern Pacific areas. Several Florida dealers raise it, and the wiry, shiny, three-pinnate leaves, offer no serious problem in storage-—RaLPH 8. Benepict, Pilot Knob, New York. Notes and News Boston Fern Connection Frxps 4 New Home.—In March 1964, the collection of Boston Fern varieties and related types which had been assembled at the Brooklyn Botanic Garden, be- ginning in January 1914, was shipped air freight to the Bo- tanical Garden at the University of Michigan, Ann Arbor. It will be available for research studies under the direction of W. H. Wagner, Jr., Professor of Botany and Curator of the Fern Herbarium. The collection was started at the Brooklyn Botanie Garden in connection with preparation of a revision of the section on ferns for edition two of L. H. Bailey’s Cyclopaedia of Horticulture. Beginning about 1895, commercial growers had introduced doz- ens of new varieties obtained as bud mutations from the Boston Fern. There was no adequate literature covering their character- istics or their derivation. In 1914 a period began when visits were made to practically every U. 8. grower who had introduced a new variety, and many variants of the Boston Fern were ob- tained and housed at the Brooklyn Botanical Garden. From a few square feet of bench space, the collection expanded to sev- eral sections in the greenhouse, with outside lath-shaded space 208 AMERICAN FERN JOURNAL used during some summers. Eventually the collection included cultivars and wild species of Nephrolepis from botanical gardens and commercial growers in Europe as well as in the U. S., to- gether with a number that originated at the Brooklyn Botanic Garden. The collection shipped to the University of Michigan consists of forty types which comprise all but a few of the most signifi- cant forms obtained during the years of study. The missing variants will be sought and sent along later, if found. Acknowledgements have been made in various publications for the aid which made possible the assembling and study of the Nephrolepis collection through the years since 1914. These in- cluded particularly Dr. C. Stuart Gager, Director of the Brook- lyn Botanic Garden during his incumbency. Grateful acknowl- edgement is made now to the present Director, Dr. George 5. Avery, and to his staff for continued maintenance of the collec- tion and for the final work in preparing and shipping it to the University of Michigan—Ra.pu C. Benepict, Pilot Knob, New ork, Dr. W. C. DkumMonp, a member of the American Fern Society since 1956, died recently. An alumnus of St. Louis University, Dr. Drummond practiced dentistry in East St. Louis, Illinois. After his retirement 31 years ago he moved to Los Angeles, Cali- fornia, where he became interested in horticulture, and espe- cially in ferns. He was one of the founding members of the Los Angeles Fern Society and its President for a number of years. Under his leadership the Society prospered and is now the largest local fern group in the United States. Dr. Drummond assisted in setting up the Fern Dell in Griffith Park, Los Angeles, particularly in preparing the fern exhibit in the Fern Dell Nature Museum. This educational display shows 140 fern specimens that were collected, pressed, and identified by Dr. Drummond and mounted in 5 by 10 foot permanent cases. For this work he was honored in 1958 by Los Angeles civic lead- ers. Dr Drummond’s fern library consisting of over 250 books, AMERICAN FERN SOCIETY 209 many of them rare, has been presented by his widow, Mary Hazel Drummond, to the University of California (Los Angeles) Bio- medical Library, where it will be available for consultation by interested persons. Proressor Epcar T. WHERRY was honored on September 10, 1964, his 79th birthday, at a reception in The Academy of Nat- ural Sciences, Philadelphia, when Cranbrook Institute of Science, Bloomfield Hills, Michigan, bestowed upon him the Mary Soper Pope Award. This award was established to recognize persons who have made distinguished contributions in the field of plant science. His was the 10th award. The ceremonies were attended by about 150 persons gathered chiefly from the Philadelphia area, but with representatives from Massachusetts, North Carolina, and Michigan. Professor Warren H. Wagner, Jr., was chairman of the Mary Soper Pope Award Committee. Other members were Dr. William Campbell Steere and Dr. Warren P. Stoutamire. American Fern Society Fern Foray The fern foray began at 9 A.M. Friday, August 21, when 24 members assembled in front of Kittredge Hall, the dormitory at the University of Colorado, Boulder, where most were rooming. After driving barely out of Boulder to the beginning of the Mesa Trail, we walked up the trail to the Flatirons where we did a considerable amount of rock-hopping. We came to see Aplenium septentrionale which erew sparsely in the rock crevices. We also Saw a beautiful colony of the Male Fern as well as other species. This area belongs to the City of Boulder which is attempting to preserve it in its natural state. Returning to the cars, we drove back to the campus where we broke for lunch. In the afternoon we drove eight miles northeast of Boulder to White Rocks, an impressive formation ab si Where the horizontal surfaces are marked off in an unexplained 210 AMERICAN FERN JOURNAL turtleback design of geometric shapes resembling the plates on a turtle shell. This area belongs to Mr. and Mrs. Bruce Weiser who are preserving it and who were kind enough to allow our visit. Here we saw Asplenium adiantum-nigrum, Cheilanthes feet, and Equisetum hyemale, and a number of interesting flow- ering plants. We were also interested in seeing the ubiquitous pigeon or Rock Dove nesting on the cliff ledges as it formerly did in its native Europe. Too bad most of them have abandoned their natural way of life. Returning to Boulder we drove a few miles up Boulder Can- yon, stopping along the way to see two Selaginellas, S. wnder- woodui and S. weatherbyana. We turned off on a dirt switch-back road which climbed the wall of the canyon and onto Falstaff Mountain where we saw Cheilanthes fendleri and other ferns. This first day’s foray was interestingly and ably guided by Dr. William Weber. On Saturday we assembled at 7:30 and drove to Bear Lake in Rocky Mountain National Park where we were met by Mrs. Ruth Ashton Nelson and Mr. Gilbert Culter who led us on the wonder- ful climb past Bear Lake, Nymph Lake, and Dream Lake to over 10,000 feet elevation. Mrs. Nelson was familiar with the plants of the area, having written books on them, and Mr. Culter, the Park naturalist, furnished us with much general information on the park. During the walk, fifteen species of ferns and fern allies were seen, only five of which duplicated those seen the first day. Notable among these were Isoctes bolanderi, a new record, growing in Nymph Lake, Lycopodium annotinum, Botrychiwm lunaria, Dryopteris disjuncta, another new record, and Athyriwm alpestre which was observed at Dream Lake and higher. After a late lunch in a park pienic area, the group broke up with some driving up Trail Ridge Road mainly for the view, though they did see Cystopteris fragilis growing in crevices at 12,150 feet. Thirty persons participated in all or part of the foray, and this was felt to be a good number in view of the location and the AMERICAN FERN SOCIETY 211 fact that several regulars were attending the Botanical Congress in Scotland. Many were surprised that, in such an arid region, as many as twenty-four species, subspecies and varieties of ferns and fern allies were seen during the two days. Members participating in the foray were Mr. Robert Aborn (Penna.), Dr. and Mrs. William H. Baker (Idaho), Dr. and Mrs. Ralph H. Benedict (Illinois), Dr. and Mrs. Donovan 8. Correll, Charles and Selena Correll (Texas), Mr. and Mrs. Lawrence Dillon (Texas), Mr. Frederick Dunlap (Missouri), Mr. David L. Emory (Penna.), Dr. E. 8. Ford (Florida), Dr. and Mrs. L. K. Henry (Penna.), Dr. Walter H. Hodge (Maryland), Dr. Donald G. Huttleston (Penna.), Mr. and Mrs. James Leach (New York), Mr. and Mrs. Blake Saunders (Colorado), Dr. and Mrs. Ira L. Wiggins (California), Mr. and Mrs. Walter W. Willis (Ohio), Mr. Robert Willis (Connecticut), Dr. William A. Weber, Mrs. Ruth Nelson and Mr. Gilbert Culter, our leaders, from Colorado. amerie Ipine Ladyfern), Athyrium filix-femina (Ladyfern), Bo- trychium lanceolatum nee-leaf Grapefern), Botrychium lunaria v lunaria (Moonwort), Botrychium lunaria var. minganen trychium tricariifolium esperium (Matrieary Grapefern), Cheilanthes feei (Slender Lipfern), Cheilanthes fendleri (Fendler Lipfern), Cryptogramma acrostichoides Fern), Cystopteris fragilis (Fragile Bladderfern ) Dryopteris dilat (Mountain Woo ), Dryopteris disjuncta (Oak fern), Dryopteris filix-mas (Male fern), Pteridium aquilinum var. pubes: ce racken Fern), sia mexi xican Woodsia), Woodsia moss), Selaginella densa (Compact Spikemoss), Selaginella underwoodii (Underwood Spikemoss), Selaginella weatherbyana (Weatherby Spikemoss). D. G. Hurrieston, Longwood Gardens, Kennett Square, Penn. REGISTRATION For A. I. B. S. Meetinas.—Some members of the American Fern Society, who were not individual members of the A. I. B.S., were displeased when A. I. B. 8. personnel at Boulder refused to accept their applications for housing unless they also paid the registration fee ($15.00 if application was made prior to August 1, $17.50 if made later) to attend the entire series of 212 AMERICAN FERN JOURNAL meetings. The Council of the Fern Society considered the mat- ter at its regular Annual Meeting, August 23 The Council agreed that the registration fee seems excessive for one who plans to attend only the Fern Foray and/or the ses- sions of the Fern Society. Therefore, the policy of the Council with regard to attendance and payment of registration fees will be that it recommends compliance with the A. I. B. S. scheme when practical. But it believes non-members of A. I. B. S. may arrange for their own housing in a motel, hotel, or private home in the vicinity of the annual meetings, and may attend the Foray and/or the formal sessions of the Fern Society without register- ing for the whole series of meetings. Owing to an editorial error the Honorary Members of the American Fern Society were not listed in Vol. 54, No. 3 of the Journal. They are given here, with the Editor’s apologies! Honorary Members Benedict, Dr. R. C., Pilot Knob, N. Y. (First joined) poe Looser, Dr, Haars pers 5542, Santiago, Chile Manton, Prof. Irene, Department of Botany, The University, Leeds, England 1927 Stokey, Prof. Alma G., Emeritus, Dickinson House, South Hadley, Mass. 1949 Wherry, Dr. Edgar r T., Botany sp aa University of Pennsylvania, Philadelphia 4, 1918 It was impossible to Soa last minute changes in addresses after the list of members had b set in type. Consequently, kno errors occur. The Editor will ‘Ghee having any and all errors pone in this list brought to his attention. A note or post eard will be sufficient, and corrections will be made as early as possible—I. LL. 213 Index to Volume 54 Geer phere: 68, yah abie s, 69 bip » 68; haalilio- 9; 69; ancien aay 69; Pedal egy fy Adiantum att (4 veneris, 77, 198; his- pidulum, 129, 135, i ale diadchila 1, 3-5; aquilina, 6; aspera, 4; australis, 4; ee, 6 An American Spee es of Stegnogramma, 41 ieph radenium, 68; gaudichaudii, 68 Aspidrum lindheimeri, 146; sclerophyl- Aspen Ste peremstbaa 5 9 innati , 26, 32; platyneuron, 26, 32, 33, 42, 149, f. platyneuron, oe os serratum, 32; pumilum, 14 anthrisci- foli tek resiliens, “26, 33, 199; rut 42, brunfelsii 124; a iniie ‘199, 200, 204, ne 209, 211; serratum, 137; trichomanes, 26, 42, 81, 202; trudellii, 152; vir ride, 42, 90, 124" Asplenium xX ebenoides R. R. Scott in Kentucky, 149 Athyrium, 99; alpestre, 118, 210, var. americanum, 211; angustum, 90; net ea 26, 83, 42, S115 var. michau 83, var. sitchense, 79; pyenoca a 26, 338, 42; thelypteri- oides, 26, 33, 42 iactis mexicana, 26, 32 saat Soc G. d R. White. Reten of Viabitity in Lyophilized Spores of Fiddlehead Fern, Matteuccia ohee 1 aiivanive: ones pene Experiences in Raising Ferns fro 152 B ‘ ‘om Spores, enedict, ry C. Boston Fern Collection nds a ome, 207; Ferns in the Florists’ Trade 1964, bo pin Mat rial used in Research Morphogenesis and Pastopeviedian ( seal 151; Observations on e ensi ® Fern: Supplement, 93 f BT: "geltis ag 5 ‘ “ey 57; heter ar 57 r vtiverat ihe a 14; presliana, 9, 57; semi- rag i is 18, 57, 60, var. semicor- 9, var. incisa, 13-16, 18, tie aie renata, 9, 14, 57, 60; sub- sim 57; virens, mf 14, 57 Bomingets “hispida 198, shee Pome Fern Collection oFinds a New Rokr moae dissectum, 31, 42, var. dis- 31, var. obliquum, 26, 31, 0. var. ‘hesperium, meant multifidum, 42: simplex, 91; irginianum, 26, 31, 42, Brainea insignis, 100 British Pteridological Society, 99 Maou vordgon® rhizophyllus, 26, 34, 42, 147, f. boycei, 147, 148 bali ‘shizophy llus forma boycei Wilson, 147 Campyloneurum, 13 36,1 137; angustifolium, 138; costatu 138, 139; latum, i 8-140 i phstitdis, 136-138 Ceterach officinaru akan 7 nies Species of ma boycei ©. Boeck y sertbynid "ot Polyp odi pet vir m form pracienternal (Rid- lo . rnald, 126 chelgnthes cowie. cle 0 ne acre 205, 198; i 0 bey ~ 118, 122 3-24: wootonii, ii, 19 Correll, Donovan S. An from t esident, 155 Cc 145; heterophylla, Cryptogramma ac — a 211; crispa var. ponica, stelleri, 42, 90 Ctenopteris, 70, 183 Cyathea, 1-8, 48; alternans, 7; arbor 6; ‘as ssimilis, 8; celebica, 4; crassipes, 8; subgen . Cyathea, 2-5; ‘sect. Cya- b 145 ‘118, 122, 125; ps, 8; P 4; pse' i Le bry gg PR creel : sect. Schizocaena, 3, 5; subsect. a ; - 3. 5 e, a n sumatrana 5; tripinnata, “4s Biches mium falen ‘fe mn 36. oa. 42, 43, 90 Cc tise rosin bulbifera, ' texas 26, 34, 42, 90, 210, 211, 8, 214 var. sebbransesl 34, var. mackayi, 34, var. protrusa, 34 Danaea, 5 Darling, ies Jr. tion Death Takes Noted Pteridologist, 99 Dennstaedtia punctilobula, 35, 42, 80 Development of the St vet ar Cylinder in og df apie of Bolbitis and Egenol- Bickess 5 Diohipoxiphtum, og a be: Yadb eins: 125 Diellia a, 185 Southwest Vaca- rummond, W. (Obit.), 208 Drymoglossum, Po Dryopteris, 37, 42, 105; arguta, 106; austria var. intermedia, 26, 35, var. spinulosa, , 385; augescen var. Orga 146; campyloptera, 42, 48, 90, 109, X intermedia, 107- 109; clintoniana, 42, 106, 109, 110, 4 Seager oy 106, 10 7 11( 211, marinas lindheimeri, “145; harperi, yeh Pilosa, 141; polypodioides, 142; es, 143; sclerophyila, 144; spinulosa, 42, 106, 10 Edwin Bingham Copeland ‘iene flint and his Contributions to Pterid ology. The Effect Pag Sucrose on Apogamy in yrtomium falcatum Presl, 20 rs 17, 57-59, 61; appendiculata, 60; anpeniftia, 57; helferiana, 77; eattacatad: 41, Bthnobotanical” hag California niga phytes b stern American in Raising Ferns 152 Fern Foray (1964), Fern Aad to the Mieotehabie duck Area i Vermat (1963), 39 from AMERICAN FERN JOURNAL Fern Material Used Morphogenesis (Rev 151 ph ort Associated with solcoaheasey Rocks the Pacific Northwest, Poien. and Fern Allies on icnanaia of Pine Hills in Research on and Photoperiodism ee Strap Ferns and Their Cu iture, The br aa aie (sensu lato) in Ma- a, a, 32: linearis, 125 Goniophlebium, 144; brasiliense, 144; triseriale, 144 Goniopteris, 144; eclenep iin Oite 144 Grammitis, 70, 18 Growing Berry : der Fern, 43 Growing Mother Ferns, 94 r re ryopteris, 42, 90; Gpukerrans pilges, 141, var. major, 142 5 Gymnosphaera, 2 Hawkes, Alex Florida Strap and Their Cuture, te Litbrochi in Florida, 38 A Not glossum crinitum, Bi Hemitelia, 1, 5-7; capensis, 6; decipiens, Holttum, R. The (sensu lato} in Resa ih Fee aa fessor A bys Pls ” Study of Fern Hittiteion. Yih d é. For (1964), 209: Fern ani to aha fessrg tock-Dorset Area of Vermont Reid acaes A. Asplen xX eben oides R, eel Scott in piel 149 Hymenodium, Hymeno er a 139, 183 Isoetes, 37, 95, 96; bolanderi, 210, 211; is tle ri, 29; echinospora, 42; melano- 29 Iwate "Kunio. An So pal ag Species of gnogramma, Pie gona Neville. "Phytageoranhy of Selaginella douglasii, Kaur, Surjit. cote of the Stelar Cylinder in the Rhizome of Bolbitis and Egenolfia, 57 Knotioch, Irving W. Some Hints for the Fern Culturist, 73 Kruckeberg, Arthur R. Ferns Associated with ultramafic Rocks in the Pacific Nort Fae ais s8e Lepti a Bobi pilosa, 145; ensis, 145 eee! chia a in Plo rida, age Sig oe gy rygie "38, Robert Ethnobotanical Uses of Nutiieas Tevaianearee y Western American Indians, 76 pilosa INDEX TO VOLUME 54 cabeay ote 37, 71, 83; annotinum, 42, 91, 210 211; arcturi, 72; arthurii, 72; bonae-voluntatis MoT capellae, 72; ichinntes 72; pen tela, i eleon, 72; clavatum, 42 76, 91; complanatum, 29 ; costaricense, ae -australis, cuatrecasasii, 72; denticulatum, 85: ie, 12; = ster i fee maaiee re” 42; ulixis, 73; ohlberedtii Marengo, Norm P. Sporogenesis in Pter ope bree es ipacial Refer- one o the Cytoplasmic Inclusions, Marsilea pe 32 Matteucia p msylvanica, 87, 88 3 struthiop- 90 eterophylla, 145 orium heterophyilnm, 145 Mihiokeot Robert H. and Jane Hin- ners Engh, yh aa Fern Allies of Pine Hills Ps eat and Environs C. V. New Combinations in Lycopodium, 71 Nayar, B. K. and Prakash Chandr ‘bee Ww ene and Variety of Belbitis i Nephr odium m olle, 129 Hephsolent pis, 208, exaltata, 206 7 Ww ombinations in Lycopodium New Species of Pyrrosia oo india, A New aotesine aoe Variety of Bolbitis fro: ee gee on Elaphoglossum crinitum, 9 on Notes e Hawaiian Fern Cente Adenophorus, 68 Notholaena aurea 200; grayi, 200; oe aes ee 128: sinuata, 200; stand- Observations on Drought (es in Obes ign verse Rydb., oe : he Sensitive Pern: Sup- 151; sensibilis, 26, 35, 13, 185, f. obtusilobata, 94 o es Test m the President, p logiostum sherpa wide? nii, 30; eicabin see 42, var. pseudopodum, 30, ,, Pyenostichum, 80 51; cinnamomea, 42, ’ ar. cadioee omea, ¥ viet 135, , 42, 91; regalis, 31, 42, 129, var, regalis, be var. spectabilis, Pai, Dominck J, The a pape my of Isoetes (Rev. ); 215 Pellaea atropurpurea, 26, 36, 42, 78; ata es longimucronata, 198; fae tundifolia, 154; rete cena var. Na cng 201 Phanerophlebia auriculata, 198 Phlebodium Phymatodes exiguum, 145; lum, 137, 145; swartzii, 145 Phytogeography of Selaginella douglasii. aces Pityrogramma triangularis, 78, 199 aca! flee 183 Polypodium, 136; Bi in “im 69; brasili- goes — alifornicum, 80; exi- dat 1 Pobyetreas acrostichoides, 26, 36, bo adiantiforme, 207; -aunii, 42, 90; lonchitis, 118; eye devi 123, var 8, 22, “oy lemmonii, 113, 11 sites tum, 79, A preiaiiaiy omatographic Study Chr of Eastern American Dryopteris, 105 Professor . Ching and the Study of rn 99 m, 138 teridium aquilinum, 26, 42, 89, 90, i var. lanugi var. aquilinum, 37, 118, eyes 78, var. latiusculum, 37, 125, r, pubescens, Senta _retica, 20, 23, 47, a 131, ss Pyr 62, Sa, 00 63, “focculoss, 63, 65; ae 63, en, ariana, 62, ‘63, 67 a: of Polypodium virginianum — brachypteron (Ridlon) Fern- - Auditing Committee, 52; cfs dge of Elections, wt omen , 46; Treasurer, iabili in oonaed iddleh Fern, parva pensylvanica, Rhipidopteris peltata, 93 St. John, Harold. Flora of Southeastern Wa nagar a and of Adjacent Idaho Rev. 15 graphic "Study a Sronen American Dry ris, 10 eiacaee “37, et 83; arizonic 00 densa, 189-19 g & Zon xtasil, $2- 84; galeot tii, 195; _lepidoptyes 190, 95; rupestris, 193; rupincola, 216 200; selaginoides, 84, 90, 91; subg. Stac 1 geet x, c nderwoodii, ~op at ate tong 210, 211 Some Hints for the Fern Culturist, 73 Some Rave Rams arise nes fox ’South- eastern Ferns, 1 Southwest Mating ve Sperka, Mar Growing Berry Bladder Fern 43. hiue west in Pteris cretica with Spe- cial Reference to the Cytoplasmic Inelusions, 129 Stegnogramma, 141, 142. agony Moor ey pea _— 142, va 141: 14 Nein ag “alien A. Flora of Missouri (Rev.), Storer, Poker ow. Ferns and Fern ag on Bonaventure Island, Quebec, Tetley, M. I. Growing Mother Ferns, 94 Thelypteris, 99; atrovirens, 143; augescens var. lindheimeri, srossifolia teris, 143: gracilis, 143; | hexagonoptera, 26,387: lindheime linki 29, 37: pilosa, Lae var. ‘alabamensis, 142, 145, var. Major, 142; sclerophylla, 144 Todea ba rbara 15 Trichomanes, 139, 183; boschianum, 31 Trudell, Harry Ww. (Obit.), 152 AMERICAN FERN JOURNAL Tryon, R. M. The British Pteridological Society, 99 Urostachys, 71; arcturi, 71, 72; pa gles 2; bo nae-voluntatis, 72, buesi eee censis, 72; crucis- -australis, "71, 72° cuatrecasasii, 72; dianae, 71, 72, ewanii, 725, hy ain cep ge to je TROD macbridei, 72; poseidonis, 71, 72: socratis, 72; _tellae polaris 71, 72; trachyloma, ney ‘wohl- beredtii, 73 Wagner, War win Bing- ham Gopaiana ein: rier: ) and his tributions to Pteridology, 17 d Taylor A. §S Sela a densa Rydb., Wisers: Edgar arry Ww. _ Trudell, 152; Some New N for Southeastern Fern bane Dean P. The ita a Sucrose n Apo pogamy in Cyrtomium falcatum Wilson, oKemveth A. Notes on the Hawai- A P Fern Genus Ade horus, 68 Ww aoitina ilvensis, 42; mexicana, 211: obtusa, 26, 37, 42; oregana, 118; 3 scopulina, 118, 122, 211 Woodwardia fimbriata, 80, 207 Xiphopteris, 70, 1838 ERRATA Page 40, line Page 57, line 18: For “Gwyne” r Page 63, line 18 Page 68, line 44 is pa Page 68, singe stor ma age 68, Page 69, line i ‘For Page 70, line 9 Page 70, line Page 78, line 2 -age 79, line 12 Page 85, line 3: Page ine 1 : For “i ‘> read “a 17: For “1855” rea 25 i) co © . jest =) Le z F age 90, line 5 from age 98, line 19: For A age 124, line 25: For, “ ; i 125, line 13: For “Des.V” Page 146, line 28: For “pte ridop Salis tay ni the List of Msibers i : lysti Page 96, line 22: For “thelpyterioides” read “thelypteri Ls”? Dor brun ed rea ad “brunfeslsii. ” 3: For “elifi” read ie s ead ‘ For developeivnt? pri caeaivianant: fd 154. ipinnata” read Sioa ining ceal : For “minuta ” read “m unicellar” ap “unicellalar. fe : For “Chestnut” ey “Chesnut. nf : For “Chestnut” read ‘Chesnut. 7 F Ta. ”? chium” read “Polystichum.” ioides.” pent ia” read ‘ ‘pteridographiae.” will appear in Volume 55). Long-awaited... Sq a complete book yee on native and gam exotic ferns gf THE GARDENER'S FERN BOOK By F. Gordon Foster This definitive book on ferns for the horticulturist and nature-lover is also an immensely i i ork t practical guide e er. It is the pene complete information on bot ve and exotic species. 1 identification gs and descriptions carefully, simply oy beautifully show th parts of fern—in silhouettes of the entire leaf enlarged es of its special features. 16 color pictures show si ape of structure ag ee = ope = white photographs cific information ne propagation, “lighting, soil, and rare fe both ind aE GARDENERS @“""" FREE EXAMINATION COPY! ---~~ and out. THE rpoagpree seg eg BOOK also a ! D. VAN NOSTRAND COMPANY, INC. ] ere, and aye descriptive tables, 20 Alexander Street, Princeton, N. J. ake kok on = It ' Picase send me _. THE GARDENER'S sn FERN BOOK at Hs .95 per copy. Within 10 days I will remit $____—__—_—» plus small delivery aire. or return book(s) ot Foster has lectured at —— and owe nothing Rstit hi itute ne Technology, is a micro- ea ee oe PY specialist — the Bell Tele- Phone Laborato es, and gives lec- Adites tures f Audubon and ___ State. Zip Code_—— 204 be. cAVET 1 — wih order ade e pay de- Pages. 6”x9”. Illustrated $7.95 ! livery. Same return privilege _quaranterd American Fern FYournal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY EDITORS IRA L. WIGGINS Cc. V. MORTON ROLLA M. TRYON JOHN H. THOMAS VOLUME 55 1965 MONUMENTAL PRINTING COMPANY, BALTIMORE, MARYLAND Contents VoLtumeE 55, NuMBER 1, Pages 1-48, IssurpD Marcu 25, 1965 Serrenk Studies in the erway of the Dryopteris villarsii Woynar Complex in Europ G, Panigrahi Abortion of Spores and Sporangia as a Tool of the Detection of Dryopteris Hybrids...H. H. Wagner, Jr. and Katherine Lim Chen OW eMnival Cyatien eo ss ae C. V. Morton The Gametophyte of Cystodium Lenette R. Atkinson New Cytological Records for Cystodium oes Dickson . K, Roy oe R. EF, Holttwm American Fern Seah Annual Meeting in 1965; Report of the Presi- mur gine: f the Secretary; Report of the Treasurer; Report of the Judge a Elections; New Members _ VoLUME 55, NUMBER 2, Pages 49-96, IssurD JUNE 9, 1965 A Peréel of Cameroon Permnsy as oe er ek SN Alice F. Tryon Cup Ferns (Dennstaedtia) Cultivated in California Barbara Joe A Pteridophyte Botanical Garden Helen Ross Russell The Essentiality of Boron for Dryopteris dentata and Selaginella BpOCa een ee ane John EB. Bowen and Hugh G, Gauch Preliminary Field Studies on the Fern Flora of Oaxaca, Mexico hn T. Mickel Shorter Notes: Return to Panther Creek; Schizaea pusilla—Peren- nial Gametophyte?; Osmunda regalis var. spectabilis in India; Cystopteris bulbifera (L.) Bernh. New to Texas Reeent Fern Literatire, 2 ota NODES and NeWe! oo os: American Fern Society _. — we On VotumE 55, NuMBER 3, Pages 97-144, Issurp SEPTEMBER 9, 1965 County Distribution of Ferns and Fern Allies in Rhode Island othy L. Crandall 97 Vernation in Some Species of the Genus Cheilanthes Irving W. Knobloch 113 Preliminary Field Studies on the Fern Flora of peng Mexico (Concluded) ohn T. Mickel 117 An wulg of a Variable Population of ees arvense and E. X, litorale ard L. Hauke 123 mecaut Wwern Literature to Hoe ee ee ee 136 BNO LGS. ATE ON OWS ig 2 ehh eter pear ie eae aan ene iatae pal Nd Ee 142 American Fern Society, 6.26 O86 ee ee 143 VoLumeE 55, NumBeErR 4, Pages 145-188, IssueD DECEMBER 30, 1965 Observations on Some Ferns in Georgia Wilbur H. Duncan and Donald Blake 145 Cytological Observations on Ferns from Southern China S. K. Roy and R. E. Holttum 154 K. Roy and R. EF, Holttum 158 Cytological and ELE ee? Observations on Metaxya rostrata H.B.K.) Pre S Observations on Cultivated Ferns, VIII. Stenochlaena — C. V. Morton 164 Dispersal of Marsilea mucronata by Water Birds arles R. Malone and Vernon W. Proctor 167 A New ae a of Pellaea glabella var. glabella from Missouri H. Wagner, Jr., D. R. Farrar and Katherine L. Chen 171 Shorter Notes: Japanese Climbing Fern Spreading in South Carolina; A Simple Method for Growing Ferns from Spores. 179 Recent err biteratupes Woe a Oy oe ee 181 American Fern Society FS Pei edls SONS tab Cor cael cre Qe mane Peas ecr ito Tndex to GB ee a ek 185 Vou. 55 JANUARY-MarouH, 1965 No. 1 American fern Fournal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY ts IRA L. WIGGINS 0. V. MORTON ROLLA M. TRYON JOHN H. THOMAS ty CONTENTS Pre reliminary Studies in the Cytotaxonomy of the eee villarsii (Bell.) Woynar Complex in Europe..__G. PANIGRABI 1 TL. Waener, Jr. AND Katuerine Lim CHEN 9 A New Jamaican iii C. V. Morton 30 The Gametophyte of Cystodium _____ __._Lenerre R. ATKINSON 32 New Cytological Records for re ~~ imeem 8. K np R. E. Hotttum 35 an Fern Society: Annual Meeting in a ; Report of the "Provident; Report of the Secretary; Repo: the Treasurer; Report of the Judge of Elections; New Me pn dbeRSavscccessevsnecsscsves 37 Missoup, BoTANicag APR ¢ 1965 GARDEN Linrapy Che American Fern Hocietp Council for 1965 OFFICERS FOR THE YEAR Donovan 8. CorrELu, Texas Research Foundation, Renner, Texas President Irvine W. KNOBLOCH, Department of Botany & Plant Pathology, Michigan State bre dos East Lansing, Mich. Vice-President LENETTE TKINSON, 415 S. Pleasant Street, Amherst, cpmsumcagr bo cretary RICHARD L, Havuxe, Department of Botany, University of Rhode per Kingston, Rhode Island asurer Ina L, Wiaains, Dudley Herbarium, Stanford University, a Cat ie Watters H. Hopez, National Science Foundation, i lesane . Rep dasatar te to A.A.AS. Warren H. Waener, Jr., Dept. of Botany, Univesity of Michigan, Ann Arbor, Mich. epresentative to A. I. B. 8. OFFICIAL ORGAN American Fern Journal EDITORS Ira L, Wiaeins. eee ee, Stanford University, Stanford, Calif. - V. Mortoy.___.____..___ Smithsonian Institution, Washington 25, D. C. Gray Herbarium, Harvard University, Cambridge 38, Mass. Joun H. THomas__ Dudley Herbarium, Stanford University, Stanford, Calif. An illustrated agar devoted to the general study of ferns, owned b the American Fern Society, and published at 3110 Elm Ave., Baltimore il, Md. Sesondstiae oan paid at Baltimore, Md. Matter for publication should be addre ssed to Tra L. Wiggins, Dudley Herbarium, Stanfor University P eppncn Californ Subscription $4.50, esueaive s FS geney handlin, ing na ages coe = esas bers of the American Fern Soci “hg tancast dues, $4. 00; bership, $8.00; life membership, $80.00). Ss reprint Ts paces in advance, will be furnished authors at cost, plus pos Back volumes $5.00 to $6.25 each; single back saachacd # 64 pages or less, $1.25; 65- ° Pages, $2.00 each; over 80 pages, $2.50 each; a tive Index to Volum 1-25, 50 cents. Ten percent discount oe orders of six volumes or Changes of aie applications for membership, subscriptions, orders for back numbers, and other business communi _—— should be addr = - a ace AND CURATOR or THE HERBARIUM gner, Unive ersity of M ichigan n, Ann Arbor, Mich. A regular loan reanadin is maintained in connection with the library and herbarium. Members may bavroe books and specimens at any time also are open to members who wish to arrange exchan nges; a membership list is published at ree to assist ae interested in obtaining specimens nae different localities American Fern Fournal VoL. 55 JANUARY-MarcuH, 1965 No. 1 Preliminary Studies in the Cytotaxonomy of the Dryopteris villarsii (Bell.) Woynar Complex in Europe G. PANIGRAHI The discovery of diploid forms with n = 41 and tetraploid forms with n — 82 (Figs. le, 4e) in Dryopteris villarsii (Bell.) Woynar (Manton, 1950) raised interesting cytological and taxo- nomic problems. The relationship of the exclusively tetraploid forms with restricted distribution in the British flora with diploid alpine forms in Switzerland and France needed study to throw light on the past history of D. villarsii in Britain, as Suggested by Manton (1955). MorePHoLogy or CyTOTYPES A diploid stock plant from France had fronds nearly tripin- nate, green, herbaceous ; lowest pair of pinnae not reduced, or only slightly so; pinnae fewer and more crowded than in the tetra- ploid, those of adult fronds ascending; pinnules with larger humber of teeth (12-13), with a corresponding number of bi- fureate veins ending short of the margin; stipe and rachis scaly (Fig. 3b) ; spores small, 40.3 » broad by 48.6 » long (Fig. 1d). On the other hand, tetraploid plants from the Pennines in England had fronds bipinnate only, straw-colored, stiff; lowest pinnae reduced (Fig. 3c), occasionally equal to subbasal pair; pinnae more numerous than in the diploid form, almost at right angles to rachis; pinnules much less toothed (3-5), with veins equally prominent, some of them bifureating and running to the margin ; stipe and rachis scaly; spores larger, 53.5 » long, 44.6 broad (Fig. le). It should be noted that the characters ascribed above to the cytotypes are based on only one plant of each, and I am not sure 2 AMERICAN FERN JOURNAL “get i : °? = ts ect dey * é all ™“s Ay. ’ ata 2) a & .* CYTOTAXONOMY OF DRYOPTERIS VILLARSII a that all of them are constant for each cytotype. Nor do I know how much these characters vary with age, habitat, seasonal fluc- tuations and edaphic conditions. The venation may be a more or less constant character, and with the size of spores, may serve as a character of some taxonomic value. I have no clue as to whether the tetraploid form occurs in the continental flora, al- though I have positive evidence that the British population of the species is tetraploid, and that the diploid form occurs in both Switzerland and France (Manton, 1950). EXPERIMENTAL HYBRID . With a view to establishing the genetic relationships between the two cytotypes, the British tetraploid form was crossed with the French diploid (Panigrahi, 1954). As a result of about 25 attempts in each direction, only one F, triploid hybrid was pro- duced artificially with the tetraploid as the female parent. The hybrid showed pronounced heterosis (Fig. 2c) and was inter- mediate morphologically between its parents (Fig. 2a). Although it had bipinnate fronds, the serration of the pinnules and the number of teeth on a pinnule were like those of its female parent. The disposition of the pinnae on the rachis was like that of its male parent. It became sporangiferous in 11 months from the date of crossing, but it was sterile and formed only abortive spores. Study of its meiosis showed 33 bivalents and 57 univa- lents; no multivalents were seen. (Two meiotic cells are illus- trated j in Figs. la, 1b, 4a, and 4b). The formation of 33 bivalents indicates homologous pairing between 33 chromosomes of each parental genome, the remaining 57 from both gametes forming univalents in the absence of their homologous partners. Since n = 41 in the genus Dryopteris, there is therefore some, although ‘apparently not complete, homol- ee Fig. 1. a aNp b: PHOTOMICROGRAPHS OF CYTOTYPE OF F, TRIPLOID HYBRID SHOWING 33n + 571. e: CYTOTYPE OF TETRAPLOID FROM GREAT BRITAIN WITH 821, d: SPORES FROM DIPLOID PLANT. e: SPORES FROM TETRAPLOID PLANT. (a, b, e, X 800; d, e X 80). 4 AMERICAN FERN JOURNAL ogy between the chromosomes of the diploid and the tetraploid forms. There is no suggestion of autopolyploidy in the British tetraploid. DISCUSSION These findings raise important nomenclatorial points which - c Fig. 2. FRONDS FROM YEAR-OLD SPORELINGS: a: FROM TETRAPLOID SPORE LING. b: FROM DIPLOID SPORELING. e: From F, TRIPLOID HYBRID. (ABOUT 4/5 NAT. SIZE) CYTOTAXONOMY OF DRYOPTERIS VILLARSII 5 Fig. 3. FERTILE FRONDS FROM ADULT PLANTS. a: FROM F, TRIPLOID HYBRID. O/- r b: From prptom pian, e: FroM TETRAPLOID PLANT. (ABouT 2/5 NAT SIZE), 6 AMERICAN FERN JOURNAL 8 xe %, p & Fe elder. 6." GY ) “9 xs Pre yep Moss oe Oy 4 0 cae Fait On) SF ¥ Fev © 8, ¥o0¢ ae M2 Se Sy RS 6.0 Fy - “eo oe CR Peale us ae Oy DD “ Fy. : b Nes » . Bete’ Be so 9 * NA a mw Ne oe, « . tT CYTOTAXONOMY OF DRYOPTERIS VILLARSII 7 y be pursued profitably a little further. Bellardi (1792) founded Polypodium villarsit on material collected from the pla- teau of St. Nicolai on Mt. Cenis along the Italian-French bor- der, 45.2° N., 6.8° E., and described his plants as having “frondi- bus bipinnatis, alis horizontalibus, foliolis serratis, dentibus aris- tatis saepe bifidis.” It is from this type locality that my diploid stock plant came. Comparison of the description of the diploid cytotype with Bellardi’s description given above shows some dis- erepancies, whereas the description agrees well with the tetra- ploid cytotype, which may mean that both the diploid and tetra- ploid forms occur at the type locality and that Bellardi’s descrip- tion was based on a tetraploid rather than on a diploid plant. It might, however, mean that it was drawn from a young diploid plant that had bipinnate fronds, or that Bellardi’s description is broad enough and vague enough to include both diploid and tetraploid cytotypes. Which alternative is the more probable I cannot say. We may further note that Hoffmann (1795) founded Poly- podium rigidum apparently on continental material, but did not specify the locality from which his plants came. However, his description of the species (viz. “fronde bipinnata ; pinnulis ovato- lanceolatis incisis; laciniis conniventi 2-3-dentatis, stipite paleaceo, colore ex flavescente viridi” ete.) fits the morphological ehisactars of the tetraploid from Britain. It is clear, therefore, why all British taxonomists with the exception of Clapham, Tu- tin, and Warburg (1952) have described British materials as Hoffmann’s P. rigidum (such as Lastrea rigida, Aspidium rigi- dun, Dryopteris rigida, ete.). But the latter authors eall the British species D. villarsii (Bell.) Woynar, and cite rigida as a Synonym. Rela pean mmenrec ns Fig. 4. DIAGRAMS OF PHOTOMICROGRAPHS OF CYTOTYPES ILLUSTRATED IN FIG. 1. a: AND b: CHROMOSOMES OF F, TRIPLOID HYBRID SHOWING 33 PAIRS IN BLACK AND 57 UNIVALENTS IN OUTLINE. ¢: CHROMOSOMES OF TETRAPLOID PLANT SHOWING 82 BIVALENTS. (X 1600). ta AMERICAN FERN JOURNAL In view of the allotetraploid nature of the British form, it will be necessary to treat the two cytotypes as two taxonomically different species if the morphological characters present in my stock plants also characterize the populations in the wild. In such a case the specific epithet villarsit (or villarii) of Bellardi may be used for the diploid form if there is no tetraploid form in the type locality for Polypodium villarsti Bellardi. The specific epithet rigida on the other hand may be used for the British form if only the tetraploid form is cytologically established elsewhere on the continent. If the tetraploid is found also in the type lo- eality of D. villarsii, this will be the correct name for the British form and another name has to be provided for the diploid species. This nomenclatorial problem and past history of the British form need more extensive investigation of material from the European continent. Grateful thanks are due Professor I. Manton, F. R. S., The University, Leeds, for supervision and guidance. LITERATURE CITED Betiarpi, L. 1792. Appendix ad Floram Pedemontanam. Augustae Taurinorum, p. 255. CrapHam, A. R., T. G. Turin, and E. F. Warpure. 