American Fern Journal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY ea EpITors DAVID B. LELLINGER C. V. MORTON ROLLA M. TRYON IRA L. WIGGINS e VOLUME 58 - 5‘ Been \1)4 MONUMENTAL PRINTING COMPANY, BALTIMORE, MARYLAND Missouri EOTANICAL GARDEN LIBRARY Contents Votume 58, Numper 1, Paces 1-48, Issupp Marcu 26, 1968 Edith Seamman (1882-1967)..... Rolla M. Tryon and Alice F. Tryon A Study of Spore Characteristics of the Ferns of Indiana Jeanette C. Oliver Rate of Gametophyte Maturation in Sexual and — Forms of NR MMNONR Sec. cp tory eh, os ee eee ee Dean P. Whittier The Gametophyte and Juvenile Leaves of be Sorcha .B. K. Nayar Shorter Notes: Polystichum acrostichoides X lonchitis versus P. X _ marginale; A New Bolivian Collection of the Rare Elaphoglossum eget Trichomanes petersii in the Boston Mountains of Arkansas. ee Notes and } American Fern Society: Report of the President; Report of the Secretary; a of the Treasurer; Report of the Auditing Committee; Report of the Judge of Elections; Report of the Spore Exchange; Report of the 1967 Fern Foray Os 60 hale ely en Win OR TNE ae wid hk ee ee pee en Pet. ee eS Barbara Joe Hoshizaki The Valid Varieties of Pellaea andromed aefolia....... ide omas R. Pray . Troop pa dae T. Mickel Marsilea maheshwarii, a New Species a Poadishars: sage 17 Gopal _ Phloem Studies in the Pteridophytes, Part I. Equisetum. e ru N. Agashe ANew — of Salviniaceae and a New Species of Azolla from the late Cretaceo John all Re ee ee eRe eR ME eee ge 5 gt gl Ca neers eed munitum in South Dakota; Trichomanes es petersii Revisited at Sara- sc acl Madi, oo en SN Ie are ne Oe 8 ee ew tee ited hath te oe ee bens Eh Sadat Mh ADA det dee Sa A ay ee ee ee ee PS Fe Oe wie. lee ele St ee OO SF Oe ee ee mee Sim oan a Ce al _ Shorter Notes: Notes on Woodwardia areolata and Other Rare Illinois Ferns; A Spiny Dennstaedtia from Central America; Polystichum toga, Mississippi; Lygodium microphyllum, Another Fern Escaped VOLUME 58, NUMBER 3, Pages 97-144, Issuep SerTeMBER 20, 1968 A Culture Chamber for Tropical Rain Forest Plants.. Donald R. Farrar A Presumable Asplenium Hybrid from Kentucky. ..Thomas N. McCoy The Correct Name of a Common Tropical American Oleandra . Morton Vv a Reproduction in the Ferns I. Leaf Buds ot Grammitis NN a oR ON EN PSO OREN COR phe ea s Richard R. ite Seana Revision of the Polypodiaceous Genera of India-III. Pleo- peltis S. S. Bir and Chander K. Trikha Trends of Specialization in the Stipe Anatomy of Dennstaedtia and mated Glnern i ses 55590 6 i Richard C. Keating phorter vote: A. NOte on ASpmotie) aco. 080s ci oe ees eee eI RN ae Fs a ie a os Ls als eae ee EN eR EpeCeiy MGrh: LARGO eo oot i lee ees Oe Oe PAUP IORE PPI, OES is a ois ok cao ee dl we bo ve oe ee ee Se ee ek 8 ee we 6 we 8 ee eel 8 6 ote Voutume 58, Number 4, Pages 145-188, Issuzp DecemMBER 26, 1968 Alma Gracey Stokey (1877-1968)............ Lenette Rogers Atkinson Polypodium vulgare on Long Island.................. F. R. Fosberg Pee Ge PM PND lg cs is we os David B. Lellinger The Fern Collections in Some European Herbaria...... . C. V. Morton The Polypodium pectinatum—plumula Cait in ee A. rray Evans Exclusion of the New York Fern and the Hay-scented Fern from the Me OE AON ct, po te Re ares T. 8S. Cooperrider Shorter Notes: The Identity of Polypodium salicifolium Vahl; Asplenium trichomanes New to Newfoundland; The Correct Name for the Button Fern; Mecodium wrightii on Vancouver Island Notes WO Og aid sg Vos oo cele se Pees ieee ees cree eeee Mk Pe ee lapa puns © ake Oued S0b > kee Ceo Vou, 58 JANUARY—Marcu, 1968 No. 1 American fern Journal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY ts DAVID B. LELLINGER 0, V. MORTON ROLLA M. TRYON IRA L. WIGGINS ty CONTENTS Edith Scamman (1882-1967)....Roiua M. Trron anp Arice F, Tryon A Study of Spore Characteristics of the Ferns of Indiana JEANETTE C. OLIVER Rate of Gametophyte Maturation in Sexual and Apogamous Forms of IMINO ig os Si Sec Ac i ees Dean P. WHITTIER The Gametophyte and Juvenile Leaves of Loxogramme..B. K. Nayar Shorter Notes: Polystichum acrostichoides x lonchitis versus P. X marginale; A New Bolivian Collection of the Rare Elaphoglossum cardenasii; Trichomanes petersii in the Boston Mountains of Arkansas SOFC HPTLC SCR HOCK OSES SHO TOK SSC CTO SSE HEC OD HOC HES O eRe eee American Fern Society : Report of the President; Report of the Secretary; Report of the Treasurer; Report of the Auditing Committee; Report of the Judge of ——— Report of the Spore Exchange; Report of the 1967 Fern Fora Pewee eee eeee ses eeseteeee seers sneseeeonereeerseeee ree Miesour: Botan: APR 22 1968 iy or _ bw Che American Fern Hocietp Council for 1968 Invinag W. Knostocu, Department of Botany & ont Pathology, Michigan State University, East Lansing, Michigan President Warren H. Waaoner, Jr. , Department of alee: University of Michigan, Ann Arbor, Michigan 48104. ee Lenetre R. ATKINSON, 415 S, Pleasant Street, Amherst, Massach gp etary y K. HENRY, Division of Plants, Carnegie Museum, ues ~~ —_— vicaain 15213. Davip B, LELuingeR, Smithsonian Institution, Washington, D. c. “20560. Editor-in- Chief HMational Society Representatives Wageen H, WaGnesr, sn Dept. of Botany, University of Michigan, Ann Arbor, a * 8104, Representative to A. A. A. 8. H. Wagner, Jz., Dept. of Botany, University of Michigan, Ann Arbor, Michigan 48104, Representative to A. I. B. 8. American Fern Journal EDITORS B. LELLINGER____ Smithsonian Institution, Washington, D. C. 20560. 4 ‘v. ae. Seatteoaten Institution, Washington, D. C. 20560. Rotua M. Tryon Gray Herbarium, Harvard University, Cambridge, Mass. 02138. Ira L, acacaammein ey Herbarium, Stanford University, Stanford, Calif. the Pace ig Fern Society, and published at 3110 Elm Ave ertemeee _ 21211. Second-class postage paid at Baltimore. The pages es of the _Jour- are i : ‘eren Matter for publication should be addressed to Dr. David B. Lellinger, Smithsonian Institution, W. ashington, D. C. 20560, Changes of address, applications for membership, subscriptions, orders — ees and other business communications should be addressed bers of the American F, song coment dues, $4.00; mem- bership, $8.00; life membership, $80.00). Gevasted i ar if ordered in adv au ors at cost, plus Back volumes $5.00 to $6.25 each; single back numbers of 64 pages or $1.25; 65-80 00 3 over 80 pages, ; ulative Index to Volumes 1-25, 50 cents. Ten percent discount on orders of six volumes or more, Library and Herbarium Dr. W. H. Wagner, Jr. Department of Botany, Universi: —_ Ann Arbor, Michigan £8104, in js Nbrarian and curator. te aig borrow books and specimens at ang robacctagien paying all pat Bip or express charges. Spore Exchange Mr. Neill D. Hall, 1225 Northeast 95 ost, Seatti, caeaies is Director of the 7 Viab bed Spore Exchange. Viable spores and dis patehed, and lists of the Midinmes tian: American Pern Journal Vo. 58 JANUARY—Marcn, 1968 No. 1 Edith Scamman (1882-1967) Rouua M. TrRYon AND ALICE F. TrYON “When the list in my pine-lot had reached sixteen, I climbed my neighbor’s fences and wandered farther afield.”” This remark in Edith Scamman’s (1923) delightful account of the ferns in her woods in Saco, Maine, sounded the prelude to long adventure and travel far afield in Alaska, Iceland, Europe, Costa Rica, Porto Rico and other Caribbean islands. Her botanical pursuits are most remarkable in that they followed studies in Middle English literature leading to a Master’s degree from Radcliffe College and years of activity on church mission boards, both on the Prudential Committee, and as President of the Missionary Council of the Congregational Church Women of Maine. Miss Scamman joined the American Fern Society in 1937 and served as its Secretary in 1950 and 1951. She was a member of the Overseers Committee to visit the Gray Herbarium, Harvard University, from 1942 to 1946 and held an appointment as Re- search Associate in the Gray Herbarium from 1949 to 1962; since that time she was an Honorary Research Associate. Her death on November 4, 1967, less than a month before her 85th birthday, was in Cambridge, Massachusetts, where she kept residence for many years. Edith Scamman was born in Saco, Maine, on November 30, 1882, the only child of Francesca Allen Scamman and Henry Scamman; her grandfather, John Allen, was the attending phy- sician. Her father, after years in California during the Gold Rush Period, returned to Maine and settled his family in the large home built by his grandfather. He instilled in his fond daughter a sense of adventure and pioneering spirit which were to carry her, Volume 57, No. 4 of the JourNAL, pp. 145-188, was issued December 22, 1967. y. AMERICAN FERN JOURNAL decades later, to new frontiers. His death, when she was less than ten, brought her closer to her mother. They traveled to San Francisco, where she attended school for a short period. She entered Thornton Academy, in Saco, after returning to Maine. Evita SCAMMAN WITH PLANTS OF BLECHNUM BUCHTIENII NEAR LA CHONTA, IN THE SIERRA DE TALAMANCA OF CosTA Rica, in Aprix 1951. Except for her college years and a brief time in New York she lived with and devotedly cared for her mother. She graduated from Wellesley College in 1907 with a major in English. Two years later she received a Master’s degree from Radcliffe College, and her study in Middle English, “The Alliterative Poem: Death Eprra SCAMMAN a and Life,” was published in 1910, in Radeliffe College Mono- graphs, Studies in English and Comparative Literature. Her botanical activities centered at the Gray Herbarium, Harvard University, when she came there in 1935 to study with Professor M. L. Fernald and enrolled in his classes as a Radeliffe student. She became acquainted with many students and eminent botanists who were to influence her work. Professor Fernald at that time was interested in plant distributions in unglaciated Alaska; Miss Scamman became a willing ally and promptly embarked on a program of botanical exploration in this region. She collected there on nine separate trips (the last in 1954) and she made a total of some 5000 collections which are deposited at Harvard. Her research on these collections was published in 1940, “A list of plants from interior Alaska.’? Many of her specimens were sent to specialists and several new species, a Spring Beauty (Claytonia scammaniana), a Poppy (Papaver scammaniana) and a Crucifer (Lesquerella arctica var. scammanae), were named for her. She published an account of the ferns of the Yukon in THIS JOURNAL in 1949. During this period she was also closely associated with Mr. C. A. Weatherby. With his encouragement and guidance she enlarged her botanical experience during the preparation of ‘“The Ferns and Fern Allies of New Hampshire.” This useful reference for students and botanists interested in New England ferns was published by the New Hampshire Academy of Science in 1947. It is an accurate and well written account with many illustrations. Her deep appreciation of botanical specimens is revealed in the introductory remark: “As one studies the beautiful specimens gathered and preserved with so much care and reads the hand- writing of many old labels of former years one can picture the delight and sense of achievement which came to many early fern students.” With her interests concentrated on the ferns, she embarked in a wholly new direction, in tropical America with the study of Costa Rican ferns. She collected there in 1951, 1953, 1955 and 1956. During these trips she made about 1400 collections representing 4 AMERICAN FERN JOURNAL about 500 species. Specimens of these are also deposited at the Gray Herbarium. The extensive collections thus accumulated formed the basis of several publications, with technical keys, illustrations, and descriptive material. The first of these was on the Maidenhair ferns (Adiantum) of Costa Rica, in 1960. Although she was at this time in her 78th year, she continued with similar treatments of other genera: Pteris, Oleandra, and Eriosorus. A paper on the genus Asplenium was in preparation at the time of her death. Her collections from Costa Rica as well as those from the Caribbean area have been cited in other recent studies, and a rare fern from the high mountains of Costa Rica, Jamesonia scam- manae, commemorates her studies and collections in the American tropics. Her appreciation and concern for the American Fern Society, as a member for thirty years and as its Secretary, is thoughtfully expressed by her bequest of three thousand dollars to the Society. BiBLioGRAPHY OF EpiTH SCAMMAN 1923. Ferns in my pine lot. The Maine Naturalist 3: 18-3 1940. A list of Plants from Interior Alaska. Rhodora 42: aie Reprinted in Contrib. Gray Herb. 132: 309-349. 1940. 1947. The Ferns and Fern Allies of New Hampshire. New Hampshire Acad. Sci. Bull. No. 2 —96. 1949. Ferns and fern alice of the Central Yukon Valley. Amer. Fern J. 39: 1-12, 45-58. 1960. The mre ferns (Adiantum) of Costa Rica. Contrib. Gray Herb. 87: 1961. The genus eg of Costa Rica. Rhodora 63: 194— ge 1961. The genus Oleandra of Costa Rica. Rhodora 63: 335- 1962. The genus Eriosorus in Costa Rica. Contrib. Gray fact 191: 81-89. Gray Herpartum, Harvarp University, CAMBRIDGE, Mass. 02138. | SPORE CHARACTERISTICS 5 A Study of Spore Characteristics of the Ferns of Indiana JEANETTE C,. OLIVER Most taxonomic work on the ferns in the past has involved gross morphology, stelar anatomy, and the structure and position of the sori. Recently attention has been directed toward the diagnostic values of spore characteristics (Brown, 1960; Crane, 1953; MeVaugh, 1935; and Marengo, 1956). My work has been carried out to assess the value of spores as a means of identification of the ferns indigenous to Indiana. As a result a diagnostic key was devised using spore characteristics. The spores examined were fresh, except for Dennstaedtia punc- ttlobula (Michx.) Moore, Woodwardia virginica (L.) J. E. Smith, shares atropurpurea (L.) Link, and Cheilanthes lanosa (Michx.) . Eaton, which were Obie from herbarium specimens. oe four ferns have been collected in Indiana, but are rare (Deam, 1940). Untreated spores were observed for color and size. Fifty spores of each species were measured using an ocular micrometer. Spores of each species were taken from several specimens chosen randomly. For detailed study, spores were treated according to the acetolysis procedure of Erdtman (1943). A set of reference slides was prepared by mounting spores in Hoyer’s Medium. This clears the spores and acts as a preservative. Slides may be kept for several years. Most species can be distinguished by differences in shape and in the structure and markings of the perispore and exospore layers. However, the species of Botrychium and Osmunda are very similar, with the main difference being size. The spores of Athyrium pycnocarpon (Spreng.) Tidestr. closely resemble those of Asplenium. Further, the spores of Athyrium asplenioides (Michx.) Desv. have ridges much like A. pycnocarpon, whereas Athyrium angustum (Willd.) Presl, does not have distinct wings and ridges, but is tuberculate. VotuME 58, PLATE 1 AMERICAN FERN JOURNAL DrawinGs or Spores or INDIANA FERNS SPORE CHARACTERISTICS t The species of Thelypteris are placed in the genus Dryopteris by some authors. On the basis of spore morphology, it is difficult to distinguish characteristics which are confined to either of the genera. Thelypteris hexagonoptera (Michx.) Weath., 7’. phegop- teris (L.) Slosson, and 7’. noveboracensis (L..) Nieuwl. are distinct in that there are no obvious perispore wings. Thelypteris palustris Schott has perispore wings similar to Dryopteris. The gross, rough- ened spores of 7’. palustris, however, are similar to other members of the genus. Some authors place JT. hexagonoptera and T. phegopteris in a different genus, Phegopteris, but spore morphology does not support this concept. KeryY TO THE GENERA AND SPECIES 1. Perispore (perinium) absent or obscure.....5.....0..0.00- 00 cence eee 2 Ty POrepore Greene oe ck ke i a ee 15 2. Spares clear oF nearly sao ee ee Pts ic ee 3 a. Spores yellow of brown 6 ede a aa 10 By SIOERE RUGUNAD ac ee ag ee ee a ee 4 SOONER OPW Sy eee toy as hire ca Sa 7 4, PYOLOpIAst Broel; @xine DIOrOUE or o.o. Vi 6 eal eo ks hd es 5 4. Protoplast not green; exine reticulate. Ophioglossum engelmannii Prantl (30-40 u; Fig. 4) o. Sporen tee than G0 o i WIAINOtel. F336 ok ol cost Cie Ge sce we 6 5. Spores 60—69 uw in diameter................ Osmunda regalis L. (Fig. 1) 6. Spores 36—45 uw in diameter............ Osmunda claytoniana L. (Fig. 2) 6. Spores 46—58 yw in diameter........... Osmunda cinnamomea L. (Fig. 3) dq. poles Wetranea ra to a. bo ie Oa oe 8 7. Spores bilateral..... Polypodium vulgare L. (38-45 X 53-60 »; Fig. 5) Re RU SAA a ae dct la wie 9 8. Exine: smooth 20. oe Adiantum pedatum L. (32-41 y»; Fig. 7) 9, Spores 42-44. 38-40 gic bes oe ak Botrychium dissectum Spreng. 