1952. Flora of the British Taled Cambridge Manron, I. 1950. Problems of ayokley and evolution in the Pteridophyta. Cambridge 1955. Cosi ntributions of the tropical ferns to understanding of te British Flora. “Species studies in the British Flora.” London. PANIGRAHI, G, 1954. Cytogenetic and taxonomie studies on certain species land. Woynar, H. 1918. Betrachtungen iiber Polypodium austriacwm Jacq. Osterr. Bot. Zeit. 67: 268. BOTANICAL SURVEY OF Inp1A, ALLAHABAD, INDIA. DETECTION OF DryopTEeRIS HyBrRIps 9 Abortion of Spores and Sporangia as a Tool in the Detection of Dryopteris Hybrids W. H. Waener, Jr. AND KATHERINE Lim CHEN! It is practically axiomatic that interspecific and intergeneric hybrids in ferns are, with rare exceptions, “sterile.” The situa- tion found by Trevor Walker (1958) in certain hybrids of Pteris remains our only example of interspecific hybridization that yields fertile progeny directly. Because sterility is otherwise so nearly universal among fern hybrids, it seems desirable to de- scribe some of the signposts of this condition. What is meant by “sterile hybrids” requires explanation, because there have been some misunderstandings. The mere absence of sori, for example, is not necessarily diagnostic for hybridity. So far as has been determined, fern hybrids have just as many sori as do normal species. Species as well as hybrids both tend to lack sori when growing under conditions unusual for the plants involved, such as exceptionally deep shade and especially wet, rich substratum. The point is that production of sori and of sporangia per se are not inhibited in interspecific crosses, so that a hybrid plant which is unable to reproduce itself may nevertheless be abundantly soriferous. A more serious problem, as will be exemplified by alleged “hybrids” described and figured by Crane (1953) and Tryon (1942) to be discussed below, involves misinterpretation of the criteria for the detection of hybrids. We hope, therefore, that the following notes and illustrations will prove useful to those who are engaged in clarifying the variations and taxonomy of American woodfer ns, Dryopteris. DETECTION OF HYBRIDS Interspecific crossing usually expresses itself in terms of what is traditionally called “abortion.” There are all degrees of abor- Research supported by National Science Foundation Grant G-10846. We Wish to thank the persons whe np contributed spe sre and field eae unee to this research, especially D. J. here A. Murray Evans, E. Wherry, and (. vy. eos AMERICAN FERN JOURNAL ¢ COG ST C) O60 COGIC OCOUO. oO 00000 OUOOC Jee He OOO “000 So OCOOCn O OOCa¢ 0000e C0000 JUd0OO mm m 8D 00000 80006 * OOQ00 00000 60000 00 weer Y 2 , LP OD F GBQC 9G BU @> 32 ay. V 3Q &> ° aad - D Fig, 1. A-E, BXOSPORE OUTLINES OF NORMAL SPECIES EXCEPT FOR ac SPECIMENS IN WHICH EXOSPORE COULD NOT BE SEEN. A. DRYOPTERIS WHERRY?, 63172 ER 76. Micu., D. J. Hagenan (MICH). H. D. ritix-mas MARGINALIS, ALPENA SINK, MICH., Hacenan H-27. (UNLESS OTHERWISE INDICATED ALL SPECIMENS DEPOSITED IN MICH, AND NuMBERS OF W. H. WAGNER). hee + Sela DETECTION OF DRYOPTERIS HYBRIDS 11 tion—from the state of only a few defective spores to the con- dition in which all of the spores appear to be distorted and inviable, and the spore-cases themselves fail to mature normally. Normal species usually have at least some abortive spores (as shown in Fig. 1, E) ; and sterile hybrids usually have at least a few spores that appear normal (Fig. 1, G@). Experienced field workers can usually tell with a good hand lens (15x or 20x), or sometimes even with the naked eye, whether a plant shows spore or sporangial abortion; at least in the genus Dryopteris. The sorus often appears darker in color in hybrids as a result of the presence of numerous deformed spores with blackened walls and because the majority of sporangia fail to open and appear darker than a corresponding mass of normal sporangia from which the spores have already been discharged. The mature sorus of a hybrid, no matter what its size, usually seems to be more com- pactly constructed than in normal species, with the sporangia appearing as more or less globular objects tightly grouped to- gether. When the sporangia are later examined under the com- pound microscope it is seen that even those capsules which de- veloped to full size contain spores of various sizes and shapes. Such irregular spores, when seattered on an appropriate growth medium, germinate in extremely low percentages or not at all If a few prothallia are obtained from them in culture, these will generally be peculiar in form and fail to complete their growth or to reproduce the plant. The signs of abortion will be described in detail below. The phenomenon of spore abortion may be traced back to the Process that produces the spores. This process is described in detail in familiar cytology and genetics books (e.g., Sharp, 1943, PP. 216-219) so needs no repetition here. The important point is that in hybrids the chromosomes tend to be distributed irregu- larly to the young spores, and the spores differ genetically among themselves. Under the microscope one can tell, even in very young spores, that they are abortive. Often the sporangium dies 48 a whole at a stage of early spore development. When the 12 AMERICAN FERN JOURNAL spore-cases do reach their full size and do discharge their con- tents, the spores are highly irregular. The assignment of interspecific hybrid status to a plant found in nature should not rest alone on whether the spores are abortive. There is always the possibility that an individual of a normal species had been damaged by sudden environmental changes which injured the developing sorus, or that the plant was geneti- cally abnormal in the first place and that abortion was due to some peculiar internal factor. Hybrid status should be assigned first and foremost on the basis of intermediacy. During recent years botanists have become more and more familiar with the basie species of Dryopteris in temperate North America, thanks to such workers as R. C. Benedict, E. T. Wherry, and Stanley Walker. We now know the ranges of the species and their normal spectra of variation to a high degree. And where popu- lations of two well known species grow together intermixed in the same habitat and an unusual plant is found which unques- tionably combines the characters of the associated species, there is no reason not to interpret the plant as a cross. Presumably all or most of the species differences between the parents are multi- genic, involving many interacting factors. If they cross, their hereditary factors influence each other in such a way as to pro- duce intermediacy in all or most characteristics. Hybrids which closely resemble one or the other parent in all their characters would not only be unlikely, they would be extraordinary. Hybrids in Dryopteris are practically all first generation crosses since the vast majority are sterile and unable either to breed with themselves or with other plants. Thus crosses in these ferns are expected to be almost exactly intermediate in most of their characteristics because recombination of parental traits or backcrossing is not likely. Only when the cross is a hybrid be- tween a 4X species (in which the chromosome number is 164) and a 2X species (with the usual diploid condition of 82 chromo- somes) do we expect the situation to be otherwise. In the latter DETECTION OF DRYOPTERIS HYBRIDS 13 case, the hybrid will still be intermediate between its parents, but its characteristics will tend to be inclined toward the 4 species. For example, the well known D. X boottii, although obviously a compromise between its parental characters, resembles its 4 x parent (D. cristata) more than its 2 parent (D. intermedia). The actual steps involved in the detection of an interspecific hybrid found in nature usually follow this pattern: (1) Recognition that the majority or all of the characteristics of lant in question are intermediate between those of two distinct and well-understood species. (2) Observation that the sori contain abortive sporangia and spores (3) Determination that the chromosomes, when studied in eyto- logical preparations under the compound microscope, do not be- have normally in the process of spore production It is easy enough to state that no taxon has been “demon- strated” to be of hybrid origin unless the same kind of plant has been produced experimentally under controlled laboratory conditions. This is true, of ecourse—and in questionable situa- tions the artificial production of a hybrid is mandatory. How- ever, the array of knowledge of hybridization among ferns now available is so extensive that the basic patterns and principles may be considered as firmly established. So many cases, indeed, have been experimentally confirmed that one can say with con- siderable certainty that if the three criteria given above are met, the specimen in question is with very little doubt an interspecific cross. In most cases, moreover, it is entirely sufficient to meet only criteria (1) and (2). However, precautions must be observed. In the present study of the authors involving a survey of the natural occurrences of wild hybrids in Dryopteris in the United States, we have some- times been misled in making interpretations. Others have like- Wise come to erroneous conclusions. Accordingly, the problems of recognition of hybrid abortion will be examined in more detail, to be followed by specific examples. 14 AMERICAN FERN JOURNAL RECOGNITION OF ABORTION ‘lo greater or lesser degrees one uses in the field the following points as aids in determining hybridity: indusial texture, spor- angial size, sporan ngia 1 dehiscence, sporangial bow (so-called Nennulus ’) thickening, and spores. The latter two characters are best studied under the compound microscope. However, a good 15x or 20 hand lens may often reveal the necessary in- formation about all of these characters at the time of collection if the light used to illuminate the sori is bright and the specimen is held in the full rays of the sun. The spores can frequently be scattered by pressure from a fingernail and observed on the leaf surface around the sorus. In some groups of Dryopteris the indusia tend to appear more or less the same whether the specimen is a hybrid or a normal species. In the complex of D. filix-mas, D. goldiana, and D. mar- Fic. 2. HALFPTONE DRAWINGS OF SPORES. A. DRYOPTERIS CELSA (NOTE OUT LINE OF EXPOSPORE WITHIN PER renin > WAGNER 8234. B. D. CLINTONIANA SARGENT IN 1957. C. D. X LeEpsm (NoTE THAT I 5 O08 8 Se THE EXOSPORE CANNOT BE pi a ait th WAGNER 63172 AND WHER DETECTION OF DRYOPTERIS HYBRIDS 16 ginalis, however, we find that there seems to be a tendency for the indusia of the hybrids to become rigid and persistent, with a more or less dark pigmentation, especially as observed in August and September during the latter half of the season. The parents tend to have more papery indusia which may or may not fall off; if they persist, the indusia are more or less curled and the now discharged normal sporangia are conspicuous, radiating in all directions from the receptacle of the sorus. The hybrid spor- angia, on the other hand, are inclined to be tightly clustered and only partially exposed in a ring under the margins of the stiff indusia. The sporangia in all of the hybrids which we have examined tend to average smaller, roughly two-thirds or less the size of those of the parents. What this means is probably that the same process of abortion which influences the spore development, as described above, extends to the behavior of the whole spore ease. When a certain percentage of the spores within die, the whole Sporangium stops development altogether. Associated with this smaller size of the capsules is a failure of dehiscence and dis- charge, so that the sporangia are mostly unopened. The dehis- cence of the spore cases is usually absent except for some of those around the periphery of the sorus. In normal species, in con- trast, dehiscence is the rule except for a central small group of sporangia near the receptacle which tend to remain rudimentary. The majority if not all of the sporangia in a hybrid thus remain unopened even at the end of the season, while at the same time the majority of sporangia in a typical species are opened and have released their spores. One must be warned, however, that early in the growth season, even the spore-cases of normal species are closed, even if there are young viable spores within. Not until the sporangium is mature will the capsule rip open to release the spores, and this may take until late June or July. As might be expected from the foregoing, the sporangial bow (the string of cells along the side and top of the capsule which activates the opening and discharging mechanism) is imperfectly 16 AMERICAN FERN JOURNAL developed in hybrid spore-cases as a rule. In terms of the com- ponent cells, this means that the process of secondary thickening which characterizes the cells of the bow is curtailed sooner or 6 6 C fe re oe OC OOC G G Pea ae cc BOP0O UOC0 9100 0000 Od ‘EOC 0 “p60e80 00000 “20 ie) B200G8 COO oo 000 20000 OOOO NG Oeee apoGD UGOO0UG ¢000 G60)s0 JOO0CO OD VO 0008 OOIGO 060 ‘QOHEO O80 O O Soroe GCGOQOO SOOO 100° “9 a O OO0BO8 C000 O BOo0o00 GOOQC0 OO = — D. “D. CAMPYLOPTERA SPINULOSA,” CRANE 5025 (AFS). E. “D. CAMPY- LOPTERA” (= “D. pILATATA”), CRANE 5022 (AFS). F. D. spINuLosA, 631658. G-J, SPORES OF INTERSPECIFIC HYBRIDS INVOLVING D. INTERMEDIA AND D. SPINULOSA (EXOSPORES ONLY, UNLESS SHADED). D. INTERMEDIA ULOSA, MICHIGAN, HaGeENAH H-35c. D. DILATATA > INTERMEDIA, Huron , Micuigan, Hacenan H-20. I CRISTATA X SPINULOSA, ROSA AND WaGNeER 103. J. D. crISTATA X INTERMEDIA, Huron Mrs., MICH., HaGEnan H-2a, (UNLESS OTHERWISE INDICATED ALL SPECIMENS DEPOSITED IN MICH AaNp NUMBERS or W. H. WAGNER). DETECTION oF DrYoPTERIS HYBRIDS Ag later, so that under the compound microscope the thickening of the cells is seen as less than that of normal sporangia, and many of the sporangia do not seem to be provided with a secondary wall layer at all. Hybrids commonly show the lateral walls of the sporangial capsules to be covered internally with more or less numerous aggregations of globules of blackish or dark-brown substance. Such materials are rarely seen also to some extent in normal sporangia, but they become abundant and striking in the sporangial walls of many hybrids. The abortive spores have various shapes and sizes, and all the forms are mixed together in the same spore-cases (Fig. 2, C). Many of them are very dark in color, even with bright illumina- tion under the compound microscope. The outer jackets or “peri- Spores” which surround the exospores are often much more irregularly crested than usual. The contents of many hybrid sporangia contain numerous small undefined blackened objects, some of them probably very imperfectly developed spores, others related to the globular materials mentioned above. Normal spores can be measured easily by determining the exospore dimensions as seen through the transparent, jacketing perispores (cf. Fig. 2). The advantages of measuring the exospore dimensions rather than the perisporial jacket are obvious. The perispore is ex- tremely rugose, but the exospore is smooth. The exospore is the smooth and immediate boundary of the spore protoplast, so it gives a good measure of the volume of the living cell within. Even in young sporangia, so long as the spores have separated and the *xospores have reached their normal dimensions, it is possible to make measurements, even though the later-developed perispores have not yet been laid down. Unfortunately, in hybrids it is often impossible to make out the exospores within, if the peri- Sporial coverings have formed, so that the drawings and measure- ment data in this article are not exactly comparable between hybrids and Species. We have tried to indicate this in Figures 1 and 3 (which were designed only to show exospores) by trac- ing the entire spore outline of the opaque spores, including peri- 18 AMERICAN FERN JOURNAL spore, and then shading the entire outline. In Figure 3, for ex- ample, G, H, I, and J contain outlines of whole spores including perispores because we could not make out the exospores within. 1 micrometer unit = 1,6 microns D. MARGINALIS Ohio, Wagner 63207 D. FILIX-MAS Crane Herb. 54144, AFS D. F-M X MARG. c Hagenah-27 ok D "D, FRAGRANS X INT." Tryon 4621 (WISC) "D. CAMPYLOPTERA" Crane 5022 (AFS) "D. CAMP, X SPIN." is CRANE 5025 (AFS) Tl D. INT. X SPIN. G Hagenah 35¢ (MICH) D, INTERMEDIA Michigan, Wagner 63170x i Saat ee ee ee ours 5 10 1s rae) 25 30 si DETECTION OF DrRYoPTERIS HYBRIDS 19 To illustrate the appearance of samples of normal species and hybrids of Dryopteris, the figures were prepared by scraping the sori into hot lactic acid and drawing the first fifteen spores in outline under a camera lucida. The perispores were not drawn (except as indicated above) but only the boundaries of the actual protoplast as represented by the exospore. To indicate the meas- urements of exospores in Fig. 4, 50 spores each of the plants in- volved were measured with an eyepiece micrometer at 450 X magnification. The slides were mostly diaphane mounts. If frequency histograms of maximum exospore length are plotted, the sizes of normal spores of sexual species tend to ap- proximate rather steep normal curves (ef. Fig. 4, A, B, H) of variation, with the majority of spores close to the numerical mean. Spores which are abnormal, either much larger or much smaller than the median, will tend to stand out. Sterile hybrid Spores will tend, on the other hand, to have much broader curves (ef. Fig. 4, C, G). (Many hybrid spores cannot be measured at all, of course, and this is indicated in Fig. 4 by the number at the right of each histogram. Each histogram therefore contains the first measurable spores). Maximum exospore length by itself is not sufficient to express completely the situation, however, because there are at least two other factors involved in abortion—outline (ef. Figs. 1 and 3), and exospore definition (as shown by the stippled spores in Which the exospore cannot be distinguished by an ordinary light microscope). The major abnormalities of the morphology of ak Fig. 4. FREQUENCY HISTOGRAMS OF EXOSPORE LENGTHS IN DRYOPTERI = NUMBER OF UNMEASURABLE (OPAQUE OR OTHERWISE) GRAM, A. G 63207 (MICH); B. D. ritix-mas, AS MONT, CRANE 54144 (AFS); C. D. FILIX-MAS X MARGINALIS, MICHIGAN, HaGeNAH H-27 (MICH); D. ayerr' oO wera RAS? g Simcoe TRY reel 4621 (WISC); E. “D. campynoprera” (= “D. piLaTaTa”), MinNgsota, CRAN 5022 (AFS); F, sad D CAMPYLOPTERA x Falco tocol MINNESOTA, NE 5025 (AFS 13 . INTERMEDIA X SPINULOSA, MICHIGAN, HAGENAH H-35C (MICH); H. D Feiner eis. W. AGNER 631760 (MICH). 20 AMERICAN FERN JOURNAL sterile hybrid spores in Dryopteris as known to us may be sum- marized as follows: (1) Size: Range large, with abnormally large numbers of unusually small spores present (ef. Fig. 4). (2) Shape: Some spores not typically “bean-shaped,” but either spherical, twisted, square, or triangular (ef. Figs. 1 and 3). (3) Color: Some spores not transparent enough to enable defini- tion of the exospores within (shaded objects in Figs. 1 and 3). (Non-measurability of exospores may be due also to (2), when the spore is so twisted as to obscure the internal structure. ) In a series of interesting papers, Crane (1955, 1956, 1960) presented much of value to our understanding of spore diversity in American Dryopteris. However, in these papers one cannot obtain a clear idea of the quantitative conditions involved be- cause, in general, single spore specimens were selected to illus- trate the taxa. The curves and drawings presented in this paper should help to allay this deficiency, and make it possible for a student to have an operational basis for ultilizing abortion of spores and sporangia as a tool in the detection of Dryopteris hybrids. As shown in Fig. 1, A-E, the spores of presumably normal species are rather conspicuously uniform in outline and size. Occasionally there is a somewhat larger one (Fig. 1, C, middle), occasionally a smaller one (A, bottom row). Sometimes (although rarely) we find a small, dark “spore” like that so frequently seen in hybrids in which the exospore cannot be seen at all through the irregular, pigmented perispore, as shown in the shaded object in the top of the third column of Fig. 1, E. It is our impression that reproductive polyploid species (4X; 6X; such as D. campyloptera and D. clintoniana) often show some- what more irregular spores than diploids, although we have as yet no statistics to support this. In Fig. 1, F-H, are shown 15-spore samples of three hybrids, D. cristata X marginalis (F), D. clintoniana X goldiana (G), and D. filir-mas X marginalis (H). These spores show much greater irregularities, of course, than we would expect in normal species. The histogram in Fig. 4, (, illustrates the size variation of a 50- DETECTION OF DryopTeRIs HysBrips 21 spore sample of D. filiz-mas marginalis. The most extreme condition includes very large, nearly spherical spores; more or less normal-appearing spores; and many very tiny spores—as well as opaque objects in which the exospore cannot be distin- guished. Some other illustrations of spore abortion may be found in Wagner and Boydston (1958, Fig. B, and 19), Morzenti (1962, Fig, 2), and Wagner (1962, Fig. 1, C). The following descriptions will illustrate the application of spore abortion data to the detection of Dryopteris hybrids. EXAMPLES 1. Some “hybrids” involving Dryopteris filiz-mas: It has become widely recognized over the past three or four decades that wherever D. filiz-mas co-exists with D. marginalis in Ver- mont, Ontario, and northern Michigan, numerous crosses may be expected. There seems to be little or no barrier to hybridization between these species in nature, and the situation is quite unlike most other species pairs when they oecur together. After argh ing numerous herbarium collections of so-called “D. filix-mas” we have come to the conclusion that many of them are actually hybrids that have been overlooked. Why taxonomists have failed to recognize that these collections are actually D. filix-mas < mar- ginalis is related, we believe, to two factors: (a) Dryopteris mar- ginalis is one of the most abundant and familiar of the woodferns of eastern North America, and practically every field botanist is familiar with its variations. D. filiz-mas, on the contrary, is a rarity, highly localized in a few localities. Its variations are not well known to most field botanists. Thus any hybrid plants will tend to be referred to the less familiar, rare taxon. (b) D. filix- mas is a tetraploid species, with 164 chromosomes (possible formula A,A,AsA> ), but D. marginalis is a diploid, 2X=82, (BB). Not surprisingly, therefore, the interspecific crosses (A;A,B) tend to simulate D. filix-mas more than D. marginalis. The displacement of sori in the hybrids toward the margin of the segments that we would expect from the influence of D. marginalis is not so strong as to be a precisely intermediate con- 22, AMERICAN FERN JOURNAL dition; the soral position is accordingly somewhat more like D. filiz-mas than D. marginalis. We recommend that field workers and herbarium botanists re- examine their specimens of “D. filiz-mas” carefully for speci- mens showing traits in the direction of D. marginalis. Suspicious specimens should be examined for spores. The spore data gives an excellent test, for all of the now numerous hybrids we have studied in the field under natural conditions have shown remark- ably irregular and variable spores (cf. Fig. 4, A, B, and C; Fig. 1, Our studies of D. filiz-mas lead us to the conclusion that con- siderably more work is needed to understand the variations of eastern American populations of this species. For example, in our joint investigations with Dale J. Hagenah, we have found circumstantial evidence that unusual temperature conditions that “burn off” the early leaves of the season by freezing lead to the formation in this species of peculiar fronds, sterile and fertile, characterized by much broader and more deeply divided pinnae and segments. These injured specimens are very confusing because they are so unlike the ordinary nar- row form of D. filix-mas. Some are so divided as to suggest hybridization with D. intermedia in their morphology, but at least one striking specimen illustrating such strong division had normal spores of D. filix-mas. It should be noted that most of the unusually broad and dissected specimens of the latter descrip- tion show damage to the frond tip, involving incompletely de- veloped and abbreviated segments. A very robust specimen kindly lent for our study by the New England Botanical Club was annotated in Mr. C. A. Weatherby’s handwriting as “Dryopteris goldiana x ?” and by someone else as “> filix-mas?.” When we examined the spores, however, Wé found no special evidence that the plant was of hybrid origin. A 30-spore sample showed only slightly more irregularity (Fig. 1, E) than a more typical specimen of D. filiz-mas (cf. Fig. 1, D); and the majority of the spore-cases have dehisced in the normal DETECTION OF DRYOPTERIS HYBRIDS 23 way. We interpret this specimen, therefore, not as a hybrid but another of the forms of D. filix-mas. . Some “hybrids” involving Dryopteris intermedia: Our “faney fern” or “evergreen woodfern,” D. intermedia, is the most important species in the eastern part of North America in terms of breadth of range, abundance, and economic value. It is known to hybridize with practically every other species of woodfern with which it grows. On October 19, 1963, the senior author in com- pany with Dale J. Hagenah, A. Murray Evans, and Herman Becker, studied a woodfern locality in Kalamazoo Co., Michigan (T48S, R12W, n. side of sect. 20, swamp along “W” avenue), which is strongly recommended to students of this genus. Here such species as D. clintoniana, D. cristata, D. goldiana, D. inter- media, and D. spinulosa grow in luxuriance, and many hybrids are found as well as variations of the basic species. Crosses of D. intermedia with D. clintoniana, D. cristata, and D. spinulosa are common and are readily found over a large part of the swampy woods. While studying these hybrids we found a colony of a peculiar fern that resembled D. X boottw (D. cristata & intermedia) but which seemed more dissected and more like D. intermedia than usual. In the Great Lakes region, D. spinulosa and D. intermedia, the two most common “spinulose” ferns, develop narrow-leaved forms that more or less resemble D. & uliginosa and D. X bootti, the respective crosses of these species with D. cristata. Figure 9 shows the typical broad form of D. intermedia from the Kalamazoo County locality beside fronds from the apparent “hybrid” in question. Our later study of the peculiar plant showed that if the details of cutting were compared carefully, the narrow- leaved plant matched D. intermedia in spite of its frond outline. Furthermore, its spores are entirely normal. Thus a specimen which we might have carelessly identified as the hybrid D. X boottii turned out in fact to be one of the narrow-leaved extremes of D. intermedia. The narrowness of the fronds of this form 1s genetically fixed, as evidenced by the fact that it grows side-by- 24 AMERICAN FERN JOURNAL side and completely intermixed with the more typical, broad In 1942, Tryon published a “New Dryopteris Hybrid” involving what he called “D. fragrans and D. spinulosa var. intermedia.” His figure ( 1942, Pl. 8, Fig. 1) suggests a narrow form of D. in- termedia, Materials of this “hybrid” sent to us for identification from the Herbarium of the University of Wisconsin provide an illustration of the need for careful study before publishing such interpretations. In spite of a rather elaborate description as sum- marized in a two-page chart (op. cit., pp. 82-83), the evidence is unconvineing. As the author hicaaclt wrote, his “hybrid” is “definitely more closely related to” D. intermedia than to D. fragrans. In our opinion the only possible character which might suggest D. fragrans is the narrowness of the leaf blade. But the actual shape of the frond is like D. intermedia only more nar- rowed, The alleged absence of “spine-tips” on the acute teeth is more apparent than real, and is due to the immaturity of the fronds which had not yet completed their foliar growth at the time of collection (July 4, 1940). Obviously, because of the season, the sporangia of this speci- men were young and undehisced. In fact, the perisporial jackets of the spores were not yet formed. However, by merely placing the sporangia in lactic acid, the exospores (which were fully de- veloped) were separated out easily, and they proved, as expected, to be entirely normal. Tracings of 30 exospores of the supposed “hybrid” were compared with a random sample of 15 each from D. intermedia and D. fragrans in Fig. 3 (ef. A with B and C). The spores of the “hybrid” are obviously more like D. intermedia than D. fragrans. Frequency histograms (Fig. 4, D and H) con- firm their essential identity. Accordingly, the citation of this “hybrid” by Wherry (1961) must be revised. Although D. fra- grans X intermedia may possibly be found or synthesized in the 2 Since tI the above was written we ag pete orn the chromosomes of the narrow form to be nor rmal, with — pairs at meiosis, as in typical D, intermedia, DETECTION OF DRYOPTERIS HYBRIDS DAS future, this interesting biotype has still not been described validly to our knowledge, and has yet to be obtained. 3. Some “hybrids” involving D. dilatata: Reeently it was pointed out that there are two taxa passing under the name “D. dilatata” in the eastern United States (Wagner and Hagenah, 1962). One of them, tentatively designated as “Lake Superior D. dilatata,” is known to occur in the western Great Lakes region and is characterized by its very broadly ovate blade, the petiole nearly equalling the length of the midrib, the blade texture her- baceous, the scales narrowly triangular and dark-brown or chest- nut in color. “Lake Superior dilatata” has not yet been found east of the Upper Peninsula of Michigan and the corresponding part of Ontario. The other taxon included in “D. dilatata,” D. campyloptera, extends from eastern Canada to the Carolinas in the mountains, and is distinguished by its relatively narrower blade, shorter petiole, firmer texture, and broader, paler scales. The “Lake Superior D. dilatata” is a relatively large-spored diploid like D. fragrans, with n = 41. Eastern D. campyloptera has very slightly larger spores, but is a tetraploid with twice as many chromosomes, n — 82. It was hypothesized by Wagner and Hagenah (op. cit.) that the latter species might actually be an allopolyploid derivative of a hybrid between “D. dilatata’ and D. intermedia. Accordingly, in the summer of 1962, a field study was made of populations of D. campyloptera in the Mountain Lake area of Giles Co., Virginia, in the hope of finding examples of the hybrids between D. campyloptera and D. intermedia. If our hypothesis were correct, such hybrids should show that D. intermedia (AA) Shares its species genome with one of the two genomes of D. campyloptera (A ABB) and at spore production the chromosome behavior should conform to the formula AAB, ie., 41 pairs (2 genomes of D. intermedia) and 41 singles (1 “dilatata”’ genome). This field experiment provided an excellent illustration of the use of spores in determining hybrids in the field and although described elsewhere (Wagner, 1963), the details will be repeated 26 AMERICAN FERN JOURNAL here. As it turned out, the variations of D. campyloptera in south- western Virginia were so extensive that the characters of D. im- termedia and those of D. campyloptera seemed to run together. Practically every feature of D. campyloptera overlapped with D. intermedia, and at first we had considerable difficulty in separating the two species (cf. Wagner, op. cit., Table 1, and mee, Mig. 5. gh pate OF DRYOPTERIS INTERMEDIA FROM KALAMAZOO COUNTY MICHIGA A. eras vn WAGNER 63162A. B. Narrow “D. X BOOTTII- LIKE” FORM, 63159A , DETECTION OF DRYOPTERIS HYBRIDS 27 Figures 5, 6). Not until we hit upon the procedure of using the spores as a basis of identification, in fact, could we separate the intermedia-like plants of D. campyloptera from the other species. The method is highly recommended to all concerned with these plants. The spores of D. campyloptera are so much larger than those of D. intermedia that the difference can be immediately recognized under the microscope. The actual difference, as measured by the longest diameter of the exospore, is almost exactly 20 per cent. Under the circumstances, the discovery of a hybrid between these two taxa turned out to be difficult. Every time we foun what seemed to be an intermediate specimen, careful examination showed it to be an intermedia-like D. campyloptera or a very- broad, large D. intermedia. Finally we were forced to take a dissecting microscope into the field and examine literally hun- dreds of plants in the mixed populations. In this way we finally found what we sought. When we ultimately discovered the plants which had both morphological intermediaey combined with highly irregular, obviously abortive spores, we later confirmed in the laboratory that all of the presumed hybrids were in fact triploids with three genomes (3 123) and at spore production there are 41 singles (presumably the unpaired chromosomes of “D. dilatata”) and 41 doubles (the pairs of chromosomes of D. inter- media). At present we know the following natural crosses among the “spinulose” woodferns of the eastern United States: D. campy- loptera < intermedia, D. dilatata X intermedia, and D. inter- media X spinulosa. All three have been examined cytologically by S. Walker (1961), Wagner and Hagenah (1962), and Wagner (1963) ; and all three are being maintained as living plants for research purposes by D. J. Hagenah of the Cranbrook Institute of Science. We were greatly interested in the plant which was figured by Crane (1953, Fig. IX, Crane 5025) as “Dryopteris campyloptera x spinulosa,” taken at Beaver Bay, Minnesota. The specimen 28 AMERICAN FERN JOURNAL preserved in the American Fern Society Herbarium was indistin- guishable to us from forms of “Lake Superior D. dilatata” from that area (including Crane 5022, op. cit., Fig. VII, taken at the same time and place). Since D. spinulosa is a tetraploid species and “D. dilatata” is a diploid species, we would expect any hybrids to resemble D. spinulosa somewhat more than D. dilatata rather than the reverse. However, the pinnae and segments of the specimen of the “hybrid” in question are like those of the 2X species: The pinnae are finely divided, the segments more trun- cate, the angles of divergence wider, and the whole frond more open and ample—as in D. dilatata. The scales on the stipe are very dark, nearly blackish, rather than pale tan as in D. spinu- losa. Nothing about the specimen suggested influence of D. spinulosa. When the sori of the questionable specimen were examined the following facts were revealed: (a) the sporangia are all of full size; (b) all except those in the center have opened fully (as seen in the dry condition) ; and (c) the spores (as shown in the 30-spore sample in Fig. 3, D) are evidently normal, with only a few reduced ones. The spore abortion in this specimen is by no means comparable with that known in the hybrids of “spinu- lose” ferns listed above. Yet the plant, if it were truly the hybrid “D. dilatata” spinulosa, would be expected to have the triploid chromosome condition and the spores should be equally abortive. Our measurements of 50-spore samples of the “hybrid” (Crane 9025) and the “species” (Crane 5022) gave remarkably similar frequency histograms that probably belong statistically to the Same curve (Fig. 4, E and F). There is probably no good reason, from what we know about these plants, why true hybrids between “Lake Superior D. dilat- ata” and D. spinulosa should not be found in the great Lakes area. They do occur together occasionally in wooded swampy places, and might be expected to generate some hybrids. Thus far, however, we have not seen any good evidence that such a cross has been found. DETECTION OF DRYOPTERIS HYBRIDS 29 LITERATURE CITED CRANE, FERN Warp. 1953. Spore studies in Dryopteris. I. Amer. Fern Jour. 43: 159-169; 1955. II. aah pita celsa and D. separabilis, Ibid. 45: 14-16; 1956. ILI. Ibid. 46: -130. 1960. A key to American i Bg Asta on characters of the perispore. Ibid. 50: 270-275. MorzENTI, VIRGINIA M. 1962. A first report on pseudomeiotic sporogenesis, a type of spore i dda aap by which “sterile” ferns produce gametophytes. Ibid. 52: 69-78. Suarp, Lester W. 1943. Fowinaebs of Cytology. MeGraw-Hill Book Co., New York Tryon, R. M., Jr. 1942. A new Dryopteris hybrid. Amer. Fern Jour. 32: 81-85. Waener, W. H., Jr. 1962. Cytological observations on Adiantum tracyi C. C. Hall. Madrofio 16: 158-161. . 1963. Pteridophytes of the Mountain Lake Area, Giles County, Virginia, including notes from Whitetop Mountain. Castanea. 28: 113-150. AND Karuryn E. Boypston. 1958. A new hybrid spleenwort from artificial culture at Fernwood and its relationship to a pecu- liar eet from West Virginia. Amer. Fern Jour. : 146-159 LE J. HaGENAH. 1962. acces in the Huron Moun- pee area of Michigan. Brittonia 14: 100. WALKER, Beek 1961. Cytotaxonomic studies in gs Dryopteris spinu- losa complex.—II. Amer. Jour. Bot. 48: 607-614. WALKER, TREVOR G. “1958, Hybridization in some species of Pteris L. volution 12: 82-9 Wuerry, E. T. 1961. The tas guide. Doubleday and Co., Garden City, Ne OY. Botany DEPARTMENT, UNIVERSITY OF MICHIGAN, ANN ARBOR, MicHican. 30 AMERICAN FERN JOURNAL A New Jamaican Cyathea C. V. Morton A study of type specimens in the British Museum (Natural History) has shown that one of the Jamaican tree-ferns is cur- rently passing under an untenable name, Cyathea nigrescens (Hooker) J. Smith. Since there is no published name available, the species is described below as new. However, the plant itself has been known for over 150 years; it is not uncommon in the Blue Mountains of Jamaica, where it appears to be endemic. CyATHEA hystricosa Morton, sp. nov. x erectus usque ad 3.5 m. altus, hystricosus ; stipites numeros paleacei, paleis rigidis fere pacino t lucen tibus fuscis; page foliorum usque ad 2 m. longa, subtripinnata, coriacea, subtu plus minusve glauca, rhachibus fuscis aieiralie basi pa aullo econ’ pinnae usque ad 65 em. longae et 20 cm. latae, lanceolatae, sub- sessiles, apice acuminatae, rhachillis basi muriculatis, furfuraceis supra suleatis et hirsutis ; pinnulae ca. 30-jugae, subsessiles, longae et m ; venae 9-12-jugae, basi fure atae; sori pauci, saepe LL Sa 2 (vel 3)-jugi, basales; indusia harephaerite mem- branacea persistentia brunneseentia cupuliformia, margine regu- laria coe paraphysibus pat Type in the herbarium of che British Museum (Natural His- tory), tod collected in Jamaica by Wiles about 1805 (Morton photograph 8097). This species has passed under the name Cyathea nigrescens (Hook.) J. Smith, e.g. in Maxon, No. Amer. Flora 16: 74. 1909, and also in the Index Filicum. However, C. nigrescens is based on C. arborea (L.) J. E. Smith var. nigrescens Hook. (Sp. Fil. 1: 17. 1844). Hooker divided C. arborea into two varieties, var. 4 New JAMAICAN CYATHEA $1 nigrescens (of which Polypodium arborewm L. is cited as a synonym) and var. 8 pallida (based on C. elegans Hew.). Thus it is clear that var. nigrescens is only a varietal name for the typical variety, which should now be called var. arborea, and the species C. nigrescens (Hook.) J. Smith, which is based on the variety, becomes a straight nomenclatural synonym of C. arborea (L.) J. E. Smith. Hooker doubtless included several species in his concept of C. arborea var. nigrescens, for he cited specimens from Jamaica, Hispaniola, Martinique, St. Vincent, and Brazil, and possibly saw the one described above originally collected in Jamaica by Wiles, although this collection is not mentioned. The type of var. nigrescens must be the same as the type of Polypodium arboreum 1., and this Wiles collection has nothing to do with that. It appears that the species represented by the Wiles collection has never been described under a tenable name, since when it has been recognized at all, it has always received the incorrect name C. nigrescens. ADDITIONAL SPECIMENS EXAMINED: Jamaica: Beyond Vinegar Hill, vicinity of Cinchona, Under- wood 3253. East of Cuna Cuna Gap, Maxon 9426, 9460. Mac- casucker Bump, Mazon 9557. Stone Hole Bump, Mazon 9492, 9512. Gossamer Peak, Maxon 9219. Vinegar Hill, Maron & Killip 827, John Crow Peak, Maron & Killip 984. Moreces Gap to Vinegar Hill, Johnson 1804. Trafalgar Gap, Stoudt 674. Pleasant Hill, Harris 7723 (BM; fide Maxon). Above Fort Stewart, Purdie (K; fide Maxon). Without specific locality : J. P. 46; Sherring; Jenman; Hart 80; Home (BM; fide Maxon) ; Bancroft (K; fide Maxon). So far as the labels indicate, this 1s a common species in the Blue Mountains at elevations between 725 and 1650 meters ae AMERICAN FERN JOURNAL mens. He scarcely ever took the living plant into view in draw- ing up his descriptions; such appeared to confuse him. If he had looked at the Garden ens | of this and C. arborea he would have paused before he united them as one species.” It is not possible to check Smith’s pies ations regarding the “articulation” of the fronds from herbarium material; it is a fertile field for investigations in the wild. However, the species that he separated as “C. nigrescens” differs not only in this character but in several others, notably in the smaller darker scales of the caudex and stipe-bases (those of C. arborea being large and whitish), scaly secondary rhachises, yellow and only subbullate scales on the costules beneath (rather than white, bul- late scales as in C. arborea), few pairs of sori, and especially @ deeper indusial cup. SMITHSONIAN INSTITUTION, WasHINGTON, D. C. The Gametophyte of Cystodium LENETTE R. ATKINSON One of the rare and little known ferns of the world is Cysto- dium sorbifolium (Sm.) J. Sm., a monotypic tree fern species of Malaysia. It is variously placed in taxonomic works, depend- ing upon relationships postulated. The characteristics used are those of the sporophyte. Holttum (1963) includes Cystodium in his family Cyatheaceae, sub-family Cyatheoideae, tribe Dick- sonieae, along with Dicksonia but says that more information is needed about this fern. Copeland (1947) places Cystodium as the third genus in his Pteridaceae, following Thyrsopteris and Dicksonia, and states that Cystodium is close to Dicksonia and to nothing else. Christensen (1938) places Cystodium tentative- ly in the family Dicksoniaceae, sub-family Dicksonioideae, as a probable derivative of Dicksonia. These authors, however dif- ferent their views, agree on an affinity of Cystodium with Dick- sonia. Cystodium sorbifolium grows in wet lowland forests at eleva- tions up to 1300 feet in contrast to tropical Malesian Dicksoma GAMETOPHYTE OF CYSTODIUM 3 +5" which grows in the high forest between 3300 and 9900 feet (Holttum 1963). A chromosome count from root tips, 2n = 112, and at meiosis, n = 56 has been made recently at Kew Gardens, England (Roy and Holttum, 1965). This fern was collected in August, 1963, in New Guinea by R. E. Holttum, to whom I am indebted for the spores. Germination was excellent and the cultures grew well but slowly on peat in covered dishes. The spore is tetrahedral, yellow in color and ornamented by a delicate but irregular network of ridges. The outer coat is fragile and parts of it crack and fall off on handling, yet it was two weeks after planting before the spore wall opened. Part, at least, of the reserve food is oil and a large droplet can be ob- served in the basal cell during the early stages of growth. A filament of from 2-6 cells is produced, although often in later stages only one undivided cell is found at the spore end of the plate. Divisions in the terminal cell of the filament are oblique and an apical cell with three cutting faces is developed in a cordate plate. Some irregularity was observed in a few individuals: the apical cell may divide transversely, initiating an apical meristem at 4 weeks, or an apical cell may still be present in the thallus at 7 weeks. The mature thallus (nine months) is characterized by a large unruffled wing and a heavy midrib 6-9 cells thick. The first rhizoids contain chloroplasts which soon disappear. The wall is colorless and may remain pale or become golden brown with age. The rather short rhizoids are few and form behind the archegonia on the cushion. No hairs of any sort have been seen. It was 3 months before sex organs appeared. Antheridia were produced sparingly and were observed on small ameristic thalli growing among the rhizoids on larger thalli. The antheridium is of the less advaneed type, that is, with a several-celled wall. Most, perhaps all, thalli in my cultures produced archegonia without having produced antheridia. The necks of the archegonia are massive with very little curvature. 34 AMERICAN FERN. JOURNAL DISCUSSION The great difference between the gametophytes of 1) Dicksonia and the associated Culcita and Cibotium and 2) Cyathea, in the broad sense (Holttum 1963), is in the matter of hairs. Stokey (1930) found that the thallus of the Cyathea group bears hairs and that of the Dicksonia group does not. The hair develops from a special initial cut obliquely from the anterior face of a cell of the midrib at or near the notch. It becomes multicellular and is produced in numbers sufficient to be seen by the naked eye, at least in Cyathea. These are gametophytic structures and not associated with embryo formation or apogamy. Dicksonia, Culcita, Cibotium and now Cystodium are found to lack these hairs. Stokey reported minor differences between the two groups which are reflected also in Cystodium: the thallus is thicker and the antheridia are larger than those in Cyathea. Thus morphological characteristics of the thallus of Cystodi- um suggest a close relationship with that of Dicksonia. The chromosome number, however, of Cystodium, n=56, does not agree with that of Dicksonia, since in the 5 species counted, n=65 (Brownlie 1957, 1961; Manton 1958, Roy and Holttum 1965). Nor does it fit that of the genera associated with Dick- sonia: Cibotium counted in 2 species, n=68 (Manton 1958) ; Culcita macrocarpa, n=66-68 (Manton 1958). The chromosome number which comes closest to that of Cystodiwm is that of Culcita (subg. Calochlaena) dubia where n=55-58 (Manton in Holttum 1963) It is perhaps too much to hope, in view of the great similarity among Dicksonia species, that a chromosome number nearer that of Cystodium will be found among the yet uncounted ( about 22) species. The fact remains that the evidence from the gametophyte does not contradict an affinity of Cystodium to Dickson. LITERATURE CITED BRowNLIE, G. 1957. Cyto-taxonomie studies on New Zealand Pteridaceac. New Phyt. 56: 207-209 ————, 1961. Additional chromosome numbers—New Zealand Ferns. Trans. Roy. Soc. N. Z. (Bot.) 1: 1-4 CytoLogicAL REcorpDs FoR CYSTODIUM AND DICKSONIA 35 CHRISTENSEN, C. 1938. In Verdoorn’s Manual of Pteridology. CoPELAND, E. B. 1947. Genera Filicum. Waltham, Mass. Hourrum, R. E. 1963. Flora Malesiana, Ser. I Pteridophyta, pt. 2. Manton, I. 1958. Chromosomes and fern phylogeny with special reference to “Pteridaceae.” Jour. Linn. Soe. London B. : 73-90. Roy, 8S. K. and R. E. Hoirrum. 1965. New cytological records for Cysto- dium and Dicksonia. Amer. Fern Jour. 55: 35-37, Stoxry, A. G. 1930. Prothallia of the Cyatheaceae. Bot. Gaz. 90: 1-45. AMHERST COLLEGE, AMHERST, MASSACHUSETTS. New Cytological Records for Cystodium and Dicksonia S. K. Roy anp R. E. Hourrum Young plants of Cystodium and Dicksonia were among col- lections made by Holttum in New Guinea in August, 1963, through the cooperation of the Division of Botany, Department of Forests, at Lae. Mr. J. S. Womersley kindly arranged for the despatch of the plants to the Royal Botanic Gardens, Kew, where they have made good growth. Roy has made eytological prepa- rations from these plants. Cystodium is a monotypic genus. Further notes upon it and references to previous literature are given in the accompanying paper on the gametophyte by Lenette R. Atkinson. Cystodium sorbifolium has been found in lowland forest in Sarawak, North Borneo, North Celebes, Ceram, and at many localities in New Guinea and neighboring smaller islands. Holttum found it common on steeply rising ground near two small streams. The stock is prostrate, covered with long hairs as in Dicksonia; the fronds may attain 250 em in length. The young plants sent to Kew had fronds about 10 em long; by July 1964 they had fronds about 50 em long which were fertile. Before fertile fronds were produced, Roy made cytological observations from root-tips; these were treated with paradi- chlorobenzene before fixation in acetic-aleohol, and then with snail enzyme to separate the cells. Clear preparations showing n=112 resulted (Fig. 2). Subsequently, when fertile fronds were produced, preparations were made from sporangia at 36 AMERICAN FERN JOURNAL CHROMOSOME COMPLEMENTS IN CYSTODIUM SORBIFOLIUM. Figure 1. DIA- KINESIS SHOWING n=56 BIVALENTS. * 1000. Figure 2. Root TIP MITOSIS SHOWING 2n=112. * 1000. AMERICAN FERN SOCIETY BT meiosis, showing n=56 (Fig. 1). This result gives confirmation to the status of Cystodium as a genus distinet from Dicksonia. At the same time, there can be little doubt that, as judged by characters of the sporophyte, it is more nearly allied to Dick- sonia than to any other genus. The difference in chromosome number between Dicksonia and Cystodium is very like the dif- ference between the two subgenera of Culcita (66-68 for subg. Culeita, 55-58 for C. dubia in subg. Calochlaena which comprises the species of Malaysia, Australia and the Pacific; the approxi- mate numbers are due in both eases to difficulty in obtaining satisfactory preparations). It would be not unreasonable to raise sube. Calochlaena to generic status. Dicksonia hieronymi Brause grows at 4500-9500 ft. im moun- tain forest in New Guinea. A young plant in cultivation at Kew has yielded root-tip preparations showing 2n=1: his agrees with the counts for three species in New Zealand by Brownlie, and for D. arborea (St. Helena, type species of the genus) in cultivation at Kew by Manton. It should be noted that Dicksonia scabra Wall., reported by Fabbri (1963) as a possible tetraploid (nm — ¢.123) is a species of Dennstaedtia, wn- der which genus Fabbri gives further records. LITERATURE CITED Faspri, F. 1963. Primo supplemento alle Tavole Chromosomiche delle Pteridophyta di Alberto Chiarugi. Caryologia 16: 237-335 Kew GARDENS, RICHMOND, SURREY, ENGLAND. American Fern Society ANNUAL Mretine IN 1965—The annual meeting of the Society will be held in conjunction with those of the American Institute of Biological Sciences at the University of Tllinois, Urbana, Il. The A. I. B. S. meetings are scheduled from August 15 to 20, and our formal meeting sessions, including the luncheon, will be held on August 16th. The American Fern Society will hold two or possibly three sessions for the presentation of papers, and may cooperate with other societies in sponsoring symposia. 38 AMERICAN FERN JOURNAL A two day Fern Foray will be held in southern Illinois, on Friday and Saturday, August 13 and 14, leaving Sunday the 15th free to drive the 175 miles north to Urbana. Dr. Robert H. Mohlenbrock will be the leader of the Foray and he has sug- gested the following schedule: Thursday afternoon, August 12th Arrive in Carbondale Thursday, 8:00 p.m. Illustrated lecture Friday, 8:00 a.m. to 5:00 p.m. Field Trip Friday, 8:00 p.m., August 13 Lecture Saturday, 8:00 a.m. to 5:00 p.m. Field Trip Dr. Mohlenbrock writes as follows :—‘‘The field trip should be divided into two full days to see the maximum number of ferns. I will also be happy to discuss other vascular plants encountered on the trip. I shall arrange a University bus for transportation and will equip myself with a battery-powered megaphone to describe the areas we drive through. “Box lunches for the field trip may be purchased from the University cafeteria. “There is a good possibility of finding 43 species of ferns and fern allies on the trip.” To register for the Foray, write direct to Dr. Robert H. Mohl- enbrock, Department of Botany, Southern Illinois University, Carbondale, Ilinois 62903. Tell him the number in your party, and whether you wish box lunches from the University cafeteria on the days of the Foray. This letter should reach Dr. Mohlen- brock by August Ist. Request for motel or hotel rooms should be sent direct to the manager. The following, all in Carbondale, Illinois, are suggested : Franklin Hotel, Heritage Motel, Holiday Inn Motel, Plaza Motel, Uptown Motel. Write for reservations before August 1st if possible. Report of the President for 1964 It is gratifying to be able to report that the Society is in good shape. The steadfast and energetic work of the various officers, and especially of the Editor-in-Chief of our Journal, is to be commended, AMERICAN FERN SOCIETY 39 Our annual field foray preceding the American Institute of Biological Sciences meetings at the University of Colorado, Au- gust 23-28, was well-attended. We were most fortunate to have Dr. William A. Weber, who was also our local representative, and Mrs. Ruth A. Nelson, serve as our leaders. Dr. Weber guided us over the Flat-irons, some magnificent promontories above Boulder, where we saw several interesting ferns, including Asplenium septentrionale, and to several other choice areas in the vicinity of Boulder. Of especial interest were the great sheets of Selaginella weatherbiana, a species honoring Mr. C. A Weatherby, that covered precipitous ledges in Boulder Canyon. Mrs. Nelson, assisted by a Park Ranger, guided us to some fine fern habitats in the Rocky Mountain National Park. At the formal meeting in Boulder, on Monday, August 23, a fine program of contributed papers was given in the morning and part of the afternoon. Our Vice President, Donald Huttle- ston, who served as Program Chairman, had assembled some interesting papers for the occasion. The luncheon held at noon was attended by a small but enthusiastic group. Considering the fact that many of our professional members were unable to be with us at the meeting because they attended the International Botanical Congress held earlier in the month in Edinburgh, Scot- land, we had a better attendance for all our functions than had been anticipated. The council also held its annual meeting in Boulder where various and pending items of business for the Society were dealt with. During the year Dr. Warren H. Wagner, Jr., in addition to his several other duties for the Society, assumed the responsi- bility of being our representative to the Governing Board of the American Institute of Biological Sciences. He sueceeded Dr. Walter H. Hodge, who is to continue as our representative to the American Association for the Advancement of Science. The long-delayed publication of our membership roster was accomplished during the year. It evidences a strong Society and attests the sincerity and loyalty of its individual members. Our membership in 1964 remained essentially the same as that 40 AMERICAN FERN JOURNAL for 1963 although a substantial increase in dues occurred for the year. It is hoped that every member read my “open letter,” pub- lished in the July-September 1964 issue of the Journal, and, if so, that my appeal will not go unheeded. I omitted there the fact that since our Society has a tax exemption status with the Internal Revenue Service, all gifts to the American Fern Society are tax exempt. Special service to the Society has been rendered this year by Dr. LeRoy K. Henry, who served as Judge of Elections, and by Professor Elmer A. Palmatier, who served as Auditor. The Society is indebted to both of them and appreciates very much their services. T am sure that everyone is pleased that the Society bestowed upon Dr. Edgar T. Wherry an Honorary Membership. This is the very least it could do to show its appreciation to one who has been so magnanimous to the Society. Recently, I felt a great personal loss in the death of Dr. H. L. Blomquist, a member of the American Fern Society since 1934, who was my good friend and major Professor when I attended Duke University many years ago, It has been a distinct honor to me to serve as President of the American Fern Society during the past year and it is my desire to do my best to serve it well during the year 1965. Respectfully submitted, Donovan S. Correwu, President. Report of the Secretary for 1964 The American Fern Society at the close of 1964 has 754 mem- bers living in all states of the union excepting Nevada, New Mexico and North Dakota, and in 27 countries outside the United States. I regret to report the death during 1964 of eleven members (year in parentheses indicates date of joining the Society) : Dr. W. C. Drummond (1956) ; Mrs. Gifford Ewing (1948) ; Mrs. W. S. Learned (1953); Mrs. Fay MacFadden (1953); Mrs. Elizabeth Sawyer (1961); two members of long standing—Mrs. AMERICAN FERN Society 41 Arthur W. Doubleday (1928) and Mr. Henry Trudell (1919) ; three life members and members of long standing—Dr. H. L. Blomquist (1934) elected President of the Soicety in 1962, Mr. Francis Hunnewell (1915), Miss Gertrude Phair (1916); and the distinguished pteridologist and Honorary Member of the American Fern Society, Dr. E. B. Copeland (1948). The annual meeting of the Society, held August 24 with the AIBS on the campus of the University of Colorado at Boulder, was attended by about thirty people. The president of the Society, Dr. Donovan Correll, presided at both the formal sessions and at the luncheon. A varied and interesting program was arranged by Dr. Donald Huttleston, Vice-President of the Society. Titles of papers follow in the order of presentation: “The gametophyte of Cystodium (Cyatheaceae),” by Lenette R. Atkinson ; “Distributional records of the true ferns of Idaho,” by William H. Baker; “Recent data on ferns of Georgia,” by Wilbur H. Duncan; “Variation in Equisetum L.,” by Richard L. Hauke (read by D. S. Correll); “Sporogenesis in Cyrtomium faleatum,” by Norman P. Marengo and Mildred M. Marengo (read by Mrs. Marengo) ; “Leaf anatomy in some of the species of the genus Cheilanthes,” by Paul Volz and Irving Knobloch (read by Paul Volz) ; “The phytogeography of some Galépagos Island ferns,” by Ira L. Wiggins; “The phylogenetic position of Anemia colimensis,” by John T. Mickel; “The log fern complex (Dryopteris celsa and its relatives) in eastern United States,” by Warren H. Wagner, Jr. Four of the five members of the Council were present at a meeting on August 23 when it regretfully accepted the resigna- tion of Dr. Walter H. Hodge as representative of the American Fern Society to the Governing Board of AIBS. Dr. Warren H. Wagner has agreed to represent the Society for a four year term. Respectfully sumbitted, LENETTE R. ATKrnson, Secretary. Report of the Treasurer for 1964 The Society has finished another year in the black. The in- crease in dues caused some loss of membership, and as a con- Sequence the receipts this year were not quite as high as antici- 42 AMERICAN FERN JOURNAL pated. However, there was some increase over last year. With the increase in institutional subscription rates, effective January 1965, and with the increase in price of back issties, we can prob- ably look forward to a further improvement in total receipts next year. The Council decided to use some surplus cash-on-hand to re- print five additional back numbers of the Journal. This gives us a sufficient quantity of all numbers to meet the anticipated rise in orders for complete sets as new colleges open and college libraries expand. It is hoped that the sale of back numbers will be an increasing source of income for the Society, which in turn will be reflected in a larger number of pages in each year’s issue of the Fern Journal. Tax-exempt status was granted the Society by the Internal Revenue Service. This will permit members, of course, and others, to make gifts to the Society, and take income tax deduc- tions in their own tax reports. It is hoped that this will bring about an increase in the number and in the size of bequests and eifts to the American Fern Society. At present, gifts to the Society consist mainly of the royalties on Dr. Wherry’s “Fern Guide.” However, these can be expected to decrease year by year, and something else must replace them for the Society to remain solvent. Since this is to be my last year as Treasurer, thought must be given to a successor. I wish to repeat a suggestion made earlier, that a retired member of the Society who has had experience in some segment of the financial world volunteer his services! The financial report for the year follows: Receipts Cash on hand, January 1, 1964 $1,597.82 Membership dues Arrears and Renewals $1,939.02 Sustaining 392.10 230.70 Advance payments 108.97 Life AMERICAN FERN SOCIETY 43 $2,750.79 Subscriptions Arrears 6.15 Current 235.30 Advances 573.53 814.98 Sale of back numbers 539.12 Sale of reprints 260.52 Gifts 382.22 Extra charges, Journal articles 30.00 Advertising 4.00 Miscellaneous 3.11 4,780.74 $ 6,378.56 Disbursements American Fern Journal Vol. 53, No. 4 $ 869.21 Vol. 54, No. 1 3) Vol. 54, No. 2 825.64 Vol. 54, No. 3 1,234.89 $3,863.33 Reprin 324.91 “a Mailing Journal 117.00 Printing, Stationery 72.2 Treasurer’s expenses 127.41 Secretary’s expenses 102.44 Shipping sa handling back numbers 68.87 Editor’s nses 5.92 Life Meutenahig Fun 80.00 Reprinting back numbers 930.00 B. 8. membership 400.00 Fern Foray 55.06 Miscellaneous 10.51 6,157.65 Cash on Hand, January 1, 1965 $ 220.91 Statement December 31, 1964 Assets Cash in Industrial National Bank 220.91 44 AMERICAN FERN JOURNAL Cash in Greenpoint Savings Bank issell Herbarium Fund 850.76 Life Membership Fund 1,330.67 Una therby Fund 3,595.23 Accounts receivable 119,25 Inventory, American Fern Journal 6,202.50 Library 396.0 $12,715.32 Liabilities Advance dues 108.97 Advance subscriptions 573.53 Fund Balance s Herbarium Fund 850.76 fe auigiiete Fund 1,330.67 i Weatherby Fund 3,595.23 General ae 6,256.16 $12,715.32 Respectfully submitted, RicHarp L. HauKkg, Treasurer. Report of the Auditing Committee I hereby certify that I have seen the books and accounts of Dr. Richard L. Hauke, Treasurer of the American Fern Society, Inc., and have obtained confirmation of the correctness of the Society’s balances on hand as set forth in detail in the accom- panying report of the Treasurer. ELMER A. PALMATIER, Auditor. Report of the Judge of Elections The results of balloting for officers of the American Fern So- ciety are as follows: For President Tiondtan & Cotrell 342 Siidved Wate 3 re ee 1 D. G. Huttleston Se oe 1 For Vice-President Irving W. eee Co 337 Wilbur Duncan 1 David Emory 1 AMERICAN FERN Society 45 John Mickel C. V. Morton = Thomas Pray Kenneth Wilson i. For Secretary Lenette R. Atkinson . 842 For Honorary Member Edgar T. Wherty 3 339 I therefore declare the following candidates elected to office: Donovan S. Correll, President ; Irving W. Knobloch, Vice-Presi- dent; Lenette R. Atkinson, Secretary. It is also my privilege to declare Dr. Edgar T. Wherry elected to Honorary Membership in the American Fern Society, Inc. Respectfully submitted, L. K. Henry, Judge of Elections. New MEMBERS Mr. A. H. Arndt, Edelweiss Haken s, P. O. Box 66, Robbinsville, N, J. Mrs. L. C. Bock, Jr., 823 Tanglewood Drive, Bryan, Texas 77 He were » Cali f eae ee Mrs. William Hebert, 223 E. Canterbury Drive, Stockton, Calif. 95207 Mrs. B. W. Heitkotter, 306 Irene St., Bake sa Calif. 933 Miss Mary Howard, Box 345, Bolinas, Calif. Miss Juanita Hi phutbiand Route 1, St ee Cas Mr. Norman B. Parsons, 91 Glengrove Ave., West, ans 12, Ontario, Canada Mrs. Muriel Perz, 2141 West 29th St., Long Beach, ei ss Dr. Richard H. Sharrett, 1190 Park Ave., Plainfield, N. 7060 Mr. William R. Smith, 20 vgs Millstone Lane, ng ee “alt 94549 Mrs. Arthur Stucky, P, O. Box 306, Stateline, Calif. Mrs. Joseph T. W est, 109 eh County Line Road, chalet Illinois Mr. Stillman Wright, 5822 Conway Road, Bethesda, Maryland 200: CHANGES OF ADDRESS Miss Harriet E. Baker, 1508 Campbell Ave., Thousand Oaks, ope aod Mr. Richard A, Benedict, 63-145 Alderton St., Rego Park, N. Y. Mr. Fred ©. Boutin, 605% - Duarte Road, Monrovia, Calif. Mr. Floyd L, Brown, P. O. Box 764, Clarksville, Tenn. Mrs. Kenneth L, Clarke, 1420 E. Orange Grove Ave., Orange, Calif, 92667 46 AMERICAN FERN JOURNAL Mrs. Blanche E. Dean, 2242 Arlington Ave., Apt. 6, Birmingham, Ala. 35205 Miss Gertrude E. Douglas, 175 Jay St., Albany, N. Y. 12210 Mr. Dara E. Emery, c/o Santa beebae Botanic ae 1212 Mission Can- yon Road, Santa Barbara, Calif. Dr. Mildred E. Faust, 1057 Lancaster Ave., were N. Y. 13210 Mrs. John Zell Gaston: Box 57160, W eae Tex Dr. Richard Hevly, Dept. of Biology, Wisconsin Dat Univ., Superior, Wise. Dr. Herbert H. Hollinger, as Cleveland Road W., Huron, Ohio 44839 Mr. Ralph x Hudson, Rt. 2, Box 1265, Nokomis, Florida 33555 Mr. Thomas A. Hutto, Box 583, Edt South Carolina os Mr. John P. Knable, IT, 807 Wallace St., Birmingham, Mich. 09 Mr. Gunther Kunkel, Camino Viejo 9, Tafira Alta, Las Beni Gran Cc nd Mr. Leroy B. Lee, ine McKinley, Madison, Wisconsin 53705 Mr. W. E. Liggett, 700 Swarthmore Lane, St. Louis, Mo. 63130 Dr. Blanche MeAvoy, 370 N. Cincinnati Ave., Tulsa » Oklahoma 74106 fi s . 47t i 5 right J 8 4 2g el , Ark ank E. Nejdl, 5510 Cumner Road, Downers Grove, ea 60515 Mr. Alfred L. Poe, Jr., 215 Rose Drive, Milpitas, Calif. Capt. Peter G. Root, 775 Country Club Rd., Waterbury, C Prof. Dale M. Smith, Biologieal Sciences, Univ. of ae Santa Bar- bara, Calif, Prof. Alma G. Stokey, ee meng South Hadley, Mass. 01075 Mr. Stephen Snow estes 174 Sunset Ave., Amherst, Mass. 01002 Mr. H. Harwood Tracy, aE Ww. aad Dies accent. ey 91712 ERRATA IN 1964 List oF MEMBERS Pg. 157, line 17: For “817” read “617” Pg. 158, lines 2 and 3 from bottom: omens bape wks Pg. 160, line 6 from bottom: For “Ann” read “Anna Pg. 162, line 10: For * Rtn cca sia? rea Da ou sse” P A vy”; for 1942” re a panies Pg. 163, line 4: For “Ortego” read “Ortega’”’ Pg. 163, line 17: Change address to: “6382 London, Detroit, Mich. 48221” Pg. 164, line 23: For “San Jose 38” read “San Jose 28 Pg. 164, line 2 from bottom: For “Mrs. Claude L.” read “Mrs. Claude E.” Pg. 165, line 22: Change address to “3441 So. Peck Ave., San Pedro, Calif.” Pg. 165, line 3 from bottom: For “Texas” msn oe Pg. 165, last line: For “Flagstaff” read “Tueso Pg. 166, line 1: for “Kruger, Mr.” read eae, Dr? ia AMERICAN FERN SOCIETY 47 Pg. 166, line 30: For “Liggette” read “Liggett. Pg. Be following line 12: Insert, Wiehe Mrs. W. E., 3547 Richmond , Jacksonv qe ane ida 19 Pg. af line 3 from bottom: Delete entire line; insert in its place, ‘“Mar- nier- Lapotal, Mr. Julien, Jardin Botaniqué, St. Jean-Cap-Ferrat, Fra 2.170; hie SE 8: Insert “Poole, Dr. James P., Dept. of Botany, Dartmouth College, Hanover, N. 1940” Pg. 170 eileen line 7 fron ee tton a sert eect Marqua Zoe, 5602 Clifton Ave., Jacksonville, hare 32211 1964’ Peo173, tines 8 ana 9: Delete “Dept. of Plant isis Mt. Holyoke Col- lege Pg. 173, line 27: For “Shaw Road” read “Shwa Road” Pg. 174, line 4: For “Tryon, Mrs. Alice F.” read “Tryon, Dr. Alice F.” Pg. 174, ie ves line 10: Insert “Van de Water, Dr, C. M., 264 Hillside Ave., Chatham, N. J. 1947” (Note: "The Editor is aware - there are feminine names without the appropriate ‘Miss’ also probably are masculine names listed to whie when the title “Dr.” should have been used. Please call such errors, or others, either of omission or commission, to his attention in order that in the list corrections may be made.—I.L.W.) a Exotic and Hardy Ferns BOLDUC’S GREENHILLS NURSERY 2131 Vallejo Street St. Helena — California Begonias Open Saturdays and Sundays 10 A.M. to 4 P.M. or by appointment Phone 963-2998—Area Code 707 Mail orders accepted _—_t+erhaceesteumnsseseeusnimnnanetunneeamemeees Our member, Mrs. R. M. Gunnison, is interested in exchanging or purchasing herbarium specimens of unusual or uncommon ferns. She prefers them unmounted, and pressed carefully so as to avoid withering or appreciable discoloration while drying. Her address is: The White Sands, 7450 Olivetas, La Jolla, California. 48 AMERICAN FERN JOURNAL Statement of Ownership, Management, and Circulation In accordance with the rules and regulations a bs United States Post one as neetablishel under the Act of October 23, 1962; Section 4369, Title 39, United States Code, the following statements are published. itle: AMERICAN FERN JOURNAL , of Issu Quarterly Sie pies March 31, m i June 3 er Location of Office of Publication aaa: 3110 Elm Ave- nue, Baltimore, Marylan 211 Location of Business gs of Publishers : Not printers): sso of Richard L. Hauke, Department of Botany, Uni- ersity of Rhode Tan iri oktok., hot Island 02881 Name of de sher: AMERICAN FERN Soc , Inc., Depart- f Botany, University ra pies Tie: Me lcouphes sah teed 028 Name Ms Edit Ira L. Wiggins, Dudley Herbarium, Stan- ord She siey ee. California 94305 eace: Editor: None Owner: AMERICAN ane Society, INc., Department of nan isi ond of Rhode Island, Kingston, Rhode 1 and AEC arabs ines and other Security Holders: None Average No. each Single issue issue during pre- nearest to ceding 12 months filing date Total No. Copies Printed 1200 1200 (Net Press Run) Paid pia ak term subseri Sih 940 2. Se through agents, ete. None None sis Distribution Gn samples) 24 24 1 No. Copies Distribute 964 920 Richard The Statements made above certified to be correct, and signed by: Hauke, Treasurer of the American Fern grin and Business Manager of the American Fern Journal, on October it Long-awaited... | a complete book | on native and exotic ferns 2 THE GARDENER’S FERN BOOK By F. Gordon Foster This definitive book on ferns for practical guide for the gardener, It is the scat se to give bo i xOtIC 'speries 71 identification show the enl habi poy deta of structure — and = bene: and white photographs fant fern her Ith of specific information on propagation, “lighting, _ oo ‘ersilizing and other important pointers for growing c indoors po—m——— FREE EXAMINATION COPY! ---~-~ Cc. ol ae THE GARDENER’S vega BOOK also contains a * VAN NOSTRAND COMPANY, IN oye e and gm . —o 120 iterander Street, Princeton, N. J. Rtstokcotu.e t | Please send me ____ THE GARDENER’S : FERN soe at eed 95 per copy. Within een: 10 days I will remit $—_____ » plus small delivery care or return 5 heoktn) and owe nothin ai — er has lectured at Stevens itute of Technology, is a micro- scopy specialist with the Bell Tele- — — Laboratories, es gives lec ures cig to Audu si ad Solty oo eee ip SAVE! Remit with order and we pay de- return privilege guaranteed. c.. 224 pages. 6”x9”, Illustrated $7.95 1 livery. Same Vou, 55 APRIL-JUNE, 1965 No. 2 American Fern Journal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY ts IRA L. WIGGINS C. V. MORTON ROLLA M. TRYON JOHN H. THOMAS A Parcel of Cameroon Ferns. ALICE F,. TRYON 49* Cup Ferns (Dennstaedtia) Cultivated in California BARBARA JOE 58 A Pteridophyte Botanical Garden. HeLen Ross Russent = 63 The Essentiality of Boron i~ ae on dentata and Selagi ON BowEN AND HuaH G. Rite 67 Preliminary Field Studies on the Fern Flora of ome Mexico HN T. MICKEL 74 Shorter CSacosrarg Return to Panther Creek; Schizaea pusilla— mnial Gametophyte?; Osmunda regalis var. spectabilis in ‘Sudiny Cystopteris bulbifera (L.) Bernh. New to Texas 80 Recent Fern i i 85 i ie i ee 94 Missour: Bosrican Fern Society Re a cc I IUN 18 y9g6 Che American Fern Hocietp Council for 1965 OFFICERS FOR THE YEAR Donovan S. CoRRELL, Texas Research Foundation, Renner, Texas sident IRVING ba Kwnosiocu, Department of espe & Plant Pathology, Michigan State University, East Lansing, ice-President ae °R. ATKINSON, 415 s. Pleasant — Amherst, amma 50 cretary RicHarp L. HauKE, weirs of Botany, University of Rhode _—— Kingston, Rhode I and Treasurer Ira L. oye dlg Dedler Herbarium, Stanford University, “Eaiorn alit ie Watter H., Hopae, National Science Foundation, Washingt Repre aa oh ‘to AAAS, WakrEN H. Waener, Jr., Dept. of Botany, University of erence Ann Arbor, Mich. Representative to A. I. B. 8. OFFICIAL ORGAN American Fern Journal EDITORS Tra L, \ essa ....-Dudley ces tertcan: Stanford University, Stanford, ee Vs MORON so ode hsonian Institution, Washington 25, D. C. Rotia M. Teron Gray Herbarium, Harvard University, Cambridge 38, Mass. Joun H. THomMas... a Herbarium, Stanford University, Stanford, Calif. An illustrated apc devoted to the general study of ferns, owned by the American Fern Soc and Seer at 3110 Elm Ave., Baltimore 11, Md. Seccndehiad. po gi " paid at B , Md. of 0 , $4.00; susta bership, $8.00; life membership, $80.00). Ex actu ‘reprints if ordered in advance, will be furnished authors at cost, plus postag Back rep $5.00 to $6.25 each; single back Sua of 64 pages or less, $1.25; 65-80 ee 2. 00 each; over 80 pages, $2.50 each; Cumulative Index to Volum 1-25, cents. Ten percent discount on orders of six volumes or Changes of siiens, applications ~~ membership, subscriptions, orders for te —* and other business communications should be aa. to the Treasury, Dr. = hard L. Hauke Departmen * of Botany, Univers of Rhode Saeed. Kingston, Rhode Island. Member - subscribers peer! allow two months for aac in address to take e oo AND nb ce OF cho Emnenysrres Dr. W. gner, Unive ity of Michi Ann Arbor, Mich. A regular loan hinartoaak is pole tained in connection with zo library and herbarium. Members may borrow books and specimens at any time, the borrower paying all postal or ag Meese pearl The pages of the . S carnal also are open to members who wis range exchanges; a membership list is published at Reavy to assist pti latareatad in obtaining specimens from different localities American Fern Fournal Vou. 55 APRIL-JUNE, 1965 No. 2 A Parcel of Cameroon Ferns ALICE FF’. TRYON Botanically, Africa is one of the brightest of continents, for collectors and botanists from many nations have worked on the flora over 300 years. American botanists! have done relatively little in the area, but speed and convenience of modern travel grant new opportunities to see the unique vegetation of this continent. Our opportunity came with an invitation to atten the 50th Jubilee celebration of the National Botanic Gardens of South Africa, at Kirstenbosch, near Cape Town in South Africa. The program included a 3200 mile trip to the Cape Peninsula, the Podocarpus forest at Knysna, the Karroo deserts, and the eastern provinces of Natal and Transvaal. We combined with this a trip to Mount Kenya to observe the montane flora of east Africa. We made several trips out of Nairobi and camped on Mount Kenya with Dr. and Mrs. B. Verdcourt. His work is primarily on flowering plants but he has published also on mollusks and on some African species of Adiantum. We visited Cameroons Mountain, the highest land mass in west Africa, to sample the montane flora of west Africa. Schelpe (1951) pub- lished a paper in this journal on the Pteridophyta of Mount Kenya and is currently preparing a treatment on the ferns of South Africa. An account of the ferns of former French Cam- eroun has been published by Mme. Tar dieu-Blot, H. Nicklés and 1 Two members of the a hr S aol aE collected and Piegpr has West African plants. Mrs. Win ek Harley (19 a as published an illustrated Handbook of Libe oa i pon and Pr of. J. T. Baldw rin traveled extensively in Liberia in 1947-48 and has Rar gre al aig on native and cultivated plants relating tbe his travels in West Afri Volume 55, No. 1, of the JOURNAL, pp. ae was issued Pa 25, 1965. 