9. Spores 35-36 X 35-36 n..... Botrychium virginianum (L.) Swartz (Fig. 6) Fic. 1. OsMunDA REGALIS L. Fic. 2. OSMUNDA CLAYTONIANA L. Fic. 3. Os- MUNDA CINNAMOMEA L. Fig. 4. OPHIOGLOSSUM ENGELMANNII PRanTL. Fig. 5 PoLyropium vu 2 LF OTRYCHIUM VIRGINIANUM (L.) SwWARTz 1G. 7. ADIANTUM PEDATUM L. Fic. 8. PreRipIuM AaQuILinuM (L.) KuHN Fic. 9. DENNSTAEDTIA PUNCTILOBULA (Micux.) Moo I CysTo- PTERIS FRAGILIS (L.) BERNH. Fic. 11. CysTOPTERIS BULBIFERA (L.) BERNH. AMERICAN FERN JOURNAL VotuME 58, PLATE 2 DrawineGs oF Spores oF INDIANA FERNS SPORE CHARACTERISTICS 9 10: Sporeateirehedral nS ook a er ee ee es 11 10. ppordetawieral or ovals: so ee. SS ee es a 12 11. Exospore minutely scaly. Pteridium aquilinum (L.) Kuhn (28-34.5 yw; Fig. 8) 11. Exospore verruco Deiievaiints eae: (Michx.) Moore (80-35 & 20-25 yu; Fig. . IZ. Sporesaiiateral, offen spiiwicees soo. os eae eee 12. Spores TOUMG-OMOVAL ORCA os is mon a i ee ee i 13. Spores spinulose. Cystopteris fragilis (L.) Bernh. (23-32 34-41 yu; Fig. 10) 13. Spores granular with a few prominent ridges Cystopteris Sooty (L.) Baral (17-26 X 34-38 u; Fig. 11) 14. Spores oval, yellow-brow Pellaea ie (L.) Link (43-46 X 53-56 uw; Fig. 12) 14. Spores round, yellow Chedlanthes lanosa (Michx.) D. C. Eaton (88-42 yw; Fig. 18 ib. Pernspor cweey adherent oo. is oe a is Ved es 16 15; Perispore sen die attached. wardia virginica (L.) J. E. Smith (36-40 XK 42-60 u; Fig. 14) 16. Spores Sie to he hd aan arte Bek ayy ep Cee Suara ony aM tome e 17 16. Spores clear to grayish. Polystichum pen aac (Michx.) Schott (35-45 & 48-53 yu; Fig. 15) i, Perispore wig nat opeiotiess 666 2006205 Se a es ee 18 iinPeringors Wey miatinet ges a a a ee 24 18. Exine with subtle, reticulate or oval ridges..................-..-4- 19 18. Exine waberotiate, often with rides. 5. Gen. i Fee US ee eS 22 19. Ridges elongate, distinct; spore surface granular or oe Mai a 20 19. Ridges limited, inconspicuous; spore surface gran Thelypteris phegopteris (L.) Slosson (0-47 x 30-60 u; Fig. 16) 20. Ridges broken; spore surface granular................+..02.-00005 al 20 Ridges anastomosing, reticulate; spore surface smooth. Thelypteris hexagonoptera (Michx.) Weath. (00-27 * 39-50 w; Fig. 17) Fig. 12. PeELLAEA ATROPURPUREA (L.) Link. Fic. 13. CHEILANTHES LANOSA (Micux.) D. C. Eaton. Fig. 14. Woopwarpia virGINica (L.) J. E. Smita. Fie; 15. ee ea ACROSTICHOIDES (Micux.) Scuorr. Fic. 16. THery- PTERIS PHEGOPTERIS (L : ony eee Fic. 17. THELYPTERIS HEXAGONAPTERA (Micux.) Wratuersy. Fic A fuecversnii PALUSTRIS ScuorTT. Fig. 19. THELYPTERIS NOVEBORACENSIS gs Niguw.. Fig. 20. ArHyRIUM ANGUSTUM (Wixip.) Presu. Fig. 21. ATHYRIUM PYCNOCARPON (SpRENG.) TipEst. Fig. 22. ATHYRIUM ASPLENIOIDES (Micux.) Desv. AMERICAN FERN JOURNAL VoLUME 58, PLATE 3 DrawinGs oF Spores oF INDIANA FERNS SporRE CHARACTERISTICS 11 21. Wings distinct. . ..Thelypteris palustris Schott (34-45 X 53-63 y; Fig. 18) 21. Wings absent. Thelypteris noveboracensis (L.) Nieuwl. (26-34 X 34-45 p; Fig. 19) 22. Wings distinct; spore surface ridged 2... 35 biG ls. oe Soe 23 22. Wings obscure; spore surface tuberculate. Athyrium ae (Willd.) Presl (27-38 X 32-39 u; Fig. 20) 23. Ridges reticulate, anastomos Athyrium pycnocarpon (Soren ) Tidestr. (832-41 XK 38-49 y; Fig. 21) 23. Ridges sisca tie little anastomos Athyrium. asplenioides (Michx. ) Desv. (33-35 X 39-42 yu; Fig. 22) 24, Perispore wing wide or continuous : 065 ohh A wy be 25 24. Perispore wing narrow or brokemeiis. <2iii5 es aE PA cis 26 25. Ridges prominent, primarily longitudinal. amptosorus rhizophyllus (L.) sea (24-32 X 34-39 uw; Fig. 23) 25. Ridges few, both longitudinal and transve Woodsia obtusa (Spreng.) ae (30-34 X 38-48 p; Fig. 24) au. Peripore Wik Wille, broke. 20500009) oe ee es ee ee 27 26. Perispore wing narrow, broken or not. Onoclea sensibilis L. (36-57 X 58-65 pw; Fig. 25) 27. Ridges regularly anastomosing, the pattern reticulate.............. 28 27. Ridges occasionally anastomosing, mostly not reticulate............ 29 28. Major ridges 3-4 in lateral view; spore surface granular between ridges. Asplenium pinnatifidum Nutt. (80-38 & 41-53 y; Fig. 26) 28. Major ridges many; spore surface not granular between ridges. splenium —- (L.) Oakes (32-41 X 38-52 yu; Fig. 27) 0. Badges cloned; anastomoses, iii. bebe Fred CA eS. ee, oe 2 30 29. Ridges Sdnwkis. Hite ANAEUOINOSING: ye oi shoe wh ee ne Ce BL 31 30. Spores entirely ridged. Dryopteris marginalis (L.) Gray (26-38 X 38-53 »; Fig. 28) 30. Spores sparsely ridged. Dryopteris cristata (L.) Gray (30-40 X 40-60 u; Fig. 29) 31. Ridges loosely reticulate. Dryopteris — (Muell.) Watt (26-35 & 38-45 yu; Fig. 30) 31. Ridges elongate, not reticulat Dryopteris aia, (Hook.) Gray (25-30 & 30-40 u; Fig. 31) Fig. 23. CAMPTOSORUS RHIZOPHYLLUS (L.) Link. Fic. 24. Woopsia OBTUSA (SpreNnG.) Torr. Fic. 25. ONOCLEA SENSIBILIS L. Fic. 26. ASPLENIUM PIN- NATIFIDUM Nutt. Fig. 27. ASPLENIUM PLATYNEURON (L.) Oakes. Fic. 28. DRYOPTERIS MARGINALIS (L.) Gray. Fic. 29. Dryopreris cristata (L.) Gray. Fic. 30. Dryopreris spINULOsSA (MugE.uL.) Watt. Fic. 31. Dryo- PTERIS GOLDIANA (HooK.) GRAY. 12 AMERICAN FERN JOURNAL LITERATURE CITED Brown, C. A. 1960. What is the role of spores in fern taxonomy? Amer. Fern J ~ 502 6-14. Crane, Fern Warp. 1953. Spore studies in Dryopteris I. Amer. Fern J. 43: 159-169 Dream, C. C. 1940. Flora of Indiana. pp. 37-59. Dept. of Conservation, Indianapolis, Ind. Erpmman, G. 1943. An introduction to pollen analysis. pp. 26-31. Ronald Press, New York. McVavan, R. 1935. Studies on the spores of some northeastern ferns, Amer. Fern J. 25: 73-85. Marenoo, N. P. 1956. The microscopic structure of the mature spores of the sensitive fern, the ostrich fern, and the royal fern. Amer. Fern J. 46: 97-104. DEPARTMENT oF Biotocy, Batu Strate University, MUNCIE, Inp. 47306. Rate of Gametophyte Maturation in Sexual and Apogamous Forms of Pellaea glabella Dean P. Wuirrrer’ Wagner et al (1965) have reported the occurrence of a diploid sexual form of Pellaea glabella var. glabella in Missouri. Previously, this variety was known only to be obligately apogamous at the 4x chromosome level (Tryon and Britton, 1958). A comparison of the haploid and tetraploid gametophytes of the sexual and apogamous forms respectively is of interest because artificially produced polyploid series of gametophytes in other fern genera (Manton, 1950; Whittier, 1966) demonstrate a number of quan- titative variations. The sexual and asexual methods of sporophyte formation have different requirements, e.g., water for sperm movement, which is required for sexual reproduction, is un- 1T wis i of thls aptciee AAS RE De hana tok Tat nena ices Te encase blake at DeSoto, Missouri. This stud i ’anderbilt University Recah Comet y was supported in part by the Vander GAMETOPHYTE MATURATION IN PELLAEA 13 necessary for apogamy. Modifications in gametophyte growth and development possibly accompany the method of sporophyte formation. The gametophytes of the sexual and apogamous forms of P. glabella var. glabella provide an opportunity to investigate prothalli of the same species with variations in chromosome number and mode of sporophyte formation. Spores of the sexual (Whittier 3) and apogamous (Whittier 10) forms of Pellaea glabella var. glabella were obtained from DeSoto, Missouri (See Wagner et al, 1965) and Rome, Tennessee, respec- tively. Voucher specimens have been deposited in the Vanderbilt University Herbarium. Aseptic techniques (Whittier, 1964) were employed to sterilize and sow the spores on 25 ml of nutrient medium in petri plates. The medium consisted of Knudson’s solution of mineral salts, FeXEDTA, minor elements, 1% or 0% sucrose, 0.6% agar, and had a pH of 6.0 (Whittier, 1964). In one experiment, antheridial hormone (Naf, 1958) at a 1/10 dilution was incorporated into the medium to insure antheridium formation. The areas of the gametophytes were determined with a polar planimeter (Whittier, 1964). The measurements reported are the means of 64 individuals, and the standard errors of the means are included. Over 200 gametophytes were examined to determine the percent with archegonia or initial stages of apogamy. Spores of the apogamous and sexual forms are trilete with ow ridges on the exine. Spores of the apogamous form exceed in diam- eter (73.2 + 0.6u) those of the sexual form (55.7 + 0.5 yu). The development of the gametophytes was observed using prothalli grown on the nutrient medium lacking sucrose. The early development in both forms is very similar. The spores become swollen and crack at the triradiate ridge in three days for the apogamous form and in four days for the sexual form. The spore divides to produce the first prothallial cell and first rhizoid which enlarge to rupture the spore coat (Figs. 1, 2). Generally, two or three cell divisions give rise to a filament of three to four cells (Figs. 3, 4). A small plate of cells arises as the cells divide parallel to the filament axis (Fig. 5). In some instances no filament is formed, as irregular cell divisions of the first prothallial cell form AMERICAN FERN JOURNAL VoLuME 58, PLatr 4 es a fe l0 9 DEVELOPMENT OF APOGAMOUS (SHADED) AND SEXUAL (UNSHADED ) GAME- TOPHYTES OF PELLAEA GLABELLA, X 300. Fics. 1, 2. SPORE GERMINATION. pe 7 EARLY FILAMENTOUS STAGE. Figs. 5-8. YOUNG PLATE STAGES. FIG. ¥. {LATE STAGE WITH APICAL CELL. Figs. 10, 11. ANTHERIDIA. GAMETOPHYTE MATURATION IN PELLAEA 15 a plate of cells directly (Fig. 6). With additional cell divisions and growth, larger plates develop (Figs. 7, 8) and ultimately an apical cell arises (Fig. 9). The plate continues to grow by the activity of the apical cell, but eventually a marginal meristem replaces the apical cell (Fig. 12) and a small cordate prothallus' develops. y” d i 4 ae Fy ? f . : Le i Arica, Notcw Region or Mature Corpate GAMErTopnuytes, X 200. Fie. 12. SEXUAL FORM SHOWING MARGINAL MERISTEM AND ARCHEGONIA (ARROWS). Fic. 13. APOGAMOUS FORM SHOWING EARLY APOGAMOUS DEVELOP- MENT WITH SPOROPHYTIC HAIRS (ARROWS). In the presence of antheridial hormone, the young gameto- phytes form antheridia (Figs. 10, 11). The diameter of the anther- idia are 48.6 + 0.4 and 36.2 +0.3y for the apogamous and sexual forms respectively. No antheridia were observed unless antheridial hormone was added to the medium. After the initiation of the small cordate prothallus, the develop- ment of the gametophytes differ. In the prothallus of the apoga- 16 AMERICAN FERN JOURNAL mous form, cells behind the notch divide to produce a mass of meristematic cells (Fig. 13) which develop into an apogamous sporophyte. The cells behind the notch in the sexual form give rise to a cushion with archegonia (Fig. 12). The archegonia and an- theridia of the sexual form are functional because diploid juvenile sporophytes have been collected at DeSoto. Fertilization was not accomplished in culture, apparently due to technical difficulties. TABLE I. Size Comparison (IN mm?) OF SEXUAL AND APOGAMOUS AMETOPHYTES OF PELLAEA GLABELLA Sexual form Apogamous form 0% sucrose 1% sucrose O% sucrose 1% sucrose Size on 21stday 0.2040.01 0.2840.01 0.39+0.02 0.52 + 0.02 Size at first apo- gamy or archegonia 1.31 + 0.04 0.83 40.08 1.18 +0.04 0.64 + 0.02 The rate of prothallial development was compared between the two forms. The growth of the apogamous gametophytes is faster with or without sucrose (Table I) than is the growth of the sexual. In both forms prothallial growth is greater with sucrose in the nutrient medium than without it. The prothalli of the asexual form initiated apogamous sporophytes in a shorter period of time and on smaller prothalli than the gametophytes of the sexual form produced archegonia (Table I). On the 26th day after sowing the spores, 50% of the gametophytes of the asexual form on 0% sucrose had initiated apogamous sporophytes and none of the prothalli of the sexual form had produced archegonia. Five days later only 21% of the gametophytes of the sexual form had archegonia on 0% sucrose. With 1% sucrose, 79% of the gameto- phytes of the asexual form had apogamous developments and 21% of the gametophytes of the sexual form had archegonia on the 26th day. The presence of sucrose in the medium caused apogamous sporophytes and archegonia to be initiated sooner and on smaller gametophytes than without sugar (Table I). Thus gametophytes of the apogamous form were more efficient than those of the sexual form with or without sugar in the medium. 2c oS pty pa GAMETOPHYTE MATURATION IN PELLAEA 17 The findings of the present study on spore size and sculpturing confirm the reports of Wagner et al (1965) and Pickett and Manuel (1925). The growth of the gametophytes of the apogamous form in sterile culture is somewhat different from the growth on soil as reported by Pickett and Manuel (1925). The gametophytes in sterile culture have a shorter filamentous stage and the apogamous plant is mitiated two months sooner than the gametophytes in soil culture. The development of the apogamous sporophyte from the gametophytic cells behind the apical notch appears to be the same under both conditions. Spore germination, gametophyte growth, and apogamous sporophyte initiation were faster in sterile culture than on the soil cultures of Pickett and Manuel. The size differences between the spores and antheridia support the findings of Manton (1950) and Whittier (1966). Manton has reported that the diameter of spores and antheridia increases with an increase in the level of ploidy in Osmunda. Size differences between the apogamous and sexual form appear to be due to the difference in the chromosome number and not to the mode of sporophyte reproduction. The promotion of gametophyte growth and the formation of apogamous plants on smaller prothalli earlier with sugar in the medium has been reported by Whittier (1964, 1965) in other fern genera. Since the initiation of archegonia was also accelerated by sugar, it appears that sugar similarly affects gametophyte develop- ment in both forms. The earlier conclusion (Whittier, 1964, 1965) that sugar brings about the conditions necessary for apogamy, and in this case archegonia, by affecting carbohydrate metabolism and the availability of respiratory substrate appears also true for P. glabella var. glabella. In discussions of apogamy in ferns (Tryon & Britton, 1958; Stokey, 1948; Whittier, 1965) it has been noted that water for sperm movement is not required for apogamous reproduction. Gametophytes of an apogamous species can grow and produce sporophytes using only capillary water from the soil. Sporophytes cannot form by fertilization on gametophytes with mature sex 18 AMERICAN FERN JOURNAL organs with only capillary water (Duncan, 1941) because a film of water is necessary for sperm movement to the archegonium. Farlier investigators (Hayes, 1924; Stokey, 1948; Tryon & Britton, 1958) noted the advantage that apogamous species have in drier habitats because this additional water is unnecessary. besides the advantage of only requiring capillary water, apogamous reproduction occurs faster than sexual reproduction in P. glabella var. glabella. Since the gameto- phyte is More susceptible to desiccation than the sporophyte, more rapid sporophyte formation is important because less time in the life history of the plant is spent as a gametophyte. The conclusion that apogamous reproduction is more rapid and more advantageous than sexual reproduction in P. glabella var. glabella 18 Supported by the greater range of the apogamous form. An explanation of the early maturation of the 4z of the apogamous form is impossible at this time because in P. glabella phytes GAMETOPHYTE OF LOXOGRAMME 19 Literature CITED Duncan, R. E. 1941. Apogamy in Doodia caudata. Amer. J. Bot. 28: 921-931. Hayes, D. W. 1924. Some studies of apogamy in oe atropurpurea (L.) ink. Trans. Amer. Micro. Soc. 43: 119-13! MANTON, 1 1950. Problems of Cytology and Bvelation in the Pteridophyta. mbridge University Press, Cambridge Nar, U. 1958. On the physiology of antheridum tendinitis in the bracken fern (Pteridium aquilinum (L.) Kuhn). Physiol. Plant. 11: 728-746 Pickett, F. L. and M. E. Manvgt. 1925. Development of the stall and apogamous hee ly in Pellaea glabella Mettenius. Bull. Torrey ee Club 52: 514. Stoxry, A. G. 1948. me eametophyte me Actiniopteris australis (L. fil.) bea Link. J. Indian Bot. Soc. 27: Tryon, A. F. and D. M. Brirton. ae pean studies on the fern genus Pellaea. Evolution 12: 137-145. Waener, W. H., Jr., D. R. Farrar, and K. L. Cuen. 1965. A new sexual form a Pellion glabella var. glabella from Mixa: Amer. Fern $552 .171-178. Wuirtier, D. P. 1964. The effect of sucrose on apogamy in Cyrtomium alecatum Presl. Amer. Fern J. 54: 20-25 1965. Obligate apogamy in Hiring tomentosa and C. alabamenss Bot. Gaz. 126: 275-2 . . Induced dlp in nap gametophytes of Pteridium. Bee st Bot. 44: 1717-1 DEPARTMENT OF GENERAL BroLoGy, VANDERBILT UNIVERSITY, NASHVILLE, TENN. 37203. The Gametophyte and Juvenile Leaves of Loxogramme B. K. Nayar Some years ago I described spore germination and prothallus development in two Indian species of Loxogramme, based on materials collected in the field along with samples of the sporo- phyte (Nayar, 1955). The prothalli were collected on the tufted, sponge-like root masses of L. involuta and L. lanceolata. It was soon evident that reporting mature prothalli as cordate possibly was a mistake. Since then I have made several attempts to raise pure cultures of Loxogramme prothalli in the laboratory and to review the nature and development of these prothalli. 20 AMERICAN FERN JOURNAL For this study spores of L. involuta (Blume) Presl and L. lanceolata (Swartz) Presl were collected during different years from various parts of Assam, mostly from the Khasi Hills. The spores were cultured on sterile nutrient agar at 24 + 2°C and 600 ft-c. light intensity (Nayar, 1962). Prothalli at different stages of growth also were collected attached to their substratum and transferred to the laboratory. These were maintained in the laboratory under the same conditions of light and temperature as the agar cultures. No marked difference was noticed between the prothalli raised from spores and those collected in the field. Spore GERMINATION AND PROTHALLUS DEVELOPMENT Morphology of the spores of several species of Loxogramme, including L. involuta and L. lanceolata, has already been described in detail (Nayar, 1963¢, 1964; Nayar and Devi, 1964). The spores of L. involuta are monolete-bilateral, whereas in L. lanceolata both monolete-bilateral and_ trilete-tetrahedral spores occur mixed together (Nayar, 1963c). A perine is absent in both. Fresh spores contain many large plastids and a few deep yellow oil globules. As can be expected of thin-walled, chlorophyllous spores, those of Loxogramme cannot withstand desiccation. The spores remain viable for only a short time, and are difficult to culture in the labo- ratory. However, when sown fresh and when contamination 1s avoided, they germinate profusely in about two weeks. The oil globules in the spore become prominent and often coalesce into a centrally placed, large droplet ; the plastids become deep green. Soon the spore swells and the exine breaks open at the laesura, partially exposing the deep green prothallial cell. A proximal rhizoid initial is soon differentiated, which grows into a short, achlorophyllous rhizoid. The rhizoids have a distinct violet- brown tinge in L. lanceolata. The prothallial cell, meanwhile, enlarges and protrudes as a thick papilla lateral to the rhizoid. The protruded portion is cut off by a transverse wall from the bulbous basal region, and by successive transverse divisions and elongation of the daughter cells develops into a uniseriate germ filament (Fig. 1). The basal cell of the germ filament is bulbous, GAMETOPHYTE OF LOXOGRAMME 21 and the oil globule persists for a long time in the basal cell. Oil globules are not generally found in the other cells of the germ filament. Rhizoid development is sometimes delayed on both species, for germ filaments devoid of rhizoids are quite frequent in the cultures. By about two weeks after spore germination the germ filament is usually 3-5 cells long and is composed of deep green, barrel- shaped cells. Some of the proximal ones bear lateral rhizoids (Fig. 1) which are brownish in both species, and are usually very short in L. lanceolata. The germ filaments of Loxogramme are slow growing. Increase in length ordinarily results from the division of the terminal cell, or less frequently by division of the older cells. There is a strong tendency for the cells of the germ filament to elongate markedly. There are many growth irregu- larities in the germ filaments of L. lanceolata. Simple, elongate germ filaments, typical of L. involuta, are rare in those of L. lanceolata raised in culture. In many cases the germ filaments are branched; the branches develop laterally from some of the median cells (Figs. 3-4). In most cases the basal cell, which is the prothallial initial, bears several germ filaments, and in some cases the filaments are so crowded that the spore appears to germinate into an amorphous mass of cells (Fig. 6). However, the basal cell can easily be distinguished by its characteristic oil globule and larger size. In some cases the germ filament terminates in arhizoid and stops growth when it is 2-5 cells long. Then lateral branches grow out into secondary germ filaments (Figs. 6 and 9), similar to those reported in Christiopteris tricuspis (Nayar, 1967). These secondary germ filaments may be branched in turn (Fig. 7) Rhizoids are produced profusely; often the basal cell bears 2-4 of them. Another peculiarity of the germ filaments of L. lanceolata is that they are moniliform. The cells usually have bulged sides (Figs. 2 and 5), but especially when crowded they may be pyriform (the anterior end broad and the posterior narrow). Most germ filaments grow erect or oblique to the substratum; branches form in all directions, giving the cultures a characteristic appearance. Formation of a prothallial plate is initiated in the cultures JOLUME 58, PLATE 5 AMERICAN FERN JOURNAL V : SAME, CA. 8 WEEKS OLD SHOWING PROTHALLIAL PLATE FORMATION. FiG. 9. SAME, CA, 10 WEEKs OLD. Figs. 10-12. PROTHALLIAL PLATE FORMATION IN L. 7 GAMETOPHYTE OF LOXOGRAMME 26 about 6-8 weeks after spore germination. The germ filaments are usually 5-8 cells long by then, but sometimes, particularly in L. lanceolata, there may be an extensive development of the fila- mentous stage. The distal cells of the germ filament, including the terminal cell, divide longitudinally, and the daughter cells expand laterally to form a dorsiventral prothallial plate (Figs. 8 and 10). Individual branches of the germ filament develop into separate prothalli (Figs. 8 and 9). About two weeks after the initiation of plate formation an obovate prothallial plate with a smoothly rounded or somewhat flattened distal end is formed (Figs. 11 and 12). Cells in the distal region are more actively meristematic and thus progressively smaller. The prothalli of L. involuta are comparatively quicker growing, and may develop into a prothallial plate 6-8 cells broad (Figs. 14 and 15) about two months after spore germination, whereas those of corresponding age in L. lanceolata are often only half as large (Figs. 8 and 9). Soon the prothalli become spathulate, and later the distal region expands markedly, making the prothallial plate broader than long (Figs. 13 and 14). Rhizoids are produced in profusion from the marginal cells at the posterior end. e young prothalli of Loxogramme, unlike those of most leptosporangiate ferns, never develop a regular meristem or meristematic cell. In some cases, however, an obconical marginal cell is developed which acts for some time as a typical meristematic cell by cutting off daughter cells regularly on the oblique sides (Figs. 14 and 15), but its activity is never prolonged. About three months after spore germination the prothalli become nearly circular and about 2 mm in diameter, and are composed of small, densely chlorophyllous cells (Figs. 16 and 17). The cells are progressively smaller towards the distal margin. Superficial rhizoids similar to the marginal ones are produced on the posterior half. In L. lanceolata the prothalli exhibit a marked tendency to INVOLUTA. Fig. 13. SPATHULATE PROTHALLUS OF L. LANCEOLATA CA. 11 WEEKS OLD. Figs. 14 anp 15. SPATHULATE PROTHALLI OF L. INVOLUTA CA. 2 MONTHS OLD SHOWING OBCONICAL APICAL CELL. THE SHADED AREAS REPRESENT OIL GLOBULES. > yet ee ee ae OED res 2 AMERICAN FERN JOURNAL VoLuME 58, PLATE 6 Tw F IATL Ty 200 * | REO Se LUN UOG 4 ES cht e | Siler RES nN S2 3 | q . : 3 | \S E 4 Pe : ) Bi : yf pees bine one oF Loxocramme. Fig. 16. L. INVOLUTA, CA. 12 WEEKS ' or L iat SAME, CA. 15 WEEKS OLD. Fic. 18 HAPED PROTHALLUS ‘ eg Betbicheaan OVER 3 MONTHS OLD. Fic. 19 SAME, APICAL PORTION. FIG wiht aeetie NEARLY 4 MONTHS OLD SHOWING BRANCHING. Fia. 21. SAME, ’ MONTHS OLD. Figs, 22 anp 23. LONGITUDINAL SECTION OF ARCHE- ; GA TE OF LOXOGRAMME 25 elongate at this stage and soon become strap-shaped (Fig. 18). Rhizoids are produced all over the lower surface and are mostly aggregated in irregular patches; marginal rhizoids are developed profusely on the posterior half. In L. involuta, on the other hand, the prothalli expand and become irregular in outline and even irregularly lobed due to unequal marginal growth. The lobes develop into broad branches, often 3-10 at the distal end (Fig. 21). Occasionally only two lobes develop and the prothalli appear cordate (Fig. 20). There is no well defined meristem in these branches either; like the unbranched prothalli, the cells at the distal margin are smaller and divide more actively. Rhizoids develop on the lower surface and sometimes on the margins of the branches (Fig. 21). All prothalli are one cell thick through- out; in L. lanceolata they bear characteristically short rhizoids. ApULT PROTHALLI Further development of the prothalli is by diffused meriste- matic growth. The prothalli become ribbon-like and highly branched. Prothalli of both species grow slowly. In culture they remain one cell thick for nearly a year, and by then may be over 1 em long. Sex organs are produced in patches superficially on the lower surface of the branches behind the growing apex. Antheridia are very sparse and are nearly always mixed with clusters of rhizoids. They are of the common polypodiaceous type, having a central mass of sperms surrounded by a 3-celled wall which is composed of a short, saucer-shaped basal cell, an annular cell, and a cap cell. Archegonia are found mixed with antheridia. Where archegonia are borne the thallus is 2-4 cells thick, forming a Cushion subtending the sex organs. When archegonia are crowded, the cushions subtending individual archegonia merge to form a single broad patch usually two cells thick. Archegonia are initiated behind the growing apex of the branches where the thallus is still only one cell thick. The archegonial initial becomes slightly larger and its lower wall protrudes markedly from the surface of the prothallus. This cell divides transversely, making the prothallus two cells thick. The daughter cell on the lower surface of the prothallus divides again and develops into the archegonium in 26 AMERICAN FERN JOURNAL the usual way, forming a row of three cells. The outer one pro- trudes and develops into the neck, the middle one into the egg and neck canal cell, and the lower one into the basal portion of the venter. Mature archegonia (Figs. 22 and 23) are small and have a short, nearly straight, papilla-like neck composed of four tiers of three cells each. The neck canal cell is narrow and binucleate at maturity. The basal row of neck cells divides vertically towards maturity, making the neck two cells thick at the base, as in most advanced leptosporangiate ferns. 25 27 JUVENILE Leaves or LOXOGRAMME INVOLUTA. Figs. 24-32. PRoGRESSION OF VENATION PATTERN. Fig. 33. PoRTION OF LAMINA FROM SPOROPHYTE, CA. 6 MONTHS OLD. HATCHED AREA REPRESENTS MIDRIB. Fic. 34. Harrs OF THE FIRST JUVENILE LEAF. Mature prothalli of Lozogramme are highly branched and ribbon-like. The branches usually have irregular, sometimes lobed sides. They presumably grow for years and form extensive patches. Some prothalli grown in cultures formed masses nearly 2.5 em broad and had several crowded, lobe-like branches growing nearly erect after about two years. Much bigger masses were collected in the field. The prothalli remain one cell thick, except for irregularly placed areas where sex organs are borne. Midribs are absent, as are trichomes of all sorts. GAMETOPHYTE OF LOXOGRAMME Ze JUVENILE SPOROPHYTES As in other polypodiaceous ferns, the juvenile laminae of Loxogramme are simple and entire. The following description is that of L. involuta alone because juvenile plants of L. lanceolata were not available for study. The first few leaves have spathulate or strap-shaped laminae which taper downward to a narrow base (Fig. 24). A single, unbranched vein traverses the lamina and ends well behind the rounded apex. In succeeding leaves the vein is unequally 1- or 2-forked at the apex, and a midrib is differen- tiated by successive unequal forking and overtopping (Fig. 25). Such pinnately branched veins sometimes are found as early as the second or third juvenile leaf. Soon the basal lateral veins become forked (Fig. 27), and the branches (which face each other) of successive veins fuse to form areoles (Figs. 26-29). Sometimes the laminae are inequilateral with areoles only on one side of the midrib (Fig. 28). In some small leaves the single vein entering the lamina forks at the tip and the two branches form a terminal loop as in Vittaria (Wagner, 1952). A series of elongate, costal areoles develops on either side of the midrib of the laminae of larger, successive leaves (Figs. 29 and 30). Leaves from the fifth to seventh onward are supplied by a pair of parallel vascular strands, each of which bears the lateral veins of its side, and which fuse distally. Regular vascular commissures between the two main vascular strands form a row of narrow areoles which constitute the midrib of the leaf (Fig. 31). Meanwhile, the lateral veins become prominent, a second row of smaller areoles forms beyond the costal row (Fig. 31), and short, free-ending veinlets which point toward the midrib develop within the costal areoles from the outer vein of the areole. Larger juvenile laminae develop several rows of areoles beyond the costal ones, and main lateral veins become more conspicuous. The midrib in these leaves has three main vascular strands (Fig. 32). A venation pattern quite similar to that of adult leaves is formed by plants nearly six months old (Fig. 33). In some cases early juvenile leaves are forked, with each lobe receiving a branch of the forked vascular strand entering the leaf base. Early juvenile 28 AMERICAN FERN JOURNAL leaves, including the first leaf, bear a few clavate hairs (Fig. 34) on the lower surface and on the margins. These hairs are bicellular, with a slender, elongate stalk cell and a rather swollen, probably glandular, densely protoplasmic terminal cell. They are similar to those reported on adult leaves (Nayar, 1955). Discussion This study corrects an earlier report on prothallus morphology and growth in Loxogramme (Nayar, 1955). The prothalli are of the ribbon-like, branched type and develop not by the activity of an obconical meristematic cell, but from a diffuse meristematic area. Although the prothalli of L. involuta may approach a sub- cordate shape (Fig. 20), this is due to diffuse development of lateral branches by the cessation of the meristematic activity of the medianly placed cells at the apex, and not by the development of a median apical meristem as reported earlier; the branches ultimately become elongate and ribbon-like. The elongate, ribbon-like, mature prothalli are much like those reported in some microsorioid genera of the Polypodiaceae, e.g., Kaulinia (Nayar, 1963a), Leptochilus and Paraleptochilus (Nayar, 1963b), Colysis (Nayar, 1962), and Christiopteris (Nayar, 1967). So also are the juvenile leaves, with their simple, entire laminae traversed by a solitary vein, but which later develop a midrib by over- topping. These facts appear to support Holttum’s (1947, 1949) belief that Loxogramme is polypodiaceous. In addition, the germ filaments of L. lanceolata occasionally branch profusely and stop terminal growth, while the lateral branches develop into individual prothalli, similar to those of Christiopteris tricuspis (Nayar, 1967). In many cases L. lanceolata produces several aggregated, branched germ filaments from a single spore. Loxogramme is sometimes considered a grammitid fern. But the prothalli of Grammitidaceae are basically cordate albeit narrow, elongate, and often strap-like (Stokey and Atkinson, 1958). They also bear characteristic trichomes, which are highly branched and glandular or elongate and acicular. A midrib, which is sometimes interrupted, is found in all. Branching of the prothalli GAMETOPHYTE OF LOXOGRAMME 29 is uncommon. An apical meristematic cell is regularly formed during early stages of prothallus development. This is replaced by a multicellular meristem. The prothalli multiply vegetatively by fragmentation of the germ filament. The antheridia possess slender, elongate, barrel-like basal cells. The early juvenile leaves of the grammitid ferns, insofar as known, have lobed laminae. All of these features differ from Loxogramme. However, the very slow prothallial growth rate, the protracted filamentous stage of L. lanceolata, the characteristic branching of the germ filaments, the tendency for one spore to produce multiple germ filaments, and the tendency toward moniliform germ filaments recall the prothalli of grammitid ferns. Probably these features are adapta- tions to similar environments. LITERATURE CITED Houttum, ee E. 1947. A revised classification of leptosporangiate ferns. J. n. Soc. London, Bot. 53: 123-158. 1949 The Giethimbion of ferns. Biol. Rev. 24: 267-296. Nayar, B. K. 1955. Studies in Polypodiaceae~IIT: Loxogramme (BI.) Presl. J. Indian Bot. Soc. 34: 395-407. 2. Mo arpholokey of the poeire and prothalli of some species of the Polypodiaceae. Bot. Gaz. 123: 223-232. - 1963a. Contributions to the morphology of some species of Microsochan. Ann. Bot., n.s. 27: 89-100. . 1963b. Contributions to the oo of Leptochilus and Passhestéahilae. Amer. J. Bot. 50: 301-309. - 1963c. Spore morphology of Loxogramme. Grana Palyn. 4: 87-94. Se 1084: egcertact of Modern Pteridophytes. Chapter vi in P. K. K. Nair, Advances in Palynology. United Block Printers, Lucknow. - 1967. Spores and prothallus of Christiopteris tricuspis. Amer. Fern J. 56: 5-27. aera S Devi. 1964. Spore morphology of Indian ferns—III: Po = aperaens Grana Palyn. 5: 314-367. Sroxey, A. G. . R. Atkinson. 1958. The gametophyte of the Gram- Be ducks Phytomorphology 8: 391-403. Waener, W. H., Jr. 1952. Types of foliar dichotomy in living ferns. Amer. J. Bot. "30: es 92. —______. — _ Diese ee NATIONAL BoTaNic GARDENS, Lucknow, Inpra. 30 AMERICAN FERN JOURNAL Shorter Notes POLYSTICHUM ACROSTICHOIDES X LONCHITIS VERSUS P. X MAR- GINALE'—In the spring of 1952, W. H. Wagner, Jr., and D. J. Hagenah discovered the hybrid Polystichum acrostichoides X lonchitis in the Bruce Peninsula, Ontario. They described the morphology of this hybrid in great detail.? Since the hybrid does not have a formal taxonomic binomial, I propose the following in honor of Dale J. Hagenah, longtime student of ferns: Potysticuum X hagenahii, hybr. nov. Verisimiliter hybridus P. acrostichoides X lonchitis, major et ad P. acrostichoidem ver- gens. Frons lanceolata, pinnulis infernis paullum tantum reductis. Frons fertilis pinnulis parum differentiatis, vix sporiferis et gradatim minoribus et angustioribus quam sterilibus. Polystichum acrostichoides X lonchitis W. H. Wagner & D. J. Hagenah, Rhodora 56: 1-6. 