50 AMERICAN FERN JOURNAL H. Jaeques-Felix (1949). We collected on Cameroons Mountain during the last week of October and the first in November of 1963. To my knowledge, my husband, Rolla Tryon, and I have been the first American botanists to collect extensively on the mountain, although David Fairchild visited it on one of the Armour voyages in 1927. There are about 119 species of pteridophytes reported from the mountain and our collections, deposited at the Gray Herbarium and at Kew, include nearly half of them. The treatment on the Ferns and Fern-allies of West Tropical Africa, prepared by Alston (1959), which is a supplement to the second edition of the Flora of West Tropical Africa, has been my most useful reference and I have depended much upon it for ranges and records from Cameroons Mountain The Federal Republic of Cid soon is centrally located at the sinus on the west African coast. It is one of the newly indepen- dent countries formed from a British mandate, south of Nigeria and a somewhat larger French mandate, north of Gabon. The mountain is a few miles inland from the coast of the Gulf of Guinea. It is a massive form reaching 13,350 feet altitude with a somewhat elongated base about 28 by 30 miles. Meteorological observations are practically non-existent on the mountain itself but at the coast the monthly average temperature ranges between 70 and 90° F. Snow may occur on the upper slopes of the mountain but does not persist. Cameroons Mountain is in the belt of converging trade winds and an average annual rainfall of some 10 yards, one of the heaviest in the world, is reported at the coast near the mountain. At Buea, 3000 feet on the moun- tain, the average rainfall is 114 inches annually and 122 inches is reported for the summit. Cameroons Mountain is the only active member in the chain of voleanie mountains extending from Lake Chad, in interior Africa, through Cameroon, and out into the Gulf of Guinea to include the islands of Fernando Po, Prin- cipe, Sao Tomé and Annobon. The most recent eruption 0¢- eurred in 1959. It is estimated that the eruptions began in the Cretaceous and about 300 cubic miles of material has been CAMEROON FERNS 51 ejected. A Plio-Pleistocene basaltic layer about 1,000 meters thick caps the summit and slopes. Eight eruptions have been reported since 1800, the most spectacular of these was in 1922 when six craters erupted, forming a lava-flow 3000 feet wide which produced great clouds of steam as it entered the Gulf of Guinea. There are some accounts of early Greek and Portuguese voy- ages down the west African coast as far as Cameroon but there is no formal record of explorations near the mountain until 1841. The first ascent was made by the botanist, Gustav Mann, of Hanover, sent out by the British Admiralty. He collected first on Fernando Po, and made the remarkable discovery of Podo- carpus on the Island of Sao Tomé which was the first record of the Coniferae in west tropical Africa. Mann began his ascent of the mountain in December 1860 and part way up was met by the party led by the British explorer Richard Burton, the Bap- tist missionary Alfred Saker, and a Portuguese judge named Calvo. Together they reached one of the summit peaks, which they named Mount Albert, in January 1861. Mann botanized in the region for the next three years and collected about 3000 flowering species and these specimens came to the attention of Sir Joseph Hooker, Director of the Royal Botanic Gardens at Kew. The vegetation of Cameroons Mountain is rather broadly de- lineated by R. W. J. Keay (1955) and includes the following three major vegetational zones. Montane Forest extends up to 6000 feet, the upper portion of which is eovered for long periods by clouds and is characterized by many epiphytes. Montene Woodland occurs between 6000 and 8500 feet, is drier and much of the zone is occupied by grassland which has been affected by burning. Above this is a transition to Montane short-grassland occurring above 10,000 feet. The woodland grass areas are of much the same composition as the lower portion of the short- grassland, Our collections are from the southeastern side of the mountain, largely near Buea at an elevation of about 3000 feet, and following the road to the high frequency station at 8500 aye AMERICAN FERN JOURNAL feet. In cleared areas and at the forest edge, around Buea epiphytes were abundant, especially on palm trunks, among them Oleandra distenta, Polypodium (Phymatodes) Scolopendria and Asplenium homuittnie: Selaginella Soyaurti was weedy in gar- dens and appeared especially aggressive in taro patches. Pityro- gramma calomelanos var. calomelanos, an introduced American species, was planted as an ornamental and was spontaneous along road cuts. We also found it in areas apart from human habitation where it was well established and integrated in the native flora. Our richest collecting was in a ravine about 200 feet above Buea. This was approached along the dry stream bed and was bordered by steep walls of voleanic rock with some open places affording growth of large trees and lianas. Three species of Selaginella occurred together in soil pockets in the rocks, 8. Kalbreyeri, 8. versicolor and 8. Vogelii. Large plants of the lat- ter had a distinctive, irridescent-blue overcast to the delicate sprays of leaves. Robust plants of Tectaria camerooniana grew in the humus near the stream bank and had aves neal two yards long. Another coarsely cut fern here was Coniogramme africana with leaves once pinnate and with less than 10 pinnae— each of these about 10 inches long—and pale, straw colored petioles. The genus is placed near Pityrogramma as the spor- angia are exindusiate along the veins and the rhizome bears seales. It is peculiar in being almost exclusively Old World in distribution except for a species in Mexico. The most imposing fern in the ravine was Marattia fraxinea. Each plant had about a dozen leaves approximately eight feet tall with bright maroon colored petioles striped with white pneumatophores ( ventilating tissues), arising from the massive rhizome. We also collected the rare orchid Manniella Gustavii in this ravine. In open places in the forest at about 5000 feet we found a large colony of the tree fern Cyathea Manniana (Fig. 1). This 1s one of the species in which the base of the scales on the petioles and croziers become indurated into prickles and make collecting difficult. At the edge of the forest at 6000 feet we collected the CAMEROON FERNS Se large epiphyte Drynaria Volkensii. This is similar to Platycerium in habit—the old leaves persisting and forming humus upon which new portions develop. A second epiphyte, a small plant of Aspleniwm, had become established on the Drynaria. Lycopodi- um ophioglossoides also hung from the tree in long, lax, hanks. In the montane woodland at 7800 feet we found Anogramma leptophylla and this is the first record of its occurrence in west Africa. It grew luxuriantly on bare soil under a rock ledge much as we found it on Loma Lachay, near Lima, Peru. It has a relatively wide distribution and Alston particularly noted its absence in west Africa. It might be easily overlooked, for plants usually grow under some rock shelter and mature ones are from 5 inches to less than an inch tall. Except for this and another species on Ascension Island, the species of Anogramma are wholly American. Anogramma leptophylla is reported to be diploid in New Zealand and tetraploid in the western Himalayas. The biology of the species may be unusual for it is reported to produce perennial gametophytes which can survive dry periods and produce lobes from which new sporophytes arise. In the grassland at 8500 feet there were numerous rocks with crevices and soil pockets in which leathery leaved Dryopteris Pentheri and Athyrium Schimperi were abundant and depauperate plants of Pellaea quadripinnata were less common. A few species we collected apparently have not been reported from Cameroons Mountain. In addition to Anogramma, men- tioned above, these are Asplenium hemitomum which grew on the trunk of an oil palm at 3000 feet, near Buea, Ctenitis Bar- teriana on a lava wall in the ravine above Buea, and also in the same locality, Thelypteris cruciata, which is not reported from Cameroon. The first account of the flora of Cameroons Mountain was by Sir Joseph Hooker (1864). He was primarily interested in the geographic relationships of the species occurring above 5000 feet and observed that the montane flora is relatively poor; and that there is a preponderance of Ethiopian (Abyssinian) forms —most of the genera and over half of the species are natives of AMERICAN FERN JOURNAL . AA GROVE OF TREE FERNS, CYATIEA MANNIANA IN RIGHT IN A FOREST OPENING, NORTHWEST OF BuEA, AT 3200 FEET. Fic. 2. (BeLow). ADIA at 2000 FEET IN ABOVE) CENTER, TRUNKS ABOUT SIX TO TEN FEET HIGH, PHILIPPENSE ON A BANK NEAR MU'TENGENE, TUM CAMEROONS. THE CAMEROON FERNS 55 Ethiopia. These and the other relationships he noted have been evaluated by Keay and considered still applicable. His obser- vations on the poverty of the flora and its relationships with the flora of east Africa can be exemplified among the Pteridophyta. In the following paragraphs I have taken up some of the floristic relationships mentioned by Hooker and to illustrate these I have cited examples mainly of the Pteridophyta we studied and collected in the Cameroons. There appear to be no endemic species on the mountain aside from a few, as Aspleniwmn Adamsii, recently described from limited collections and which may prove to be more widely distributed. Frequent vulcanism resulting in extensive burning and the porous volcanic rocks which retain little moisture are undoubtedly factors affecting the poverty of the whole Cameroons Mountain flora. The lower alti- tude of Cameroons Mountain, about 6000 feet lower than the high eastern peaks, Kilimanjaro and Kenya, has also been con- sidered in explanation of the more limited flora. The lesser height may have been critical during warm periods of the Pleis- tocene when the lower summits may have been too warm to sup- port the montane vegetation. The island of Fernando Po about 32 miles southwest of Cameroons Mountain has the nearest high mountain, reaching 9800 feet. Exell (1944) in his work on the islands in the Bight of Biafra drew some interesting comparisons with adjacent areas. He indicated that 160 species not on the other islands occur on both Cameroons Mountain and Fernando Po. One of these, Polystichum fuscopaleaceum, we found growing at 8500 feet on the mountain; it is also reported from Bamenda in the eastern Cameroon highlands. Others, such as Blotiella (Lonchitis) gracilis, Pteris camerooniana and Nephrolepis pumicicola have been reported from the Cameroons and some of the adjacent islands. The slightly richer pteridophyte flora of Fernando Po includes several species that are not on Cameroons Mountain, such as Osmunda regalis, which is widely distributed in Europe, Asia, North and South America and in eastern, southern and western Africa. It occurs in the eastern highlands near Bamenda 56 AMERICAN FERN JOURNAL in the Cameroon and on Fernando Po but it has not been re- ported from Cameroons Mountain. The floristic relationship that Hooker remarked upon as strong- est is with Ethiopia (Absyssinia). This, and relationships with the east African mountains, are shown by many genera of africana, which also occurs in Kenya, Diplazium zanzibaricum from the east African mountains and Zanzibar, Selaginella Soyauxti which occurs eastward in Uganda and S. versicolor which extends from Sudan to Nyasaland in east Africa. Most of the species we collected on the mountain, among them, Dryop- teris Pentheri, Pellaea quadripinnata, Trichomanes borbonicum, Asplenium abyssinicum and A. aethiopicum, occur also in east- ern and southern Africa. Some species we found on the mountain, as Drynaria Volkensit, Polypodium (Phymatodes) Scolopendria and Asplenium wni- laterale, are widely distributed in the tropics of Africa and Asia. Comparative study of these plants throughout their range is much needed as shown by the work on the Pteris quadriaurita complex by Walker (1958). The name Pteris togoensis is used by Alston for the Cameroon plant of this complex although the relationship of American material is still a problem. Cytological reports have been made for some of the widely distributed species, such as Cheilanthes farinosa, which occurs in Asia, Arabia, east and west Africa, Malesia, Mexico, Central and South America, and plants from east Africa were reported to be dipioid, tetraploid, and apogamous triploids. Adiantum philip- pense (Fig. 2) was reported to be an apogamous triploid in Ceylon and in south Africa, but has not been studied in other areas of Africa or Asia nor in Australia, Mexico, or Central America. The distribution of disjunct species, particularly those showing relationships between Cameroons Mountain and Britain, inter- ested Hooker. He examined the kinds of propagules produced by plants showing these relationships and found them to be structurally adapted for clinging or to have minute seeds. The ~l or CAMEROON FERNS manner of dispersal is one factor of these distributions. Specific ecological and climatic requirements of the plants are likewise involved for such epiphytes as Lycopodium ophioglossioidcs, pendulous on trees usually above 3000 feet, and Trichomancs borbonicum also on trees at higher altitudes, undoubtedly have particular growth requirements. Geological events, as Pleisto- eene glaciation which influenced migrations of floras, and the climatic changes of the period are additional critical factors. These problems of biogeography which concerned Sir Joseph Hooker and his correspondents, including Asa Gray and Charles Darwin, are still live topics. New information on rock mag- netism, migration of land masses, topography of ocean bottems and new records from the radiocarbon dating of fossil material provide additional data. The explanation and resolution of the problems of biogeography will be implemented by these but are also dependent upon our knowledge of the kinds and dis- tribution of ferns, among other plants, and upon our knowledge of the plants on Cameroons Mountain, among other mountains. LITERATURE CITED ApaMs, C. D. 1957. Observations of the Fern Flora of Fernando Po. Journ. Eeol. 45: 479-494. Auston, A. H. G. 1959. The Fern-Allies of West Tropical Africa. Flora of West Tropical Africa. ed. 2. Suppl. EXELL, A. W. 1944. Catalogue of the Vascular Plants of 8. Tomé (with Principe and Annobon). British Museum. London. Haruey, W. J. 1957. Handbook of Liberian Ferns. Ganta, Liberia. Hooker, J. D. 1864. On the Plants of the Temperate Regions of the Cameroons Mountains and Islands in the Bight of Benin. Jour. Proe. Linn. Soe. [Botany] 7: 171-240. Keay, R. W. J. 1955. Montane Vegetation and Flora in the British Cameroons, Proc, Linn, Soc. Lond. 165: 140-148. ScHeupr, E. A. C. L, E. 1951. The Pteridophyta of Mount Kenya. Amer. Fern Jour. 41: 65-74. TarpDIEU-Bior, M., H. Nickiis and H. Jacqurs-Feix. 1958. Contribution a la Flore et a V’Reologie des Fougéres du Cameroun. Etudes Cam. 2: “112. WALKER, T. G. 1958. Hybridization in some Species of Pteris L. Evolution 2: 82-92. Gray Hersartum, HARVARD UNIVERSITY, CAMBRIDGE, Mass. 58 AMERICAN FERN JOURNAL Cup Ferns (Dennstaedtia) Cultivated in California BARBARA JOE! The cup ferns cultivated in southern California are finely divided ferns of large to medium size, and sometimes are used as background foliage. Little known to the gardening public, they may be recognized by the characteristic cup-like marginal structures on the under side of the fronds. The sporangia are borne within these cups. With the exception of Dennstaedtia punctilobula, a temperate North American species, all are trop- ical species. Once these plants are established in gardens they require little care except watering. Today they are most frequently found in older gardens where they were planted before the era of philo- dendrons and other foliage plants of present popularity. The Huntington Estate in San Marino, California, has a well estab- lished bed of D. davallioides in its Japanese tea garden. Denn- staedtias rarely are found in the nursery trade and plants are primarily acquired through the generosity of fern gardeners. New plants are started easily from division of a mature plant. Some gardeners object to the wide-creeping habit of D. cicutarva and D. davallioides. However, I have found them easy to con- trol by occasionally pulling them out of trespassed areas. Dur- ing the winter months these plants undergo a period of rest. They are at their best in the spring and summer months. Al- though capable of withstanding more abuse than many other ferns, their requirements are basically the same; shade, a moist friable ies some protection from wind, and fairly good drain- age. It is doubtful if these ferns wpaad survive in areas with long cool winters. Dennstaedtias in general are large to medium sized ferns with creeping, branched, hairy rhizomes. The fronds usually are broadest at the base and may be once to four times pinnate. The veins are free, forked, and on fertile fronds bear sori at their tips. The indusium and leaf-margin form a cup-like structure 1Mrs. Barbara Joe Hoshizaki. Cup FERNS IN CALIFORNIA 59 holding the sorus. There are about 70 species native to the tropics and one species native in temperate areas. The following key distinguishes the species known to be cul- tivated in California. Hairs present on upper surface of frond; surface not glossy; axis on upper tl of tertiary pinnae not flanked by an herbaceous pleat-like win Many of eet on ie surface gland- -tipped ; frond narrowly deltoid o lanceolate, bipinnate-pinnatifid D. punctilobula Hairs ue gland-tipped; frond deltoid, tripinnate-pinnatifid : Hairs densely concentrated in rachis groove; frond dull green, to 15 dm. long; rhizome 12 em. in diameter or more, 1 cultivated plants mostly wide- -creeping on surface BOLL er ee eee et D. cicutaria Hairs more or less evenly distributed over surface of rachis; rk nd medium green, to 1 m. long; rhizome 5-6 mm. in meter, creeping beneath soetats of soil _ D. davallioides Hairs essentially Sat from upper surface of frond; surface glossy; axis on upper surface of tertiary pinnae flanked on each side by pleat- like wing almost perpendicular to leaf surface — D. bipinnata DENNSTAEDTIA BIPINNATA (Cav.) Maxon. (D. adiantoides (Willd.) Moore) Rhizome about 6 mm thick or more, aS pera beneath the surface of the soil; frond deltoid, 1 m. long, tripinnate; larger tertiary pinnae pinnatifid into 5-7 lobes, each lobe : lapith “toothed, the axis on uppe 4 herbaceous pleat-like wing continuously to the next axis; surfac ronds essentially ee glossy above; sori cylin- drical to a manal borne in sinues of ultimate segmen nts. American tropics, including anes Florida. Semi-hardy. Semi-evergreen. This fern is distinguishable from the other three species by the glossy foliage and the small pleat-like wings on the upper surface of the tertiary axes (Fig. 1). DENNSTAEDTIA CICUTARIA (Sw.) Moore. (D. rubiginosa (Kaulf.) Moore) Rhizome 12 mm, thick or more, in cultivated plants mostly wide-creeping on surface of the soil wom surface in native plants; fronds deltoid, dullish green, to 15 dm. long, tripinnate; larger tertiary pinnae inti pinnatifid into about 11 lobes, each jobe apically bluntly toothed; upper surface of frond bearing whitish or reddish brown hairs, the hairs particularly abun- 60 AMERICAN FERN JOURNAL dant on rachis and in the rachis groove, the main rachis frequently zig-zag; sori borne in sinuses of the ultimate segments, American tropics. Semi-hardy. Semi-evergreen. The long-creeping, thick rhizomes bearing large, somewhat drooping fronds, is the most conspicuous feature of this species. Recently Tryon (1960) has merged D. rubiginosa and D. cicu- taria, stating that insufficient differences exist between the two > FIGURE 1. DENNSTAEDTIA BIPINNATA, PART OF A PINNA. FicureE 2. D. CICUTARIA, PART OF A PINNA, Cup FERNS IN CALIFORNIA 61 to warrant separation. Traditionally D. rubiginosa has been separated from D. cicutaria by the abundance of hairs on the un- der surface of the leaves. whereas such hairs are few on the fronds in D. cieutaria. Our cultivated plant is the one formerly ealled D. rubiginosa (Fig. 2). DENNSTAEDTIA DAVALLIOIDES (R. Br.) Moore. Lacy Ground Fern. Rhizome about 6 mm. in diameter, wide-creeping under the soil; deltoid, medium-light green, to about 1 m. long, tripinnate; leaf blade rachis and major veins with seattered whitish hairs; sori borne in sinuses of ultimate segments. Australia and Tasmania. Semi-everereen. Semi-hardy. 3 FIGURE 3. DENNSTAEDTIA DAVALLIOIDES, PART OF A PUNCTILOBULA, A PINNA. in; larger tertiary pinnae pinnatifid into about 9 lobes, these apically PINNA. FIGURE 4. D. 62 AMERICAN FERN JOURNAL This fern is more delicately cut and with a thinner blade than those of D. cicutaria and D. bipinnata. The broad deltoid frond and gn of glandular hairs separate it from D. punctilobula (Fig. DENNSTAEDTIA PUNCTILOBULA (Michx.) Moore. Hay-seented Fern. Rhizome about 6 mm. in ao ereeping under the soil; frond narrow- ly triangular to lanceolate, to dm, long, eae -pinnatifid; larger secondary pinnae pinnatifid ae about 13 bluntly toothed segments; rachises with acute, simple hairs intermingled with ae pha hairs; sori borne in sinuses of ultimate segments Newfoundland, eastern United States. Hardy. Deciduous. The narrower fronds and presence of gland-tipped hairs readily separates this plant from the other cultivated species. At present I know of no specimen of this species established in southern California, although living plants have been purchased from Oregon and the eastern states from time to time (Fig. 4). Ferns most likely to be misidentified as Dennstaedtia are the closely related Microlepia, which has a hood-shaped indusium that is placed submarginally and opens on the side facing the apex of the segment. Dennstaedtia has a cup-shaped indusium located marginally, and opens at the top. Other species of Dennstaedtia reported to be cultivated in the United States are D. smithii Moore from the Philippines, and D. dissecta from the American tropics. I wish to acknowledge the kind help of C. V. Morton, of the United States Museum, and of R. M. Tryon, of the Gray Her- barium, Harvard University during preparation of this paper. LITERATURE CITED Tryon, R. M. 196 A review of the genus Dennstaedtia in America. Contr. Gray hase 187: 23-52: DEPARTMENT OF Botany, UNIVERSITY OF CALIFORNIA, Los ANGELES, CALIF. PTERIDOPHYTE BOTANICAL GARDEN 63 A Pteridophyte Botanical Garden HELEN Ross RUSSELL When we purchased our home we acquired also a trapezoidal piece of land at the end of the backyard about 65 X 20 «K ‘68: xX 15 feet. The northeast corner of the plot was a good 2 feet lower than the back yard. The southwest corner—6 inches higher. Midway across the plot was a bank with a 45 degree slope. The plot was surrounded by trees on four sides with a dead and decaying wild cherry tree in the center. The ground was cov- ered with vegetation: mostly Broad-leaved dock (Rumex obtust- folius) interspersed with Gill over the eround (Nepeta hedera- cea) and Burdock (Arctiwm lappa) on the higher southern sec- tion, and an almost pure stand of Celandine poppy (Chelido- nium majus) on the lower northern section. Poison ivy flourished along two sides of the property line. It seemed like “a good piece of land to own” because it would serve as a buffer between our property and a potential building lot. But it looked like a miserable piece of land to own if you cared about yards and growing plants. Even after we cut down the old and dying wild cherry trees on our property, pushed over the dead tree in the middle of the plot, pulled the hundreds of celandines, poisoned the poison ivy and dug out the dock, the area looked hopeless. The trees on our neighbors’ properties shaded one part or another all through the day and the terrain was neither suited to gardening nor mowing. So we dubbed it “our woods” and decided to specialize in ferns and fern allies. Our first ferns were common ones of the area. Sometimes they were snatched from destruction by bulldozer and_ backhoe. Sometimes they were carefully selected from abundant patches in woodland areas. At all times they were lifted with soil on the roots and planted in holes well supplied with leaf mold. The second spring we watched Royal fern, Osmunda regalis, Interrupted fern, Osmunda claytoniana, Cinnamon fern, Osmun- la cinnamomea, Lady fern, Athyriwm asplenioides, Christmas 64 AMERICAN FERN JOURNAL fern, Polystichum acrostichoides, Ostrich fern, Pteretis nodulosa, and Senstive fern, Onoclea sensibilis, unfold their various fronds and produce spores along the northern edge and across the center of the area. At the same time Hayscented fern, Dennstaedtia punctilobula, Eastern bracken, Pteridwm latiusculum, New York, fern, Thelypteris noveboracensis, started to compete for space on the one consistently sunny slope at the edge of the yard. Clumps of Rock polypody, Polypodiwm virginanum, grew on two granite glacial erratics. On the siope which divided the area, in the shade of the bush honeysuckle, Maidenhair, Adiantum pedatum and Fragile fern, Cystopteris fragilis grew in neutral soil. We had established 13 species of ferns in four micro-habitats. The first moist, shady and semi-acid, the second sunny and acid, - third rocky and the fourth shady and neutral. Rattlesnake fern, Botrychium virginianum and Northern Beech fern, Phegopteris connectilis had been planted but did not reappear. Since all the pinnae of the rattlesnake fern had been eaten by some insect the previous summer it seemed reasonable to conclude that it did not have enough food reserves to survive. We could not account for the death of two clumps of Beech fern. Out of four species of Lycopodium only L. obscurum remained healthy. We decided that probably soil conditions were at fault here so we dug trenches 18 to 24 inches deep and filled them with leaf mold before reintroducing L. complanatum, L. lucidulum, L. clavatum and L obscurum. All four species are now holding their own, and L. complanatum and L. lucidulum have produced _ fertile strobili and added new growth. That summer we added three species of Dryopteris, the Toothed Wood fern, D. spinulosa, the Marginal Shield fern, D. marginalis, and the Narrow Swamp fern, D. cristata. Then we went to Bucks County, Pennsylvania on our vaca- tion and visited the Wherry Fern Trail. We knew we couldn't grow as many species or have as many micro-habitats as the PTERIDOPHYTE BOTANICAL GARDEN 65 Pennsylvania State Forest had established under Dr. Wherry’s supervision but there were undoubtedly other Massachusetts ferns we could grow. That fall we found Flora of Worcester County, written by Joseph Jackson and published by the Wor- cester Natural History Society in 1909, and learned that at that time, Worcester County, Massachusetts had 38 species and six subspecies of ferns, seven species and two subspecies of Lycopodium and six species of Equisetum. We again tried Northern Beech fern, Phegopteris connectilis and all three clumps flourished. We planted another Botrychium virginianum. It did not grow. The chipmunk who lives in “our woods” dug his tunnel under its roots. We added the Northeastern Bog fern, Thelypteris palustris to the moist, shady semi-acid habitat, as well as the Massachusetts fern, Thelypteris simulata. Equisetum hyemale was added at the fence corner, bringing the count of Pteridophytes to 24. Probably because it is in too dry a habitat EZ. hyemale has increased in size slowly adding six new stems in a three year period and losing two. The next year we collected a Bulblet fern, Cystopteris bulbi- fera, between limestone strata in a New York state gorge. On the same trip we had collected examples of Pennsylvania limestone. When we returned to Massachusetts we set up a new micro- habitat—the first one that was truly foreign to our particular area of the Commonwealth. We decided to use a part of the sloping area for ferns and other plants requiring Or tolerating lime. Under the surface we buried small pieces of limestone. On the surface we placed several pieces of limestone whose ir- regular shapes testified to the dissolving action of water and mild organic acids. We then added a short flight of steps of limestone flagstones. In the curve of one of the rocks we planted C. bulbifera. Be- fore the summer was over we added Maidenhair Spleenwort, Asplenium trichomanes, and Brownstem Spleenwort, Asplenium platyneuron. All three grew well. In two years C. bulbifera has 66 AMERICAN FERN JOURNAL increased greatly in size. Its 40 em fronds are laden with bulblets. About this time we discovered the Garden in the Woods in Sudbury. This delightful spot was founded and is maintained by a retired landscape gardener. Along its five miles of trails flourish hundreds of species of native and exotic plants including a bog area overrun with Climbing fern, Lygodiwm palmatum. We learned that if we returned in spring we could purchase a specimen of this fern. So we again set about creating a new micro-habitat. We dug a hole 214% feet deep, five feet long and three feet wide and lined it with several layers of heavy polyethylene. We filled the hole with black peat from a swamp area which was being drained. On this we planted Sphagnum in which sundews were grow- ing. Our habitat was ready, providing it could survive the activity of the small mammals who thoroughly enjoyed digging and burying things in the damp earth. The next spring we fought succession as we pulled out the oaks and cherries that had been planted there. But otherwise our habitat was flourishing not only with sphagnum and sundews but with a very healthy T. simulata and the Bog clubmoss, Lycopodium inundatum. We made another trip to the Garden in the Woods and re- turned to “our woods” with a climbing fern for the bog and a Walking fern, Camptosorus rhizophyllus, for the limestone slope. At present our seven microhabitats boast 31 different species of pteridophytes. They add charm to our yard, they introduce a note of expectancy and exploration to our travels, they pro- vide an excellent teaching area for college classes and other groups who hear of it and ask to come visit. Someday, if the Climbing fern and some of the less common ferns continue to flourish and multiply, they may serve as a souree of specimens to reintroduce into areas which have lost them through cutting and mismanagement. STATE CoLLeGe, FrrcHpure, Mass. ESSENTIALITY OF BORON 67 The Essentialty of Boron for Dryopteris dentata and Selaginella apoda' ° Joun E. BowEN AND HueH G. GAUCH The essentiality of boron for angiosperms has been recognized for about 50 years, but, despite a diligent search of the litera- ture, the authors are unaware of any investigation of a possible requirement of boron for Pteridophytes. In order to determine whether any of the Pteridophytes re- quire boron, a true fern, Dryopteris dentata (Forsk. ) Cy nr., and a club moss, Selaginella apoda (L.) Fern., were selected. For the former, very small sporophytes, with well-developed roots and a degenerating prothallus still present, were obtained in the University greenhouses. For the latter, terminal pieces of growth, approximately 5 em. in length, were collected, taking care to use pieces whose aerial roots had not made contact with the soil. This precaution avoided the necessity of removing soil particles which might have contained boron. Both types of plants were raised on quartz sand in plastic containers, The sand had been treated with 10% sulfuric acid, and then thoroughly leached with demineralized water and, later, quartz-distilled water. Quartz-distilled water was also used in the nutrient solutions. The cultures were placed under a plastic-covered frame in the greenhouse to increase the hu- midity around the young plants and to prevent possible con- tamination of boron from dust or from the tap water spray ap- plied to nearby plants. The four stock solutions contained 0.5M concentrations of Ca(NO3)o, KH»PO,, and MgSO,7H20, and 500 ppm of boron as H3BOz.. Each liter of nutrient solution contained 2, 0.5, and 0.5 ml., respectively, of each of the first three stock solutions. ' Scientific Artiele No. A1130, Contribution No. 3581 of the Maryland Agricultural Experiment Station (Department of Botany). - Based on a portion of a thesis submitted by the senior author to the ulty of the Graduate School of the University of Maryland in partial fulfillment of the requirements for the degree of Master of Science, 1963. 89 NVOTUAWN VY IVNYNOf Naa FIGURE 1: DRYOPTFRIS DENTATA 126 DAYS AFTER 2” HIGH SPOROPHYTES WERE TRANSPLANTED TO SAND CULTURE ANI GIVEN NUTRIENT SOLUTIONS. ‘TOP ROW: PLUS-BORON CULTURES; BOTTOM ROW: MINUS-BORON CULTURES, ESSENTIALITY OF BORON 69 For each liter of nutrient solution which contained boron, 1.0 ml. of the stock solution of H3BO; was added to give 0.5 ppm. of boron. Trace elements, all as salts of ethylene-diaminetetraacetic acid (EDTA), were added to give the following concentrations for iron, manganese, cobalt, copper, and zine: 5 ppm. for iron and 1.0 ppm. for the other elements. The pH values of both the plus- and minus-boron nutrient solutions were brought to 7.0. Each of the nutrient solutions was stored in a five-gallon polyethylene container and placed in the dark to prevent the growth of algae. Five sporophytes of D. dentata, each approximately 40-45 mm. in height, were placed in sand in each of ten plastic containers, 12 em. square and 16 em. deep. The plants in five containers received nutrient solution with no added boron, and, the other five, nutrient solution containing 0.5 ppm. of boron. A poly- ethylene tray under each container prevented the loss of nutrient solution and any possible contamination of boron from the boards on which the containers rested. Each culture received a total of 100 ml. of nutrient solution per week. Selaginella apoda was similarly grown except that the plastic containers were 21 & 9 em. and 9 em. deep. After receiving the two types of nutrient solutions for three weeks, the sporophytes of D. dentata receiving no boron were Statistically (5% level) smaller than those receiving 0.5 ppm. of boron; from the fourth week on, the difference in heights was highly (1% level) significant (see Table I). During the 15th week after initiation of the treatments, sporophytes receiving no added boron showed necrosis of above-ground meristematic tissues and, during the 18th week, some of the plants receiving no boron were dead or dying (see Fig. 1). By the 8th week after initiation of the treatments, sori were macroscopically visible on the underside of the fronds of plants receiving 0.5 ppm. of boron in the nutrient solution, but they were not visible on plants receiving no added boron. At the end of 18 weeks, there were numerous sori on the plus-boron plants, but very few on the minus-boron ones (see Fig. 2). In addition, AMERICAN FERN JOUR ESSENTIALITY OF BORON fi: the sori of the latter plants had either incomplete or no indusia and the sporangia appeared to be partially aborted. No attempt was made to determine whether the few spores in the aborted sporangia would germinate, and hence a report of their viability is not possible. For Selaginella apoda, receiving the two nutrient solutions for 10 weeks, there was a highly significant (1% level) difference between the lengths (heights) of the sporophytes receiving no boron and those receiving 0.5 ppm. of boron. The minus-boron plants were smaller and were a yellowish-green, as compared with the normal green of the plus-boron plants. The most strik- ing differences, however, between the two sets of plants were the number and condition of the strobili. The number of strobili on the plus- and minus-boron sporophytes averaged 2.7 and 0.8, respectively. The strobili on the minus-boron plants were only about one-fourth as long as those on the plus-boron plants and appeared to have aborted (Fig. 3). Inasmuch as young sporophytes ( Dryopteris) or pieces of sporophytes (Selaginella) were used as starting material, obvi- ously boron was “carried over” at the time of “planting” and placing of- half of the plants of the two species on “minus boron.” If low-boron plant material could have been used as starting material, it is possible that boron deficiency would have shown up earlier, been more severe, and, possibly, would have shown that plants of these two species could not complete their life cycles in the absence of boron. The essentiality of boron was not proven for either species, since no attempt was made to substitute other elements for boron and both species may have been able to complete their life cycles. That is, the spores of both species may have been viable, but —_—_---—— Figure 2: UNDERSURFACES OF LEAVES OF DRYOPTERIS DENTATA SPOROPHYTES N LEAVES FROM PLUS-BORON <0 IN ROW HESE REPRESENTATIVE LEAVES WERE TAKEN FROM THE PLANTS SHOWN IN THE PREVIOUS FIGURE. 12 AMERICAN FERN JOURNAL no attempt was made to germinate them. As with higher plants, however, boron appeared to have special significance for D. dentata and 8. apoda with regard to the reproductive phase of growth. FIGURE fe en )\SE-UP OF STROBILI OF SELAGINELLA SPOROPHYTES RECEIVING am NO BORO EFT) OR 0.5 PPM. OF BORON (RIGHT) IN THE NUTRIENT SOLUTION. rasan caine a STROBILI FORMED ON PLANTS RECEIVING BORON, de peng VERY ABNORMAL, APPARENTLY ABORTED STROBILI FORMED ON PLANTS sedans NO BORON. (TEN WEEKS ON TREATMENTS). ESSENTIALITY OF BORON 73 [aBLE I—AVERAGE HEIGHTS AND STATISTICAL SIGNIFICANCE OF DRYOPTERIS DENTATA SPOROPHYTES RECEIVING PLUS-AND MINUS-BORON NUTRIENT SOLUTIONS Avg. Ht. of Sporophytes (mm) Treatment Statistical Date +B —B significance! 5/5/62 50.9 a ed N.S. 5/12 60.7 58.2 N.S. 5/19 67.5 60.7 . 5/26 72.9 64.9 ** 76.9 67.0 ** 86.3 74.3 ** 6/16 92.9 74.7 “* 6/23 95.3 81.7 ** 6/30 98.8 84.1 “* /7 101.7 86.4 *# 7/14 104.2 87.2 ** 7/21 104.5 91.2 ** 7/28 105.3 95.2 ** + 105.3 - 8/11 106.3 100.7 * 8/18 106.9 102.5 * 8/25 106.9 101.82 * 107.6 101.22 * 1N.S.—Not significant. * Significant at the 5% level. ** —Significant at the 1% level. 2 Decrease in length was due to necrosis and resultant dieback of the meristematic tips of the fronds. EPARTMENT OF BoTANyY, UNIVERSITY OF MARYLAND, COLLEGE Park, Mp. 74 AMERICAN FERN JOURNAL Preliminary Field Studies on the Fern Flora of Oaxaca, Mexico JOHN T. MIcKEL The state of Oaxaca, Mexico, occupies a strategic phytogeo- graphic position. The eastern and western mountains bordering the central plateau of Mexico meet in Puebla and northern Oaxa- ea, and the Sierra Madre del Sur in turn dwindles to 300 feet elevation on the Isthmus of Tehuantepee and rises again to form the mountains of Central America. Plant species migrating along these mountains are funneled through Oaxaca. The great diversity of habitats here is to be equaled in few, if any, other state in Mexico. Oaxaca has slopes facing both the Gulf of Mexico, and the Pacific Ocean, with quite different floras on each. Rainfall in the state varies from about ten inches in the central dry valley near Mitla to over 250 inches in some of the rain- forests on the Gulf slope. Relatively few botanists have visited the state, and most of these have been general collectors. They include Galeotti, Pringle, Reko, Mexia, and Conzatti. Several fern floras have included the ferns of Oaxaca, such as those by Conzatti (1939), Hemsley (1885-1886), Fee (1857), and Martens and Galeotti (1842), but these are very incomplete. Many fern species, for example, are recorded from the neighboring states of Veracruz and Chiapas, but not from Oaxaca. There are good reasons, however, why the plants of Oaxaca have been incompletely collected. The roads have been few and poor until recent years, and the only means of access to most areas of the state has been by foot. In some parts it is difficult for a stranger to engage porters or guides, and bandits have, at times, made travel dangerous. Several recent developments make this a propitious time to concentrate on the ora of Oaxaca. The state is building many new roads, which will be passable most of the year. Airplanes are visiting more and more villages, and there are reports that helicopter service is available to some areas. Fortunately, for the present investigation, [ have a friend, ed FrerRN FLORA OF OAXACA RE: Mr. Boone Hallberg, living in Oaxaca who has been very co- operative in acting as my guide in several areas of the state, and without whose help much of the progress to date would not have been possible. I have begun a study of the pteridophyte flora of Oaxaca. The following is a brief report on a reconnaissance trip made in 1962 to sample the pteridophyte flora of the state. My field work ex- tended northeast approximately 70 miles from the dry area of Mitla (see map), across the continental divide, to a coffee ranch owned by Mr. Hallberg in the rainforest of the Gulf slope, an area with nearly 250 inches annual precipitation. The expedition was supported by the Alumni Achievement Fund of Iowa State University and by a grant from the Society of Sigma Xi. On July 16, 1962, I flew from Des Moines, Iowa, to Mexico City. After spending a day visiting the Instituto de Biologia and obtaining the permits necessary for collecting and sending plants from the country, I made the one-hour flight to Oaxaca. The new airport is quite an improvement over the old one, but it is now much farther from the city, much to the consternation of the Indians who fly into Oaxaca from the outlying districts. The rest of the day was spent in attending to last-minute de- tails for the trip, such as obtaining a raincoat for me (a 6-foot square piece of plastic), bottles for pickling cytological material, and preparing packs and presses for the two boys who were to carry the equipment. The boys, Jacinto and Ernesto, were each to carry a pack of about 50 pounds, for 56 cents per day plus their meals. Late that afternoon we took a bus to Mitla, where we stayed overnight at the Posada La Sorpresa, an inn associated with the Frisell Museum of Zapotec Art. The area around Mitla is quite dry, the tallest vegetation be- ing Agave, Yucca, and various cacti. Even so, as we started up the slopes, we found several species of pteridophytes. Selaginella pallescens, 8S. wrightii, Cheilanthes kaulfussti, and C. myriophyl- la were frequently found, huddled at the bases of exposed rocks. We followed a dusty, dirt road which ran from Mitla east and 6 AMERICAN FERN JOURNAL ri a > p Q o ®o a ; he oo : s - ae ‘ef pe . 2 Saat, a’ 2} fas] Sal = 8 & 8 ~ by 8 = 79 2, 3 I. riparia var. canadensis Engelm. I. tuckermanii Br. In this family four species are known from herbarium speci- mens but the genus is poorly represented in collections. There are only a few specimens of each species. The only specimen of I. tuckermanii seen is in the Gray Herbarium. A majority of the few specimens available are in very poor condition and in- adequate for accurate identification. These four species were recorded from the Diamond Hill area in Providence County by Ernest Palmer (1947). Perhaps because of their aquatic habitat, their resemblance to tufted grasses, and because they are diffi- cult to identify, these plants have been overlooked or avoided by many fern collectors. Ophioglossaceae—Adder’s Tongue Family Botrychium dissectum Spreng var. dissectum _ dissectum f um (Gilbert and wna Weath. B, dissectum f., ea (Muhl.) ooh B, dissectum f. oneidense (Gilbert) C B. Lomas (Gmel.) Angstr, var. ice REE Pease and Moore. Ferns IN RHODE ISLAND 103 . matricariifolium A, Br. multifidum (Gmel.) Rupr. var. intermedium (D. C. Eat.) Farw. . simplex E. Hitche. . virginianum (L.) Sw. Ophioglossum vulgatum L.—Adder’s Tongue Fernald (1950) lists many forms of B. dissectum that Wherry (1961) calls varieties. One of these forms, B. dissectum f. oneidense is considered a species by Wherry (1960, 1961). Studies by Wagner (1959) indicate that B. ternatum, not in- cluded in the above list, occurs in the United States. Since this fern resembles small-lobed forms of B. multifidum var. inter- medium which occurs in Rhode Island, it may be found ulti- mately that B. ternatum also occurs here. Six species and three forms of Botrychiwm have been collected in the state with Provi- dence and Washington counties best represented. I have seen no specimens from Kent County. B. dissectum f. obliquum is the only grapefern known from Bristol County, The only observed specimen of Ophioglossum vulgatum was * collected from Providence County and is in the Gray Herbarium. by by by Fern The Cinnamon, Interrupted and Royal ferns occur in all five counties. O. cinnamomea var. glandulosa was collected in 1906 from Barrington, Bristol County, and according to Fernald (1950) occurs “on or near Coastal Plain, R. I.” If herbarium specimens indicate accurately its frequency, it is relatively rare in the state. These ferns are among the largest that grow within the state. The Cinnamon Fern occasionally attains a height of six feet in swamps and at margins of bogs in Washington County. Schizaeaceae—Curly-Grass Family Lygodium palmatum (Bernh. ) Sw.—Climbing Fern This interesting fern with its twining rachis is known from only two counties. During the past year a sight record was Tre- ported in Washington County but it has not been verified. The 104 AMERICAN FERN JOURNAL most recent specimens from Providence County are dated 1906, while specimens from Kent County were collected in 1920. Polypodiaceae—Fern Fami Adiantum pedatum I Northen Maidenhair Fern This beautiful fern is known in Rhode Island only from Wash- ington and Providence counties. Most specimens were collected in the last decade of the 19th century and probably several of the early stations are no longer in existence. E. J. Palmer (1947) found this species in a few places in rich and moist woods in Providence County. Asplenium montanum Willd.—Mountain Spleenwort A. platyneuron (L.) Oakes—Ebony Spleenwort A. trichomanes L.—Maidenhair Spleenwort The Mountain Spleenwort was reported from Rhode Island first by Champlin (1961). The specimens were discovered in West Glocester, Providence County. It is surprising that a spe- cies (unless introduced) should be discovered so recently in an area in which there has been intensive collecting for many years. Although the Ebony Spleenwort is not recorded from Kent County, it is found so commonly in the other parts of the state that a search in this county may reveal its presence. It is well represented in the herbaria checked. The Maidenhair Spleenwort is an inhabitant of rock crevices in Providence, Kent and Washington counties. The only speci- men from Kent is dated 1897 and the two from Washington County, 1908, Athyrium filix-femina (L.) Roth var. asplenioides (Michx.) Farw.— y Fern A. filiz-femina var. michauxii (Spreng.) Farw.—Lady Fern A. thelypteroides (Michx.) Desy.—Silvery Athyrium The first named of the above ferns is maintained as Athyrium asplenioides (Michx.) A. Eaton, and the second as A. angustum (Willd.) Presl., by Wherry (1961). The latter name is found in many of the early fern lists, including that of E. J. Palmer (1947). Fernald (1950) deseribes this species as “discouragingly variable” and numerous forms of var iety michauzii are labeled in FERNS IN RHODE ISLAND 105 the herbaria. Among these are f. clatius, rubellum, elegans and laciniatum. The variety michaucit is widely distributed in the state, having been collected from all counties and from Block Island. Only a few specimens of variety asplenioides were ob- served, none from Kent or Bristol. The Silvery Athyrium is known from Providence, Kent and Washington counties. Camptosorus rhizophyllus (.) Link—Walking Fern The Walking Fern is rare within the state, known only from the towns of Lincoln and Smithfield in Providence County. In the five herbaria checked, the most recent specimen was collected in 1910. Cystopteris bulbifera (L.) Bernh.—Bulblet Fern C. fragilis (.) Bernh.—Fragile Fern C. fragilis var. mackayi Lawson Two specimens of the Bulblet-fern were observed from Provi- dence County. One was collected on September 18, 1899, from “Dr, Field’s Grounds,” indicating that it may have been culti- vated in his garden. The other specimen was collected on July 9, 1873, in East Greenwich, Kent County. Whether or not this fern may still grow native in the state is uncertain. Numerous records of the Fragile Fern are known from Providence and Newport counties but only one from Washington County. Dennstaedtia punctilobula (Miehx. ) Moore—Hay-seented Fern This fern, growing on Block Island and in all five counties, 1s one of our most common ferns in pastures and open woodlands, frequently spreading rather rapidly along stone walls and edges of woods. Dryopteris cristata (L.) Gray—Crested Wood-Fern D. disjuncta (Ledeb.) C. V- Mort.—Oak Fern . hexagonoptera (Michx.) Christens.—Broad Beech-Fern . marginalis (L.) Gray—Marginal Shield-Fern . simulata Davenp.— Massachusetts Fern . spinulosa (O. F. Muell.) Watt—Spinulose Wood-Fern spinulosa var. intermedia (Muhl.) Underw. ae . thelypteris (L.) Gray var. pubescens (Lawson ) Nakai—Marsh or Meadow-Fern See eS Soe = D S ° 8 ~ o 2 ” on ae > : Ww Q = 3g fe pai oO =] 7 Co = 99 ies) is*] is] oO ca 7) oOo 4 106 AMERICAN FERN JOURNAL Hybrids She clintoniana X cristata Wherry D. cristata * intermedia D ] D. cristata * marginalis Beek: D. intermedia * spinulosa Wherry The Crested Wood-Fern has been found in all five counties. The only three specimens of the Oak-fern observed were from Washington County. Wherry (1961) and Gleason (1952) ac- cept the name Gymnocarpium dryopteris (.) Newman, but Fernald (1950) retains the genus Dryopteris. Nomenclature of the Broad-Beech-fern is also questioned and this species is as- signed to the genus Dryopteris by Fernald (1950), to Phegop- | teris by Wherry (1961) and to Thelypteris by Gleason (1952). — Herbarium specimens of the Broad Beech-fern are known only from Providence County. Specimens of the Marginal Shield-fern have been collected in all counties, with f. elegans (J. Robins.) F. W. Gray known from Providence County. One specimen is filed from Bristol, three from Newport and twenty-four from Providence County. Examples of the New York Fern, a relatively common inhabi- tant of acid woods and boggy or swampy places, have been col- lected from all five counties. Both Gleason (1952) and Wherry (1961) assign this fern to Thelypteris. The Long Beech-fern, known from Providence, Kent, and Washington counties, is included in Dryopteris by Fernald (1950) but has been placed in Phegopteris by Wherry (1961). Gleason (1952) includes both of the Beech Ferns in Thelypteris. The Massachusetts or Bog Fern is known from all five counties and Block Island and is well represented in the various herbaria. Both Gleason (1952) and Wherry (1961) assign this fern to Thelypteris. Several varieties and/or hybrids of the Spinulose Wood-fern occur throughout much of the state. Dryopteris spinulosa var. spinulosa grows in all five counties. The most frequently en- countered variety, intermedia, is given species status by Wherry (1961). All except Newport County and Block Island are rep- Ferns IN RHODE ISLAND 107 resented by specimens in the herbaria. The Marsh or Meadow-fern is represented by numerous speci- mens from each of the counties and from the vicinity of Harbor Pond on Block Island. Much confusion exists concerning Dryopteris hybrids and this paper will not attempt to verify the records and names for them. For uniformity the four hybrids that presumably occur within the state are listed under the names of the two parents although not all of these are included in Fernald (1950). The hybrid D. clintoniana X cristata was found in Washington County and identified as D. clintoniana var. australis Wherry. Fernald did not recognize the species clintoniana but considered this a variety of D. cristata. Today this fern is thought to be a hybrid between clintontana (hexaploid) and cristata (tetraploid ) and has been given the epithet D. X australis (Wherry, 1961). However, the absence of one of the parents of D. clintoniana, namely D. goldiana, from Rhode Island raises doubt in the mind of the author that this hybrid actually occurs here. Perhaps on the basis of glandular indusia and other similarities, these specimens may prove to belong to the next hybrid. Boott’s Shield Fern (D. cristata X intermedia), one of the commonest and best known of the Dryopteris hybrids, is inter- dia and D. cristata and probably occurs over much of the state ents are found. It was recognized by Fernald as D. X bootiit (Tuckerm.) Underw. Records are known from all counties ex- cept Newport. The only specimen of D. nae Wherry was collected in Bristol County in 1894. Two specimens collected by J. F. Collins from Barrington in Bristol County and labeled D. spinulosa var. f ructwosa (Gilbert) Trudell, appear to be the hybrid, D. intermedia X spinulosa, or D. X triploidea Wherry, since upon examination most of the spores appear to be aborted. Wherry (1960) notes that most of the specimens in herbaria labelled “fructuosa” are merely lux- cristata X marginalis or D. X slosso- 108 AMERICAN FERN JOURNAL uriant intermedia but this is presumably not the case in this in- stance Onoclea sensibilis L.—Sensitive Fern Dr. Wherry prefers the common name “Bead Fern’ which refers to the bead-like fertile pinnules rather than the more familiar name (at least to the author) of Sensitive Fern which is in reference to the blackening of the fronds at first frost. This fern occurs in all five counties and on Block Island. Pellaea atropurpurea (L.) Link—Purple Cliff-brake The Purple Cliff-brake, one of the rare ferns in the state, is known only from limestone cliffs in Limerock, Providence Coun- ty, where it was collected in 1906 and 1913. Polypodium. virginianum L.—Rock-polypod The Rock-polypody or Rock-cap fern is known from all coun- ties, where it occurs chiefly in humus on rock outcrops. Polystichum acrostichoides (Michx.) Schott—Christmas Fern The Christmas Fern is common in some of our wooded areas and specimens have been collected from all counties and there are representatives in all five herbaria. Newport County is repre- sented by one specimen from the town of Portsmouth and Bristol by one specimen from the town of Barrington. Pteretis pensylvanica (Willd.) Fern.—Ostrich F Although included in the genus Pteretis by Heeialg (1950) the accepted name today is Matteuccia pensylvanica Raymon (Wherry, 1961; Gleason, 1952). The Ostrich fern is rather rare in Rhode Tatend, Kiving been collected only from Providence County in 1914 and 1925. Pteridium aquilinum (L.) Kuhn var. latiusculum (Desv.) Underw.— Brake or Bracken The Bracken Fern may cover extensive areas in dry woods and pastures especially following fire or other disturbance. Although this is perhaps one of the more common ferns in the state, it was not represented from aoe County in the herbaria studied. Woodsia ilvensis (L.) R. Br. —Rusty or Fragrant Woodsia W. obtusa (Spreng.) Torr.—Blunt-lobed or Large Woodsi The only two specimens of the Rusty Woodsia seen were col- FERNS IN RHODE ISLAND 109 lected in 1908 from Copper Mine Hill in Providence County and are filed in the Gray Herbarium and Brown University Her- barium. Many collections of the Blunt-lobed Woodsia are known from Providence County but only two sites have been recorded for Washington County and one for Kent County. In 1937 this Woodsia was found growing profusely in the rock walls of a well in Washington County. This well has since been covered over and the station no longer exists. Woodwardia areolata (L.) Moore—Netted Chain-fern W. virginica (L.) Sm.—Virginia Chain-fern Both species of Woodwardia have been collected from all five counties and from Block Island. They are especially prevalent in the. Great Swamp and the cranberry bogs in Washington County. The Netted Chain-fern is placed in the Genus Lorin- seria by Wherry (1961) but is included in the Genus Wood- wardia by both Fernald (1950) and Gleason (1952). TABLE 1. County Distribution of Ferns and Fern Allies in Rhode Island ea COUNTIES Sk wW Bi Be EQUISETACEAE Equisetum arvense var, arvense x ipa ies age a E. fluviatile x . E. hyemale var. affine = na - E. sylvaticum var. pauciramosum x x E. sylvaticum f. multiramosum x LYCOPODIACEAE Lycopodium clavatum var. clavatum See sy L. clavatum var. megastachyon x : L. complanatum var. flabelliforme 4 See . Ae L. inundatum (?% var. x * ag L. lucidulum alee oats . L. obscurum “Dg ee : L, obscurum var. dendroideum x zs L. tristachyum , ae a SELAGINELLACEAE Selaginella apoda et oe . S. rupestris x ¢ z 110 AMERICAN FERN JOURNAL ISOETACEAE TIsoetes engelmannii x I. muricata I, riparia var. Yr I. riparia var. canadensis x I. tuckermanii x OPHIOGLOSSACEAE v elt aaipet dissectum var. dissectum . diss m f, elon. ngat um lane eolatum Mh angustisegmentum matricariif oliu multifidum var. asian by by by Be by By by by Rm virginianum Ophioglossum vulgatum HARK KKK KK OSMUNDACEAE riigteneat cinnamomea x 0. meee var. glandulosa 0. eatin x O. regalis ie spectabilis x SCHIZAEACEAE Lygodium palmatum x POLYPODIACEAE eacesinnl pedatum Asp A, eaate fliz-femina @ var. michaux ide Camptosorus rhizophyllus Cystopteris coun ifera C. fragilis r. fragilis C. feast lis mackay Viowistasaila \ punetilob ita Dryopteris cristata mt Saal >s a) = = es HMMM MM KBR MK S - 8 Q =‘ Sr aS i) 3 Ss nw spinulosa var. pus inulosa nulosa var. intermedia iaipvioie ti var. lpehesien ns HR KH AK KH ww HARK HK KH Kw KK a a a | es KR A RK Ferns in Ruope IsubAND 111 Dryopteris iota ee cristata ¥ D. cristata X inter Coe er ee ok De wipsiches x marg site x D. inter vedia x spinulosa x w 4 w tal “ ~ Ono tie sensibilis Pellaea ibaa trate P odi S \ olypodium virgini OAS Ue a me Polystichum cerostchoides eer tes chai Cutan shee. | Pteretis pensylvanica x Pteridiu a aguitinn var. latiusculum x > ee 4 Woo asia ee x W. obtu ee ae Woodwardia areolata Beek a eo ee ee W. virgin gg ge i eg Te “Not listed by Bailey and Collins (1893) Sumary.—Exclusive of varieties, forms, and hybrids, the ex- isting herbarium specimens represent 16 species of fern allies and 40 species of ferns. These records are summarized by counties as follows: Fern Allies Ferns Providence 16 39 Kent 9 24 Washington 11 29 Bristol 2 19 Newport 8 19 There are known collections of all ferns and fern allies from Providence County except Dryopteris disjuncta which has been collected only in Washington County. In order to have as com- plete a list as possible, | would appreciate hearing of additional verified reports of county records of ferns and fern allies within the state. LITERATURE CITED Baruny, W. W. 1893. Notes on the Flora of Block Island. Bull. Torrey lub 20: 227-231. 1895. Among Rhode Island Wild Flowers. Preston and Rsecile Providence, R. I. , and J. ¥F. CoLLans. 1893. A list of Plants prey on Block Is- land, R. 2 in July and August. Bull. Torrey Club : 231-239 Bennert, J. 1888. Plants of Rhode Tsland. ids Press, Provi- dence, R. " 128 pp. 12 AMERICAN FERN JOURNAL BuakKeE, S. F. 1950. State and Local la Floras of the United States. ib! emia I. Amer. Fern Jour. 408 148-165. Braun, E. Lucy. 1950. Deeidndus peers - Eastern North America. The Blakiston Co., Philadelphia. 596 pp. 9 CHAMPLIN, RicHarp L. 1961. A Fern New to Rhode Island. Rhodora 63: 46 CHURCHILL, J. R. et hy 1928. Reports on the Flora of Massachusetts I. Rhodora ne 933: hock on the Flora of Massachusetts II. Rhodora 35: 351-359. DoE, ELEAZER, J., ed. 1937. Botanical Club XIv +35 FENNEMAN, N. M. 1938. Phosiograry of Eastern United States. MeGraw- Hill Book nei shone .. New venrneee MM Ee aoe Flora of Vermont. 3rd rev. ed. Vermont . Gray’s ‘lean of ee 8th ed. American Book Co., oa oe p. Bika siee. ae A, 1952. New Britton and Brown Illustrated Flora, vol. I, pp. 1-57. atgite Gardens, New York. GRAVES, C. B. al. 1910. Catalogue of the Flowering Plants and Ferns of eihadioee Growing without Cultivation. Bull. Conn. Geol. and Nat. 14. 56 pp. Hareer, E. B. et al. 1930 (1931). First Supplement. Additions to the Flora of Connecticut. Bull. Conn, Geol. and Nat. Hist. Surv. 48. pp- Nogug, G. 1920, Ferns, Fern Allies and Flowering Plants of Rhode Island. Snow and Farnham Co., Providence, R. I. pp. 2-4. OapeN, Epiru B. 1948. T Maine. University of Maine Studies. N Sacarueal EpitH, 1947, The Fern and rape Allies of New Hampshire. New Hampshire eure Sci. Bull, No. , H 19 American ‘rape rns Resembling oa ternatum: a Pram ary Report. Amer. Fern Jour. 49: 3. . Progress in the nae of Dryopteris Hybrids. Amer. -92. ————. 1961. The Fern Guide. Northeastern and Midland United oor Canada. Doubleday and Co., Inc., New York. 318 pp. Wrieut, K. E. and Dororny CRANDALL. 1941, Rhode Island Ferns. Torreya 3-7 DEPARTMENT OF BIOLOGY RaANDOLPH-Macon Woman’s CoL- LEGE, LYNCHBURG, VIRGINIA. VERNATION IN CHEILANTHES Lis Vernation in Some Species of the Genus Cheilanthes! Irvine W. KNoBLOCH Circinate vernation, or the coiled condition of the leaves prior to expansion, or within the bud, is a well known characteristic of many species of ferns. Modern work on this phenomenon is summarized by Steeves and Briggs (1958) and by Briggs and Steeves (1959). The coiling is due to more active cell division and cell elongation on the abaxial (lower) side of the blade until the crozier beomes dormant. The unequal growth on the abaxial side may be termed hyponasty. The unrolling of the croziers is due to more rapid and more total growth on the adaxial side of the blade. The unrolling is an epinastic phe- nomenon. Both coiling and uncoiling are independent of light and gravity. Thus mitosis and elongation seem to be the factors involved directly. However, auxins, diffusing from the young developing pinnae, play an important part in elongation. The following available species of the genus Chetlanthes ex- hibit circinate vernation: C. viscida (Fig. 7), C. wrightu (Fig. 8), C. siliquosa (Fig. 13), C. lanosa (Fig. 14), C. californica (Fig. 15), C. feet (Fig. 16), C. intramarginalis (Fig. 17), €. pyramidalis (Fig. 18) and C. parryi (Fig. 21). Other species in this category which are not figured are C. cooperae and. kaulfussii. Non-circinate vernation has been known at least since 1894 (Newman, 1854). This was noted at that time in the bracken fern and has been studied more recently in the same species by Webster and Steeves (1958). In the Ophioglossaceae, one finds non-cireinate fronds in Ophioglossum and doubtfully so in Botrychium. Some species in the Hymenophyllaceae, according 1 Aid from the piace al a (Grant No. G. B. 1851), a contribution from The of the Sigma Xi, and the photographie skill of Philip Coleman are es ised els 114 AMERICAN FERN JOURNAL VERNATION IN CROZIERS OF CHEILANTHES. Figures 1-6, 9-12, 19, 20, ANP 22. NON-CIRCINATE: FIGURES 7, 8, 13-18, AND 21, CIRCINATE. Fic. 1, C. ALA- BAMENSIS, Fic, 2, C. AEMULA, Fig. 3, C. EATONII. Fig. 4, C. MICROPHYLLA. Fig. 5, C. CLEVELANDIL. Fic. 6, C. GRACILLIMA, Fic. 7, C. VISCIDA. Fia. 8, C. WRIGHTII. Fie. 9, C. HORRIDULA, Fig. 10, C. NOTHOLAENOIDES. Pig. di, C. TOMENTOSA, Fig. 12, C. LEUcOPODA. Fig. 13, C. SILIQUOSA. Fig. 14, C. LANOSA. Fig. 15, C. CALIFORNICA. Fic. 16, C. FEEI. Fie. 17, C. INTRA MAR- GINALIS. Fig. 18, C. PYRAMIDALIS. Fic. 19, C, FENDLERI, Fig. 20, C. WOOTONTL, Fic. 21, C. PARRYI. Fig. 22, C. vitLosa. Fig, 23, C, LEUCOPODA. 7 VERNATION IN CHEILANTHES 115 to the 1930 observations of Goebel as reported by Steeves and Briggs (1958), are non-circinate. Du Buy and Neurnbergk (1938) noted the same characteristic in Pteris serrulata(?), P. wmbrosa and P. cretica. Wherry (1926) reported Cheilanthes tomentosa as non-circinate and Weatherby (1926) added C. eatoni. Generally, in this type of development, the blade is not coiled but is directed straight downward, and the upper part of the stipe, at varying distances below the blade, exhibits an approxi- mately 180° bend. It would thus appear that the mitotie activity and subsequent elongation are, in respect to bending and un- bending, more localized in non-circinate than in circinate verna- tion. This has yet to be confirmed. Bean seedlings develop a hook similar to the non-cireinate “bend” in some ferns. Dr. R. J. Downs (1964) of the Crops Research Division, Agricul- tural Research Service, U.S.D.A., quoted the findings of W. H. Klein on this subject, “The opening of the excised hypocotyl hook of beans is brought about by the stimulation of cell elonga- tion on the inside (concave) of the hook and not by cell division, since no mitotie stages have been detected in any prepared slides. The elongation is proportional to the incident energy.” Dr. Downs added his own observations that “Opening and closing of hooks in dicotyledons is controlled by light. Although TAA, GA, and Kinetin can affect the response, there appears to be no interaction between the radiant energy and the additives.” The following available species of Cheilanthes exhibit the non- circinate condition: C. alabamensis (Fig. 1), C. aemula (Fig. 2), C. eatonii (Fig. 3), C. microphylla (Fig. 4), C. clevelandu (Fig. 5), C. gracillima (Fig. 6), C. horridula (Fig. 9), C. notholaen- vides (Fig. 10), C. tomentosa (Fig. 11)—(dark eolor is due to over-fixation), C. leucopoda (Fig. 12), C. fendlert (Fig. 19), C. wootonii (Fig. 20), and C. villosa (Fig. 22). Other non- cireinate species are (. covillei, C. lendagera, C. lindheimert, C. myriophylla and C. pringlei. Cheilanthes villosa does not have, however, as well-developed a hook as occurs in some species. Cheilanthes leucopoda (compare Figs. 12 and 23) actually is 116 AMERICAN FERN JOURNAL non-cireinate but the lateral pinnae develop so rapidly shortly after emergence from the ground that a resemblance to a coil can be noted. The two types of vernation possessed by these ferns is un- doubtedly a fixed genetic response and not changeable by envl- ronmental factors. A scrutiny of both the normal habitats of the ferns and of the creeping or non-creeping nature of the rhizomes, reveals no apparent correlation with the type of vernation. One might suggest that the buds on widely-creeping species would emerge more intact from the ground if the stipe-hook preceded the more delicate frond. It may be, however, that the abaxial bend on a normal circinate crozier is equally effective in pro- tecting the bud against abrasion. LITERATURE CITED Brices, W. R. and T. A. Steeves, 1959. Morphogenetic studies on Osmunda cinnamomea L.—The mechanism of ecrozier uncoiling, Phytomorphology 9: 134-14 Downs, R. J. 1964. (Personal communication). Du Buy, H. G, and E. L. Nuernse owth, tropisms and other ERGK. 1938, Gr Sioveadegth in F. Verdoorn, Manual of Pteridology. The Hague, M. Nijhoff. NEWMAN, races 1854. A history of British ferns. London, John van Ep Voors STEEVES, A. Saal W. R. Briaas. 1958. Morphogenetic studies on Osmunda cinnamonea L.—The origin and early development of vegetative fronds. sciiceih lags 8: 60-72, WEATHERBY, C. A. 1926. Imperfectly circinate vernation in ferns. Amer. Fern Jour. 17: 109-111. Wesster, BARBARA D., and T. A, Sreeves. 1958. Morphogenesis in Ptert- dium aquilinum (1.) Kuhn.—General morphology and growth habit. Phytomorphology 8: 30-41. Wuerry, E. T. 1926. Observations on the Wooly fern. Amer. Fern Jour. 17: 107-109. DEPARTMENT OF BotaNy AND PLaNt PatHoLoay, MICHIGAN State University, East LAnsine, Micu. FERN FLORA OF OAXACA 117 Preliminary Field Studies on the Fern Flora of Oaxaca, Mexico JouN T. MickeL (Continued from p. 79) The city of Villa Alta is located at 4000 feet elevation. The trail from the summit was quite wide and it took little more than an hour to hike down to the village. The next day, Monday, was market day in Villa Alta. This enabled us to obtain extra boys to carry the additional presses brought in by plane from Oaxaca. Mr. Hallberg secured three boys without difficulty, and late in the afternoon we were off on the trail again. This time it was to get across the valley (down 1500 feet and up 2000 feet) on very steep trails to the next and last village of Yetzecovi for the night. This tiny village of 130 people has official jurisdiction over Mr. Hallberg’s coffee ranch. By making this start on the trail to the ranch, we were able the next day to make a more leisurely hike to the ranch, collecting ferns along the way. The trail from Yetzecovi to the ranch rose steadily but not too steeply (compared to the other side) for three hours to the highest point of the ridge at 7200 feet. In abandoned fields near Yetzecovi we passed through a “forest” of Bracken fern, Pteridium, the fronds reaching 10 to 12 feet in height. Pine and oak were still the principal trees on this side of the mountain. The average rainfall was conspicuously heavier here than earlier on the trip, as indicated by the many epiphytes. For example, filmy ferns (Hymenophyllum myriocarpum) made their first appearance. Also species of Gleichenia became common in open areas on the upper parts of the slopes (G. palmata, G. pectinata, G. bancroftii, and G. underwoodiana). When we reached the peak, it was like stepping into another world. There was a nearly constant cover of clouds and the vege- tation showed their effect. The epiphytic mosses and ferns were so dense as to cover completely all trunks and branches. Here the true rainforest began. (Weather instruments kept by Mr. Hall- 118 AMERICAN FERN JOURNAL berg in the next valley beyond recorded about 240 inches of rain per year. ) The woods were dark, cool, and moist. This might truly be called “the fern-hunter’s paradise.” Epiphytic species outnum- bered the terrestrial species by about three to two. The former included several species of Elaphoglossum, as well as Asplenium harpeodes, A. fragrans, large polypodiums (P. loriceum, P. lowei), and many tiny ferns that are easily overlooked among the bryophytes, such as species of Hymenophyllum (H. fucoides, H. microcarpum, H. elegantulum, H. polyanthos), Cochlidium rostratum, Xiphopteris delitescens, Polypodium anfractuosum, P. pilosissimum, and P. cultratum. Conspicuous among the ter- restrial species were several large-leaved ferns: H ypolepis nigres- cens with its lax 16-foot fronds, Polystichum denticulatum, Cul- cita coniifolia, and Marattia weinmanniifolia. The Marattia is quite common throughout the forest. It is called “maiz del monte” by the Indians, who in times of poor corn crops have ground the fleshy stipules of Marattia as a substitute. We searched for the large, ribbon-shaped gametophytes of Marattia but collected by mistake only great quantities of the liverwort Symphyogyna. (Symphyogyna is quite interesting in its own right, however, possessing well-developed tracheary tissue.) Also on the forest floor we frequently saw banks of Sphagnum bearing many spor- ophytes. Another interesting find in this area was the first tree fern of the trip, Cyathea fulva. This species was of particular interest since it had not been collected in Oaxaca since the type collection made by Galeotti, who had also found his specimen in the District of Villa Alta. From the peak of the ridge we continued for about an hour on a gentle downward slope through clouded forest that here in- cluded the peculiar angiosperm Weinmannia (Cunoniaceae ) and the conifer Podocarpus until we came to a small spring, El Pozo. From this point the trail dropped markedly and ranged over steep, wet rock stairs (La Escalera), bare tree roots, and slick, wet clay. It is impossible for any beasts of burden (other than FrerN FLORA OF OAXACA 119 man) to traverse this trail, so all supplies and equipment to come to Mr. Hallberg’s ranch must be carried on someone’s back. This rich valley is the valley of the Yelagago River. The trail ran down the west side of the valley, and after two more hours we arrived in Mr. Hallberg’s ranch. The ranch con- sists of a clearing of several acres, a house and a couple of out- buildings. The ranch house is made of hand-sawed boards. The roof is thatched with bunches of Setaria paniculifera, a broad- leaved grass common in the clearing. The clearing itself is used primarily for the cultivation of coffee, corn, and beans. Here I stayed for ten full days of collecting. Ernesto, who is a Mixe Indian, worked for me full-time, and was an excellent as- sistant. He was quick, eager to learn, and seemed to be interested in the project itself. He remembered just which ferns we did and did not have, and he would go to great lengths to get a new one whether it was high in a tree or hanging from a cliff. When I saw Ernesto again in 1963, he still remembered the scientific generic names that I had taught him. The Yelagago Valley is the richest area for pteridophytes I have seen in Mexico. During my ten-day stay at the ranch I col- lected nearly 100 species that I had not seen previously on the trip. The tree ferns were of course the most conspicuous. (See Fig. 1.) Several of these columnar plants were common, inelud- ing Alsophila schiediana, Cyathea princeps, C. mexicana, and C. jurgensenii. The latter species, known only from Veracruz and Oaxaca, has been collected only rarely. ; Besides the omnipresent Marattia, another member of the Mar- attiaceae, Danaea cuspidata, is common in the woods. The filmy ferns are some of the most interesting and at the same time frus- trating ferns found. I found 14 species of Hymenophyllaceac, largely epiphytes, but they are so abundant and interwined among the mosses on the tree trunks that in spite of my efforts several mixed collections resulted. The genus Elaphoglossum, the stag-tongue ferns, is represented by approximately 15 species, but due to the great difficulty in identifying them, most of them 120 AMERICAN FERN JOURNAL remain unnamed. The other best-represented genera in numbers of species are Asplenium, Thelypteris, and Selaginella. Many of the species which were found can now be reported for the first time for Oaxaca, and several are first records for Mexico. The greatest single range extension found was for Blechnum di- vergens (Struthiopteris exaltata), which previously had not been reported north of Costa Rica. Only two species of Pteris were found. Pteris quadriaurita is common in southern Mexico, but P. podophylla is quite striking with fronds over ten feet long. FASS MER (Fe FIGURE 1. CYATHEA PRINCEPS IN YELAGAGO VALLEY. FIGURE 2. PTERIS PODOPHYLLA, SAME AREA. (See Fig. 2.) Around the edge of the clearing and in exposed areas in the forest the bramble fern, Hypolepis, forms dense thickets. With prickles on the backs of the rachis and costae, this plant is as great a nuisance as Rubus in the northeastern United States. In the clearing itself Pityrogramma calomelanos and Histiopteris incisa are common weeds. Fern FLoRA OF OAXACA 121 One of the most interesting groups of ferns both taxonomically and morphologically is that of the radicant aspleniums, which act like our walking fern, Asplenium (Camptosorus ) rhizophyl- lum. Asplenium rutaceum is common and several taxa ascribed to A. radicans are also found. Plants placed in A. radicans vary in their dissection from once pinnate to nearly four times pinnate, each form having a separate varietal name. Miss Shirley Nordahl of Iowa State University is currently studying this group to de- termine whether this is indeed a single species with distinct vari- eties, or a series of closely related species. The Lycopsida are also abundant in the Yelagago Valley. Seven species of Selaginella have been identified, including the large, plume-like S. martensii ; one or two others are probably as yet un- described. Lycopodium is represented by six species: the terres- trial L. reflecum, L. complanatum, and L. clavatum; the pendant epiphytes L. taxifolium and L. | inifolium; and the erect epiphyte L. pithyoides. I nearly missed finding the latter species. While I was at the ranch, trees were being felled to make new land avail- able for corn. I took advantage of these huge fallen trees (100- 150 feet tall) to explore for epiphytes. Tt was here that I ran across a few plants of L. pithyordes. This species has the appear- ance of a pine seedling; the plant grows erect with a thick, deep red stem and leaves over an inch long that resemble conifer needles. Collecting in the rainforest presented the obvious problem of how to dry specimens. The blotters from my presses had to be dried over a fire each day. The air was far too humid for effee- tive drying of the blotters in the sun (and it was, in any event, a waste of sunny weather not to be out collecting). On August 4, after ten days of collecting in the Yelagago Val- ley, we hiked back to Villa Alta. Ernesto and three other boys carried the dried and living specimens while I hiked ahead of them, collecting additional ferns. After waiting a day in Villa Alta, I took the light plane from there to Oaxaca, and soon de- parted for the United States. 122 AMERICAN F'ERN JOURNAL The pteridophytes in the collection made during this field ex- ploration include over 200 species, most of which were found in the Yelagago Valley. Of this number between 15 and 20 species have apparently not been described heretofore, and include spe- cies in Asplenium, Selaginella, Thelypteris, Grammitis, i a dium, and Diplazium. Since the expedition described above, I now have made subse- quent collections in four other parts of the state: central Oaxaca, on Cerro San Felipe and other parts of the valley of Oaxaca; north-central, north of Oaxaca on the new road through Ixtlan toward Tuxtepec; southeast, on the Isthmus of Tehuantepee and towards the Chiapas border; and south, from Oaxaca to Pochutla on the Pacific Coast. My own collections from Oaxaca now in- clude over 300 species and represent what I feel is still a rela- tively sparse sampling of the state. Most of the previous collec- tions have been from only four of the 30 state districts: Central, Pochutla, Tuxtepec, and Cuicatlan. Ten of the districts appar- ently have never been touched at all. Further collecting in all parts of Oaxaca will undoubtedly increase considerably the num- ber of known species. LITERATURE CITED Conzarri, C. 1939, Flora Taxonomiea Mexicana I. Mexico, D. F. A Fr, A. L. A. 1857. Catalogue basi bs des Fougéres et des Lycopodi- acées du ack Strasbur Hems.ey, W. B. 1885-1886. Fata Centrali-Americana 3. London Martens, M., and a GALEOTTI. 1842. Mémoire sur les Fougéres du ‘Mexi- que. Brussels, DEPARTMENT OF BoTANy AND PLANT PatHouoay, Towa STATE University, Ames, Iowa. Dr. Mickel has found a fern student’s paradise! Many isolated valleys and ranges in the American tropics and Eph oy are pes to yield treasures such as he reports in this aper. May he return to the Oaxacan localities as often as his teaching duties permit, ar Be a more complete listing as soon as he ean make the difficult determinations required.—Editor. ANALYSIS OF POPULATION OF EQUISETUM. 123 . An Analysis of a Variable Population of Equisetum arvense and E. X litorale. Ricuarp L. HAUKE IntRopuction—The first species in the genus Equisetum sus- pected to be of hybrid origin was E. litorale. Ruprecht described it as a species in 1845 from material supplied by Kuehlewein. He noted that it was intermediate between E. arvense and E. fluvia- tile. Laseh independently described the same plant under the name E. inundatum a year later. Lasch, according to Milde (1852) suspected it to be a hybrid between E. arvense and FE. fluviatile, and Milde, after studying its internal structure and noting the constant spore abortion, decided that it was such a hybrid (1851). He later (1867, p. 368) was impressed with how common E. litorale seemed and decided on that basis that rather than a hybrid, it was a species becoming extinct (hence the steril- ity). The acceptance, otherwise, of EZ. litorale as a hybrid be- tween E. arvense and E. fluviatile was general. There were but few exceptions, such as DuBuysson (1888, p. 204) who considered it a true species, and A. A. Eaton (1901), who reserved judg- ment pending further information (p. 84, “At present it is an open question, but those holding the hybrid theory are certainly wrong in saying the burden of proof rests with those who reject it, for hybridity being contrary to the natural mode of propaga- tion, it certainly cannot appertain to those who hold the natural method to prove that it is violated.” John Schaffner (1923) dismissed E. litorale as being semi- sterile shoots of E. palustre and also hybrids between E. palustre and E. arvense, and perhaps between E. fluviatile and E. arvense- He later (1926) decided it was only semi-sterile shoots of E. palustre. Victorin (1927) wrote that “E. littorale est une eate- vorie trés artificielle, une sorte de (depotoir ) taxonomique’”’ into which are placed specimens combining the characters of the three species E. arvense, E. limosum (E. fluviatile) and E. palustre. He later (1935) considered it a hybrid between E. arvense and E. palustre. 124 AMERICAN FERN JOURNAL 6 a Figure 1. AREA IN WHICH EQUISETUM ARVENSE AND E, X LITORALE WERE GROWING. FIGURES 2-3. SprecIMENS COLLECTED IN PLOODED DITCH NEARBY ; 2. E, PLUVIATILE, 134; 3. E X LITORALE, 105B. Figures 4-9. SPECIMENS COLLECTED IN AREA PICTURED: 4-6, E. X LITORALE; 4, Hote, WET WOODS ; 5. 105B2, WET FIELD; 6. 105Bs, DRY FIELD; 7-9. E. ARVENSE; 7. 133B1, WET Woops; 8. 133Be, wer FIELD; 9. 