1954. Typr: Ontario, Bruce County, west of Cape Crocker Indian Reservation, base of low hummock about 20 yards from edge of woods, 1 large plant not far from parents, Fall, 1953, W. H. Wagner & D. J. Hagenah s. n. (DAO; isotype MICH). Another specimen collected from the same plant in the spring of 1952 by Wagner and Hagenah is preserved in the Herbarium of the University of Toronto (photo DAO). About thirty years before Wagner and Hagenah’s discovery, W. R. McColl described P. lonchitis f. marginale (Amer. Fern J. 14: 107. 1924). There are four specimens of this in MceColl’s herbarium, which was given to the University of Toronto. Three of the specimens are labelled Polystichum lonchitis f. marginale McColl. They were collected in 1925 and 1937. The fourth is labelled Polystichum lonchitis marginale?; this specimen, col- lected in 1923, carries the data: Woodford Road [10 miles from Owen Sound], in crevice, in sun and dust on roadside. These data are similar to those given by McColl when he described f. ? Contribution No. 564 from the Plant R h Institute. Research Branch, Canada Department of Agri pee ela es crunk * Rhodora 56: 1-6, pl. ci Ottawa. Se SHortTER Notes dl marginale. Although the 1923 specimen was not designated as the type by McColl, it is the only specimen in that herbarium which was collected prior to 1924, so it must be regarded as the holotype. At the time I discovered McColl’s specimens I suspected that they might also be the hybrid described by Wagner and Hagenah. This I mentioned to Bernard Boivin, who, in his ‘Enumeration des plantes du Canada,” published’ the following under “In- novations Taxonomiques”: ‘‘X Polystichum marginale (W. R. McColl) Cody stat. n., P. lonchitis (L.) Roth f. marginale W. R. McColl, Rhodora 56: 3-6. 1954.’ Because Boivin cited the Rhodora reference instead of the reference to McColl’s paper, it is clear he intended to name Wagner and Hagenah’s hybrid. I have recently examined the McColl specimens and found that among other characters, the spores are full and of similar size and shape. Therefore, neither the holotype of f. marginale upon which P. X marginale was based nor the other three MeColl specimens are of hybrid origin, and it follows that the correct binomial name for the hybrid is Polystichum X hagenahii.—W. J. Copy, Plant Research Institute, Central Experimental Farm, Ottawa, Canada. A New BotiviAn CoLLEectTion or THE RARE ELAPHOGLOSSUM CARDENASII.—During their recent visit to the Field Museum in Chicago, I showed Dr. and Mrs. Tryon a number of unidentified ferns, among which was a peculiar Bolivian specimen collected in December, 1966 by Roy F. Steinbach “entre los musgos sobre tallos de Arboles, Km. 104, Camino Chapare, Depto. Cochabamba, 3100 m.” Dr. Rolla Tryon thought it could be Elaphoglossum cardenasii, a curious species described by Dr. Warren H. Wagner, Jr. (Bull. Torr. Bot. Club 81: 62. 1954). Although neither frond on the plant which we examined is fertile, further study proved without question that it is indeed E. cardenasii, a strange pedately-lobed species first collected by Dr. Martin Cardenas in November, 1940, along the “Way from Cochabamba to Chimoré, about Km. 120; Province of Chapare, 3 Nat. Can. 93: 253-273. 1966. 32 AMERICAN FERN JOURNAL Department of Cochabamba, Bolivia, 2000 m.”? Although both collections come from the same general area, the new station Is distant enough and the lobing of both collections uniform enough to make it unlikely, as Dr. Wagner concluded, that the fionds of the original plant represent an abnormality. Our Steinbach 630 has two fronds arising from the suffrutescent rhizome, along with several bases of broken stipes. The larger frond has a stramineous stipe 42 em long, with a central lobe 20 em long and 2.1 em wide, at the base of which are borne two smaller, lateral lobes, one on either side, each of these being thrice-lobed, nearly to base. The second frond, which apparently is an immature fertile one, has a stipe 50 em long, and is divided much like the sterile one, but the central lobe is only 5 em long and 0.4 em wide and the lateral lobes are proportionately reduced. Dr. Wagner points out that “it would be desirable ... to find the early leaf stages which show at what stage in the progression of leaves the unique foliar organization arises.”’ Unfortunately the Steinbach specimen !n hand does not help in solving this problem, but perhaps now we can be optimistic that more E. cardenasit will turn up in future collections—Rosertr G. Srouze, Field Museum of Natural History, Chicago, Ill. 60605. TRICHOMANES PETERSIE IN THE Boston MovuNTAINS OF ARKANSAS.—Field studies (under National Science Foundation Grant GB-4095 to P. L. Redfearn) of the bryophytes of the In- terior Highlands of North America have resulted in the discovery of Trichomanes petersii A. Gray in Arkansas. Like 7’. boschianum, which was reported from Arkansas by Clark,! this species 1s associated with relic mixed mesophytie forest common there. The population of 7. petersii occurred at the base of a massive sandstone boulder along the bottom of a narrow ravine that is a tributary to Indian Creek, ca. 3 miles SW of Sandgap in Pope County, see. 16, T12N, R20W (Redfearn 21412, MICH, NCU, SMS, US, UT). Plants grew near the base of the boulder and con- sequently were subject to inundation by rapidly flowing water ? Amer. Fern J. 52: 85-86. 1962. ee Notes AND NEWS 33 during periods of excessive rainfall. The narrowness of the ravine also suggests that they are never exposed to direct sunlight. This first record of Trichomanes petersii from Arkansas con- stitutes a very large range extension, the nearest locality being in Franklin County in northwestern Alabama, more than 300 miles away. Although these plants belong taxonomically to 7. petersii as presently understood, they do have characteristics peculiar for this species and quite different from other populations of this species from the eastern United States. The blade bases approach a subcordate condition, as opposed to the typically more acuminate base. The length/width ratio of the blade is smaller than in the typical form; and there is a strong tendency for the midrib to branch dichotomously so that the blade often has two sori, or rarely more. Although these differences in the Arkansas plants may be genetic, it is possible they are due to environmental factors, perhaps damage of the blade apices. Consequently, culture studies of these plants are in order, and living materials are now undergoing investigation at the University of Michigan Botanical Gardens.—Dona.p R. Farrar and Paut L, REDFEARN, Jr., University of Michigan Botanical Gardens, Ann Arbor, Mich. 48105 and Southwest Missouri State College, Springfield, Mo. 65802. Notes and News Tue AmeERICAN Fern Society ANNUAL Meerine this year will be with the A.I.B.S. at Columbus, Ohio, home of Ohio State University. Dr. Jane Decker will be our local representative. We plan a foray on September 1 and 2. Foray headquarters will be at the Holiday Inn, Chillicothe, Ohio 45601. For reservations, write to the Inn (identifying yourself as a foray participant) or to Dr. Clara Frederick, Urbana College, Urbana, Ohio 43078. On September 3 we will have a Society luncheon, which will be followed by a program of papers. Dr. W. H. Wagner, Jr., eo Gardens, University of Michigan, Ann Arbor, Mich. 48105, 1 the program chairman. Contributors should send titles, a and projection equipment needs to him immediately. Kk. 34 AMERICAN FERN JOURNAL 18tH ANNUAL SPRING WILDFLOWER PILGRIMAGE.—Plan now to spend April 25-27 in the Great Smoky Mountains at the peak of the spring wildflower season. Although wildflowers will be the principal attraction, Dr. A. Murray Evans, of the University of Tennessee, will lead several fern walks. Motorcades and trail hikes under expert leadership will be offered. Early morning bird walks are planned, as are evening lectures, one of which will be Dr. Evans’ “Ferns of the Appalachians.” Participants should register beginning at 9 am on April 25 in the Gatlinburg Civic Auditorium. The registration fee is $2.00. There are no advance registrations. Descriptions of the various events will be furnished at registration. For lodging information, write to Department W. P., Box 527, Gatlinburg, Tenn. 37738. Tue Firra AnnuaL Tropica, FLtower & FERN SHow of the Los Angeles International Fern Society will be held in Brookside Park at Pasadena, California, on Saturday, May 18 (1:00 to 10:00 pm) and on Sunday, May 19 (10:00 am to 6:00 pm). For further details write to Bee Olson, 13715 Cordary Ave., Haw- thorne, Calif. 90250.—D.B.L. Recent Fern Literature THE SouTHERN FERN GuipeE, by Edgar T. Wherry. Doubleday & Co., Garden City, New York, 1964. 349 pp. $4.95.—Through an oversight this important book by one of our Honorary Members has never been reviewed in the Fern Journal. The users of Dr. Wherry’s well-known “The Fern Guide” (1961), which covered the northeastern United States, will find this new work familiar, for it follows exactly the same format, and even the same drawings are used when the species concerned occurs both in the north- eastern states and in the south. The new illustrations have been drawn by the same artist, are very good, and will serve for the ready identification of all but the most critical species. In my opin- ion, one of the defects of the new work is that it does not cover all Oe On ene He Te REcENT FERN LITERATURE 35 of the southern states as is implied by the title. The ferns treated are those occurring south of the Fall Line, from North Carolina south to Florida and west to Texas. Dr. Wherry notes that the ferns of the uplands of North Carolina, Georgia, and Alabama are the same as the more northern ferns treated in his previous “The Fern Guide.” This is true, but nevertheless it would have been convenient to have a complete treatment of all the ferns of the southern states. It is true that Wherry mentions, and in some cases illustrates, these northern ferns in an ‘“‘Appendix,” but they are not included in the generic or specific keys, and are not even listed in the Index to the book. This will be a drawback, especially to the amateur. It is possible that this limitation was imposed by the Doubleday Company rather than being voluntary on Dr. Wherry’s part, because even as it is the book must be rather expensive to print, with all of its illustrations. At least some workers have been desirous of having a conserva- tive fern guide to replace that of Small, but they will be disap- pointed with this, for Wherry recognizes not only most of the segregate genera recognized by Small but some additional ones. Wherry unquestionably knows these southern ferns very well, but he has fallen into the trap of making his generic decisions on the basis of the Florida species only. It is easy enough to recognize segregate genera on the basis of the few Florida representatives of widespread tropical groups, but it is not so easy when these groups are considered from a worldwide viewpoint. It is an es- sentially provincial attitude. It might be mentioned incidentally that Dr. Wherry lost an opportunity in not segregating the two species he refers to Stenochlaena, S. kunzeana and S. tenuifolia, for Holttum has indicated that the first of these is referable to Lomariopsis, a much better founded segregate than some of those that Wherry does adopt. Commendably, the Polypodiaceae is kept in its traditional sense, but it is broken up into groups for which “family” names are suggested. Some of these appear to be highly artificial, such as Stenochlaena, Acrostichum, Trismeria, and Pityrogramma being grouped into the “Gymnogrammaceae.” A nomenclatural error 18 36 AMERICAN FERN JOURNAL the restriction of the “family” name Pteridaceae to Pteridium while the type of the family, Pteris, is referred to the ‘‘family” Sinopteridaceae. A taxonomic error is the inclusion of patens, augescens, normalis and some other more doubtful species like versicolor in Thelypteris, although in their characters (including chromosome numbers) these are surely referable to Cyclosorus, as Christensen indicated long ago; in fact at least some of them probably hybridize with Cyclosorus dentatus. Incidentally, Wherry recognizes quadrangularis as a species distinct from dentatus, for the first time, so far as I know, for any United States plants. I have tried to follow Alston’s suggested distinctions, but they always fail, with African as well as American material. One weakness is Wherry’s desire to provide “common” names for all the species, an idea that goes back to the first edition of Britton and Brown’s Illustrated Flora, where common names were in- vented for all the species, some of them manifestly atrocities. The late Paul C. Standley, when thinking of a local field trip, is reputed to have asked, “Shall we go out and collect Knieskern’s smooth-fruited beak-rush?”, referring jokingly to the name as- signed in Britton and Brown to Rhynchospora knieskernii. Bot- anists are now generally agreed that it is impossible to get the public to agree on an artificially coined “common” name, and such names have been omitted from the latest Britton and Brown and Gray’s Manual. Some of those coined by Wherry are almost unbelievable: “Free-tip star-hair fern,” ‘Toothed lattice-vein fern,” “Alabama streak-sorus fern,” “Grid-scale maiden fern, “Twin-spore-stalk fern,” or “Spready tri-vein fern.” I leave the reader to guess, if he can, which species receive these appellations. One other matter is of some technical importance. On page 346 ot the book are published seven new combinations, with the note: These combinations are also published in periodical literature.” Indeed, they were published in the American Fern Journal, vol. a4, pp. 144-146. But this particular number of the Journal came out November 12, 1964, whereas the Southern Fern Guide, which is dated merely 1964, was officially published on October 2, 1964- Thus, the new combinations date from it, not from the Journal. SEN Sh pebaais Sere AMERICAN FERN SOCIETY 37 Wherry spells some specific epithets ferrissi, engelmanni, chapmani, and so forth. He justifies this in the introduction by saying that he adopts the original spelling, not feeling competent to correct the Latin of earlier workers. This is a misapprehension, because no knowledge of Latin is required. The International Code of Botanical Nomenclature specifies that epithets that are genitives of names ending in consonants add “‘ii,” except after ‘“-er,” and further that names published originally with other endings are to be treated as orthographic errors and corrected, in these instances to ferrissii, engelmannii, and chapmanii. The amateur or general botanist does not know this and so it is up to a pro- fessional like Wherry to adopt the correct spelling. One peculiarity is the reference to the younger Linnaeus as “L. Jr.,” but I suppose that this is as good Latin as the usual “L. fil.” Wherry has included a good deal of additional information which will be of value, such as the meaning of specific names, basic chromosome numbers, references to other books on ferns, an article on the life cycle of ferns, notes on fern culture, and incidental notes on a number of cultivated species of ferns. Some of my remarks above may seem to indicate that I am very critical of this new work. I do wish that it were better in some respects, but nevertheless this is a fine book, the work of a dedi- cated botanist, and it is going to be the working guide for all students of the southern ferns for many years to come.—C.V.M. American Fern Society Report of the President for 1967 One of the highlights for members of the American Fern Society is the annual meeting in association with the American Institute of Biological Sciences in the summer, which provides an oppor- tunity for fern enthusiasts to get together in an informal manner, to discuss ferns from various points of view, and to observe them in their native habitats. The August field trip to the Edwards Plateau in Texas was exceptionally rewarding, and those who 38 AMERICAN FERN JOURNAL attended left with a much better understanding of the habits and habitats of our western ferns. Dr. and Mrs. Donovan 8. Correll, both authorities on the Texas Flora, arranged and led a magnificent trip. We were most comfortable and pleased with our headquarters at the friendly Lazy Hills Guest Ranch, where excellent food, swimming, and horseback riding were provided. We were especially thankful for their showing us, in such a limited time, a large number of Texas ferns in their varied habitats. We drove on a highway in a river bottom where we could see ferns on the wet banks, climbed a mountain with ferns along the rocky crevices, found Jsoétes in shallow pools on the mountaintop m close proximity with cactus (Opuntia), and on the very last trip were thrilled with the discovery of a colony of Ophioglossum. We were fortunate in having Dr. Frank W. Gould ay Oe representative at College Station, who arranged for a delightful luncheon and an air-conditioned program room with remote controlled projector. The planning committee which serves as liaison between the Society and the National Committee of the XIth International Botanical Congress is well under way with plans for field trips and * program on ferns. The adherent societies in AIBS are helping to finance the Congress and also to request that commerative postage stamps be issued for it. A second index is being planned for the American Fern Journal, starting with 1936. We wish to thank Barbara Joe Hoshizaki for acting as Judge of Elections and Mrs. Pearth who served as auditor. gly and efficiently for the Society, insures an excellent year for 1968. Respectfully submitted, Mivprep EF. Faust, President AMERICAN FERN SociETy 39 Report of the Secretary for 1967 Membership in the American Fern Society at the end of 1967 stands at 671, of which 83 are members from 29 countries abroad. In the United States, only the Dakotas, Nevada, and New Mexico are not represented. I report with regret the death of 7 members: Miss Edith Seam- man, a member since 1937 and secretary of the Society in 1950 and 1951; Miss Claudia Schmidt, also a member for 30 years; Mr. Thorleif Fliflet and Dr. C. M. Van de Water, both members for 20 years; Mr. Ralph S. Hudson, since 1961; Mr. George L. Kirk, since 1961; Mrs. Thomas 8. Christensen, since 1965. The annual meeting of the Society, held August 29 with the American Institute of Biological Sciences on the campus of the Texas A & M University, was preceded by a luncheon attended by 26 members and presided over by President Mildred E. Faust. There were also 26 at the afternoon session, with Dr. Faust presiding. The titles of the papers presented are: ‘‘Natural history of Equisetum in Costa Rica,” by Richard L. Hauke; ‘The game- tophytes of Taenitis blechnoides (Willd.) Swartz and Syngramma pinnata J. Sm.,” by Lenette R. Atkinson; “Asexual reproduction in two species of Hawaiian ferns,’ by Richard A. White; “Spore morphology as an aid to classification of Malaysian ferns,” by James E. Canright; “The fern foray in retrospect,” by Donovan S. Correll; “The present status of the Hart’s Tongue fern in central New York,” by Mildred E. Faust. Four of the five Council members were present at a meeting August 29. Dr. Warren H. Wagner, Jr., reported that his liaison committee between the American Fern Society and the XIth International Botanical Congress in Seattle in 1969 (Dr. Wagner, chairman, Dr. T. M. C. Taylor, and Dr. Arthur R. Kruckeberg) has made tentative plans for our society’s participation in the Congress. Dr. Wagner stressed the desirability for individual contributions toward financing the Congress, which has been noted in this JouRNAL 57: 183. 