133Bs, DRY FIELD. Ficure 10. Lert, E. FLUVIATILE; MIDDLE, E, & LITORALE: RIG HT, E. ARVEN ANALYSIS OF POPULATION OF EQUISETUM 125 Dr. Irene Manton (1950) showed eytological evidence that EF. litorale is a hybrid, and accepted Milde’s morphological evidence that the parents are EL. arvense and E. fluviatile. I had the good fortune to be introduced to an extensive mixed population of L. X litorale and E. arvense, im an area where E. fluviatile was also present. This population was discovered by Dr. Warren H. Wagner, Jr. Since the hybrid and one parent were present in large numbers over a variety of habitats, it was de- cided to make mass collections and to study them closely. Conse- quently, on June 3, 1957 Doctors Wagner, Aban J. Randeria, Robert F. Blasdell and I went there and collected a series of spec- imens. The analysis of these revealed two noteworthy things: that E. X litorale is indeed intermediate in nearly every char- acter between E. arvense, and E. fluviatile, and that the morpho- logical responses of E. arvense to environmental stress are paral- leled closely by the responses of #. X litorale. The purpose of this report is to elucidate these two observations. DESCRIPTION OF AREA AND POPULATION—The population under discussion was growing in Monroe County, Michigan, just west of Hitchingham Road, 0.2 mi. north of Darling Road. This local- ity is about 3 mi. east of Milan. The area is rather flat terrain, with dry sandy soil, the result of submersion by glacial lakes. Along the road was a flooded ditch inhabited by E. fluviatile and E. X litorale. Across the ditch, there was a dry open field, once planted in beans but later abandoned, according to the owner, because of the persistent growth of horsetails. The field abutted on a wet wooded area (Fig. 1). The two horsetails E. X litorale and EF. arvense grew intermingled in the field and into the woods. It was surprising to find E. * litorale in a dry sandy area but presumably its clonal nature enabled it to invade an area where it would not normally grow. Most surprising was to find that only in the dry part of the area was E. X litorale cone-bearing. Meruops or stupy—Mass collections were made from four dif- ferent habitats within this area, as follows: 134. E. fluviatile in Be I26.-> AMERICAN FERN JOURNAL Table 1. Mean values for ig of FE. X litorale and its possible parent Number (except arvensée % litorale* fluviatile palustre where noted ) 42 80 15 5 pert ee ae a RN SR em es Length 2.8 4.9 6.0 9 Sheath idth 2.9 3.9 5.8 4.7 Number 9.6 FS 14.7 8.6 Teeth Length 1.6 2.0 2.4 5 Number 1 fs 23:8° 31.6 — Branched 8.8 10.1° 5.3" 11.3! Inter- Above node branches 6.5” (gre 13.6! — Width 7 2.8 4.9 aH. Length 17.4 21.8 37.9 61 Plant height 216° 400* 1005! 7605 Sheath width ~ Internode width ay gs 1.34 1.19 1,52, Teeth number — Sheath width 3.45 2.96 2.60 1.86 Sheath length ~ Sheath width 0.96 1.26 1.04 1.92 Internode length —. Sheath length 5.99 5.56 6.42 6.90 “except cone- -bearing st a-2] specimene hoe ae a: 73 © 71 £-65 g-74 h-10 i-8 j-4, ANALYSIS OF POPULATION OF EQUISETUM 127 flooded ditch (Fig. 2); 105. E. X litorale, 105b (Fig. 3) with 134, 105 b, (Fig. 4) in wet woods, 105 by (Fig. 5) along edge of woods in wet part of field, 105 bs (Fig. 6) in dry part of field ; 133 EB. arvense, 133 b, (Fig. 7) with 105 bi, 133 be (Fig. 8) with 105 be, 133 bs (Fig. 9) with 105 bs. Specimens were pressed for later analysis. Measurements were taken from these specimens and recorded on duplicated forms. To standardize measurements the following procedure was observed. Sheath measurements were made on a sheath from the lower portion of the stem, but not the lowermost. Sheath length was measured from the base of the teeth to the slight constriction marking where the sheath joins the stem, Sheath width was measured just below the teeth. The base of the tooth was marked by the depth of the sinus between teeth. Internode measurements were made on the internode above or below the nodal sheath measured. Width was measured near the middle. Length was taken from the base of one sheath to the base of the sheath next above. Plant height measurements were taken from ground level, this being judged by the darker color of the below-ground portion of the stem. When all the measurements had been made, means and stand- ard deviations were calculated, and the collections of one species from several habitats were compared. Then an overall average for each species was obtained and these were compared. For eompari- son, a collection of E. palustre from the northern part of the lower peninsula was included in the study. Resuuts.—As Table 1 shows, E. * litorale does indeed fall be- tween E. arvense and E. fluviatile in most of the characters meas- ured, and E. palustre is quite divergent from these. An excep- tion is in the number of nodes bearing pranches, but this can be explained readily by the fact that E. fluviatile is initially un- branched, and becomes more branched as the season progresses. That is, the genus Equisetum is characterized by formation of branch buds at the bases of the leaves (between leaves). In the species which are normally unbranched, these buds remain dor- 128 AMERICAN FERN JOURNAL mant; in the species which are normally branched they develop vomdlly. In E. fluviatile they develop more slowly than in E£. arvense, and many of them remain dormant. Whereas this col- lection averages 5.3 branch-bearing nodes per plant, and 13.6 nodes above the branched nodes, a collection from another locality made on 11 July averaged 11.8 branched nodes and 13.3 nodes above the branched nodes. Two of the ratios calculated, sheath length-width and inter- node length-sheath length reveal that E. X litorale is not inter- mediate between its postulated parents in sheath form or rela- tive sheath length. In each case this is the result of an imperfect intermediacy of the hybrid in the two characters used in the ratio; i.e. it is more like one parent in the first character, but closer to the other parent in the second character. This non- intermediacy of ratios is of interest because in an earlier work, (Hauke, 1963, pp. 88, 90) I postulated that a hybrid might in- herit a ratio (i.e.a form). Rather than the several dimensions of a given plant part being each controlled by a separate set of genes, it was hypothesized that one dimension and a certain form might be genetically determined. For example, one species may have leaves averaging 10 inches long and 5 inches wide, or a ratio of 2/1. Another may have leaves averaging 2 inches long and 4 inches wide, for a ratio of 1/2. A hybrid between them might in- herit an intermediate length 6 inches, and an intermediate form, a 1/1 ratio. It would have leaves 6 inches wide, or wider than either parent. Recent experimental evidence (Haber, 1962; Haber and Foard 1963, 1964) has supported this by showing that the overall form of an organ is under strong morphogenetic control. In the present case, however, E. X litorale seems to have in- herited, not an intermediate form, but rather an intermediate size. One might question the parentage of the hybrid, but considera- tion of the other possible parent, E. palustre, shows that in sheath length-width ratio E. X Iitorale is intermediate between it and E. fluviatile, whereas in internodal leneth-sheath length ratio the hybrid is lower than any of the possible parents. ANALYSIS OF POPULATION OF EQUISETUM 129 Table 2. Comparison of mean values for sterile ae fertile stems of E. X litorale from the dry field ha bit. Number (except Sterile Fertile where noted) 24 25 Sch ci ae cia ERAN eis MARRS URnoA ene en Se ORT Length 4.3 5.2 Sheath Width 3 4.3 Number Lo 11.6 Teeth Length ig 2.8 Number 20.9 11.5 Branched on ie 3.0” Inter- Above ss node branches Bio, 4.0 Width 2.2 2.6 Length 18.2 19.0 Plant height ; 264 165 Sheath width + 1.62 1.69 Internode width ee oe a ee Tooth number > Sheath width vl es ee ele Sheath length take S Sheath width 1.23 1.22 ees Internode length ~ 3.69 Sheath length 4.30 z ian 5 specimens 9 specimens In constructing Table 1, I used only non-strobilate material of E. X litorale. This was done to make it comparable to the postu- lated parents. A separate comparison of sterile and fertile E. X litorale was made (Table 2). Here, for comparative purposes, only specimens from the single habitat containing fertile stems 130 AMERICAN FERN JOURNAL were used. Table 2 demonstrates that the fertile stems have larger sheaths, longer teeth, fewer branched internodes, fewer total in- ternodes, and are thicker than the sterile stems. The sheaths have T 78 . : 3 g7 i 16 Ha Ft Tt x IS s f ¥ E .* : % ae . b bLb2b3 BIB? b3 B iq flu. X lit. arv. ait 10 1 — 9 H 1100 8 000 7 900 bbl b2b3 bl b2 b3 flu. XX lit. arv. rae P 650 L of Ee tI N t 550 7 500 Hy s ingad H 6 IT 400 : § 350 H r 5 T 300 } 250 t* 4 i 200 t 3 " } 150 G 100 < . 50 x . bd blbebz3 wl b2b3 us. X Ate ary. 2 b blb2b3 bl b2b3 : , F flue X lit. ary. - Az. T. : c | at3 we? 2.3 3° Da.o z 1.9 5 / : 8 T 2. Hy wee w 1.5 I ig 13 H 2 ft 1.2 } : wi-l 2 Tio b blLb2b3.ob2 ata D oe s = ih fies F te a uae Tae Table 3. Comparison of various characters of Equisetum X litorale and itats: A. Tooth n node width ratio; flu. = E. ayo from a flooded ditch; X lit. = EF litorale; arv. = E. arvense; b = e habitat as E. fluviatile; bi = wet woods; bs = wet field, partial ane bs = dry, open field. ANALYSIS OF POPULATION OF EQUISETUM 131 fewer teeth per unit sheath, and the sheaths are relatively larger than the internodes. All of these features reveal a tendency toward the dimorphism of fertile and sterile stems which is such a prominent feature of EF. arvense and which is absent in E. flurr- atile. So here again, it would appear that E. X litorale is inter- mediate between E. arvense and E. fluviatile. An analysis of E. X litorale and E. arvense from the several habit-forms' present in the population under study was made. On Table 3 are given the ranges, means and standard deviations for several characters of E. arvense, E. X litorale, and E. fluvia- tile from several habitats. In sheath length, sheath width, and plant height there is a progressive reduction in E. arvense as the habitat becomes hotter and drier. This is paralleled closely by E. X litorale, which however, remains always intermediate between E. arvense and E. fluviatile. In tooth length and number, E. X litorale remains constant (except for specimens from the flooded ditch) even though E. arvense shows a reduction with drier habitat. In the sheath width-internode width ratio E. arvense remains relatively constant, and E. litorale in wet habitats is closer to E. fluviatile, while in drier habitats it becomes progres- sively more like E. arvense. The intermediate morphology of E. X litorale shows up in other ways not apparent in the tables. Whereas E. fluviatile has relatively short, spreading branches and E. arvense has relatively long, ascending branches, E. X litorale has branches about as long as those of EH. arvense or shorter, which tend to spread (Fig. 4). As a consequence, the long whip-like vegetative tip charac- teristic of E. fluviatile frequently shows up in E. X itorale, while the tip of E. arvense is inconspicuous (Fig. 10). The first branch internode is longer than the subtending nodal sheath of the stem in E. arvense, and shorter in E. fluviatile. In E. X : ; ; -pes because since Equisetum is 2 aidnce teas boise it rips coats 18 is assumed here that all of the E, X litorale specimens are from one clone, and hence represent one genetic individual. 132 AMERICAN FERN JOURNAL litorale the lower whorls of branches have the internodes shorter, whereas the upper have the internodes longer than the subtend- ing sheath. The stem cross-sections (Figs. 11-14) also show, as Milde pointed out so long ago, that E. X litorale is intermediate between E. arvense and E. fluviatile. The hybrid has greater variation in stem structure than either parent, particularly in its endodermal pattern. Usually each vascular bundle has its own endodermis, like E. fluviatile, and sometimes there is both an outer and an inner common endodermis (Fig. 12), which is a re- markable approach to an intermediate pattern between the in- dividual endodermis of one parent and the outer common endo- dermis of the other. Discussion.—The quantitative data given above provide de- tailed substantiation of Milde’s (1851) conclusion that E. X Uy COLLENCHY MA NN PARENCHYMA tts CHLORENCHYMA Figures 11-14. DIAGRAMS OF CROSS-SECTIONS OF NON-STROBILATE AERIAL STEMS, DRAWN FROM SECTIONS PREPARED BY GEORGE CONANT, RIPON, WIS- CONSIN; 11. E. FLUVIATILE; 12. E. X LITORALE; 13. E, ARVENSE; 14, E PALUSTRE. ANALYSIS OF POPULATION OF EQUISETUM 133 litorale is intermediate between E. arvense and E. fluviatile. His original hypothesis, that E. X litorale is a hybrid between those two species, is further supported by Manton’s (1950) cytological demonstration of its hybrid nature, and the repeated observation that the spores of this taxon are abortive. Schaffner’s (1923) statement that FE. litorale comprises semi- sterile shoots of E. palustre and hybrids between EZ. arvense and E. palustre, and E. arvense and E. fluviatile is misleading. The name E. litorale designates a specific category of plants set ac- cording to a type specimen. The fact that various specimens have been misidentified as E. litorale does not make them E. litorale. To contend that it represents semi-sterile shoots of E. palustre, as Schaffner later did, is also fallacious. I have shown that popu- lations of Equisetum do exist which are intermediate between E. arvense and E. fluviatile. Manton has shown that such plants have the irregular meiosis characteristic of hybrids. I have seen specimens collected by Kuehlewein at the type locality (“in litore marino prope Ora” 21 Jun. and 3 Jul. 1847. Herb. Norm XIIT n.99. S-PA) whieh he labeled “E. litorale m” and which bear a close resemblance to the specimens from my hybrid population. Consequently I believe that the name EF. X litorale is correctly applied to the hybrid between FE. arvense and E. fluviatile. The confusion between this hybrid and £. palustre is under- standable when one considers that E. palustre itself in some ways resembles E. arvense and in others E. fluviatile, and Linnaeus himself included a specimen in his herbarium under the name E. arvense. As the data above show, however, E. palustre is dis- tinct from E. X litorale, and this hybrid is not intermediate to any great degree between it and EF. arvense. A second conclusion to be drawn from the data presented above is that the size of E. arvense is strongly affected by environment. Thus the range of stem height, sheath length, and sheath width for specimens from within the moist, shaded habitat is completely outside that for specimens from the sunny, dry habitat, and the two collections constitute two distinct statistical populations. 134 AMERICAN FERN JOURNAL Yet, the population was continuous from the woods into the field, showed intergradation, and presumably represented a single clone. Even though superficially quite different, the plants from different habitats were, upon close inspection, found to be quite similar in many respects, such as sheath shape, tooth number, branch morphology, and anatomical structure. In fact, the only characters strongly affected by environment are size and habit, i.e. upright, reclining, or decumbent stems with simple or com- pound branches. Ahernian of the main-stem apex and elabora- tion of the branches is also of frequent occurrence. Identical manifestations of morphological plasticity are exhibited by EL. X litorale. It is indeed unfortunate that these and other obvious morphological responses to environmental conditions should have been named taxonomically, as they have by various authors. The morphological plasticity of Equisetum has long been recog- nized. Alvah A. Eaton wrote (1898, p. 49), “As applied herein, however, the term ‘variety’ includes all forms and monstrosities of a species, without any claim that they are constant,” (1902, p. 43) “all forms being readily accounted for by environment alone, and may be produced from one root-stock.” W. N. Clute stated (1928, p. 47), “None of these [named forms of EF. arvense| seemed fixed, and it is likely that all can be changed to the nor- mal form by a change in the soil and surroundings.” Despite this general recognition of the ease with which Equisetum is modifie by environment, many authors still retain the varietal and form names. This practice should be abandoned since these names are purely descriptive and do not indicate natural taxa. Perhaps we should reconsider Schaffner’s ( 1928) proposal for the use of descriptive polynomials, such as “EZ. fluviatile 1. f1 (fluctuation ) multiramulosum proliferwm polystachyum,” which he considered, however, an exercise in organographic ecology, not taxonomy. “For if taxonomy is not naming and establishing larger and smaller groups which reproduce themselves after their kind, then it has no legitimate basis as science.” ANALYSIS OF POPULATION OF EQUISETUM 135 I thank Dr. Warren H. Wagner, Jr. for his assistance and en- couragement. The work was supported in part by N. 8. F. Grant GB 1703 LIvERATURE CITED CLuTE, W. N. 1928. The Fern Allies. Joliet, Illinois. DuBuysson, M. R. 1888. oma fa at des eryptogames vasculaires d’Eur- ope. I. Equisetaceae. Revue Scientifique du Bourbonnais et du centre de la France. I: 1-7, 38-45, 112-121, 201-210, 245-25 pl. EATON, A. by spn enus Equisetum with h -refaranee to the North American vie ny Fern Bull. 6: 45-59. Bull. 81- 9B. nha aa Equisetum in North America. X. Fern Bull. 10: 43-45. Haper, A. H. 1962. erpnnanlivng: § of concurrent cell divisions for degree of polarization of leaf growth. I. Studies with radiation-induced mitotic inhibition. Amer. Jour. ‘Bot. 49; 583-589, ites A. H. and D. E. Foarp. 1963 PAE com liunt of concurrent cell di- isions for degree of See ization of leaf growth. vidence from Sutrented plants ey om oe ae induced changes of the degree of polarization. A ier 0: ee ere eu Os Further studie s of Oued. aa! ted wheat and their relevance to use of mitotic inhibition for dev iecantia studies. Amer 0 151-159. Jour. Bot. 51: Hauke, R. L. 1963. A ears bikie ee of the genus Equisetum sub- enus Hippochaete. J. Cramer, Weinhe g Manton, I. 1950. Problems of poe ogy Ea evolution in Pteridophyta. Cambridge University Pre Minpe, J, 1851. Uber Equisetum inundatum Lasch. Bot. Zeit. 9 . 1852. Zur sa icke langegesehicht der Equiseten and iGiikony: peen. Nova ‘Acta L 23; 557-612 sie 67. Monograph Equisetoru um. Nova Acta Leop. Me ScHAF PNER, Jd: a4 923. How to oe inguish esr North American species of Bquisetum: eae Fern Jou : 33-40, 67-72. 926. On the trail of ee ae He four thousand miles, Amer. Fern Sey 16: 81-92. 1928. Fluctuation in Equisetum. Amer, Fern Jour 8: 69-79. Mies © VICTORIN, Fri RE MARIE. Nae Les Equisétinées du Québec. Contrib, Lab. Bot. Un niv. Montreal No. 1935. Flore Henson De LaSalle Press, Montreal. DreparTMENt or Borany, UNIVERSITY OF RHODE ISLAND, KiNGs- TON b 136 AMERICAN FERN JOURNAL Recent Fern Literature NOTES ON TWO RECENT BOOKS.—Two books have recently ap- peared which might be of interest. Each, though primarily con- cerned with other things, does make some reference to ferns. The first is FLOWERING EARTH by Donald Culross Peattie. Originally published in 1939, it has been reissued by Viking Press as a handsome paperback with several attractive woodcuts by Paul Landacre. Price, $1.45. As we expect from D. C. Peattie, the book is enjoyable to read, yet instructive. After several chapters about plants in general, such as, “What a plant is,” and “Protoplasm—the body of life,” the author presents a survey of the kinds of green plants. Chap- ter 9, “The fern forests,’ could be more aptly titled “The origin and early development of the land flora.” It actually says little about present day ferns, but presents a fascinating insight into the science of paleobotany. Since this group of plants has had such a prominent place in the fossil record, fern enthusiasts should have some appreciation for the role Fornd played in the history of the world’s flora CHERRY, Euamne C. 1965. Fluorescent Light Gardening. D. Van Nostrand Co., Princeton, N. J. Price, $6.95.—This book con- tains much Denial haloniontion for those interested in indoor gardens. Especially useful to the neophyte may be the rather technical information about fluorescent fixtures and bulbs, and about light. Of particular interest here, however, is the section in which specific recommendations are made for various groups of plants. Ferns are mentioned on pp. 182-183, and apparently F. Gordon Foster is the source for Mrs. Cherry’s information, for she merely repeats what he has suggested concerning sub- _~ temperature, light level, and ferns easy to grow.—RIcH- RD L. Hauke, Dept. of Botany, University of Rhode Island, aeeae Rhode Island. Russian Fern Spore Srupres.— An important series of studies of the spores of Russian ferns, by A. N. Sladkov, which has not been noticed in the Journal is the following : RECENT FERN LITERATURE 137 POLYMORPHISM OF SPORES IN CrETAN Pteris.1—The majority of the spores of Pteris cretica as it occurs in the Caucasus Moun- tains are typically tetrahedral, with a 3-rayed fissure proximally, the distal side being dome-shaped and covered with tubercles of different sizes. However, there are also many bilateral spores with a single fissure, as well as some 4-rayed. This polymorphism is attributed to differences in the reduction division of the spore mother cells. MorPHoLogicaL FEATURES OF THE SPORES OF THE FERNS OF THE SUBFAMILY PTERIDEAR DIELS IN THE Fora or USSR.*—De- scriptions of the exospore, perispore, and epispore of Anogram- ma leptophylla, Pteris cretica, Cheilanthes sp., Pteridium aquil- inum, Coniogramme fraxinea, Adiantum sp., and Cryptogramma sp., with new definitions of true perispore, false perispore (or pseudoperispore), and double perispore. On THE MorPHoLoaicaL SIMILARITY OF DIFFERENT SPORES OF Species or Cryptogramma R. Br. and Botrychium Sw. IN THE FLora of THE USSR.2—The spores of both Cryptogramma and Botrychium are round-triangular, with the distal side dome- shaped and nodular, the proximal side concave, flat, or slightly convex. Details for the identification of three species of Crypto- gramma. and six of Botrychium are given. Typrs of DEFORMATION IN THE FossiL SPORES OF Ferns Hav- Inc A RapiaL SrructuRE.A—Comparison of spores of the fossil Anogrammites imperfectus Bolch and the living Anogramma leptophylla (1u.) Link indicates that these represent close or identical species. It is suggested that another subgroup Hetero- lateritriletes be added to the morphographic system of S. N. Naumova (1937) to include spores with radial structure and ob- 1 Doklady Acad. Nauk SSSR 117 (5): 900-903, figs. 1957. (In Russian) ery oes Pek SSSR 125 (1): 219-222, figs. 1959. (In Russian ) SS De Re eeot RP SSSR 125 (2): 414-416, figs. 1959. (In Russian) ge gecabrag peice PP. TF BSSR 129 (5): 1176-1179, figs. 1959. (In Rus- sian) (AIBS translation pp. 290-292). 138 AMERICAN FERN JOURNAL servable differences in the surfaces of the distal and proximal sides of the spores, as in Anogrammites. SPORES OF THE FERNS OF THE SuBFAMILY PrEeRIDEAE DIELS IN THE Fora or tHE USSR. I. Sertes PrermDEAE—GYMNOGRAM- MINAE’—Descriptions and illustrations of Anogramma lepto- phylla, Coniogramme fraxinea, Notholaena marantae, and Cryp- togramma crispa. SPORES OF THE FERNS OF THE SUBFAMILY PTERIDEAE DIELS OF THE FLora or SSSR. II. Serres Prermear—CHEmANTHINAE.o— Descriptions and illustrations of the spores of Notholaena ma- rantae, Cheilanthes argentea, C. kuhnii, C. pteridioides, C. per- sica, Cryptogramma crispa, C. acrostichoides, and C. stelleri. SPORES OF THE FERNS OF THE SUBFAMILY PTERIDEAE DIELS OF THE Frora or SSSR. III. Serres Prertiear—ADIANTINAE AND PrerwinaE.’"—Descriptions and illustrations of the spores of Adiantum pedatum, A. capillus-veneris, Pteridium aquilinum, and Pterts cretica. Key For IDENTIFICATION OF SPORES OF THE FERNS OF THE SUB- FAMILY PTERIDEAE DrELs IN THE FLorA or THE USSR.8—A key to all the species mentioned in the three papers listed above, plus Pteridium tauricum (Presl) V. Kreez. ON THE Spore TETRADS oF PTERIDOPHYTES.®—Pteridophyta have either tetrahedral or bilateral spores, the type usually being characteristic of a genus or a whole family. The globular spores of Equisetum do not have a suture. Heterosporous pteridophytes _ usually have the spores alike, but in Selaginella the megaspores are tetrahedral and the microspores bilateral. Among tetrahedral spores of Lycopodium and Pteris cretica are oceasionally found 5 Bull, State Univ. Moscow, Ser. VI, 6: 45-62, figs. 1961. (In Russian). 6 Bull. State Univ. Moscow, Ser. VI, 4: 48-59, figs. 1962. (In Russian). ‘Dept. (Ministry) of Higher and Middle Special Education, SSSR Sci. a” Grad. Schools, Biol. Sci. 1961 (3): 112-119, figs. 1961. (In Rus- * Dept. (Ministry) of Higher and Middle Special Education, SSSR Sci. Reports Grad. Schools, Biol, Sci. 1962 (1): 129-134, figs. 1962. (In Rus- Sian), ® Doklady Aead. Nauk SSSR 143 (2): 464-466, figs, 1962. (In Russian) (ATBS translation pp. 436-437), RECENT FERN LITERATURE 139 bilateral spores, and among the bilateral spores of Marattia may be found some tetrahedral ones. Transitional types are possible. Paleobotanical research indicates that bilateral spores are rela- tively more recent than tetrahedral. Tue Pouar AXES OF SPORES AND THE SEPARATION OF THE NUC- LEI IN THE Metosis OF TETRADS OF DIFFERENT TYPES OF FERN Puants.!°—Theoretical and geometrical discussion of the forma- tion of tetrads of fern spores. In the tetrahedral tetrad the pairs of polar axes lie in two mutually perpendicular planes, and any axis may form a pair with any second axis; in the cruciform tetrad the pairs of axes lie the same way but the axes of identi- cally oriented tetrads will be paired; in the isobilateral tetrad all the polar axes lie in one plane and the angle between adja- cent axes is 90°. ON Spore AND PoLLEN MorpHoLocy oF RECENT PLANTS IN THE USSR.1—The study of fossil spores and pollen by statistical methods is a special branch of paleobotany that should be dis- tinguished from the morphological study of recent spores and pollen, which can be called “nalynomorphology.” The n norpho- logical study of fossil spores and pollen can be called “paleopaly- nomorphology.” The study of the external envelopes of spores and pollen can be termed cot seas ret ’__CiypE F. Re&ep, Baltimore, Md. An Amareur’s EvaLuaTion.—The two best-selling, modern fern books for amateurs in Northeastern U. 8. are Cobb! and Wherry.2 The review? of Cobb in this journal was extremely critical. Aside from the illustrations, the reviewer found nothing 10 Doklady Acad. Nauk SSSR 146 (1): 995-228, figs. 1961. (In Russian) (ATBS peat pp. 1121-1123) 1 Repo oviet fangs to the og: eer ec Palynologica ess ence gels Nauk SSSR, Geol. Inst. pp. 5-16, 1962. (In Russian, hee En lish summary . as 1 Bo nak Cobb. ‘A as Guide to the Ferns. Houghton Mufflin Com- pany, ee Mass. 1956 : 6 2Kdga T2) Wo nod The Fern Guide. Doubleday and Company, Ine., Garden it yoy Sn oe aes 161-167. 1956. (See also vol. 47: 28-29, 1957). 140 AMERICAN FERN JOURNAL worthy of praise. The book was called “a disgrace to the Peter- son series of Field Guides, and a serious setback to fern study by the amateur.”” The review* of Wherry was very favorable. oth reviews were made by professional botanists. The worth of a book to professionals is judged differently from its worth to amateurs. The present review (written by an amateur with two years experience using both books side by side in the field) is intended to supplement these professional reviews, for the bene- fit of other amateurs. The first need of beginners is always to learn to recognize the various species. Cobb will easily lead anyone to the identification of most ferns, even rare ones. The figures are (and were acknowl- edged* to be) probably the best done to date. Points of difference between similar-looking species are emphasized, and the features of each species are listed conveniently. However, Cobb is not complete; although rare species are covered, several ferns from neighboring regions that grow slightly inside the region covered are not given full treatment in the text, and only the most com- mon hybrids are even mentioned. On the other hand, Wherry is complete in describing rare species and gives extensive listings of hybrids. However, the novice may find Wherry frustrating at first because of the botan- ical style of writing. Botanical terms are used (and defined in a glossary), and some topics, such as cytology and synonymy, that are given are of little interest to the beginner. A novice who is confronted with Wherry is apt to decide that the study of ferns is too complicated and technical for amateurs, whereas Cobb would never give that impression. Keys are given in both books. Wherry’s key to genera is based largely on the sori. With fruiting material at hand, and with a strong lens, this is probably the most reliable key. Cobb, on the other hand, gives three “keys.” One is a series of frond silhou- ettes, arranged by overall shape and size. Another is based on poor botanical criteria, but excellent ones for the beginner, name- 4 Thid. 52: 89-90. 1962. RECENT FERN LITERATURE 141 ly the size, shape, and cutting of the fronds. The third is based on the sori, and is recommended when fertile material is avail- able. But even this key is better, for the beginner, than Wher- ry’s because it is illustrated. It was said in the review® that Cobb’s “key characters are rather poorly selected and crudely stated.” On the contrary, I believe one of Cobb’s strong points is the keys, which as an amateur I have found more useful (though occasionally less reliable) than Wherry’s key. Wherry lists in one category, “features,” the same information about each species that Cobb lists in separate categories, “style,” “leaves,” “leaflets,” “axis,” “stalk,” “roots,” “fruitdots,” ete. These features, which are so useful for field identification, are better displayed in Cobb, for easier reference. On the other hand, Wherry gives additional information about each species, on its culture, nomenclature, and cytology. And Wherry’s data on range are better presented than Cobb’s. All this evidence leads to my main conclusion: For beginners, Cobb is the easier and simpler book. The serious amateur and the professional, however, may outgrow Cobb and find Wherry more suitable. The two books thus supplement each other, and beginners would be well advised to use Cobb at first and then gradually shift to Wherry (or use both books) if their interest in ferns continues. Much emphasis was placed in the review? on errors in Cobb. Certainly it is regrettable to find such errors, and I hope correc- tions can be made in the next edition. However, these errors do not, by themselves, justify a wholesale condemnation of the book. Paun PENFIELD, JR., Weston, Mass. CoMMENT BY A REVIEWER OF “FIELD GUIDE TO THE Ferns.” — The author generously submitted the manuscript of the above article to me before publication. Having admired the drawings, it was no surprise to me to hear that a beginner was able to identify ferns from the illustrated keys. Had this adjective not applied, that would hardly have been so easy, since Cobb uses familiar words in novel senses and invents etymologically un- 142 AMERICAN FERN JOURNAL sound complex terms. Naturally I cannot concur with the view that “features which are so useful in field identification are bet- ter displayed in Cobb.” His statements are replete with errors and misinterpretations; thus, in the text dealing with the com- mon Christmas Fern there are at least 25 of them. One might wish that the authorship of the Field Guide had been attributed to Mrs. Foster, whose contribution to it is above eriticism.— E.T.W. Notes and News ASPLENIUM RESILIENS IN Utau.—Several vigorous patches of Aplenium resiliens Kunze were found along damp shaded hori- zontal bedding planes of sandstone rocks in the newly created Canyonlands National Park, San Juan County, in southeastern Utah. This is the first record of this fern in Utah. The closest locality from which it has been previously recorded is near Flag- staff, Coconino County, Arizona, about two hundred and sixteen air miles distant. Elsewhere in Arizona it is known from the Blue River, Greenlee County, Chiricahua and Huachuca Moun- tains, Cochise County, along the eastern border of the state, and far to the southwest in the Kofa Mountains of Yuma County. In New Mexico it is known from Santa Rita, Grant County, the Florida Mountains, Luna County, and the Organ Mountains, Dona Ana County, all in the southwestern part of the state. Eastward it ranges from Kansas to Pennsylvania and from Texas to Florida. Other ferns noted in the Canyonlands National Park are Cystopteris fragilis (L.) Bernh., Adiantum capillus-veneris L., Pellaea limitanea (Maxon) Morton, Cheilanthes feei Moore, Equisetum laevigatum A. Br. and Selaginella mutica D. C. Eat. Other species reported from San Juan County which may ulti- mately be found in the Park are Woodsia mexicana Fée, W. plummerae Lemmon, Cheilanthes eatonii Baker and Péllaea fend- leri (Kunze) Prantl, all of which are on the extreme northern AMERICAN FERN SOCIETY 143 or western margins of their ranges ——SeviLLe FLowers, Depart- ment of Botany, University of Utah, Salt Lake City. FERNWoop INcoRPORATED.—Our member, Mrs. Kathryn Boyd- ston, well known to many members through her activities in run- ning the Spore Exchange for several years, has been developing her estate, Fernwood, for a long time. Fernwood is located along the St. Joseph River, a short distance north of Niles, Michigan. It contains a variety of habitats, from low marshes and ponds, to steep, wooded hills. The natural beauty of the area has been recognized by many, and a group has been organized to maintain Fernwood as a permanent nature preserve. It contains 75 kinds of ferns and over 10,000 fern plants. It is planned to develop nature trails and to conduct educa- _ tional programs. Individual memberships to support the project are $5.00 or more annually. Membership dues and contributions are tax-deductible, since Fernwood is a non- profit corporation. Any fern lover who is interested in such a project should write to Mrs. Kathryn Boydston, Route 3, Niles, Michigan 49120. Wanrep.—One of our members, Mrs. R. M. Gunnison, The White Sands, La Jolla, California, is interested in obtaining, through exchanges or by purchase, herbarium specimens of un- usual or uncommon ferns. She prefers them unmounted. If you have any such specimens, please write directly to Mrs. Gunnison. A NEW FERN BOOK Learn of Ferns We Grow by Sylvia B. Leatherman and Dorothy S. Behrends Ferns for mild climate gardens : House Ferns : Spore Culture : Unusual ways to grow ferns : Illustrated with line drawings Price: $3.85 plus 15¢ handling. (Californians add 15¢ tax). Order from:—B & L Books on re 2637 North Lee A South El Monte, ahee dens 144 AMERICAN FERN JOURNAL Tw pocket classics by an mm honorar ue — of the Am n Fern Soc DR. EDGAR x WHERRY THE FERN CUIDE Complete, accurate and con- venient; covering 135. spe- cies of the Northeastern and Midland United States and adjacent Canada; with draw- ings of 135 species by Dr. Janies C. W. Chen. $4.95 THE SOUTHERN FERN GUIDE The only comprehensive, up- to-date guide available t a : Chen and K. C. Y. Chen. $4.95 Doubleday Nature Guides Series at all booksellers ol DOUBLEDAY | Long-awaited... a complete book on native and exotic ferns THE GARDENER'S FERN BOOK By F. Gordon Foster This definitive book on ferns for the horticulturist and nature-lover is also an immense ely practical guid e gardener. It is the first work t complete informatio both native and exotic species | identification rawi d descriptions carefully, simply and peauttully — part ern—in s 1ettes of the entire leaf sketches of its special features. 16 colo ictures show details ot structure habitat and 18 black and white photographs show ferns i There is a wealth of specific information on propagation lighting, soil, potting, fertilizing, and other important pointers s for growing c f and rare ferns—both indoors [Loo ais ~ t. in DENER’S FREE — seal ote - * i D. VAN NOSTRAN tains a Dept. AFJ FERN BOO full ay. Gactaie tables, 120 apescryehe Street, Princeton, N. J. ap a list of ype cores It Please send m THE GARDENER’S he book o FERN BOOK at $7 .95 per copy. Within 10 days I will r eg eRe ouneetantn eee lus small delivery ae or return book (s) ing. and owe nothi Foster has lectured at Stevens Institute of Technology, is a micro- 7 Sg fps ish eteameanee Onto copy specialist with the Bell Tele- ph tori and ives lec- Address ae cade pelen her eae oumennenaeineNan annem eset — tures on ferns to Audubon and City Sta Zip Code. garden clubs, SAVE! Remit “with pene “and we pay de- 224 pages. 6”x9”, Illustrated $7.95 livery. sere return Ph shea ge guaranteed. se BOOKS ON VASCULAR CRYPTOGAMS Recently published or actually in preparation The Ferns (Filicales) By F. O. Bower, 1923-28 (Reprinted 1963), Cloth. Filices of the US-Exploring Expedition (“Wilkes Report’) W. D. Brackenridge. 2 vols. 1854-55 a ready 1966). 366 pages in quarto, 46 plates in folio. Clo A Monograph of the Fern Genus Woodsia y D. F. M. Brown. Beiheft 16 zur Nova Hedwigia. 1964. 164 pages, 40 plates. Mémoires sur la famille des Fougéres B L. A. Fée, 11 parts in one volume. 1844-66 (Reprint ready 1966). Quarto. 990 pages, 181 plates. Cloth. Historische Entwicklung der Nomenklatur und Taxonomie der Gattung Iséetes L. Von H. P. Fuchs. Beiheft 3 zur Nova Hedwigia. 1962. 108 pages, — plates. A Aeotoate Monograph of the Genus Equisetum, Subgenus Hippochaete By R. L. Hauke. Beiheft 8 zur Nova Hedwigia. 1963. 128 pages, 22 plates. Section Complanata of the Genus Lycopodium By J. H. Wilce. Beiheft 19 zur Nova Hedwigia. 235 pages, 40 plates, illustrations. $ 30.00 45.00 15.00 175.00 10.00 10.00 15.00 J. CRAMER e PUBLISHER e 694 WEINHEIM e GERMANY Vou. 55 OcTOBER-DECEMBER, 1965 No. 4 American Fern Fournal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY ts IRA L. WIGGINS Cc. V. MORTON ROLLA M. TRYON JOHN H. THOMAS ts CONTENTS Observations on Some Ferns in Georgia Wi.sur H. Duncan AND DoNALD BLAKE 145 Cytological Observations on Ferns pao Southern Chin _ K. Roy anp R. . Hoirrum 154 Cytological and Morphological Otmeratons on Metaxya ros- trata (H.B.K.) Presl_. _K. Roy anv R. E, Hoirrum 158 Observations on Cijitvsied Fans; po as C. V. Morton 164 Dispersal of Marsilea mucronata by Water Birds YHARLES R. MALONE AND Vernon W. Proctor 167 A New Sexual form of + Pilies glabella var. glabella from Missouri W. H. Waaener, JR., D: R. FARRAR, AND KATHERINE L. CHEN 171° Shorter Notes: Japanese Climbing Fern Spreading in South Caro- lina; A Simple Method for Growing Ferns from Spores - 179 Recent Fern Literature _..