1967. Respectfully submitted, Lenerre R. Arxrnson, Secretary 40 AMERICAN FERN JOURNAL Report of the Treasurer for 1967 In 1967 receipts were just slightly over expenses; the large bal- ance of last year kept us well out of the red. Cash on hand at end of year was about $6.00 more than that for 1966. Receipts from sale of back numbers exceeded those of 1966 by $224.00. Royalties from Dr. Wherry’s books exceed those of 1966 by $52.00. Advance dues this year were $127.00, much less than in 1966, but advance subscriptions were $826.40, which is much more than that received in 1966 Income from advertisements in the JouRNAL was $4.00 less than in 1966, but voluntary plate and page charges, instituted this year, gave us $180.00 income. The spore exchange netted $51.49 beyond expenses. Receipts Cash on hand, peteEy 1, 1967 $2,829.79 Membership Due Renewals $812.60 Sustaining 64.00 New 392 .20 -Advance 127 .00 Arrears 28 .00 $1,423.80 Subscriptions Current $324 .90 Advance-1968 774.40 Advance-beyond 1968 52.00 Arrears 31.97 1,183.27 Sale of Back Numbers 1,437.49 Sale of Reprints 352 .07 Poe (Wherry’s Books) 438.49 os mae and Plate Charges 180.00 Advertising in Journal Contribution to Intern. Bot. Congress AMERICAN FERN SOCIETY Total Receipts Total Disbursements American Fern Journal Vol. 56, No. 4 Vol. 57, No: 1 926 .37 Volu-5e-No. 2 926 .79 Vol. 57, No. 3 947 .14 Reprints and postage Envelopes for mailing journal Printing stationery Treasurer’s expenses Secretary’s expenses Editor’s expenses Foray expenses Shipping and handling back numbers AIBS dues AAAS dues Purchase of back issues Book exhibit Miscellaneous Total disbursements Cash on hand, January 1, 1968 Statement, December 31, 1967 Assets Cash in Union National Bank Cash in Green Point Savings Bank Bissell Herbarium Fund Accounts Receivable Inventory, Journal ibrary Total $1, 120.65 $3,920.95 $2, 835.32 41 5,270.00 $8,099.79 $5,264.47 $2,835.32 $17,171.21 42 AMERICAN FERN JOURNAL Liabilities Advance Dues $ 127.00 Advance Subscriptions 826 .40 Fund Balances Bissell Herbarium Fund 977 .66 Life Membership Fund 1,529 .26 Una Weatherby Fund 4,132.01 General Fund 9,578.88 Total $17,171.21 Respectfully submitted, LeRoy K. Henry, Treasurer Report of the Auditing Committee I hereby certify that I have seen the books and accounts of Dr. LeRoy K. Henry, 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. Dororuy L. Peartu, Auditor Report of the Judge of Elections The results of balloting for officers of the American Fern Society are as follows: For President Irving W. Knobloch 316 Warren H. Wagner 2 For Vice-President Warren H. Wagner 313 Richard Hauke J. L. Edwards Clyde F. Reed Alice Tryon David B. Lellinger tet et ND AMERICAN FERN SOCIETY 43 For Treasurer LeRoy K. Henry 315 Walter 8. Phillips 1 I therefore declare the following candidates elected to office: Irving W. Knobloch, President; Warren H. Wagner, Vice-Presi- dent; LeRoy K. Henry, Treasurer. Respectfully submitted, BarsBara Joe Hosuizak1, Judge of Elections Report of the Fern Spore Exchange The Fern Spore Exchange is continuing to grow. Many new requests were made for the Fern Spore Exchange list. This list now reaches eleven National Parks or Botanic Gardens in as many different countries. The fern societies of England and Japan both contribute to the Exchange, as do other institutions and individuals throughout the world. A supplementary list will be issued in a very short time, which will include many species that are completely new to the Exchange. The current list should be retained, as eighty percent of the listings are either 1966 or 1967 collections. The two lists will contain about 380 species and varieties. The Fern Spore Exchange depends on the continuing support of all the members of the Society. Its success is due to the faithful members who consistently contribute fresh spores year after year. No matter how common a particular species is, it may be the particular species required for research in some other area of the world. All contributions to the Exchange are appreciated and sincere thanks are tendered to each contributor. Exchange lists will be supplemented in October of each year. A fresh stock of spores or fertile fronds should be collected when ripe and sent to the Exchange before that date in order to be included in the list. The Exchange is a continuing service, of course, and spores are received and disbursed throughout the year. Respectfully submitted, Nem D. Hatt, Director of the Spore Exchange 44 AMERICAN FERN JOURNAL Report of the 1967 Fern Foray The 1967 Fern Foray was held in central Texas in conjunction with the AIBS meetings at College Station. The Foray was organized and conducted by Dr. Donovan Correll. The group assembled at the Lazy Hills Guest Ranch, Ingram, Texas, on Thursday afternoon August 24, 1967. We were treated to a delicious Texas-style barbecue; then Dr. Correll took us on a botanical tour of Texas via his beautiful color slides. Karly the next morning we set out to see some of the flora for ourselves. The first day was spent on the limestone of the Edwards Plateau. The first fern we saw was the southern maidenhair, Adiantum capillus-veneris, growing profusely along the Guadalupe River. We proceeded to drive up the river bed to a parking area, where we hiked up a small side canyon. Several ferns grew on the canyon walls, and the orchid Epipactis gigantea was in fruit here also. On a ledge above the path some Selaginella wrightii was spotted. In trying to get some of it down with a stick, Dave Emory discovered that he was standing about three feet from a rattlesnake. After a rest stop at Leakey, we drove to Prade Ranch, at the source of the Rio Frio. After lunch we explored the area, finding several ferns along the river and among rocks above. In the river bed at this point are ruts in the limestone made by Conestoga Wagons. We proceeded to Garner State Park and climbed a rocky hillside. The climb was hot, but well worth it, as several new ferns were added to our list. We returned via a different route to the ranch in time for supper, after which several members showed slides and Dr. Correll led us in an enjoyable songfest. Saturday morning, fortified by a delicious cook-out breakfast of eggs, bacon, and pancakes—all you could eat, we set out for Some granite outcrops. The first stop was Enchanted Rock, a pink granite dome. In ledges on the mountain we saw several species of Cheilanthes, Pellaea, and Selaginella. At the top were pools containing Isoétes lithophila, growing like grass in a lawn. oon white rainlily, Zephyranthes drummondii, was abundant here also. AMERICAN FERN Society 45 After lunch we drove to another granite area, Balanced Rock. Here many of the same ferns as at Enchanted Rock grew to larger size. One of the highlights of the trip came when Mrs. Ixrauss discovered that she was standing in a patch of Ophioglossum engelmannii. This represented a new county (Burnet) and a new floristic region (Edwards Plateau) for this species in Texas. On Sunday we had time to explore the Lazy Hills Ranch (the more adventuresome on horseback) before leaving for College Station. Dr. and Mrs. Correll did a splendid job of planning an inter- esting trip for us. We saw 21 species of ferns, a good number for an arid region, and most of them were new to many of us. They also did a marvelous job of estimating how long it would take to cover the ground; we never felt rushed and had ample time to explore and collect. Directions printed in advance were most helpful. Dr. Correll also identified nearly any plant, fern or not, so we became well acquainted with the flora. Our thanks also to the folks at Lazy Hills Guest Ranch for providing us with very comfortable quarters, delicious food, and the facilities at their spacious ranch (that pool sure felt good after a hot day in the field!). In short it was a well organized and very enjoyable Fern Foray. . The 21 species seen were: Selaginella peruviana, S. riddellit, S. wrightii, Isoétes lithophila, Ophioglossum engelmannii, Anemia mexicana, Adiantum capillus-veneris, Cheilanthes alabamensis, C. eatonit, C. horridula, C. kaulfussii, C. lindheimeri, C. tomentosa, Notholaena candida var. copelandii, N. parvifolia, N. sinuata var. integerrima, Pellaea atropurpurea, P. ovata, P. wrightiana, Thelypteris normalis, and Woodsia obtusa. Participants in the foray were from ten states: Dr. and Mrs. Correll (Texas), Mr. Robert Aborn (N.J.), Mr. William Adams (Ohio), Mr. David Emory (Penna.), Dr. Mildred Faust (N.Y.), Dr. and Mrs. Norman Foster (Texas), Mrs. Alice Gobin (Calif.), Dr. Aaron Goldberg (D.C.), Dr. and Mrs. LeRoy Henry (Penna.), Dr. Donald Huttleston (Penna.), Mr. Ray Jordan (Texas), Dr. Irving Knobloch (Mich.), Mrs. Beatrice Krauss (Hawaii), Dr. 46 AMERICAN FERN JOURNAL James Montgomery (N.J.), Miss Eva Sobol (N.Y.), and Miss Ann Stewart (Mass.)—James D. Monrcomery, Biology Depart- ment, Upsala College, East Orange, New Jersey 07019. New Members Mrs. Arnold Abercrombie, Box 545, Atlantic Beach, Fla. 32005 Mr. Shripad N. Agashe, Botany Dept., Shivaji University, Kolhapur, India og Julian P. Boyd, Bear Tavern Road, R. D. 1, bier ere N. J. 085) rs. J. Hammond Brandt, Upper Falls, Md. 2115 Mr. Alvin R. nore: Biology Dept., State Univ. N. Y. at Albany, Albany, LT RE | Mr. Ralph E. a 5611 Maple St., Shawnee Mission, Kansas 66202 Mr. Fred L. Bussa, 6203 New Castle Dr. , Bellaire, Texas 7740 Mrs. John Carden, 4208 Corona St. Wie Fla. Sister pes D. Cardillo, gi Prof. of Biology, Ladycliff College, Highland Falls, N. Y. 1 Miss Mireya D. Correa, A 8. Army Tropic Test Center, P. O. Drawer 942, Zone Mrs. Sterling W. Edwards, Monument Road, Myersville, Md. 21773 Mr. William P. Elliott, 2 Gracewood Park, Cambridge, Mass. 02138 Mr. Raymond A. Haney, 2610 Jefferson St. , Bellevue, Nebr. 68005 Mr. agree W. Hart, Dept. of Botany, Uphark Hall, Box 108, Miami Uni- ersity, Oxford, Ohio 45056 Mrs. Walter J. Hesse, 4847 Allencrest Dr., Dallas, Texas 75234 ts. A. J. Mauney, Rt. 1, Box 70A, Biawiine Cig No. Car. 28016 Mr. Charles R. Minton, 55 Cambridge Heights, Novato, Calif. 94947 Mr. Robert Louis Muller, 1 Oregon St. , Torrington, Conn. 06790 Miss Anita M. Nicholson, 2938 Glenwood Dr. , Longview, Wash. 98632 Mrs. Harry C. Olsen, Jr., 2003-128th Ave. S.E. , Bellevue, Wash. 98004 Dr. seat ineeae, Room, A. O. Fox Hospital, Oneonta, N. Y- Mrs. G. Nettie Scorp, 84 Christopher St. , Lodi, N. J. 07644 Mr. William L. Scriven, rhe & Fruit Growers, Marriottsville Rd., iharstalisdown: Md. 2 133 Mr. Clifford W. Smith, Se. of Botany, University of Hawaii, Honolulu, Hawaii 96822 AMERICAN FERN SOCIETY 47 Mrs. Margaret Sprecher, 2621 N. Myers St., eo rea 91504 Mrs. Olive J. StClair, 428 Fairfield St., Lisonee: Pa. Miss Mary K. Story, 10371 Topeka Dr. , Northridge, i 91324 Miss Judith E. Troop, Biol. Sciences U-43, Univ. of Connecticut, Storrs, Conn. 0626 Mrs. 8. L. Whatley, 3525 Garden Ave., Oklahoma City, Okla. 73112 Mrs. W. H. Yarbrough, 8122 N. Fielding Ave., Tampa, Fla. 33604 Miss Helen M. Yeager, 3820 S.W. 91st Ave., Portland, Ore. 97225 Dr. & Mrs. Dennison Young, 3 Colonial Rd., White Plains, N. Y. 10605 Mr. John W. Zelley, 1014 Columbus Rd., Burlington, N. J. 08016 Changes of Address Mr. Richard A. Benedict, 1773 Monroe Ave., Seaford, N. Y. 11783 Dr. F. M. Breed, 4 Hadsell Court, funshumion, Ney. 4 Mr. William G. Carr, 2231 Derek Drive, Fullerton, Calif. 9263 Mr. Carl F. Chuey, pegs Dept., Youngstown State University, Youngs- town, Ohio 4 Mr. Doumhisn Cobb, ik Village, Conn. 06031 Mrs. Allan G. Davenport, Box 323, Jamestown, R. I. 02835 Mrs. W. Brooks Evert, 115 N. Lakeside Dr., Medford, N. J. 08055 Mr. F. Gordon Foster, P. O. Box 136, Sparta, N. J. 07871 Mrs. Ronald S. Gray, R.F.D. #2, Derry, N. H. 03038 Mrs. Winifred J. Harley, Bolton Road, Harvard, Mass. 01451 Dr. R. E. Holttum, 50 Gloucester Court, Kew Road, Richmond, Surrey, Mr. Paul C. Hutchison, oxi a Botanic Garden, Rt. 2, Box 5098, Escondido, Calif. 9 Miss Mary Howard, neyo Ranch, Willits, Calif. 95490 Mr. John P. Knabel II, 9117 Cherbourg Dr., Potomac, Md. 20854 Mr. Dwight M. Moore, 417 W. 8th St., El Dorado, Ark. 71730 Dr. Howard Markell Smith, 7 West Franklin St., Richmond, Va. 23220 Prof. James H. Soper, Natural banaes Branch, Nat'l. Museum of Canada, Ottawa 4, Ontario, Mr. H. Harwood Teaey, 2743 Hillerest St., LaVerne, Calif. 91750 Miss Jay Wallace, 103 Sweezey St., Patchogue, N. Y. 1 1772 48 AMERICAN FERN JOURNAL Statement of Ownership, Management and Circulation In accordance with the rules and regulations he the United States Post Office, as established under the Act of October 2 , 1962, Section 4396, Title 39, United States Code, the pimbeced aaa are published. Title: AMERICAN FERN Jour Frequency - Issue: tsisietaaly (Approximately March 31, June 30, tember 30, and December 31 Location at i of caer (Printers): 3110 Elm Avenue, more, Maryland 212 Location of reistuais Office of ‘shale (Not Printers): Dr. LeRoy Henry, Department of Botany, Carnegie Museum, Pitts- burgh, Pennsylvania 15213 Publisher: AMERICAN FERN Society, Inc. » Department of Botany, C e Editor: oe Bae cob B. Lellinger, Department Me Botany, Smith- an eager Washington, D. C. 20560 Managing “Editor : Non O aoe ae Society, Inc., Department of Botany, Carnegie Museum, Pittsburgh, Ponncdivanis 15: Bowdliaiaeie Mortelinsas and other Security Holders: None Average No. each Single issue issue during pre- nearest to ceding 12 months filing date Total mews sg copies printed: 1200 pt circulat . Sales esa = ete. 0 0 ; Mail subscription 971 982 Free distribution, inluding samples 2 2 Total distributio 973 984 Office use, iain: ete. 227 216 Total 1200 1200 The Statements made above are certified to be correct, and are signed by: ne B. Lellinger, editor-in- chief of the American Penn Journal on October 967. > Exotic and Hardy Ferns Begonias BOLDUC’S GREENHILL NURSERY 2131 Vallejo Street St. Helena, California 94574 Open Saturdays and Sundays from 10 A.M. to 4 P.M. and by appointment Phone 963-2998—Area Code 707 Mail orders accepted UNUSUAL AND RARE FERNS SHIPPED DIRECTLY TO YOU © List Available @ LEATHERMAN’S GARDENS 2637 N. Lee Avenue South El] Monte, Calif. 91733 Miesour! Boranice. JUL 15 1968 Vou. 58 APRIL-JUNE, 1968 No. 2 GARDEN LIBRARB American Fern Yournal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY ts DAVID B. LELLINGER 0. V. MORTON ROLLA M. TRYON IRA L. WIGGINS Porne and Vortilicer. 2256555 So vise eres BarBara Joe Hoswizaki 49 The Valid Varieties of Pellaea andromedaefolia....... Tuomas R. Pray 54° Petiolar Shoots in the saacagr oi and Related Ferns Jupira E. Troop anp Joun T. Micke 64 Marsilea maheshwarii, a New es from Pondichery, India Bris Gora, 70 Phloem Studies in the Pteridophytes, Part I. Equisetum. Suripap N. AGASHE 74 A New —— of Salviniaceae and a New Species of Azolla from the late Cretaceous Joun W. Hatt 77 1 be een ae eee eee See ee OS Se Be ere Oe ee Se a Notes: Notes on Woodwardia areolata and Other Rare Illinois Ferns; A Spiny Dennstaedtia from Central America; Polystichum munitum in South Dakota; Trichomanes petersii Revisited at Saratoga, eg Lygodium ss osc at Another Fern Escaped in re ene ee Me te ak Ee Se oe. eee © See oie ecb we ee ee Se 6 oe 88 8 2 Ee Oe ARES eee SRE & The American Fern Society Council for 1968 @ W. Kwosioon, a of Botany & Plant Pathology, bse State University, East Lansing, Michigan 48823. resident Warren H. Wacner, Jr., Department of Botany, University of Michignae Ann a , Michigan 48104. Vice-President TKINSON, 415 S. Pleasant Street, Amherst, Massachusetts 01002. y K. Henry, Division of Plants, Carnegie Museum, Pittsburgh, — picPs 152 Davip B. LELLINGER, Smithsonian Institution, Washington, D. C. 20560. Editor-in-Chief National Society Representatives Warren H. Wacner, Jz., Dept. of Botany, * satsoned of Michigan, Aun Arbor, Michigan 48104. Representative to A. A. A. 8. Waren H, Wagner, a+ is Dept. of Botany, University of Michigan, Ann Arbor, Michigan 48 Bepresentative to A. I. B. 8. American Fern Journal EDITORS Davin B. Letiinazer __Smithsonian Institution, Washington, D. 0. 20560. C. V. Morton______ Smithsonian Institution, Washington, D. ©. 20560. Rotta M, Trron - 02188. Hi besiege Harvard University, Cambridge, Mass. Tra > ro parca hse a erbarium, Stanford University, Stanford, Calif. An illustrated quarterly devoted to the general study o owned by the American Fern Society, and published at 3110 ag ive. "e., Baltimore, Ma. 2211. 8 1. Seeond-elass postage paid at Baltimore. The pages of the Jour- members who wish to arran e exchang eS 5 a membership list is published at st Shtervala, to assist those interested in cbtaining specimens Matter for maton should be addressed to Dr. David B. Lellinger, Smithsonian Insti ashington, D. C, 20560. Changes of nig applications for membership, subseriptions, orders Se ee and other business communications should be addressed magne tage $4.50, eae, Maanire of agency handling fees; sent free _ — bers of the Am ern Society mal dues, $4.00; bership, $8.00; life oer pec 20.00) Bxiracted rep sate ordered in advance, will be ed a rs at cost, plus Back a * of 64 pages OF ack volumes $5.00 to $6. 25. each; le back sae pa less, $1.25; 65-80 pages, over 80 $2.50 each; Cumulative Index to Volumes 1-25, 50 cents. ‘Ten percent discount on orders of sit ore. Librarp and Herbarium Dr. W. H. W: agner, J. Ty Departmen’ of Botany, University of Michigan, Ann Arbor, Michigan 48104, is nee and curator. Members may borrow books and specimens a os a tee, the Tabieaer paging ok geal or express Spore Exchange : D. Hall, 1295 N 98115, ~ all, ortheast 95th Biseet, f Seattle, Washington is the Spore Ex Viable are received and dit pelihed, ded Mala of tho caladiions aes oem en agentur Sa soiicteds ’ { American Fern Journal Vou. 58 APRIL-JUNE, 1968 No. 2 Ferns and Fertilizer BARBARA JOE HOSHIZAKI Horticultural knowledge concerning the fertilization require- ments of ferns is far greater than the literature indicates. Experi- enced fern growers have known for a long time that proper fertilization of ferns enhances their growth and beauty. Equally well known is the fact that ferns suffer readily from improper fertilization. Considering these two points, the complex reactions between plants and soil, and the reports of individual gardeners, the result is a confused potpourri of advice. Much of what is written by individual gardeners is solid, practical advice, but many times it is applicable only to a particular fertilizer or soil mix, pattern of watering, quality of the water, or other specialized condition. There is a need for trained personnel to ascertain through controlled studies more precisely the response of ferns to mineral needs. This information would be of great interest not only to the horticulturists, but also to ecologists and others who deal with ferns as experimental material. The diversity of fern habitats, which ranges from tree tops to rocks to heavy clay soils, opens up a whole field of possible investigations on soil and mineral studies. However, this paper is written with the needs of the amateur fern grower in mind, and concerns some of the basic practices and principles in providing minerals to ferns through fertilization. WHICH FERNS NEED FERTILIZATION?—All the cultivated ferns known to me benefit from proper fertilization by growing faster, larger, or both. Ferns do differ in their responses. As a whole, ferns require less fertilization than most cultivated seed plants. Volume 58, No. 1 of the JouRNAL, pp. 1-48, was issued March 26, 1968. 50 AMERICAN FERN JOURNAL WHAT HAPPENS IF FERNS ARE OVER-FERTILIZED?—Stunting, yellowing, and, more commonly, burning or browning of the foliage, wilting of the fronds due to root damage, and in severe cases death of the plant may occur. These reactions are primarily due to too much soluble salt from the fertilizer collecting on or near the plant tissue and indirectly damaging the tissue by dehydrating it. WHAT ARE THE CONSEQUENCES OF NO FERTILIZER?—If the soil is of good quality and other conditions are favorable, ferns may not need fertilizers, and growth is usually steady and adequate. Less favorable conditions may result in stunting, slow growth, poor quality foliage, and susceptibility to disease. WHAT KIND OF FERTILIZER SHOULD BE USED?—The amateur should use fertilizers that have a reputation for low burn. These fertilizers are available in forms to be applied as liquids or dry, Liquid types include fish emulsion (5-2-2), “Blue Whale” (6-2-1). “Orchid Spoonit” (30-10-10), “Hyponex” (7-619), and others. Forms applied as a powder or in granules include fish meal (9-3-0), cottonseed meal (6.4-1.5-1), “Milorganite” (6-3-0), hoof and horn (15-0-0), and others. The first of the three numbers listed after each fertilizer indicates the percentage of nitrogen (N), the second number the percentage of phosphorus (P), and the third the percentage of potassium (KX). The percentage varies from brand to brand, but is always given on the label. Fertilizers having all three of these important mineral elements are termed complete fertilizers, as opposed to incomplete fertilizers, which lack one or more elements. Complete fertilizers are generally preferred over incomplete ones, but results vary from grower to grower, depending on the soil and cultural practices. How SHOULD FERTILIZERS BE APPLIED?—Certainly the manu- facturer’s directions should be followed carefully. If you are une certain about a fern’s tolerance to fertilizer, there is no harm In reducing the fertilizer concentration to half. Fertilizers to be dis- solved in water should be thoroughly dissolved before application. Fertilizers applied as a powder or granules should be sprinkled thinly and evenly over the moist soil surface, and the plants FERNS AND FERTILIZER 51 watered thoroughly zmmediately after application. Any fertilizer inadvertently spilled on the foliage should also be washed off im- mediately. If too much fertilizer is applied in one spot, especially on or near the crown of the fern, remove as much of the fertilizer as possible and water thoroughly. Not only will this reduce the possibility of burning the fern but will also reduce conceatrations of mold growth. Fertilizers derived from organic substances may encourage microorganisms. Although such fertilizers must be de- composed by microorganisms before releasing most of their nutri- ents, very high concentrations of mold may be damaging to the fern crown and young fronds. Where warmth and high humidity occur, as in greenhouses, molds and bacteria may become particu- larly troublesome, especially with very young ferns. Under these conditions manures in particular should be avoided. Note that with ferns the surface of the soil should not be disturbed by tilling at any time. Fern roots are fine, fibrous, and close to the surface. Tilling the soil to mix in the fertilizer may easily damage the roots. This means that even with a thin even application and adequate watering, some of the fertilizer may remain on the soil surface. This should not be of consequence, as long as it is of the low burn or no burn type and is not concentrated in one spot. In time these particles will dissolve and reach the roots. Liquid applications have an advantage in that most of the fertilizer salts are dissolved before application and are promptly carried to the roots. However, any solution passing beyond the root zone is lost to the plant. Liquid fertilizers leave less residue in the soil than dry ones do. This is a disadvantage because more frequent fertilization is needed with liquid types. On the other hand, liquid fertilizer may be applied at the time of watering, which saves considerable labor. How FREQUENTLY SHOULD FERNS BE FERTILIZED?—Manu- facturers usually have their own recommendations. On the average, one application about every three weeks for liquid forms and at longer intervals for dry application is sufficient. Ferns which do not enter any noticeable rest period, but continue to grow actively the year around, benefit from fertilizers the year around. Ferns 52 AMERICAN FERN JOURNAL which are inclined toward a rest period or dormancy should not be fertilized during this time, as there is little intake of nutrients during periods of inactivity. When not fertilizing, less water is applied, which is beneficial because many tender species of ferns growing in marginal low temperatures are particularly sensitive to overwatering during periods of inactive growth. In very porous soils or under heavy and frequent watering leaching causes a great loss of fertilizer from the soil. On the other hand, soils which do not drain readily retain more fertilizer and need not be fer- tilized so frequently. WHAT CAN BE DONE FOR WILTED OR SCORCHED FRONDS?—This kind of damage is difficult to correct with young ferns and those lacking large fleshy rhizomes. As soon as there is any indication of wilting or burning, remove all fertilizer that might be on the sur- face, see that the plant is well drained, and then water copiously to leach any remaining fertilizer out of the root region. After this liberal watering, water only sparingly so that the soil is aerated and not waterlogged, which will help to reduce the growth of decay organisms on the damaged root hairs and smaller roots. Also increase the humidity of the air and remove badly burned or wilted foliage. Recovery of the plant usually depends on the re- maining undamaged roots or rhizomes which may send out new root branches and root hairs. Ferns with large fleshy rhizomes recover more easily because much food and water is stored in the rhizome. WHAT ELEMENTS DO FERNS SEEM TO NEED Most?—Nitrogen Is the element most needed by plants. It is usually taken into the plant in the form of nitrates, which are very soluble and leach readily from the soil. Plants which have abundant nitrogen re- spond by producing large, soft, dark green fronds, which though attractive to some gardeners, are not firm and may appear coarse in habit. They are also more easily damaged by wind or during watering. Fewer fertile fronds are also produced. Potassium deficiency is rare, but has been reported for maidenhair ferns (Adiantum). The fronds are said to turn an abnormal reddish- brown color. A specific treatment is to apply potassium sulfate at pO oe FERNS AND FERTILIZER 53 the rate of 1 teaspoon per 9 by 4 inch container. However, using a complete fertilizer should correct and prevent this problem. Trace elements and fern needs are not known precisely. Chelated iron compounds often added to fertilizers are reported to improve color and growth of the Holly fern (Cyrtomium falcatum). There are usually enough impurities in fertilizers to take care of the trace element needs of ferns. W4aT Is SALT INJURY?—In areas of low rainfall the water supply may contain salts which retard plant growth or cause an un- satisfactory soil structure. If these salts accumulate in the soil around roots, stunted growth, scorching of the fronds, and wilting may occur. A white crust of salt may be evident on the surface of the soil or flower pot. In many of the western states it is important to have soil mixes which drain well and thus aid in leaching out the undesirable salts. Thorough watering leaches the soil better than frequent, light watering. Fertilizers containing sodium (e.g., sodium nitrate) should not be used if salt injury is a problem. Additional information may be found in the chapter “Irrigation Water and Saline and Alkali Soils” in the Yearbook of Agricul- ture (1955), published by the U.S. Department of Agriculture. Because there are so many physiological, chemical, and physical factors entering into the relationship between growth and fertiliza- tion, it is impossible to cover every facet and discuss many of the exceptions. If proper fertilization does not bring about any im- provement in a declining plant, one certainly should look for other common causes of poor growth, such as under- or overwater- ing, poor drainage and aeration, low temperatures, low humidity, inadequate light, and salt accumulation. Lire Scrence DEPARTMENT, Los ANGELES CrTy COLLEGE, Los ANGELEs, CALIF. 54 AMERICAN FERN JOURNAL The Valid Varieties of Pellaea andromedaefolia Tuomas R. Pray! Since Pellaea andromedaefolia (Kaulf.) Fée was first described in 1824, various forms of this Californian species have received systematic recognition. In a monographic revision of section Pellaea, to which the species belongs, Tryon (1957, p. 179) regarded none of these variants as taxonomically significant. Certainly anyone familiar with this species in the field, where it grows under a variety of conditions, will recognize the plasticity of the species in response to its environment. Variety rubens D.C. Eaton and var. gracilis Summers ex Farlow undoubtedly are examples of simple, environmentally induced variants. Regarding the possible significance of other variant forms of P. andromedaefolia, some observations concerning a dwarfed specimen are in order. This specimen (Whitehead 5050) is quite fertile and is apparently a mature individual. Its leaves are fully tripinnate and it agrees with var. andromedaefolia in other respects. The leaves are only 16 cm high however, and the segments are very small, averaging only 4 mm long, compared to the usual 7-10 mm. In spite of its size, young sporophytes raised from its sexual gametophytes were all typical of var. andromedaefolia and showed no trace of the diminu- tive features of the parent plant. Thus, unusually small size, in this case at least, was environmentally induced and not genetically fixed. On the other hand, var. pubescens D.C. Eaton is decidedly distinct. The distinction between this variety and typical P. andromedaefolia is the subject of this paper. Most local floras have ignored the pubescent form of P. andro- medaefolia. The most recent, that of Munz (1959), mentioned it as occurring in the vicinity of San Diego. Tryon (1957, p. 182) recognized that the pubescent form was from the southern portion of the species’ range and stated that it was distinguished by having 32- — sporangia. The typical glabrous form was said to be he financial support of the National Science Foundation (Grant #GB- 1716) is gratefully acknowledged , t ‘s is VARIETIES OF PELLAEA ANDROMEDAEFOLIA His 64-spored. The implication was, therefore, that var. pubescens was an apogamous form of a sexual species, although at that time there was no direct evidence of apogamy. As the result of extensive field studies, transplant experiments, and the raising of numerous plants from spores, I have found that these statements represent a great oversimplification of the situation. Plate 7 illustrates the distribution of the species (see Table I for a full list of the collections cited in this study). Variety andro- medaefolia, with glabrous stipes and rachises and glaucous leaf segments, ranges from northern California—a report from southern Oregon is doubtful, as stated by Tryon (1957, p. 182)—southward through the state and along the west coast of Baja California Norte, Mexico, to the region of Bahia de Sebastian Vizcaino. In central California it extends inland to the foothills of the Sierra Nevada and in southern California it occurs along the western borders of the Colorado Desert. In the southern part of its range it occurs on some of the offshore islands. While var. andromedaefolia was well distributed at one time on Santa Catalina Island, only a single collection has been made in recent years (McNeil 76). Much searching in appropriate habitats has revealed that it is now extinct there, or nearly so. Similarly, three days of searching on Santa Cruz Island produced a single specimen of the typical variety on an inaccessible cliff where it could not be collected. Variety pubescens, with puberulent stipes and rachises and non-glaucous segments, occurs from San Luis Obispo south to Cedros Island. It appears to be a strictly coastal ecotype, a fact which has not been recognized previously. All the material I have seen is from coastal bluffs and hills and at or near the mouths of coastal canyons. The type of var. pubescens came from the San Luis Obispo area (Eaton, 1878). Its occurrence there has been re- cently confirmed (Hoover, 1966). I have not seen any material rom this most northern locality, but Hoover’s comments suggest that his material is the same as that which I have studied. The most northern collections available to me came from Southern Ventura County (Point Mugu). I know of no locality between this and San Luis Obispo, although suitable habitats in AMERICAN FERN JOURNAL VoutuME 58, Puate 7 A demonstrated apogamous populations of var. oO andromedaefolia — 64-spored collections of var. pubescens @ 32-spored collections of var. pubescens * collections of var. pubescens of unknown sporangial type Mar oF THE DistrisuTion or P. ANDROMEDAEFOLIA. SHADED INDI- CATES THE GENERAL DISTRIBUTION OF VAR. PUBESCENS AND VAR.ANDROMEDAE- FOLIA IN CALIFORNIA AND MEXICO. VARIETIES OF PELLAEA ANDROMEDAEFOLIA Bk the intervening regions have been searched carefully. Although the coastal area between Point Mugu and Los Angeles seems a likely habitat, I know of no collections of var. pubescens from there. It does occur to the south in San Diego County, where I have made several collections. On the offshore islands the presence of var. pubescens is definitely established for Anacapa, San Clemente, South Coronado and Cedros Islands. It may also have occurred on Santa Catalina, but several personal attempts to find extant material of this variety were unsuccessful. The