—--.------------------ Satie See Cision ana een 181 American Fern Society —....:-./-----—-"-~ i a ee thee wy Vile $8 ee 185 Presotm: Roranrcad mod : lm ch FEB 14 i996 Che American Fern Hocietyp Council for 1965 OFFICERS FOR THE YEAR Donovan S. CorreEvu, Texas Research Foundation, Renner, oe sident Irving W. KNosiocu, gar of Botany & Plant Pathology, Michigan State University, E ing, Mich. Vice-President LENETTE R. ATKINSON, “on : geen Street, Amherst, Massachusetts, tary RicuarpD L, Hauke, Department of Botany, University of Rhode Island, Kingston, Rhode Tslan IrA L. Wiceins, Dudley Herbarium, Stanford University, tinted C aif. wt or-in-Chief WattTer H. Hopee, National Science Foundation, Washin D Repres ada to A a. AS. WarrEN H. WaGNER, JR., Dept. of Botany, University of | Ann Arbor, Mich, Representative to A. I. B. 8. OFFICIAL ORGAN American Fern Journal EDITORS Tra L. honors eis Dudley Herbarium, Stanford University, Stanford, Calif. Vs ORO Smithsonian Institution, Washington 25, D. C. Rotua M, nice Gray Herbarium, Harvard University, Cambridge 38, Mass Joun H. THomas. Dudley Herbarium, Stanford University, Stanford, Calif. An illustrated quarterly devoted to the general study of ferns, owned by the American Fern Society, and published at 3110 Elm Ave., Baltimore 11, Matter for publication should be addressed to Ira L. Wiggins, Dudley Herbarium, S$ anford University, Stanford, Califor ubseription $4.50, exclusive of agency han dling ‘tems : “_ free to — an i e. Back volumes $5.00 to $6.25 each; single back numbers = 64 pages or less, $1.25; 65-80 pages, $2.00 each; over 80 pages, $2.50 each; Cumulative Index to — 1-25, 50 cents. Ten percent discount on orders of six $ or volume Changes of adaress — ions for bee a subscriptions, orders for rin anche , and other business communications should be ~ darts to the Treasury, Dr. Rekaed a eit Deaavann nt of Botany, University of Rhode Island, Kingston, Rhode Island. Me mbers rag subscribers ‘aioald allow two months for changes in address to take eff. oo AND CURATOR OF ts HERBARIUM ; Wee gner, University of Mich , Ann Arbor, Mich regular loan leoasinis nt is maintained m apsetener ae with the library and covhae rium. Members may Pecan Sasi and specimens at a also are open to members who wis to arrange exchanges; a membership list is published at a Ho, to assist those aneaanad in obtaining specimens from different localitie. American Fern Fournal Vou. 55 OcTOBER-DECEMBER, 1965 No. 4 Observations on Some Ferns in Georgia Wripur H, Duncan AND DoNALD BLAKE “Ferns of Georgia” (McVaugh & Pyron, 1951) summarized many of the data on ferns that were known at that time. Other publications with direct bearing on the ferns of Georgia include those of Quarterman (1953), Duncan (1954, 1955, 1960), Snyder (1955), Hutto (1960), Rossignol (1956), Bookout (1960), and McDowell and Snyder (1961). New discoveries about ferns of Georgia have continued. We have accumulated many data and specimens ourselves, and sey- eral people have sent us additional information. Their contribu- tions are gratefully acknowledged. The more significant data are presented here. New distributional records presented, un- less otherwise indicated, are documented by specimens deposited in the University of Georgia Herbarium (GA). Part of the data contributed by the senior author was accumulated during re- search supported by the University of Georgia through Dr. Robert McRorie, Director of Research. EQuiseruM ARVENSE L.—Known previously only from one sta- tion (Fulton County) in the Piedmont (MeVaugh & Pyron, 1951). The range has been extended into the Blue Ridge Prov- ince through its discovery by the junior author and Florence Montgomery (Duncan 22060, Blake, and Montgomery) in sandy deposits on a wooded terrace along Yahoola Creek, ESE of Dahlonega, Lumpkin County. The eolonies at both stations prob- ably are adventive. The colony in Fulton County is in a moist area adjacent to a railroad fill, and the other is only a few yards from an abandoned gold ore processing plant. Lycopoptum ALopEcuRoES L.—In Georgia this clubmoss has been known previously from many stations in the Coastal Plain, Volume 55, No. 3, of the JOURNAL, Ppp. 97-144, was issued September 9, 1965. 146 AMERICAN FERN JOURNAL two in the southwestern part of the Piedmont (Meriwether and Pike Counties), and one in a boggy area in the southern edge of the Ridge and Valley Province. Two colonies have been located in the eastern part of the Piedmont about 70 and 90 miles, respectively, from previously known stations. One station is among an extensive growth of Arundinaria in a bog 3.2 miles ESE of Winder, Barrow County (Duncan 21915). The other is at the edge of a small pool in a granite quarry at Echol’s Mill, Oglethorpe County (Blake, 5 November 1961 Dunean (1950) mentioned that the colony in the Ridge and Valley was undoubtedly a recent migrant, but that the distance it had migrated could not be determined. The same is true of the colony at Echol’s Mill, for the clubmoss is in a habitat cre- ated by relatively recent quarrying. The bog in Barrow County, however, is quite old, containing a considerable deposit of peat- like material. LYCOPODIUM CERNUUM L.—Previously known from five coun- ties (MeVaugh & Pyron, 1951; Quarterman, 1953; Dunean, 1955). It has been found since on moist banks of a roadcut in the south-central part of Long County (Dunean 22100B and Blake). This is the most northeastern locality known for this species. LyYcopopiuM FLABELLIFORME (Fern.) Blanchard (L. complana- tum var. flabelliforme Fern.) —“Ferns of Georgia” gives three localities for this species. Other published records for the state cite stations in Elbert County (Dunean, 1955), DeKalb and Hall Counties (Hutto, 1960), making a total of six stations re- ported previously, all in the Piedmont Province. Hutto stated that the plant was reported from ten counties, apparently mis- interpreting information sent to him by the senior author. We now have specimens from thirteen more counties, more than _ doubling the known range in Georgia. Three stations are in the Blue Ridge (Rabun, White, and Gilmer Counties). New Pied- mont records are from Banks, Oglethorpe, Taliaferro, Green, FERNS IN GEORGIA 147 Morgan, Coweta, Carroll, Haralson, Cherokee, and Stephens Counties. This clubmoss often occurs in secondary growth of pine and/ or hardwoods and may be somewhat of a weedy component in these situations. Old residents, who know this groundpine, main- tain that the species once was more abundant, got rather scarce, and is increasing in abundance again. Some say that years of extensive fires destroyed many colonies, and that now, under fire protection, the species is recovering. SELAGINELLA aPus (L.) Spring. (S. apoda (L.) Spring.).— Occurs commonly above the Fall Line in Georgia and is reported from the Coastal Plain south to Burke, Bibb, Lee, and Decatur Counties, with an outlier in Tift County. An extension in range of over 90 miles was made when Mrs. M. B. Mellinger collected it a few miles from the Atlantic Ocean in Chatham County (Mellinger, 27 April 1959). The junior author has since found it in the lower Coastal Plain in Ware (Blake, July 1963) and Charlton (Blake, 24 August 1963) Counties. The latter locality is at the head of a “bay” SE of Folkston, where the spikemoss is associated with Osmunda cinnamomea and Anchistea virgin- ica, which were under plants of Persea pubescens, Gordoma lasianthus, and Lyonia lucida, Our field experience indicates that the species is rare southeast of Burke and Decatur Counties. BorrycHium ALABAMENSE Maxon.—Collected infrequently in Georgia, but probably it is more abundant than the records indi- cate. Diligent search in the field should increase its known dis- tribution, Two such searches have revealed four new stations, including the first two localities for it in the Blue Ridge. Mr. and Mrs. Frank Snyder of Toccoa have sent specimens to the University of Georgia Herbarium from Stephens (Snyder, 17 February 1963) and Rabun (Snyder, 18 November 1956) Coun- ties, and Mrs. Elizabeth Sawyer of Demorest has donated speci- mens from Habersham (Sawyer 333) and White (Sawyer 334) Counties. All collections were from open areas or open, decidu- ous woods. This fern seems to prefer disturbed areas. 148 AMERICAN FERN JOURNAL OpHiocLossum spp.—MeVaugh and Pyron (1951) included four species of this genus in “Ferns of Georgia.’ Two species were known from two stations each, while the other two were known from one station each. Diddell (1953) reported a single station for a fifth species, O. petiolatum, in Camden County. Finding adder’s-tongues is usually a difficult job, for most plants are easily overlooked in the field unless a search is con- ducted on hands and knees. Identification to species also pre- sents some problems since keys or descriptions of all five spe- cies are not readily available, e.g., Clausen (1938) and MeVaugh and Pyron (1951), which is out of print. A key based mostly on our observations, is presented below, along with other information. Underground stem eae emo, globose, with many fine roots about 0.25 mm. diameter when dried; leaf blades obtuse at the O. cr i apex satel stem slightly globose to pense eylindrieal, with few y roots about 0.4 mm. in meter or larger when dried; — ‘ees acute to obtuse at the a Leaf blades conspicuously broadest Secasct veld above the base, acute t the apex _O, petiolatum Hook. Leaf a broadest at or near the mel or if sometimes broadest base the apex then o Hindes of thant leaves to about ey em. long, acute at the apex, ai zontal or nearly so; strobili to 10 mm. long, the stalks r less in diameter when dried._0. nudicaule L. f. Blades - inching leaves to about 9 em. long, obtuse and apiculate, or obtuse to rounded and not Hes iculate, ascending nearly erect; etrobili to 32 mm. long, the stalks usually than 1 in diameter PR dried: Sides a fruiting navi acute to rounded, apiculate, the veins in two series, larger and heavier groups of polygonal ones ve smaller — finer sets of polygonal areoles l Blades of aeiaar “iam obtuse to sa not apiculate; eins usually in one series, forming narrow polygonal eo enclosing solitary veinlets with free ends 0. vulgatum L. 3 & 3 = a= =. S kg “3 om) S > — FERNS IN GEORGIA 149 In addition to the above characteristics we have noted that O. vulgatum, O. petiolatum, and O. nudicaule form adventitious buds on horizontal roots. The leaf blades of 0. crotalophoroides are fleshy, usually horizontal, on or near the soil, and have the lowest length-width ratio. The leaf blades of O. petiolatum are usually well above the soil, often wavy margined, their surfaces not flat but irregularly undulate, and their bases cupped upward, giving them a half “funnel-like” appearance. The leaf blades of O. nudicaule are at or near the soil and their surfaces are essen- tially flat. Ophioglossum crotalophoroides has been reported previously from one station in Georgia. We report seven new stations. They extend the range widely in the Piedmont and well into the Coastal Plain. On 19 March 1961, Donald Banks found plants associated with grasses, Houstonia patens, and Draba brachycarpa on a slope overlooking a beach at a state park in Morgan County. On 24 March 1962 Dr. Gordon DeWolf found plants in closely cut grass in sandy soil on a road embankment in Bullock County. The senior author collected plants on 5 April 1962 in an undisturbed area of shallow soil at the edge of a sloping granite outcrop above the Savannah River in Elbert County. The junior author found plants on 8 April 1963 in shallow soil at a granite outcrop in Oglethorpe County, and later observed the species in shallow soils of sloping granite out- erops overlooking High Shoals of the Apalachee River in Oconee County. The senior author showed a likely habitat for the species in Jenkins County to a class in taxonomy. Some students found plants within a few minutes, and eventually all nine persons on the field trip found specimens. All plants of this locality were found in very small areas of bare, sandy soil among grass plants in an open area immediately south of Magnolia Springs. This collection was made on 10 April 1964. Finally on 6 March 1965 the junior author and Florence Montgomery found the species in an old pasture in Wilkinson County. This species probably 150 AMERICAN FERN JOURNAL oceurs in many counties south of Elbert and Meriwether Coun- ties. Ophioglossum petiolatum previously was reported from only one station in Georgia. On 16 September 1961 the senior author found a colony in Brantley County, at the base of a road fill leading to a “high water” bridge south of the Satilla River, 2.5 mi. N. of Hoboken. Ophioglossum nudicaule was reported by McVaugh and Pyron (1951) from Effingham County, and by Rossignol in Chatham County (1956). The junior author collected sterile plants in moist open flats along the Satilla River in Brantley County on 9 September 1961. These plants fruited in the greenhouse on 8 November 1961. No new stations are to be added for O. engelmannii in Georgia, but three ean be added for O. vulgatum. On 7 July 1963 the junior author and Thomas Pullen, Jr., collected late fruiting plants in a deciduous woods on a flood plain along a tributary of the Ogeeche River in Taliaferro County. On 21 August 1963 Mrs. M. 8S. Holverstott showed us plants, some of which were in fruiting condition, from a low level area under deciduous trees beside a small stream in Gilmer County. This is the first record (Duncan 22142) for the Blue Ridge of Georgia. On 25 April 1964 the senior author collected a few plants in a poorly drained area in Jasper County. The plants of this species are the most conspicuous of those known to grow in Georgia. We have looked often and carefully for other stations, so we believe the species is rare. OSMUNDA CLAYTONIANA L.—Known previously in Georgia from the eastern part of the Blue Ridge and on Sawnee Moun- tain, Forsyth County. An important extension in range was made by the discovery of this fern by Mrs. Holverstott in west- ern Gilmer County, in the western part of the Blue Ridge. CyYsTorTeRIs PRoTRUSA (Weath.) Blasdell. (C. fragilis (L.) Bernh. in part).—This fern was reported by Blasdell (1963) in Georgia from only Rabun County. It had been reported else- FERNS IN GEORGIA 151 where to occur in scattered localities south to Stephens and Newton Counties. Additional stations have been located in White (Duncan 16359) and Floyd (Duncan 15216) Counties. CYRTOMIUM FORTUNE! J. Smith.—Morton (1957) reports this taxon as one of the three true species of the genus in cultivation in the United States. Although Holly Ferns may be adventive in some localities (Degener and Hawkes, 1951) C. fortunet ap- parently has not been reported as an escape in the southeastern United States. This fern has become naturalized in Athens, Georgia. It grows in borders of yards, in crevices of rock walls in gardens, in crevices of a north-facing rock wall along Tanyard Branch, and on soil and between rocks of a steep bank in shaded areas of a railway freight yard near the business district. THELYPTERIS TORRESIANA (Gaud.) Alston.—This fern has been reported in the United States from Florida, Alabama, and Texas. Oliver (1955) predicted that it would soon be found in Georgia. This fern was shown to the senior author in 1959 in Randolph County by Martha Dean Webster and Isabel Terry, and a collec- tion made (Duncan 21655). That same year Lee Webb collected it in Colquitt County, and in 1964 Beatrice Griffin collected the fern in Thomas County (Griffin, 28 November). All three locali- ties are in the southwestern part of the state and confined to the Coastal Plain. Morton (1962) indicated that the species is naturalized. In Georgia this fern occurs in wooded habitats relatively undis- turbed by man, and some colonies are extensive. Although we know that some fern colonies are recent emigrants (Duncan, 1950) and have probably migrated considerable distances, there is little convincing evidence concerning the age of most colonies. Those of 7. torresiana in Georgia could be quite old. The species is growing abundantly locally and at present definitely is a part of our natural flora. The Randolph County colony occurs in a deciduous forest along a small tributary of Nochaway Creek about 0.5 mile above its junction with Ichawaynochaway Creek, 1.7 miles E 15° N of Shellman. The colony in Colquitt County 152 AMERICAN FERN JOURNAL occurs in a dense forest at the margin of a boggy area; and that of Thomas County is under trees along a small stream 10.2 miles N. of Thomasville. ASPLENIUM HETEROCHROUM >< RESILIENS.—Wagner (1963) has said that this is an undescribed pentaploid, apogamous fern, intermediate between A. resiliens and A. heterochroum, and ap- parently is a hybrid between them. He reports this intermediate taxon from North Carolina, South Carolina, and Florida. Geor- gia now can be added to this distribution. The junior author collected it on mortar between rocks of an old wall in the south- eastern part of the state (Blake, 8 Sept. 1963). ASPLENIUM PINNATIFIDUM Nutt.—This uncommon spleenwort has been found in Hall County (Duncan 18389), which is the seventh county in Georgia in which it is known to occur. CaMPTOsoRUS RHIZOPHYLLUS (L.) Link—The Walking Fern has been known to be common to scattered in Georgia as far south as Stephens, Bartow, and Floyd Counties. The known range is extended southward by a collection by Thomas Pullen, Jr., made in Polk County (T. Pullen, Jr. 8). A sight record is recorded for Habersham County by Pruitt (1952). PILULARIA AMERICANA A. Br.—This rarely observed species has been known from three counties in Georgia (McVaugh & Pyron, 1951). The last reported stations found in Georgia were discovered in Washington and Walton Counties in 1936. The junior author and Florence Montgomery found it growing abundantly in 1962 (11 Nov.) rooted in mud under two feet of water (and floating) in a broad, shallow quarry in granite rocks at Echol’s Mill, 9.7 miles NE of Lexington, Oglethorpe County. Plants apparently continuously covered with water freely pro- duced sporocarps. Pyron & McVaugh stated that plants “seem to fruit freely only when the water begins to disappear.” Also, Pilularia apparently had not been known to thrive in water as much as two feet deep. FERNS IN GEORGIA 153 LITERATURE CITED BLASDELL, eS F. 1963. A monographie study of the fern genus Cystopteris. Mem. Torrey Club 21: 1-102, 23 plates Booxout, Henry. 1960, A new ae for Trichomanes petarsik in Georgia. . Fern Jour. 50: -247, CLAUSEN, monies T1938... A ete of the Ophioglossaceae. Mem. Torrey Club 19 (2): 1-177 speaaes OTTO, AND ae Dz. TEawes. 1951. The Holly Fern. Amer, Fern ur, 41: 117-1 DIDDELL, vist Ww. 1953. oe Notes. Ibid. 43: 114-117. Duncan, W. H. 1950. Preliminary reports on the oe e Georgia, III. The ar ik of seven Lycopsida. Ibid. 40: 173. Polypodium aureum in Florida and potas Ibid, 44: 1955. New records for Georgia ferns. Ibid. 45: 1960. Azolla caroliniana Willd. in Georgia. Ibid. uae 266-267. Hurro, THomas A, 1960. Two new Georgia stations for Lycopodium complanatum var. flabelliforme. Ibid. 0: 240. McDowELu, GLADSTONE, AND FRANKLIN Snyper. 1961. A parcel of ferns from Stephens County, Georgia. Ibid. 51: 67-69 McVaAuGH, ROGERS, AND JOSEPH H. Pyron, 1951. Ferns of Georgia. xvii + 195, Univ. of Georgia Press, ‘Adhens, Ga. Morton, C. V. 1957. Observations on cultivated ferns, II. The ie generic name for the Holly Fern. Amer. Fern Jour. 47: 52- —, 1962. Another name Page ie the Florida fern one as Dryopteris setigera. Ibid. 52: Outver, WinuiAM F., Jr, 1955, Dry eee ae to be found in Georgia? bid, 45: 126. Prurrr, Ropert G. 1952. The alee Zone of Northeasternmost Georgia. M.S. Thesis, Emory Unive aes Este, 1953. Props from Lowndes County, Georgia. Amer. Fern Jour, 43: ROSSIGNOL, ies aa R. 1956. peter tenerum in Georgia. Ibid. 46: 107. : SNYDER, oes D. 1955. Rediscovery of Lygodium palmatwm % Georgia. d. 45: 104-105. WAGNER, re H., Jz. 1963. A biosystematie survey Preliminary abstract. Ibid. 53: 1-16. DEPARTMENT OF BoTANy, UNIVERSITY OF GEORGIA, ATHENS, of United States ferns. GEORGIA 154 AMERICAN FERN JOURNAL Cytological Observations on Ferns from Southern China S. K. Roy anp R. E. Houtrrum The fern plants which are the subject of the present paper were collected by Holttum in the Ting Hu National Park, about 0 km. from Canton, which he was able to visit through the hospitality of Academia Sinica, Peking, in October 1963. He wishes to express his thanks to Mr. Wang Chu-Hau, who ar- ranged the excursion from Canton and assisted in the identifica- tion of the species. The nucleus of the National Park is an area of 500 acres of natural forest in the valley of a stream and on the hillsides rising above it, with a larger adjacent area of hill sides planted with pine trees. Very young fern plants were mostly collected beside paths which have been made in the for- est. The plants have been cultivated in the tropical fern houses of the Royal Botanic Gardens, Kew, under the care of Mr. H. J. Bruty, and we are indebted to the Director for permission to have frequent access to them. Cytological observations have been made by Roy, both from root tips (2n) and from young sporangia as soon as these were produced. Most of the previous cytological records to which reference is made are listed, with bibliographical data, in the papers by Chiarugi (1960) and Fabbri 1963). The most important new record here presented concerns Brainea insignis. Lyeopium japonicum (Thunb.) Sw.—2n —58. The only re- liable previous cytological record of this species is by Manton & Sledge, from a plant in cultivation at Kew, of unknown origin; this plant is a tetraploid (n=58). The plant brought from Ting Hu is diploid; it is not yet fertile. CYATHEA PODOPHYLLA (Hook.) Copel—2n = ec. 140 (probably 138). This was the only species of Cyathea in the forest, abun- dant in shady gullies. It belongs to subgenus Cyathea section Gymnosphaera of Holttum’s arrangement (1963). The plants are still too small to bear fertile fronds. Root-tip preparations are more difficult in Cyathea than in most other ferns, and the CYTOLOGY OF FERNS FROM CHINA 155 count is approximate. All other Cyathea species so far exam- ined, including species of both subgenera recognized by Holttum in Asia and Malesia, have n = 69. ARACHNIODES CHINENSIS (Rosenst.) Ching.—2n = 82 (Fig. 3). The closely related and very widely distributed species A. arts- tata (Forst. f.) Tindale (better known as Polystichum aristatum or Rumohra aristata) has been reported as tetraploid in Ceylon and southern India, and plants obtained in New Guinea now in cultivation at Kew are also tetraploid. BoLBITis SUBCORDATA (Copel.) Ching.—2n = 82. No previous cytological record of this species has been published ; most other species investigated are diploids with n= 41. The plant is not yet fertile. DIPLAZIUM DONIANUM (Mett.) Tardieu.—2n = 123. The only observation of meiosis in sporangia shows 41 pairs and 41 single chromosomes; the plant is therefore a triploid hybrid. Ripe sporangia yielded some good large spores and many small shrivelled ones; this indicates apogamy in the gametophyte. The species is distributed from Assam through southern China; southward it appears to overlap with the closely related species D. bantamense Bl., which in Malaya has been shown to be tetra- ploid. Woopwarpia gaponica (L, fil.) Sm.—2n = 68 (Fig. 1). This agrees with four previous observations on W. radicans (L.) Sm. (The name W» radicans has in the past been used in @ very broad sense, and Indian records under this name are probably W. unigemmata (Mak.) Nakai). BRAINEA INSIGNIS (Hook.) J. Sm.—2n = 66, the first eytologi- cal record for this monotypic genus (Fig. 2). Brainea has a massive trunk as in some species of Blechnum, but venation more as in Doodia and Woodwardia (anastomosing similarly in both sterile and fertile fronds). Unlike both the latter genera, it lacks an indusium, and the sporangia spread along the veins of the contracted fertile pinnae. The numbers 33 and 34 have CyToLoGy OF FERNS FROM CHINA 157 been reported for several species of Blechnum (see Quinn 1961, and Chiarugi 1960). The spores of Brainea are rather similar to those of the Blechnum species for which Quinn reported n = 33. The present observation supports the generally accepted view that Brainea is closely related to Blechnum. PTERIS FAURIEI Hieron.—2n = 58. This species of China and Japan belongs to the complex of P. quadriaurita Retz. The only previous record for P. fauriet is by Walker, whose plant was an apogamous triploid from Japan. PTERIS MULTIFIDA Poir.—2n — 116. This species, widely dis- tributed in China and Japan, and also widely cultivated in other countries, has twice been reported as tetraploid. ADIANTUM FLABELLULATUM L.—2n —116. There is no pre- vious record for this species, which is widely distributed in tropical Asia. Most of the many Adiantum species investigated have a base number n= 30; one of the few with n= pedatum L. to which A. ‘Aabeliedl stim is probably related fic judged on grounds of general morphology). ADIANTUM MALESIANUM Ghatak (1963, p. 74).—2n= 120, agreeing with Ghatak’s observations. At Ting Hu this is abun- dant in erevices on limestone hills near the national park; it ae in similar situations in Malaya, and is widely Tieabta _E. Asia. It is a distinct member of the A. caudatum com- pak for the first time distinguished by Ghatak. Mrcrosortum rorTuNEr (Moore) Ching.—2n = 72. There is no previous record for this species; numerous others included in the genus have been reported as diploid with n= 386. A few species have been reported with n = 37; this is perhaps an indi- cation that the genus has not yet been ieurly defined, like others in the Polypodiaceae (s.str.). Figures 1-3. CHROMOSOMES SHOWN BY ROOT TIP SQUASHES. Figure 1. WoopwarDIa JAPONICA (L. Fit.) SM. Figure 2 BRAINEA INSIGNIS (Hoox.) J. SM. Figure 3. ARACHNIODES CHINENSIS (RosENST.) HING. ALL Na mies = hee, | AN ye a9 = : WAL Wipe TS & \y ——, W SHIGIN ZN WZ: \) JF ae 5 \ San Figure 1. Meraxya RosTRATA (H.B.K.) PRESL. DRAWINGS OF JUVENILE PLANTS, SHOWING VERY YOUNG SPOROPHYTE AT “A,’?. SUCCESSIVELY OLDER FRONDS AT “B” AND “OC.” PLANT AT “D” WITH ONE FROND APPROACHING MATURE CHARACTERISTICS. (DRAWING BY W. H. A. HEKKING). AMERICAN FERN JOURNAL 160 CytTroLoGy AND MorPHoLoGy oF METAXYA ROSTRATA 161 placed by most authors) in its creeping hairy rhizome and form of sorus and sporangia. Metaxya presents a peculiar combina- tion of primitive and advanced features. The hairy rhizome, and “simple” character of the sorus (sporangia simultaneous in development) are primitive; but the simply pinnate fronds, the sori spread a little along the veins and often two on one vein, and the form of individual sporangia, are advanced, as compared with Cyathea (to which Alsophila should be united ; see Holttum 1963, p. 76). But Bower failed to mention some other considera- tions. Bower called the vascular system of the rhizome a solenostele ; but the rhizome, though prostrate, is not dorsiventral, which is the significant feature of the solenosteles of Dennstaedtia, Dip- teris and Dicranopteris (see Holttum, 1965). The more impor- tant difference from Cyathea lies in the absence of sclerotic tissue. The prostrate, not erect, growth of the stem is not an important distinction; there are prostrate and erect stems in both Cibotium and Culcita. As pointed out by Bower, the simple pinnae of the fronds indieate by their marginal serrations that they have evolved from more amply branched fronds. This indication is further strengthened by the form of pinnae on young plants, which are quite deeply lobed, so that the fronds are almost bipinnate (Figs. 1-3). Furthermore, the pinnae are very asymmetric at their bases and the lowest pinnae are largest. These young fronds agree in shape rather with Culcita and Thyrsopteris than with Cyathea or Dicksonia. In Culeita, the deltoid shape of the frond and of its leaflets is associated with a grooved upper surface of rachises, the groove of a larger rachis open at its junctions with smaller ones. In Metarya a similar arrangement, simplified owing to the simpler branching of the frond, is found; FigurES 2 AND 3. METAXYA ROSTRATA AT FOOT OF THE TAFELBERG, SURI- 2 YouNG PLANTS WITH JUVENILE FRONDS (LENS COVER 4 CM. NAME, Fia, 2. IN DIAMETER). Fic. 3. MATURE PLANT WITH SIMPLE-PINNATE LEAVES. (PuorograPpus py K. U. Kramer, FEB. 1961). Ra ene ra 162 AMERICAN FERN JOURNAL the short stalk of each pinna is grooved, and the groove is con- tinued upwards as the midrib of the pinna, while downwards the stalk-groove opens into the groove of the upper surface of the main rachis. Bower does not mention spores. These are thin-walled, smooth, FigurE 4. Roor TIP MITOSIS oF METAXYA ROSTRATA, SHOWING 2n = 192. ix 1538), CyToLogy AND MorPHOLOGY OF METAXYA ROSTRATA 163 almost spherical, with very slight ridges indicating their trilete structure. Possibly they are short-lived. Roy’s observations of root-tips show that the sporophyte com- plement of chromosomes is not less than 190 nor more than 192 (Fig. 4), 192 being the more probable figure. Assuming that the sporophyte is diploid, this gives 96 as the haploid chromo- some number. Fixation of sporangia in February 1965 gave a count of 94-96; the preparations were not good enough to give an exact number. Comparing the information now available for the other genera of Cyatheaceae (in the broad sense of Holttum and Sen 1961), we have the following list. Cyathea (sens. lat.), several species n = 69 Cnemidaria (Jamaica, T .G. Walker, ver- bal report) aaone Dicksonia, species from New Zealand, New Caledonia, New Guinea and St. Helena Lophosoria (Jamaica, T. G. Walker, ver- bal report) Culcita (subgenus Culeita) macrocarpa n = 66-68 Culcita (subgenus Calochlaena) dubia n = 59-58 Cystodium sorbifolium (Roy and Holt- tum, 1965) Metaxya rostrata n = 65 n = 56 2n = 192 (or 190) n = 94-96 Thus the cytology of Metarya does not clearly indicate affin- ity with any of the other genera, but rather adds emphasis to the isolated position of the genus, and supports its status in a separate subfamily accorded by Holttum and Sen (1961). The evidence of the form of young fronds, and of rachis-structure, would point to an affinity with Culcita or Thyrsopteris rather than with Cyathea or Lophosoria; but the position of the sori, and especially the existence of two sori on one vein, point to a long history of evolutionary divergence from such a fern as Culcita. 164 AMERICAN FERN JOURNAL LITERATURE CITED Bower, F. O. 1913. Studies in the Phylogeny of the Filicales, III. On etaxya and certain other relatively primitive ferns. Ann. Bot. 27: 443-450. oo, 1996. The Ferns, Vol. Il, chapter 32. Cambridge Hourrum, R. E. 1963. Flora Malesiana, Ser. Leeds pti, Gyatheneese: 1965. The ca of the vascular system in Ferns, with especial reference to dorsiventral rhizomes. Phytomorphology 4: 477- “oi Hoirrum, R. E., ano U. SEN. 1961. oS pe and Classification of the Tree oe a ae 11: 406-420 Roy, 8S. K., anp R. E. Hourrum, 1965. a diy actnirieanl Records for Cystodium and Dicksonia. ‘Seer. Fern Jour. 55: 35-37. Two fig- KEw peed RICHMOND, SURREY, ENGLAND. Observations on Cultivated Ferns, VIII. Stenochlaena C. V. Morton The small genus Stenochlaena consists of a few species native to the Old World tropies. They are large, coarse ferns with simply pinnate sterile blades, climbing on tree-trunks in forests by means of thick scaly rhizomes. The modern understanding of the genus dates from L. M. Underwood, who in his paper “The Genus Stenochlaena”’ de- limited four sections—Eustenochlaena, Cafraria, Teratophyllum, and Lomariopsis. R. BE. Holttum? presented a much more detailed account, in whieh Underwood’s delimitations were accepted, except that Teratophyllum and Lomariopsis were recognized as distinct genera rather than sections. Holttum’s conclusions seem to be well taken. In Stenochlaena there is a vein on each side of the midrib of the pinnae, this forming a row of elongate loops (areoles) along the midrib. The other veins arise from the outer side of this costal vein, and run straight to the margin, 1 Bull. Torrey Bot. Club 33: 35-50. 2 eae Bulletin ee Settlements 7 a eats 1932. i OBSERVATIONS ON CULTIVATED FERNS 165 where about half of them end in sharp, spinulose teeth. In Lomariopsis and Teratophyllum all the veins are free and arise directly from the midrib. Only two species of Stenochlaena are cultivated in the United States. These may be distinguished as follows: Fertile pinnae bipinnate; sterile pinnae with numerous, close, and con- i Ss. spicuous spinulose teeth (Sect. Cafraria) -.---—----------- S. tenuifolia Fertile pinnae simply pinnate; sterile pinnae with distant, more incon- spicuous teeth (Sect. Stenochlaena) _----.-----------------—------ . palustris STENOCHLAENA TENUIFOLIA (Desy.) Moore, Gard. Chron. 1856: 193. 1856. Lomaria tenuifolia Desv. Berlin Naturf. Freund. Mag. 5: 326. 1811. Type from Madagascar. Lomaria meyeriana Kunze, Linnaea 10: 509. 1836. Type from South Africa, Drege Stenochlaena tenuifolia, the only species of sect. Cafraria Presl,? is a native of South Afriea, Madagascar, and the Mas- carene Islands, but it is apparently nowhere common. It was in cultivation in the Botanical Garden in Washington, D. C., at least as early as 1884, and was probably present in other botani- cal gardens, but it was seemingly not in cultivation by gar- deners until about 1925. The first to grow it was probably Mr. Mulford B. Foster, of Orlando, Florida, who introduced it to the trade. Mr. Foster sold about 50 plants, and so the species was not uncommon in gardens in Florida by the mid 1930’s. Since it grows well outdoors in Florida, it may be expected to be found in the wild eventually, but there are no specimens in the National Herbarium, as yet, other than cultivated ones. When forwarding a specimen to the National Herbarium, Mr. Foster commented that the species is quite slow and rather diffi- cult to start, as it does not like to be moved. He established new plants by layering the runners. However, once established it grows rapidly, providing there is good shade and heavy soil. Mr. Foster wrote further: “It seems that the most rapid growth of this fern is during the rainy season, and growth continues 3 Epim. Bot. 166. 1849 [1851]. 166 AMERICAN FERN JOURNAL until our November cool weather. I have had the fern climb seven feet up a tree-trunk in a single season. I have had many fronds as long as six feet. However, the fronds as they creep out into the sun are much shorter than those suspended from the climbing sections on the pergola. The fertile fronds are generally produced in August, but are not very numerous. In some years I’ find but two or three. The plants will stand light frost without damage, but temperatures of 24° to 26° will injure many of the fronds seriously.” In recent years Stenochlaena tenwifolia has been grown in southern California, where it was perhaps introduced by the late Mrs. Fay A. MacFadden. According to Mrs. MacF adden, it needs lots of water and attention, and does not like cold weather, which if too severe will kill it. Also, it does not fruit abundantly or regularly in California. A slightly variant form with extremely narrow pinnae has recently been introduced directly from Zululand, South Africa, through Dr. R. Rodin. This has been cultivated and distributed from the University of California Botanical Garden. STENOCHLAENA PALUSTRIS (Burm.) Beddome, Suppl. Ferns So, India & Brit. India 26. 1876. Polypodium palustre Burm. Fl. Ind. 234, 1768. Type from Ceylon. This species, common in tropical Asia and Malaysia, is hardly known in cultivation, although it was formerly somewhat grown in England as a hothouse plant under the name Acrostichum scandens. Recently it has again been introduced into cultivation by the University of California Botanical Garden from material originally grown from spores collected in New Guinea by W. H. Wagner and D. Grether. Although spores have been distributed to interested persons, this is such a large and coarse vine that not many growers are likely to have space for it, especially since it is not particularly ornamental. SMITHSONIAN INSTITUTION, WASHINGTON, D. C. DISPERSAL OF MARSILEA MUCRONATA 167 Dispersal of Marsilea mucronata by Water Birds' CuaRLEs R. MALONE AND VERNON W. PROCTOR The aquatic fern Marsilea mucronata commonly occurs in playas (temporary lakes) in west Texas and in other isolated bodies of water on the Great Plains from northern Mexico to southern Canada (Correll, 1956). The means by which this and other species of Marsilea reach isolated habitats is unknown. Guppy (1906) suggested that Marsilea might be dispersed by adhering to the external surfaces of birds. However, this appar- ently never has been demonstrated. Brown, Larson, and Bold (1964) found that spores of some terrestrial ferns may be dis- persed by wind. Both these methods of dispersal seem unlikely for Marsilea because of the size and weight of the sporocarps. Recent investigations have shown that many fresh-water or- ganisms incapable of active overland transport may be effee- tively dispersed via the intestinal tracts of migratory water birds (Proctor, 1959, 1962, 1964). In most cases resistant eggs and spores account for the ability of organisms to withstand passage through the intestinal tract. Although intact sporocarps have been found in the stomachs of various birds (MecAtee, 1939), it is not known whether sporo- earps will pass unharmed through the entire intestinal tract. Neither is it known whether the spores are sufficiently protected by the sporocarp wall to retain their viability after passage. To determine the effects of avian digestion upon the viability of Marsilea spores, sporocarps of M. mucronata were fed to 5 species of water birds. Six killdeer (Charadrius vociferus), 2 mandarin ducks (Aix galericulata) , 2 wood ducks (Aix sponsa), 2 Chiloe widgeons (Anas sibilatriz), and one mallard (Anas platyrhynchos) were used. Sporoearps fed to ducks were mixed with either chopped aquatie vegetation or small grain. Killdeer were fed sporocarps 1 Research supported by National Institute of Health Grant GM 11394- fAT. 168 AMERICAN FERN JOURNAL mixed with ground commercial shrimp. No attempt was made to determine the number of sporocarps ingested per bird. Imme- diately after ingesting the sporocarps, the birds were caged and their feces collected and examined every hour. Birds were eaged at least 24 hours and released if no sporocarps had been passed within that period. If sporocarps were passed the bird was retained for an additional 24 hours following passage of the last sporocarp. Sporocarps recovered from the feces were either placed di- rectly into sterile soil and water or were first searified. The cultures were inspected after 2 weeks and the approximate number of young plants recorded. RESULTS AND Discussion Table 1 gives the number of trials per species of bird, the number of sporocarps recovered, and the range and mean rate of passage of the sporocarps. Both the greatest number of re- covered sporocarps and the greatest time of retention resulted from killdeer. These birds are largely carnivorous and less than 5% of their diet consists of seeds (Bent, 1929). For this reason their stomach usually contains little grit. Thus, the sporocarps probably were subjected to little grinding. Although food passes rapidly through killdeer, sporocarps were retained longer than expected. A meal normally passes through the intestinal tract of these birds in about 2 hours: sporocarps were never passed under 5 hours. It seems likely that the sporocarps remained in the gizzard much as would grit. Spores from crushed sporocarps occasionally were found in the feces of killdeer. These were placed in sterile soil and water but none was viable. Ducks ingested many more sporocarps than did killdeer yet passed fewer. Feces from all the ducks contained large numbers of nonviable spores indicating that most sporocarps were crushed. Fewer sporocarps were recovered from wood ducks than from any other bird. Wood ducks ate large quantities of sporocarps but were very efficient in crushing them. The diet of these birds DISPERSAL OF MARSILEA MUCRONATA 169 TABLE 1: RECOVERY OF Marsilea mucronata SPOROCARPS FROM THE INTESTINAL TRACT OF WATER BIRDS Sporocarps Rate of Passage: Hours = N Recovered Kolideen ee ee 20 55 5-60 19.5 Mallard 8 6 2-24 8.0 Chiloe Widgeon _....... 