floras of these islands have been decimated by overgrazing of sheep, or goats, or both to such an extent that it is no longer possible to find many species where they were once collected. For Mexico, I have seen material from Ensenada south to San Quintin. From the relatively small amount of material studied it would appear that the two varieties do not usually occur in the same locality. The typical form is rarely found in the exposed maritime environment to which var. pubescens is largely restricted. The only instance of sympatry observed was at Point Mugu, where there were a few, very small individuals (2028) of the typical variety in & population predominantly of the pubescent variety. D. C. Eaton (1878) distinguished var. pubescens only by the presence of the fine pubescence on the stipes and rachises and cited a specimen from San Luis Obispo (Mrs. R. W. Summers). Apparently neither he nor anyone since has noticed that, although the pubescence provides one obviously distinguishing feature (clearly visible only when magnified), several others are equally obvious. In life the pinnules of var. pubescens are succulent and are bright green and shiny on the upper surfaces. Upon drying the upper surface looks resinous. The segments appear to be thicker than those of the typical variety; sections of the pinnules did not confirm this, however. On the other hand, the pinnules of var. andromedaefolia are consistently at least slightly glaucous and dull. The development of anthocyanins in the leaves as a response to drought or extreme exposure is common to both forms. The vein pattern is clearly evident on the upper surfaces of the pinnules in fresh condition in var. pubescens, whereas this feature is obscure 58 AMERICAN FERN JOURNAL Tas_eE I. List or ConLections! Sporangial Coll. number Locality type VARIETY iy oar eee 1668 Santo Tomas, highway #1 at Km _ 64-spored 193, “Baja Calif. N orte, Mex 1669 te Juan Canyon, Orange. Co., 64-spored 1909 Andreas Canyon, Palm Springs, 32-spored Riverside Co., Cal. 2028 Point Mugu, Nests Co., Cal. 64-spored Dunkle 1881 Santa Catalina Isl. 64-spored Dunkle 2865a Santa Catalina Isl. -spored Garth 1089 Andreas Canyon, Palm Springs, 32-spored Riverside Co., Cal. McNeil 76 Holland’s Cove, Santa Catalina Immature J. H. Thomas 8223 (DS) eins Fi Landing, Baja Cal. 32-spored ort Tryon & Tryon 5556 (GH) Redway, Haxabolat Uo. , Cal. 32-spored Wiggins 9967 (DS) 40 mi. NE of El Heonris: Baja 64-spored Cal. Norte, Mex Whitehead 5050 no data 64-spored VARIETY PUBESCENS 1665, 1666 Ensenada, S on highway #1 at 64-spored Km 66, Baja Cal. Norte, Mex. 1667 as above at Km 64-spored 1906, 1907, 2027 Point Mugu, Ventura Co., Cal. 32-spored 2056, 2057 Del Mar, San Diego Co., Cal. —_ 32-spored 5215 grown from spores of 1665 = Cooper 2172 Ensenada, Baja Cal. Norte, Mex. ag Dunkle 2043 — Gibraltar, Santa Catalina 64-spored Dunkle 4262 Johnson’s Trough, Baja Calif. se Norte, Mex Dunkle Anseapa Isl. 64-spored Elmore 383, 413 San Clemente Isl. -spored Epling t Stewart (DS) — cans Baja Cal. Norte, 32-spored Ferris 8611 (DS) San Vicente, Baja Cal. Norte, Immature Moran 8301 South Coronado Mex. -spored Moran 1065 (DS) Cedros Island, Me Immature VARIETIES OF PELLAEA ANDROMEDAEFOLIA 59 Sporangial Coll. number Locality type UCBG 62.432-1 Cultivated. Original from San 32-spored Diego Co., Cal. (Hutchison 2160) Wiggins 11974 (DS) South Coronado Isl., Mex. Immature 1 All collections of var. pubescens seen are included. Only those of var. andromedaefolia_ especially pertinent are cited; many others have been ex- Specimens are deposited in the herbarium of the Allan Hancock Foundation (AHFH), University of Southern California, unless indicated otherwise in the typical variety. Unfortunately, this characteristic sometimes becomes less evident in drying. The degree of lamina dissection is most useful in mature plants. The leaves of the typical variety are usually tripinnate (rarely quadripinnate), even in rather depauperate specimens. Variety pubescens, in contrast, has bipin- nate leaves, even in the largest plants. The largest leaves may have a very few ternate pinnules in the basal pinnae. In the typical variety many of the pinnules commonly have 5-7 (occasionally up to 10) segments; pinnules even in the upper portion of the leaf are usually at least ternate. Correlated with the simpler leaves of var. pubescens is the increased size of the ultimate segments. This distinction is well demonstrated when the two varieties are culti- vated under similar circumstances, but segment size varies with environmental conditions, so herbarium material can not be reliably distinguished on this basis. The stature of the two varieties also distinguishes them. Under the best conditions the typical form may produce leaves as much as 80 em long. The largest specimens of var. pubescens studied had leaves only 35 em long, and most specimens were much smaller. No cultivated material has been observed to exceed this height. The stouter and more rigid stipes and rachises are correlated with the shorter stature. Among the distinctive features of var. pubescens, the non- glaucous segments and bipinnate leaves can be used to identify it, even if it is not pubescent. One collection from Santa Catalina Island (Dunkle 2043) has all of the characteristics of var. pubescens except pubescence. The collection consists of several old leaves 60 AMERICAN FERN JOURNAL only, and it is possible that this latter feature has been lost with age. An effort was made to recollect this fern from the Dunkle locality, but no plants of either variety were found. This specimen is the only evidence I found for var. pubescens on Santa Catalina. A collection from Anacapa Island (Dunkle 7663) also lacks the puberulence typical of var. pubescens, but is otherwise indis- tinguishable from material of this variety. Tentatively, therefore, it is listed as var. pubescens. Perhaps these two collections are hybrids of the two varieties. It should be noted that the insular collections of var. andromedaefolia do tend to be smaller plants with less complex leaves than is typical for the variety. Whether these represent local variants cannot be decided on the basis of the meager material available. Under cultivation the differences between these two varieties were maintained and were especially obvious when the plants were compared side by side. Both varieties were also grown to maturity from spores, and the progeny retained the distinguishing features of the parents. No intermediates have been discovered in the field. At Point Mugu, the only locality where the two varieties were observed growing together, the plants of var. pubescens were apogamous, whereas the plants of var. andro- medaefolia were 64-spored and presumably had sexual gameto- phytes. Therefore, the lack of intermediates between the varieties is understandable. Tryon (1957) reported that the pubescent form is 32-spored, but further study has shown that no such correlation between spore number and morphology exists. The material from Point Mugu (1906, 1907) is 32-spored and reproduces by apogamy (Pray, ms.): Material from San Diego County also is 32-spored and apogamous (2056, 2057, UCBG 62.432-1.) In contrast, the plants on the islands that are directly offshore from these localities apparently are sexual. Among the insular collections cited, some were too immature to determine their spore number and probable repro- ductive method. Only Elmore 383 and 413 from San Clemente and Dunkle 7663 from Anacapa had spores; these are all 64-spored. Collections from northern Baja California (1665, 1666, 1667) and VARIETIES OF PELLAEA ANDROMEDAEFOLIA 61 adjacent South Coronado Island (Moran 8301) are 64-spored and produce sexual gametophytes. The southernmost collection, from San Quintin (Epling & Stewart), is probably 32-spored, although a positive count could not be made. Tryon (1957) reported a Cedros Island collection to be 32-spored. The distribution pattern for apogamy in var. pubescens based on available material is puzzling (Plate 7). If the present pattern can be taken as a reflection of the history of this variety, then perhaps var. pubescens had its origin in northern Baja California from whence it spread to the Coronado and Channel Islands. On the mainland it may have moved both north and south through the formation of apogamous races which apparently reached the islands only in the south. Although most collections of var. andromedaefolia are 64-spored (and those investigated produced sexual gametophytes), 32-spored and apogamous individuals are scattered throughout the range of this variety: Humboldt County (Tryon & Tryon 5556), Sierra Nevada foothills near Yosemite, (Tryon, pers. comm.), Palm Springs (1909, Garth 1089), and Baja California (Thomas 8223, which is the most southerly record for the typical variety). All other determinable Mexican collections are 64-spored. All the insular material for which a spore count is available is 64-spored. Thus, in the typical variety apogamy is sporadic; it seems likely that each of the apogamous populations has had a separate origin. Differences among the gametophytes from each of those that has been investigated support this assumption (Pray, 1968). ere does not appear to be any correlation between morph- ology and spore number in either variety. The only method by which apogamous populations can be identified is by spore count. This contrasts with the closely related P. ovata, which also has both 64- and 32-spored forms. All of the material of P. ovata studied thus far has shown that the 64-spored plants are glabrous and produce sexual gametophytes, whereas the 32-spored plants all show varying degrees of pubescence. All of the latter type that have been tested have produced apogamous gametophytes. It is clear that the differences between var. andromedaefolia and var. pubescens, which are summarized in Table IT, are suffi- 62 AMERICAN FERN JOURNAL ciently marked to merit the systematic recognition of var. pubes- cens. In my opinion, the two varieties are comparable to good subspecies in other groups. To a very remarkable degree the situation in P. andromedaefolia parallels a similar situation in another Californian species, P. mucronata (D. C. Eaton) D. C. Eaton. The general distribution pattern of the two species is very similar (Tryon, 1957), and in fact they are often in the same TaBLE II. CompaRISON OF THE Two VARIETIES OF P, ANDROMEDAEFOLIA Character Variety andromedaefolia Range cismontane & insular Calif.; western Baja Calif. Norte, Mex. Variely pubescens coastal & insular So. Calif.; j to Cedros slan Sporangial type 64-spored; sporadic 32- 64-spored Channel Isls. and spored populations northern Baja Calif. Norte; 32-spored, southern Calif. Norte, Mex., Cedros Isl. and southern Calif. Height up to 80 cm less than 35 em Leaf form tripinnate, many pinnules bipinnate, a ternate with 5-7 segments innules; ultimate segments Stipe & rachis Leaf color Venation glabrous, slender dull, glaucous evident on lower surfaces promine 0 on upper only surfaces, especially in living ate localities; in these P. mucronata will be found in drier, more exposed habitats. Pellaea mucronata is represented throughout most of its range by the common form, var. mucronata, character- ized by finely divided, tripinnate (rarely quadripinnate) leaves. In the higher mountains of central and southern California the typical variety is replaced by a montane ecotype, var. californica (Lemmon) Munz & Johnston, which differs in its smaller stature and in its bipinnate and more compact leaves. Thus, both of these fairly widespread species appear to have evolved specialized and reduced ecotypes to fit more extreme environments, in one case the maritime environment, in the other the montane. VARIETIES OF PELLAEA ANDROMEDAEFOLIA 63 The parallel may be extended even further, for each of these species also has a counterpart species in the southwestern region of the interior of the continent. Here Pellaea longimucronata replaces P. mucronata. Their relationship is sufficiently close to permit hybridization at points of contact (Pray, 1967). In the same region P. intermedia replaces P. andromedaefolia. In this case there is a considerable gap between the ranges of the two species. In spite of the remarkable similarity between P. intermedia and P. andromedaefolia var. pubescens it seems unlikely that there is a direct relationship because of the specialized ecology of the latter. More probably, these two are parallel developments and, if closely related, it is through var. andromedaefolia. Nevertheless, if the geographic origin of herbarium specimens of var. pubescens were not known, these would be easily confused with P. intermedia, which is also characterized by bipinnate leaves with puberulent stipes and rachises. Probably the simplest method by which they may be distinguished, as indicated in Tryon’s key (1957), is the very clear vein pattern evident on the underside of the segments of P. andromedaefolia in contrast to the unveined appearance of P. intermedia. When compared closely, the rhizome scales are also distinctive, although they are of the same general type. Those of P. andromedaefolia var. pubescens are about twice as broad at the base and appear lighter when the rhizome is viewed with a hand lens. The dark central band extends about 14 to 24 the length of the scale, whereas in P. intermedia the central band ex- tends to the scale tip. The clear margins of the scale seem a bit broader in var. pubescens, but the difference in this respect is not obvious. LITERATURE CITED Eaton, D. C. 1878. Ferns of North America. Vol. 1. Salem, Mas Hloover, R. F. 1966. An annotated list of pteridophytes of aii figie Obispo County, California. Amer. Fern J. 56: 17-26. Munz, P. A. 1959. A California Flora. Univ. 7 California Press, Berkeley and Los Angeles. Pray, T. R. 1967. Notes on the distribution of American cheilanthoid ferns. Amer. Fern J. 57: 52-58 64 AMERICAN FERN JOURNAL Pray, T. R. 1968. Interpopulation variation in the gametophytes of Pellaea andromedaefolia (Kaulf.) Fée. (manuscript). Tryon, A. F. 1957. A revision of the fern genus Pellaea section Pellaea. Ann. Mo. Bot. Gard. 44: 125-193. : . Observations on the leaves of Pellaea andromedaefolia. Contr. Gray Herb. 187: 61-68. DEPARTMENT OF BroLogicaL Sciences, UNIVERSITY OF SOUTH- ERN Cauirornia, Los ANGELES, Cair. 90007. Petiolar Shoots in the Dennstaedtioid and Related Ferns! Jupita E. Troor anp Jonn T. MIcKEL Stelar branching patterns and bud formation in the Filicales have been investigated little since the beginning of this century. Fern stems may branch by dichotomy, axillary buds, or buds not clearly in the axils of leaves. Buds from frond bases were reported and described by Gwynne-Vaughan (1903) for a number of ferns. In a monographic work on Dennstaedtia punctilobula (Michx.) Moore, Conard (1908) described in detail the anatomy of shoots from the petiole bases of this fern. Webster (1958) observed dor- mant buds on the leaf bases in Pteridium aquilinum var. latiuscu- lum, but none were seen to elongate. Further references to budding of ferns are scattered, and the phenomenon is mentioned only briefly, as in Wardlaw (1952) and Wagner (1963), for Matteuccia, Onoclea and Dryopteris. None of these workers has attributed any phylogenetic or taxonomic importance to the phenomenon, and, in view of the meager information available, it is appropriate that more comprehensive studies be undertaken. In the summer of 1967 we participated in a course in the biology of tropical pteridophytes offered by the Organization for udy was done while the senior author held a National Delewt Act Fellowship. Partial cost of publication of this paper v= ee by the Smithsonian Institution while the junior author was a National ie search Council Visiting Research Associate in the Department of Botany © the Smithsonian Institution. PETIOLAR SHOOTS 65 Tropical Studies in Costa Rica (Mickel, 1967), in which we had the opportunity to investigate a great diversity of fern genera and species in regard to their branching patterns. This study is based upon free-hand sections of rhizomes and petiole bases examined under a dissecting microscope. The results show that the phe- nomenon of petiolar branching occurs more commonly in the ferns than was thought previously (Table [). This branching is very widespread in the dennstaedtioid ferns, being a nearly constant feature in Dennstaedtia, Hypolepis, Paesia, and Histi- opteris, less constant in Pteris, and absent in Saccoloma. Also in the Cyatheaceae petiole branches are always present in Lophosoria and Cyathea mexicana, and occasionally so in Dicksonia and Metaxya. One species of Dicksonia studied had up to six buds on the lower stipe. Metaxya is reported (Bower, 1928) to bud from the petiole base, but our specimen did not. In most of the ferns studied, the vascular strand to the frond begins as a bulging of the rhizome stele (Figs. 1 and 3), followed by its breaking away from the stele and passing out through the cortex. In all the ferns studied the frond trace retains a basic horseshoe shape with the additions of hooks, corrugations, and dissections in various species. The gap left in the rhizome stele by the departing frond is then closed. The rhizomes of some Species possess a dicyclic (or rarely tricyclic) solenostele. In these plants the closure of the frond gap may be accomplished by a portion of the inner stele filling the outer. The whole inner sole- nostele may migrate to the gap, leave a filling portion, and center again as it closes its own gap (Fig. 3); or the section which will fill the outer stele may depart from the inner stele and migrate to the gap (Fig. 2). If there is a third solenostele, it repeats the pattern of the second. The bud develops well up on the stipe, at least after the leaf trace has become well differentiated from the rhizome stele (Fig. 1). (One exception to this was in Cyathea where the bud develops with the petiole trace but appears superficially to be on the stem to one side of the leaf base.) The adaxial hook of the petiolar strand lengthens laterally and separates from the AMERICAN FERN JOURNAL VouuME 58, PLATE 8 reSGtin® = (al Fig. 3 Fig. 4 Fig. 1. ExpLopep DIAGRAM OF HYPOLEPIS REPENS STELAR SYSTEM. Fic. 2. SACCOLOMA INAEQUALE STELAR SYSTEM. Fig. 3. EXPLODED DIAGRA! DENNSTAEDTIA DISSECTA VASCULAR SYSTEM. Fic. 4. EXPLODED pee OF PTERIS ALTISSIMA VASCULAR SYSTEM. The abbreviations are: ft TRACE, iS = INNER RHIZOME STELE, Irt = LATERAL RHIZOME TRACE, 1S OUTER RHIZOME STELE, S = RHIZOME STELE, and t = ABAXIAL PETIOLAB BRANCH, PETIOLAR SHOOTS. 