14 8 1-24 8.5 Mandhtin °c a ee 9 23 1-8 3.0 W 60d: iDugk= = Soe 9 2° 7 7.0 includes nuts and acorns (Mabbott, 1920) and they are known for their muscular gizzard. In only 2 cases did sporocarps recovered from birds release their spores without being scarified. One each was recovered from a mandarin and a wood duck. In both instances, half the sporoearp wall had been torn off. The spores were released within a few minutes after the sporocarps were placed in water. Seven and 12 visible megaspores were contained in the damaged sporocarps taken from the mandarin duck and from the wood duck, respectively. Apparently the sporocarp wall provides protection for the spores. Recovered undamaged sporocarps, regardless of their rate of passage, contained as many viable megaspores as did sporocarps not fed to birds. The usual number in either case was 30 to 50. Damaged sporocarps contained significantly fewer viable megaspores. Megaspores from crushed sporocarps were never viable. Proctor and Malone (1965) studied the effect of the digestive processes of 4 kinds of birds upon eggs and spores of various aquatic organisms. They concluded that because of differences in digestion, some birds are probably more effective agents of dispersal than others. This appears to be likely when consider- ing the dispersal of M. mucronata. Killdeer could carry larger numbers of sporocarps for greater periods of time than could any species of duck studied here. However, due to food habits, killdeer and other shore birds probably ingest fewer sporocarps than do waterfowl. Among the ducks, wood ducks are less likely 170 AMERICAN FERN JOURNAL than the other species to carry sporocarps. The efficiency with which wood ducks grind their food assures that most of the ingested sporocarps would be crushed. These results, together with accounts of sporocarps being found in the stomach of waterfowl, suggest that dispersal via the avian intestinal tract is of importance to M. mucronata. Although the spores themselves are not resistant to avian diges- tive processes, the wall of the sporocarp provides them ample protection. Sporoearps can remain within the intestinal tract sufficient time to allow considerable distance of dispersal by a flying bird. The wide spread cecurrence of M. mucronata attests to the effectiveness of its means of dispersal. LITERATURE CITED Bent, A. C. 1929. Life cee sen of North American he ae Order Limicolae (part 2). U.S. Natl. Mus. Bull. 146, Brown, R. M., Jz., D. A. os AND H. C. Boup. 1964. sintra algae: their ‘aidnittinite and an Sas: Science 143: -585. CorRELL, D. 8S. 1956. Ferns and fern allies of Texas. ane Research Foundation, eonige Texas, 188 p Guppy, H. B. 1906. Observations of a naturalist in the Pacific. Vol. I, Plant- ae Maemillan and Co., Ltd., New York, 627 p Masport, D. C. 192 ood habits of North American diving ducks. Tech. Bull. me U.S.D.A., Washington, D. C. McArer, W. L. 1939. Wildfowl food plants. Collegiate Press, Ine., Ames, a, 141 p. Procror, re ‘M. 1959. Dispersal of fresh-water algae by migratory water birds. Seience 130: 623-624, 962. Viability of ses oospores taken from migratory water birds. Ecology 43: 528-529, 1964. i abasty of crustacean eggs recovered from ducks. Reahigy 45: 656 = AND OC. OR Maa 1965. Further evidence of the dis- persal of small aquatic organisms via the intestinal tract of birds. Ecology 46: (in press). DEPARTMENT OF BroLogy, TEXAS TECHNOLOGICAL COLLEGE, _ Lupsock, TEXxas. A. SEXUAL FORM oF PELLAEA GLABELLA Lil A New Sexual Form of Pellaea glabella var. glabella from Missouri W. H. WAGNER, JR., D. R. FARRAR, AND KATHERINE L, CHEN! Between 5% and 10% of fern species are non-sexual. The non-sexual or apogamous type of life eyele is now known in over 80 species among such genera as Adiantum, Asplenium, Cheil- anthes, Dryopteris, and Pteris. This form of reproduction by- passes the regular sexual fertilization of egg by sperm, and the details of the pattern were worked out by W. Dopp and Irene Manton in particular (reviewed by Mehra, 1961). The sporo- phyte and the gametophyte have the same chromosome number. The typical fusion of egg and sperm that occurs in the arche- gonium is lacking, The new fern plant simply grows out of the tissue of the prothallus as a bud. If the chromosome number of the sporophyte is, for example, 87, the spore will have the same number, the prothallus the same number, and so on. The only change in chromosome number occurs just before spore forma- tion, when there is usually an automatic doubling of chromo- some number in the spore mother cells. Instead of producing the normal 16 spore mother cells with 87 chromosomes, the spore-cases produce 8 mother cells with 174. Normal spore for- mation follows, producing only 32 spores, not the typical 64. If doubling does not oceur and there are the usual 16 spore mother cells, the 64 spores which result in a regularly apogamous fern are abortive and will not germinate normally. Thus doubling is required, and the “good” spores are only those which occur 32 per sporangium. Because of these spore products it is not neces- sary to make cytological studies to Judge in general whether a given fern is apogamous or sexual. By merely looking through the microscope at sporangia from dried herbarium sheets it is fr. Farrar’s travel ex- Evolutionary Biology 1 Research supported by NSF—GB2025. Part of yenses were PERC by the aid of Michigan T e Training Progra AT AMERICAN FERN JOURNAL usually possible to distinguish which type of reproduction oc- curs. FIGURE 1, PELLAEA GLABELLA VAR. GLABELLA. SPECIMENS, OTHERWISE IN- DISTINGUISHABLE, SHOWING BOTH TYPES OF SPORES SHOWN IN FIGURE 2, THE DIPLOID-SEXUAL AND TETRAPLOID-APOGAMOUS. (E. J. Palmer 19449, MO). A SexuAL FormM or PELLAEA GLABELLA 173 Typically, if all or most of the spore-cases contain 64 uniform spores, the plant is sexual. If some of the sporangia contain 64 abortive spores and some contain 32 uniform spores, the plant is probably apogamous. This is an excellent example of one of the uses of spores as a tool in systematic studies of ferns (Cf. Wagner and Chen, 1965, on the use of spores in detecting Dry- opteris hybrids). Alice Tryon (1957) used the above criteria to assess the type of reproduction in species and varieties of cliffbrakes, Pellaea. Confirmation was made by germination tests and cytological studies by Donald M. Britton. In regard to the wide ranging smooth cliffbrake, P. glabella, three varieties were distinguished. Samples of two of the varieties showed 116 chromosomes, 32 “ood” spores, and apogamy—var. glabella (Fig. 1), occurring from Vermont to Minnesota and Texas and having large fronds (4-36 em. long), the pinnae made up of 3-7 segments; and var. simplex, from British Columbia to Washington and Utah to New Mexico, with small fronds (1-20 em. long), the pinnae usually with 3-5 segments. The remaining variety had 58 chromosomes, 64 spores, and sexuality—var. occidentalis, from Alberta to Wyoming and South Dakota, the fronds very small (1-5 em.) and the pinnae usually sessile and with only one segment. Dr. Tryon’s opinion of the varietal relationships was as follows: “Pellaea glabella var. occidentalis is undoubt- edly the source from which the polyploid members of the com- plex have been derived, either through hybridization or possibly autopolyploidy.” She further suggested (p. 147) that “Var. glabella” with 116 chromosomes may have arisen as the hybrid of the apogamous P. atropurpurea with 87 chromosomes and the sexual P. glabella var. occidentalis with 29 gametie chromo- somes. The above hypotheses will need to be rather profoundly mod- ified as a result of the facts to be reported below. We now have evidence of a 64-spored, diploid, sexual form of “Var. glabella” which oecurs in the region to the south and west of St. Louis, 174 AMERICAN FERN JOURNAL Missouri. The bearing of this new form on the interpretation of varietal relationships in P. glabella will be discussed below. The discovery of two forms of the same variety, one sexual and one apogamous, has considerable theoretical interest. As will be noted below, the two forms may even exist sympatrically, i.e., in the same habitat. On October 16, 1964, Donald R. Farrar collected a series of Pellaea glabella var. glabella on the river bluffs 10 miles west of DeSoto, Jefferson County, Missouri. The smooth cliffbrakes were growing with such other ferns as Asplenium platyneuron, A. resiliens, A. ruta-muraria, Cheilanthes feci, and Pellaea atro- SO00O|CPOCLC OPO ESC OCI ¥ Se OCaGa ee Ve LOLOLS SSO aS Saussvee -——+-+ +4 Toop FIaurE 2. OUTLINES OF RANDOM 10-SPORE SAMPLES MOUNTED IN DIAPHANE, SEXUAL SPORE TYPE A, B, C, E; APOGAMOUS SPORE TYPE D, F. ALL FROM STATE OF Missouri. A. JEFFERSON County, Farrar s.n. (MicH). B. FRANK- LIN County, Steyermark 31404 (Mo). C. Putasxt County, Palmer 39199 (Mo). D. Curistian County, Cuapwick, Steyermark 23010 (Mo.). E. Carter County, Palmer 19449 (Mo), F. SAME DATA AS * A SexuaL Form or PELLAEA GLABELLA 175 purpurea on the limestone rocks. A check of the spores (Fig. 2, A, B, CO, E) showed them to be wholly unlike those previously reported in var. glabella (Fig. 2, D, F), being uniform in size and shape, much smaller, and numbering 64 or nearly that num- ber in each spore-case. Then, on May 2, 1965, Farrar collected cytological materials to test our prediction that the plant is a sexual diploid. The results (shown in Fig. 3) fully confirmed our prediction.” In order to determine the geographical distribution of the sexual form, we examined a large number of specimens sent on loan by the Missouri Botanical Garden and the U. S. National Museum. We discovered that the morphology of the plants can- not be used to distinguish the diploid sexual form from the apog- Ficure 3. CHROMOSOME SQUASHES AT MEIOTIC METAPHASE SHOWING 1 = 29 CHROMOSOME PAIRS. ALL EXCEPT LOWER RIGHT COLLECTED IN EARLY May; LOWER RIGHT, IN LATE MAY; FROM NATURALLY GROWING PLANTS IN JEFFER- son County, MissourI. DOTTED BODIES PROBABLY NUCLEOLAR MATERIAL. 2 It is interesting to note that in all figures seen of the diploid, a small object, staining lightly and varying in size from mother cell to mother eell, ize f was observed. Possibly the object represents nucleolar material. 176 AMERICAN FERN JOURNAL amous form, so we were forced to make microscope slides of all the specimens to prepare the county distribution map in Mis- souri shown in Figure 4. (It is not at all unlikely that further studies of the sporangial contents of this fern will show that the sexual form occurs in other states besides Missouri, but such a survey was not made by us.) The small-spored sexual form is confined, so far as our present collections show, to the cokesveeer’* Figure 4. COUNTY DISTRIBUTION MAP OF PELLAEA GLABELLA VAR, GLABELLA IN THE STATE OF MissouRI. Dots — APOGAMOUS 32-SPORED FORM, STARS = SEXUAL 64-SPORED FORM. A SexvuaL Form or PELLAEA GLABELLA LTy quarter of the state which is south and west of St. Louis. The large-spored apogamous type runs much further west and north. The earliest collection of the diploid we encountered was that of E. J. Palmer in 1920 (Carter County). All of the collections of sexual var. glabella are listed as fol- lows, but it must be noted that only one specimen per sheet was examined. Thus it is possible that some collections are mixed. In four out of five of the counties where the diploid form oc- curs, the apogamous type was also observed. In one ease, indi- cated below, specimens from the same locality (Carter County, Van Buren) included both forms. Where the two oceur together interesting hybrids should be sought with 52 chromosome num- ber of 145 CoLLecTIONS: Franklin County: Semi-shaded dry limestone cliffs near Mer- amee River, State Park east of Sullivan, Julian A. Steyermark 1404 (MO). Jefferson County: River bluffs 10 miles west of DeSoto, %4 mile upstream from junction of “H” road and “Big River,” D. R. Farrar (eytological vyouncher, n = 29, MICH). Pulaski County: Face of dolomite bluffs, near Hazelgreen, E. J. Palmer 39199 (MO, US). Dent County: Bluffs along Meramec River between Howe and Sligo, J ulian A. Steyermark 15696 (MO). Carter County: Dry limestone cliffs of Current River near Van Buren, E. J. Palmer 19449 (MO)—Both of the 2 sheets of this collection have speci- mens with the apogamous type of spore and specimens with the sexual type, ef. Fig. 1, E and F. In spite of our effort to find some morphological differences other than the spores, we concluded that the sexual form is in- separable from the apogamous form. At first we thought that the diploid, sexual plant was more slender and delicate, but we later discovered that some of the large-spored apogamous specl- mens were also slender and delicate. Furthermore, two of the small-spored sexual specimens (Pulaski and Dent Counties ) are thick and robust plants. Cleared material of several sam- ples of each form showed a tendency toward thickening of the lower epidermal walls in the diploid, but there was much varia- tion in this character. 178 AMERICAN FERN JOURNAL On the basis of the above facts, the interpretations of the Pellaea glabella varietal complex outlined above must be al- tered. Our present knowledge indicates that the correlation of varietal differentiation in this species with different life cycles is not as definite as was previously supposed. As shown here, one of the varieties, namely the typical var. glabella, exists in two different forms—the one an apogamous tetraploid, and the other a sexual diploid which has been heretofore overlooked. It is conceivable that still other cytological forms will be found in var. glabella. Similar diversity in life cycles may exist also in the other two varieties of P. glabella as well. An hypothesis that var. glabella might have arisen as a hybrid between P. atropurpurea and P. glabella var. occidentalis is made very un- likely by the information presented here. More likely the apog- amous tetraploid is an autopolyploid derivative of the sexual diploid of the same variety in which chromosome doubling and modification of the life cycle took place at some time in the past. The much wider range and greater abundance of the non-sexual apogamous form of var. glabella strongly suggests that the apogamous life cycle is more efficient and successful, at least in the state of Missouri. LITERATURE CITED Meura, P. N. 1961. Cytological evolution of ferns with particular reference to Himalayan forms. 48th Indian Science Congr. Part II (Presi- dential Address): 1-24. Tryon, Alice F. 1957. A revision of the fern genus Pellaea Section Pellaea, Ann, Missouri Bot. Gard. 44: 125-193 AND DonaLp M. Brirron. 1958. Cytotaxonomie studies on the fern genus Pellaea. Evolution 12: 137-145 Waaner, W. H., Jr., and KATHERINE Lim CHEN. 1965. Abortion of spores and sporangia as a tool in the detection of Dryopteris hybrids. Amer. Fern Jour, 55: 9-29, Botany DEPARTMENT, UNIVERSITY oF MicHIGAN, ANN ARBOR, MIcHIGAN. SHorTeER NOTES 179 SHORTER NOTES JAPANESE CLIMBING FERN SPREADING IN SOUTH CAROLINA.— On November 16, 1957, we discovered a healthy plant of the Jap- anese Climbing Fern, Lygodium japomcum (Thunb.) Sw., grow- ing in semitidal unmanaged marsh along the Uernezobre Bank dike on the Savannah National Wildlife Refuge. This fern grew to a height of 12 feet and was heavily covered with fruiting fronds. On January 7, 1965, five more clumps of this fern were found on the marsh side of a dike berm at least a mile from the original station. These had been mowed off partially, but all had fruiting fronds. Much new and bright green growth was evident, even though we have had several severe frosts. These are the only records of this fern from South Carolina other than a report of a collection near Summerville in 1920 (Amer. Fern Jour. 11: 90. 1921). Previous records report Lygo- dium japonicum appearing in a wild state in Florida, southwest- ern Georgia, Alabama, Mississippi, and Texas.—Marle B. LINGER, Route 1, Hardeeville, South Carolina. A Stmpte Meruop ror Growine FERNS FROM Spores.—Start with a sparkling clean, glass, one-gallon jug; a base made of one-by-four about seven inches long, to which screen molding has been nailed along eash side to keep the jug from rolling off ; a strip of thin aluminum two inches wide and 14 inches long; and a number ten tin can (clean, of course). Place a mixture of two parts good loamy soil and three parts of peat moss in the tin can, saturate with water, cover with a piece of aluminum foil, and bake in an oven at 400° or 500° F. for an hour or two to sterilize the mixture. While the soil is being sterilized and cooled, bend the strip of aluminum upward along both sides to form a narrow, evenly rounded trough which ean be inserted through the mouth of the jug. When the soil-peat moss mixture has cooled to room tem- perature, seoop up a bit and insert the trough holding the mix- 180 AMERICAN FERN JOURNAL ture into the jug and carefully roll it over to drop the soil-peat moss mixture to the bottom of the jug as it lies on its side on the rimmed piece of one-by-four, with the handle of the jug turned down. This method of inserting the soil keeps the “top” of the jug clean. After enough soil mixture to make a layer an inch or so deep has been placed in the jug, gently pull it back and forth to level the soil. (Don’t roll the jug laterally, for you want the glass above the bottom layer to remain crystal clean and clear). Next, sift spores into the aluminum trough, insert it into the jug and gently shake the spores over the surface of the soil. Add just enough cooled, sterilized water through a piece of tubing or with a syringe, to moisten the soil. The jug, lying on its side in the base formed by the board (painted if desired), should be set in a cool, shady place. Let time take care of it for a while. Oh yes, the mouth of the jug should be closed with a firmly packed wad of sterile cotton or gauze to keep out objectionable insects and reduce the rate at which water might evaporate. It is surprising how soon some fern spores develope into the prothallia, and these produce sec- ond generation sporophytes. Bracken sometimes will do this in two or three months. This method was developed to serve as a nearly fool-proof method for growing ferns for display in school libraries, class- rooms and lobbies, and to show youngsters how ferns grow from spores. Since I am a gardener for the whole North Bend School District, which consists of eight schools, I needed something of interest for the libraries and classrooms, and such displays had to be those that required no attention for prolonged periods. Although the scheme was intended to interest the children, many adults became fascinated with watching the plants grow. The jug provides an ideal climate for growth, because the warm- er the room becomes, the more rapidly the water in the soil of the jug goes into vapor in the confined space. Growing ferns in this way has brought up a puzzling question: REcENT FERN LITERATURE 181 When the growth first appears, the soil is covered closely with the minute prothallia. Later, after the prothallia have developed and young sporophytes appear, the latter are relatively far apart. Does it take more than one spore to produce a prothallium? If not, what happens to the rest of the tiny plants initiated by the spores?! This type of unit might be ideal for rest homes and residents for elderly people. They might keep memories green for the oldsters, and would need no care after the initial preparation. They last for several years, and the small plants grow beautiful- ly indoors.—JEssE T. Martin, 2622 Liberty St., North Bend, Oregon. Recent Fern Literature THe FERNS OF PERU. POLYPODIACEAE (DENNSTAEDTIEAE TO OLEANDREAE), by Rolla Tryon. Contributions from the Gray Herbarium of Harvard University. No. CXCIV, 253 pages, 196 figures, 42 maps. Cambridge, Mass. November 19, 1964—Dr. Tryon, who began his study of the fern flora of Peru about 1954, has wisely assembled and published that portion which he has thus far finished. Since this problem is a large one and the time that will be necessary for its completion is long, this partial account serves two purposes. First, it makes available without undue delay a useful contribution, and, second, it assures the author that his work is produced within his life-span in accord- ance with his ideas. This in no way precludes the hope that Tryon will live for many years to conclude this project. Tryon summarizes his contribution better than anyone else could do when he states, “The present portion treats the species 1It requires only one spore to produce a prothallium, and the initial growth often is remarkably good. It appears that every spore germinated and started a new prothallium. But competition among the rapidly grow- ing young plants is intense, and many more die than survive this process. but such crowding is inimical to the health of the tiny plants and many 0 them fail to produce sporophytes—and die.—Editor. 182 AMERICAN FERN JOURNAL of seven tribes of the Polypodiaceae. These tribes are repre- sented by 33 genera and 176 native species. This is probably about a third of the genera of Pteridophyta to be found in Peru and about a quarter of the species. Three introduced species are also treated and eight that are to be expected in Peru, one of these in an additional genus.” With few exceptions, the work is essentially monographic, and Tryon is to be congratulated on the fact that he has made every effort to examine the holotype, isotype or authentic material of each basionym. This effort forms a solid basis for his thorough and conservative treatment of all the species in the seven tribes studied; namely, Dennstaedtieae, Cheilantheae, Pterideae, Vit- tarieae, Davallieae, Lindsaeeae, and Oleandreae. The work comprises a short introduction that includes some notes on the geography of Peru with comments on the distribu- tion and some ecology of the ferns of that country based on its four major vegetational areas: the Lomas, or coastal foothills of the Andes; the Sierra Steppe and Scrub, the generally dry mountainous region of Central Peru; the Ceja, or cool moist high eastern slopes and ridges of the Andes; the Montafia, or wet forested eastern part of Peru. The introductory matter is concluded by a synopsis of the family Polypodiaceae with a key to all of its 57 genera that Tryon knows to occur in Peru. The keys and descriptions are lucid and comprehensive. Criti- eal characteristics of practically all of the species are well- illustrated, and maps are included for some of the species to show their distribution in Peru. A feature that I find attractive is the inclusion of several vignettes that have as their motif interesting habitat views of ferns. Tryon’s work is one of the most thorough on ferns for any country in South America. It will also be very useful for the identification of ferns in these tribes found in countries adjacent to Peru. All who are interested in ferns will want a copy of Tryon’s work.—Donovan S. CorreLu, Texas Research Fownda- tion, Renner, Texas. AMERICAN FERN SOCIETY 183 American Fern Society New Members Mrs. Arnold Bentein, Route 5, Franklin, Ten Mrs. Lonnie A. Dickens, 805 E. Roosevelt St., “Dillon, So. Carolina 29536 Miss Phyllis K. Epstein, 825 Loma Vista Drive, Beverly Hills, Calif. Mr. Pierre Fischer, 1716-A Virginia St., Berkeley, Calif. Mr. Brij Gopal, Dept. of Botany, Banaras Hindu University, Varanasi-5 (U.P.), India Mrs. L. David Hanna, 4129 Fairman St., Lakewood, Calif. 90712 Mr. Arthur ms Hawkins, Sunday Telegram, Box 1460, tata Maine 0410 Dr, Charles ei Kuehnert, Dept. es Bacteriology & Botany, Syracuse University, ces N. ¥. 13210 Miss Susan Leelike, 212 E. 13th St., nae York, N. Y. poss Miss —— eae 25 Ser Street, Hamburg, N. Y. Mr. Wes . Moore, 1923 So. Washington St., Vihbure ‘alld 39180 Mrs. Bet sat ete 1643 So. Jamestown, Tulsa, Oklaho Mrs. L. Peyser, Old Sleepy Hollow Road, Br iarcliff aie “N. Y. 10510 Miss Ellen Quail, 192 Rural Ave., S., Salem, Oregon 97302 Mr. Dan Sinn, R.R. 2, Tremont, Llinois Mr. Mike Slavin, West Mountain Road, Ridgefield, ee tate Mrs. C. James Steinbach, 4811 Eppes, Houston, Texas Mrs. Paul J. Swanson, 112 8, Barr St., Craw fordsyille, bee 47933 Mr. John W. Sweet, Star Route, Ondivarat: Calif. 93644 rs. yd: Dr. William 8. Wiedorn, 1304 Mouser of New Orleans, La. 70115 CHANGES OF ADDRESS Miss Harriet E. Baker, 1508 Campbell Ave., Thousand Oaks, Calif. 91360 Mr. Bill Bauer, Botany Hill, Route 2, Box 735, Arnold, Mo. Dr. David E. Blaydes, 1301 Van Voorhis Rd., Apt. H, Morgantown, W. Va. Mr. Herbert J. Cash, R. 3, West Bend, Wisconsin Mrs. Kenneth L. Clarke, 505 Virginia Ave., Santa Ana, Calif. 9270 Mr. G. J. oe par enehees re, c/o Oranje Lijn, N. V., West Plein 11, me Box , Rotterdam, Holland Mr, ie 3. Denault, 10319 Lorenzo Dr., Los Angeles, Calif. Dr. John F. ag rqul 112 Pine Road, Nanticoke Acres, Seaford, Delaware Mrs. Robert I eee 137 Fairlawn Ave., Albany, ue 03 Mr. Robert Halbeisen, 6382 London, Detroit, Mich. 4829 Miss Muriel P. Hedgwood, Cullowhee, N.C Miss Mary Howard, aie Calif. 94924 184 AMERICAN FERN JOURNAL Mr. Paul Hutchison, Dept. of Botany, University of California, Berkeley, Calif. 94720 Mr. Thomas A. Hutt 4 B Parkway, Greenbelt, Md, 20770 t: Holi aie ey wee uce St., Philadelphia r. Emory C. Leonard, Fairfield es nnels, Owings lanl. Mr, sae H. aaa del, 41 Fift atl fib Gia. N. Y¥. 10003 Mrs. James H, Mason, Box 279, oh t Terre Haute, Indiana 47885 Mrs. aSsise Milan, 2622 Waverly Dr., ae Angeles s, Calif Mrs. Marion B. Rhodes, Star Route, Montague, Mass Capt. Peter G. Root, Hd. 15th Med. Bn., APO San Prcudleas 96224 Prof. Alma G, on c/o Lenette Atkinson, 415 S. Pleasant St., Amherst, Mass. 01002 Mrs. Carl ao. 355 Cindy Lane, garaaee Fla. 33511 Mrs. Baxter Venable, 232 Chruch Rd., Ardmore, Pa. 19003 Dr. Dean Fe Whittier, Dept. of Ge mae fires. Substation B., Vanderbilt et: Nashville, Tenn. 37203 Drira D,; eee, , 1249 Luanne “ae i Calif. 92631 Mr. John Yopp, ena Apt. H., Padueah, Ky. A NEW FERN BOOK Learn of Ferns We Grow by Sylvia B. Leatherman and Dorothy S. Behrends Ferns for mild climate gardens : House Ferns : Spore Culture : Unusual ways to grow ferns : Illustrated with line drawings Price: $3.85 plus 15¢ handling. (Californians add 15¢ tax). Order from:—B & L Books Dept. A South El Monte, Calif. 91733 Index to Volume 55 Abortion bas “ph ees and Sporangia as aie oe pomp of Soa i tis Ht Ape RIRRS ee Adiantum, 49, 78, 171; andicola, rh: bra aunii, 77, 78; eandatum 57; concinnum 77, Habel tum 157; ba hoa 1675 pedatum, 104, 157; Get peuke, An Analysis Variable Population of Reutastwnt arvense and E, X le, 123 oe . bhareeee pe eye Anemia oo TT. hirsdta, ¢773 ucaue & 58, BL: schiedana, 119 Anogramma laptophst a, Arachniodes aristata, 155; chinensis, roichas boot ttii, 99; rigidum Asplenium, whi 420; 222; ti icu ae adamsii, 55, opic 56; fragran 118; harpeodes, 118; Renitomeul 52, h ochr roum, 152, xX tr ichomanes, 65, 104; uni- laterale, Athyrium rcmnaan 7. ees asplenioides, 104, 105, var. michauxii, 104, ft; aaah ps 105, f. elegans, 105, f. laciniatum, 105, f. rubellum ~ 5; schimperi, B38: thelypte- oides, =< Atkinson, "Fase ., Sehizaea pusilla— Shean tophyte?, 81; metophyte of Cystodium, Per — Banerji, M. L., Osmunda regalis var. spectabilis in Tyeaie 84 divergen 0: nice eo 78 : ; Se eet cs 79; Blotiella gracilis, 55 Bolbitis rete 165 Bommeria ata, 7 Botrychium, 113; alabamense, 80, L475 ctum, 102, 103, f. elon- m, 102, f. obliquum, 102, ae oe 102, hick lJanceolatum angustiseg- mentum, 102; vmnatricatiifolium, 103; multifidum var. inter- medium, 103; simplex, 8: ternatum, 103; virginianum, 64, Pie EE es Bowen, John E., The pcientialtiy of Boron on et ark 103 Dryopteris dentata and Selagi- nella apoda, 67 Brainea insignis, 154, 155, 157 covillei 1 copa: 115; . rane angey 75. 1 notholae- noid 15; . m ‘115; viscida, 113; wootonii, 115; wrightii, 113 Chen, Katherine Lim (See Wagner, W. Cibotium, peli 161 Cnemidaria, 163 Cochlidium. yostra Coniogramme tenga Correll, Donovan S., Cyatopteri bulbi- - a (L.) Bernh. New to Tex- County Distribution of Ferns hae Fern Allies in Rhode Islan a. Crandall, _Dorothy ib; tag of Ferns nett in Island, Otenitis rea 6 Culcita, 34, “161, * 163; subgen. Caloch lae 34, 37, 163; eoniitolis, Oulcita, 163; 163; macrocar- Cyathea, 2s tl Pek, SEG ‘borea, 30-32, var. nigrescens, 30, 31, var, pallida, 31; sub Cya- thea, 154; elegans, 31; fulva, 118; sect. G ta) ris, 3 caro fortun 151 um, 34; sorbifolium, §2,..85 65, 80, 84, 85. Hf mg 64, 105, 150, mackayi 105; protrusa, 150 Cystopteris. “hulbifers (L.) Bernh. i eetignge and Morphological Observ: on Met coseelle (HBK) Presl, is Cytological Observations on Ferns From Sout ines China, 154 Danaea cuspidata, 119 186 Dennstaedtia, 37, 161; adiantoides, 59; bip innata, 59, 60, 62; eieutaria, 9-62: da vallioides, 58, say a es dissecta, 62; punctilobu ra 59, 61, 62, 64, 105; sac pa 59-61; smithii Dicksonia, ane 34. 35, L6i. LG34-.ar bore: 37; hieronymi, hs se Me ra, 87 Dieranopteris, 161 Diplazium, coed bantamense, 155; don- ia = 155; zanzibaricum, 56 Dipteris, 16 Dispersal vot a mucronata by Wa- Birds, 167 Doodia, 18 Drynaria volkensii, 53, 56 Dryopteris, 9, 12, 19-21, mB Be Ss By 52 Sto ustralis, 107; xX boottii, La, 23 6; campyloptera, 29, 20, 25-27, X spinulosa, 16, 19, 27; clintoniana, 20, LOT oK cristata, 106, 107, goldiana, 20, var australis, 107; cristata, LS,-23 647:105,.. 107, xX, in media, 16, 23, 106, 107, marginalis 20, 106, spinulosa, 16; dentata, 67-69, 71-73; dilatata, 16, 19, , 28 < intermedia, 16, ‘ spinu- ; disjuncta, 105, 111 filix-mas, 19, 21-23, ar- ons. 19- 21; fregrans, 16, 24, intermedia, X spinu- ios sa var. feria Heke 16; goldi- 107; hexagonoptera, 105 intermedia, 13; 746, 29% 22-27, 108, X spinulosa, 16, 19, 27, 106, 107; marginalis, 19 21, 22, 64, 105,47. elegans. 106; novaboracensis, 105; pen- ‘theri, 53, 56; phegopteris, 105; rigida, 7; simulata, 99, 105; x slosson 107; inulosa, 16, 23, 2 i es Ve fructuosa, 107, var. intermedia, 24, 105 ae By spinulosa, 106; t! hely- frpioidea, 10 ‘107; * uliginosa, 93) vill 1,- Duncan, Wilbur H., gas Donald Blake, Obse ryations on Some Ferns in Georgia, 145 Elaphoglossum 76,79, 41 um, 78; arvense, 160, tae. 125, 12 13 145; 129, 13 ii 34; myriocha cant: 78; palustre, 123," 137, 128 133 aera re var. ulti- —— ar i- The Basentiality "ot ee for Dryopteris and Selaginella apoda, Farrar, D. R. (See Wagner, W. H.) AMERICAN FERN JOURNAL Fernwood Incorporated, 143 The Gam kip of Cystodium, 32 Gauch, Hu See John E. Bowen) Gleichenia banerottii a pee bifida, 79; almata, 7 +P +I ie un iene 11% Grammitis, Gp eeeion dryopteris, 106 Hauke, gy rd L., An Analysis of a ariable Population of Equise- ped arvense and E. litorale, Ebel er ey Holttum, ey he Si.) Hymenophyitum elegantulum eee fu- oides, 118; microcarpu 18; ai jocarpum, 117; ee sicunian ss Hypolepis, 120; nigrescans, 118 Tsoetes engelmannii, 102; muricata, 102; zee ria, 102, var. canadensis, 2; tuckermanii, 102 ; Japanese » hain bine Fern ger euines in outh aah -metay rns ‘(Dennstacdtia) orgy 58 Knobloch, tee ot of the Genus Gheilenthon, Lastrea eat 7 Lomaria meyeriana, 165, tenuifolia, 165 Lomariopsis, 165, ie Lophosoria, 163; adripinnata, qu 79 Lycopodium slop ecnrsiies. | 1455 ‘cernuum 79, 14 clavatum, 64, 79, 100, 121, megastac chyon, "100, £015 souiplimatass m 64, 79, 121; var. flabelliforme, 146; flabel- liforme, 101, 146; inundatum, 66, 100; linifolium, 121; luci- dulum, 64, 100; obscurum, 64, 100, var. dendroideum, 100, 101, var. obscurum, 101; ophio- glossoides, 53, RAD pithyoides, 121; reflexum, 1 21; isco, iot. 154, aS Lygodium japonicu 179; caiiee tum, 66, 103 Malone, Charles R., and Vernon W. Proc- , Disper sal of Masaitos mu- ata by Water Birds, 167 Marattia, qT19: oe 62; weinmannii- folia, & Marsilea nape eee 167,.169, Martin, yA assead Ret A Simple * Method of ’ Ferns from Spores, 179 Woeuseig, seb, Cte brs yee McDowell, Gladstone W., Binge: to Pan- her Creek, Georgia, 80 Mellinger, Pa mi B. Japanese Climbing sg Spreading in South Caro- 159, 161- es Metaxya, 158, 161; rostrata, 163 Mickel, pore a Preliminary yg Stud- n the Fern i f Oaxa- ps micxigo; (4,117 Microlepia, 6 iisdbhovititn fortunei, 157 Morton, C. V., A New Jamaican Cyathea, INDEX TO VOLUME 55 187 30; Observations on Cultivated Ferns, ides: Capa 9 164 oo ee pumice New Gytological meeartd for Cystodium and Dicksonia, ae A New Fusriad ican Cyathea, A New —— Form of Pa alle ea Fee re ar. glabella bg a at Notholsens candida weohe PEs oy os es near a aes ralis, 78: Suanevethons oe: on lItivated Ferns, VIII. Stenochlaena, 164 Observations on Some Ferns in Georgia, 5 Odontosoria cot ta on 79 Oleandra distenta, Onoclea Sénaibiin 8, ba. 108 aabenana ve te 147; at enadag engelmannii, 150° eaiitaate 149-150; Bits latum, 148-150 ; vulgatum, 103, 148-150 Osmunda se mea, 63, 79, 103, 147, Fo Hater er 107; Sanyo a, 63, 103, 150; regalis, 55, 79, 8 r 8 ar. J nica plumieri, 84, var. ne "4, 1 Osmunda eg r" var. spectabilis in In- Panigrahi, G., hiner inary studies on the C xX at of the pert villarsii Bel .) Woyn Com plex in E a <8 A Parcel of Cam Peilace. phe ND hese grt 178; glabella, 178, "174, 178, var. glabella, 178, var. oc- cidentalis, 173, 178 , var. sim- plex =o a ovata, 77; quadri- 49 , 173, 174, pin 5 OE Bow serge 106; ee, 64, Pilularia americana, 152 rn gramma, "62: pe a 120, var. calomelanos, 52 Platycerium Polypodium, ane; anfractuosum, 118; bore 81: cultratum, iis: loriceum, 118: lowei, 118; palustre, 166; pilo- sissimum, ft : lechrum, 79; rigidum eta-rosae, 78; seolopendria, 52, ni aa ad, villarsii, (eee 64, 08 r carostishothes: 64, 108; istatum, 155; denticulatum, 1 8; fus copaleaceum, Preliminary goer paper on the Fer Flora of Oaxaca, Mexico, 74. Polystiehum, iauiauae® 3 Studies on the Oytotaxonomy eee rig ate em regs . i u Pa ayer wnihateea eridium Efe 117; aquilinum va lat- sculum, 108; latiusculum, 64 A Pteridophyte Botanical Garden, 63 Pteris, ae camerooniana, 55; fauriei, 57; ; multifida, 157; ae try 120; Naatehner ag 56, 20, 157; togoensis, 56 Report of: "Audit Committee, 44: Judge of ‘Blections, 44: Presi- oe 8; Secretary, 40; Trea- -Seepes oe Panther Creek, Geo Roy, 8. K., a ie) > Hoittum, "Croton faa ee oy * Morphologi cal yas on Metax ne = ata HBEK) Presl, aha Cytological Susarentions on om. Southern China, ise Ne ew Cyt- pie gical ie a for " Oystodium ickso Rumohra cristata 155 Russell, Helen oss, A Eueierh ys Botanical Garden, Schiza aea Ree 81-83 Schizaea —Perennial Gametophy- Selaginelly 120, 122; apoda, 69, 71, 2, 147; eee Tits “ealbreve: marten x palles is ‘02: 6; versicolor, 52, elii, 52; Sandy, +6 A Simple Stothod. “of "Growing Ferns from Spores, 179 awe anita, 164, 165; sect. Cafr 164, mi a. oat 655 St ene, T65; tenuifo a ery 166; Struthiopteris seaitela, Mentors it 164, 16 Thel ris, 106, 120, ae aertir 78; a 79 iata, 53: Sr chaeastibise 64: “oli igocarpa. 79; palustris, 65; puberula, 78; ulata, 65, 66; 5 simulat: torresiana, 15 Thyrsopteris, +61; Trichomanes ‘oorkontions: ee 67 Trismeria trifo oliata, 79 ‘ He Parcel of Cameroon Fer gn Pc oe of the Genus Wagner, W. f “ Tool in the Detection of ‘Diyopteris Hybrids, . D. BR. Farrar, and Katherine L. Chen, A New Sexual form ok Pellaca ee var. glabella Mis i") q &. Bd Been Pag 55 Be ce e32 BS ° Ey ocd $ ~~ aed ardia ae ata, 15 sid io9 Xiphopteris Paaikaseend: 118; serrulata, 79 188 AMERICAN FERN JOURNAL soon pocket-siz classics gf an a hauoeae ry htembes of the American Fern Soc ociety DR. EDGAR T. WHERRY THE FERN GUIDE Complete, accurate and con- venient; covering 135 spe- cies of the Northeastern and Midland United States and adjacent Canada; with draw- ings of 135 species by Dr. James C. W. Chen. $4.95 THE SOUTHERN FERN GUIDE The only comprehensive, up- to-date guide available to ferns of the Southeastern and South-Midland with W. Chen: Doubleday Nature Guides Series at all booksellers ef DOUBLEDAY | =e BOOKS ON VASCULAR CRYPTOGAMS Recently published or actually in preparation The Ferns (Filicales) By F. O. Bower, 1923-28 (Reprinted 1963). Cloth. Filices of the US-Exploring Expedition (“Wilkes Report”) By W. D. Brackenridge. 2 vols, 1854-55 (Reprint ready 1966). 366 pages in quarto, 46 plates in folio. Cloth. A Monograph of the Fern Genus Woodsia By D. F. M. Brown. Beiheft 16 zur Nova Hedwigia. 1964. 164 pages, 40 plates. Mémoires sur la famille des Fougéres By A. L. A. Fée. 11 parts im one volume. 1844-66 (Reprint ready 1966). Quarto. 990 pages, 181 plates. Cloth. Historische Entwicklung der Nomenklatuar und Taxonomie der Gattung Iséetes L. Von H. P. Fuchs. Beiheft 3 zur Nova Hedwigia. 1962. 108 pages, 22 plates. A Toxonomie Monograph of the Genus Equisetum, A Toxonomic Monogre*—$<—<—$__ Subgenus Hippochaete By R. L. Hauke. Beiheft 8 zur Nova Hedwigia. 1963. 128 pages, 22 plates. Section Complanata of the Genus Lycopodium By J. H. Wilce. Beiheft 19 zur Nova Hedwigia. 235 pages, 40 plates, illustrations. $ 30.00 45.00 15.00 175.00 10.00 15.00 J. CRAMER e PUBLISHER e 3301 LEHRE e GERMANY Exotic and Hardy Ferns Begonias BOLDUC’S GREENHILLS NURSERY 2131 Vallejo Street St. Helena — California Open Saturdays and Sundays 10 A.M. to 4 P.M. or by appointment Phone 963-2998—Area Code 707 Mail orders accepted