67 frond stele, gradually closing into a new solenostele. The frond stele reforms the adaxial hook. In the dieyelic condition the inner stele of the lateral shoot may repeat the ontogeny of that of the main rhizome. This was found in Pteris and some species of Dennstaedtia. In other dicyclic species, however, the inner stele forms by an invagination of the outer stele, which separates and moves into the center closing into a solenostele (Fig. 3). There are but few generic differences from the above general pattern. The species of Hypolepis investigated are fairly uniform in their anatomy with usually dorsiventral rhizomes, unicyclic solenosteles and petiolar branches. There are usually two to four petiolar buds, often well-developed and growing as lateral branches bearing fronds. Histiopteris incisa is scarcely distinguishable from Hypolepis in its stelar anatomy; the two are closely allied All the Dennstaedtias examined bore lateral shoots on the petioles (usually one per petiole), whereas only two species (D. bipinnata and D. cicutaria) displayed dichotomizing rhizomes. Dennstaedtia bipinnata is the only unicyclic species studied. Dennstaedtia cicutaria exhibits a most peculiar pattern. The rhizome appears to terminate in a trichotomy consisting of a central petiole and two rhizome branches. Since this type of branching pattern is found in no other species examined and only irregularly in D. cicutaria, it seems likely that the rhizome has branched dichotomously while a petiole was produced simul- taneously. Some ferns form buds in positions other than at the adaxial petiole trace hooks. The fronds of both Pteris podophylla and Pteris altissima produce buds from the abaxial side of the petiole strand (Fig. 4). This position also has been found in three species of Costa Rican Grammitis (L. E. Bishop, pers. comm.). The bud in Dicksonia develops from the lateral vascular are, and in Lophosoria bud formation occurs on the lateral side of the abaxial strand. The bud trace of Paesia arises from both adaxial and abaxial portions of the petiole trace, one section from the adaxial hook and two sections from the abaxial corrugated are. These three strands unite to form the lateral branch. The ability of the petiole 6S AMERICAN FERN JOURNAL trace to form a rhizome stele is not limited to a specific portion of the petiole. The petiolar branching discussed above resembles that of certain coenopterid ferns of the Carboniferous Age. Botryopteris and Anachoropteris are well-known examples in which branches often arise from the petioles. Surange (1952) interprets the anatomy of Botryopteris as a dorsiventral stem giving rise to a terete stem upon which petioles and roots are borne. Delevoryas and Morgan (1954) discuss the phenomenon of stems arising from Tas_e I. Raizome CHaRacTERISTICS IN DENNSTAEDTIOID AND R ERNS ELATED F Ae te ee Pe lar Petiolar Name cycles Dichotomies branches Blotiella lindeniana (Hook.) Tryon (3432)! 1 .u Culcita contifolia (Hook.) Maxon (3238) 1 = os Cyathea mexicana Schlecht. & Cham 1 = 34 Dennstaedtia arborescens (Will ay Ekm ex Maxon (1870, 2037, 2045, 2212, 2226. 2305, 2526, 2535, 2589, 3111, 3305) 2 ney nH D. bipinnata ae Mise (2697, 3346, 3505, 3593, 3616) 1 + vr D. eae Se ists Moore (1869, 2303, 2914, 3118) 2 + * yb 2 pete (Swartz) aoe (2209, 2210, 2217, 2224, 2301, 2 2 - + D. ue (Willd. : AS (1871, 2302, + 304, 2 2 = Pe a phe (2730) 2 = + Dicksonia — Karst. (3134) 1 is = Histiopteris incisa (Thunb.) J. Smith (2182, 2483, 3193) 1 - + Hypolepis bogotensis nae (2169, 3248) 1 = + H. nigrescens Hook. (34 1 a + H. nuda Mett. (3300) 1 = of H. pulcherrima Underw. & Maxon (3306) 1 = a H. repens (L.) Presl (1972, 2000, 2306, 2743, 3048 1 - 2: H. rigescens (Kunze) Fée (2457) 4 — ag H. viscosa (Karst.) Mett. (3318) 1 og a PETIOLAR SHOOTS 69 Stelar- Name cycles Dichotomies branches Lonchitis hirsuta L. (2293, 2590, 3557) 1 — Be Lophosoria quadripinnata (Gmel.) C.Chr. (2431 i = Metaxya rostrata (H.B.K.) Presl (2853) 1 — ieee anfractuosa (Christ) C. Chr. (3000) 1 - um aquilinum var. arachnoidewm rae ) Herter (2017, : 2 + a. var. caudatum (L.) Sadeb. (2395) 2 3% Pteris altissima Poir. 2 P. podophylla Swartz (3177) 2-3 om? P. pungens Willd. (3563 lbt++++ 44+ Retz. Saccoloma elegans K. eA1982, 87995 S. ee (Kunze) Pi 1992, 2756, 3128, 3566) 2 a re he numbers Saf = Mickel collection numbers. Voucher specimens are deposited at US, ISC, LP, and USJ. Those without vouchers cited were common species not collected at the time of the stu y- petioles, particularly in Anachoropteris clavata. They believe that Surange’s “dorsiventral stem” is actually a petiole with stems arising from it as in Anachoropteris. Anachoropteris clavata is postulated as a scrambling plant with some petioles functioning as stolons producing new stems. Other coenopterid ferns are also known to have branching petioles (T. Phillips, pers. comm.). Among the modern dennstaedtioid ferns several have a habit similar to the above. In Hypolepis, for example, the petioles are not entirely erect from the rhizome, but creep for a distance before becoming erect. This horizontal portion bears the branch shoots and closely approximates the condition described by Delevoryas and Morgan. Thus, the patterns of branching and bud formation in these ferns are perhaps more significant than previously believed, for questions are raised regarding the relation of the frond to the rhizome, the basie ontogeny of the frond, and the phylogeny of the dennstaedtioid ferns. Further work on these problems is currently in progress. 70 AMERICAN FERN JOURNAL LITERATURE CITED Bower, F. O. 1928. The Ferns (Filicales), III. Cambridge Univ. Press Engla Conarp, H. W. 1908. The structure and life history of the hay-scented fern. Carnegie Inst. | Publ. 94. Deevoryas, T., and J. Morcan. 1954. A further ao ai eas of the mor: hology cot tai acs clavata. Amer. J. Bot : 192-203. GwynneE-Vauacuan, D. T. 1903. Observations on the ne of solenostelic ferns. Part II. Ann. Bot. 17: 689-742. MICKEL, 4 ie. 1967. An advanced course in pteridophyte biology in Costa a. Amer. Fern. J. 57: 145-161. SURANGE, K. R. 1952. The morphology of Botryopteris antiqua with some observations on Botryopteris ramosa. The Palaeobotanist 1: 2420-434. Waaner, W. H., Jr. 1963. Pteridophytes of the Mountain Lake area, Giles County, Virginia, including notes from Whitetop Mountain. Castanea 28: 113-150. Warpb.aw, C. W. 1952. Phylogeny ze Morphogenesis. MacMillan, London. Wesster, B. D., and T. A. STeEves. 1958. Morphogenesis in Pteridium aquilinum “L. ) Kuhn-general Siathobes and growth habit. Phyto- morphology 8: 30-41. DEPARTMENT OF SYSTEMATIC AND ENVIRONMENTAL BIOLOGICAL Sciences, University oF CoNNEcTICUT, STorRS, CONN. 06268, AND DEPARTMENT OF BoTANy AND PLANT PaTHoLocy, LowA State University, Ames, Iowa 50010. Marsilea maheshwarii a New Species from Poedichery: India Brig GopaL The genus Marsilea is represented in India by nine species,’ most of which are restricted in distribution, M. minuta L. being the only species found throughout India. Endemism is of common occurrence throughout the genus. While examining collections of Marsilea from all over the country, I found that some collections ~ 4 Gupta, K. - New Delhi, mae: 1962. Marsilea. Botanical Monograph no. 2, C. 8. 1. MARSILEA MAHESHWARII 71 from Pondichery represented a new species, which is here named in honor of the late Professor P. Maheshwari. MarsILEA maheshwarii Gopal, sp. nov. Plantae aquaticae; rhizomata robusta, apice pubescentia; etioli longi; lamina lata, marginibus integris; sporocarpia aggregata 3 vel 4, interdum 2, petiolo adnata 2-5 mm a basi; pedicelli connati et ramosi, penitus adnati sporocarpio, eique naquilodcs vel eo paulo longiores; sh rpia 3.5-4.0 X 3.0-3.5 m, lateraliter compressa, rotundata in fro ree i ter pe vel deorsum flexa; cornua bina, distincta, ‘oboritts uidem minutum et obtusum, superius vero longum et recurvum; mail i 12-16; m mega- sporae nullae vel aberrantes; microsporae aberrantes. Type: Pondichery, India, from the paddy fields, G. Thani- kaimoni 992a (Herbarium of the French Institute, 'Pondichery, Panaty PES: loc. cit., G. Thanikaimoni 992b (US), 992c (KX), 992d (BAN). Marsilea is well known for its morphological plasticity. For example, it is very difficult to distinguish between species growing submersed. However, the characters of the sporocarp are more stable and have largely been used in specific identifications. Marsilea maheshwarii is recognized mainly on the basis of sporo- carp characters. Gupta (loc. cit.) recognized three groups in the genus which differ in the mode of attachment of the pedicel to the petiole. The Quadrifolia group is characterized by pedicels that are adnate to the petiole at one point, and until now has been represented in India only by M. quadrifolia, known from Kashmir alone. Marsilea maheshwarii from Pondichery has a similar attachment, which excludes any possibility of its being one of the Minuta group (like M. minuta or M. coromandelica Burm. f.), in which the sporocarps are always basal. The Pondichery material differs widely enough from M. quadrifolia to warrant its recog- nition as a new species. Among the various characters, the mode of attachment of pedicel to pedicel, pedicel to sporocarp, and the shape of sporo- carps in the two species are quite different (Figs. 1, 2). In addition, the shape of the horns, the aberrant microspores, and the usual absence of megaspores (aberrant if present) also contrast with AMERICAN FERN JOURNAL 72 jeuio0u 02-91 iepluts ysouye “Z 1: £0} dn soumjouros ‘T 2 Z Zuipusdose JVYMOUIOS 10 paaArnoep wu 9-G ‘peqqtt jou ‘poursivur Apart ‘posserduiooqgns ‘[eAo asuq ye Aju souITJOWOS ‘YFUI] Aley} S{ “BO B}BUTOD ayeupe Arey yos Aypared §g 10 % jUBLIagGe 10 JU8S -qe solodsefeur “yueiieqe sarodsosorul 91-<él Ayung eu 4B Uuayoig seuTjeWOS ‘peaAInoel puB Ruoy, soddn ‘yunjq pue [ews Jamoy “Z ayeupe ATNy Ayunyeur ye ayeiqeys ‘Aarey AyTesuep Lat OT et Surpueose Ajanbryqo ww F-C"g ‘peqqi you “pours -1mu jou ‘passerduioo ‘pedeys-uveq Bue] wey} &{—8; ayeuU0D eseq ejorjed eaoqe wu ¢-z ‘ayvupe Z SouTJaWIOS “F 10 E sjuajuoo dagoo10dg Los JO JaquinN susoy jo ainjeu puB JequUInN dawoos0ds 0} Jaotped jo UOTYBTOYT gonyjins davoo10dg ored diwooi0ds : aotpad uorjtsodstp Ja01pog daeoos0ds jo yySuey, puv edvyg Jaotped 03 Jasrped jo uon¥eyay ajorjed 03 Jaotped jo uonNeyey Jaquinu dagso.10dg myofiaponb * Wy MADNYSAYDUL * SLIQIDLDY,D vITOdIUavod "Jy INV ITMVMHSAHVNA “J AO SdUVOONOdS AO SAMOLV AT TVOIDOTOHAUOJ ‘T ATV T, MARSILEA MAHESHWARII te M. quadrifolia. Vegetative characters were compared in the two species and also in M. minuta grown under exactly identical condi- tions (both submersed and dry). Marsilea maheshwarii is a stouter species than the other two. The leaflet margins remain almost entire under all conditions of moisture in M. maheshwarii, but abit Fic. 1. Portion oF A RHIZOME OF MARSILEA MAHESHWARII SHOWING DIS- POSITION OF SPOROCARPS. Fig. 2. SAME, M. QUADRIFOLIA. they become crenate in the other species under dry conditions. The various characters of the two species are listed in Table I. I am very much indebted to Mr. G. Thanikaimoni of the French Institute, Pondichery, India, for his valuable help in collecting the material for my study, and to Rev. Fr. H. Santapau, Director, Botanical Survey of India, Calcutta, for kindly providing the Latin translation of the description. Thanks are also due Prof. R. Misra and Dr. K. C. Misra of the Botany Department, Ba- naras Hindu University, for encouragement and facilities accorded me. Department oF Botany, BaNarAS Hinpu UNIVERSITY, VaRANASI—5, InpIA. 74 AMERICAN FERN JOURNAL Phloem Studies in the Pteridophytes, Part I. Equisetum. SuHripap N. AGASHE Xylary tissues of the pteridophytes have been studied in detail by several workers, but phloem tissue has received very. little attention. The probable reason for this is the difficulties en- countered in studying the anatomical structure of the phloem, for example in cutting perfect radial longitudinal and cross sections of sieve elements showing the details of sieve areas and in staining the tissues. Little is known about the phloem anatomy of Equisetum. I have studied phloem in the rhizomes and aerial shoots of E. hyemale, E. telmateia, E. arvense, and E. giganteum. This paper concerns only the latter two species. The purpose of this investi- gation was to gather data on Equisetum phloem that could be compared with presumably comparable tissue described in some American calamites (Agashe, 1964). Johnson (1933), in discussing the origin and development of certain tissues in E. scirpoides, mentioned the occurrence of sieve plates with small pores on the lateral walls of the sieve cells. Golub “and Wetmore (1948) described the development of phloem in E. arvense. According to them, protophloem sieve cells may be 3 mm long or more. Their longitudinal walls have many, transversely elliptical, faintly staining sieve areas; such areas are also found on the oblique end walls, but they do not seem to form distinet sieve plates. Metaphloem sieve cells are longer, larger in diameter, and have densely granular cytoplasm and very thick walls; sieve areas are more common on the end wall in metaphloem than in proto- phloem. A There appears to be some confusion in the literature regarding the definitions of sieve cells and sieve tubes. The presently accepted definitions, according to Esau, Cheadle, and Gifford (1953), whe quote Cheadle and Whitford, state that a sieve tube consists of sieve elements joined end-to-end to form a vertical tube-like structure in which sieve areas are more highly specialized on the AMERICAN FERN JOURNAL VOLUME 58, PLATE 9 oe ee i is eres et A ‘. bal ~ : é Stem Paioem in Equisetum. Fic. 1. TRANSVERSE SECTION OF E. GIGAN- TEUM SHOWING ONE VASCULAR BUNDLE, X 160. Fic. 2. RADIAL LONGITUDINAL SECTION OF A SIEVE TUBE OF E. GIGANTEUM SHOWING PORTION OF A SIEVE PLATE ON AN OBLIQUE END WALL, X 910. Fic. 3. RADIAL LONGITUDINAL SINGLE SIEVE TUBE WITH A SIEVE PLATE ON AN END WALL, X 915. 76 AMERICAN FERN JOURNAL end walls than on the lateral ones. Sieve cells are not joined verti- cally to each other and have sieve areas on both lateral and end walls which are of similar specialization. In order to secure perfect radial longitudinal sections of phloem tissue, stem material was split lengthwise and a small piece containing only two or three vascular bundles was selected. This material was soaked in hydrofluoric acid to soften it before em- bedding it in paraffin. Harris’ Haematoxylin was the most suitable stain for sieve elements. As shown in the cross section of a portion of an aerial shoot of E. giganteum (Fig. 1), the phloem tissue consists of several cells of two types, and is flanked on either side by radial xylem strands. Just below the phloem is a large protoxylary canal. The whole vascular bundle including the protoxylary canal is surrounded by an individual endodermis marked by a Casparian strip on the lateral walls. Inside the endodermal layer is a single-celled layer of pericycle. The phloem tissue is composed of large sieve tubes and small parenchymatous cells. As shown in the radial longitudinal section of a sieve tube (Fig. 2), the end wall is oblique. However, in some sieve tubes the end wall is transverse. Most of the sieve tubes are 30 u in diameter and 430 long. Sieve areas are scattered over the radial walls and include one to three pores. The pores in the oblique end wall sieve plate are 4-5 in diameter, whereas most of the pores in the radial walls are 2 u in diameter. The most clearly defined sieve plates and sieve areas were observed in the end walls (Fig. 4) and radial walls (Fig. 3) of the sieve tubes of EF. arvense, features which have not been illustrated clearly in previous accounts. The sieve areas on the radial walls consist of many pores. In some sieve plates on end walls cyto- plasmic strands in the pores can be seen. There seems to be some deposition of callus on the sieve plates. It can be concluded that the phloem in these species of Hqutse- tum is fairly advanced among the pteridophytes because of the presence of sieve tubes and not sieve cells. I am grateful to Prof. Henry N. Andrews for his advice and encouragement and to Mr. S. K. Sutar for his help in preparing the photographs. PHLOEM Srupies: EquisETuM a LireRATURE CiTED Acasug, 8. N. 1964. The extra-xylary tissues in a calamites from the merican eis on, Phytomorphology 1 1} Esau, K., V. I. Caeapue, and E. Girrorp, Jr. 1953. Osunperiice structure nad Sieuible Saects of specialization of the phloem. Amer. J. Bot. 49: 9-19 Gouus, 8. th and R. H. Wermore. 1948. Studies of development in the getative shoot of Equisetum arvense, II. The mature shoot. Bot. 35: 767-781. Jounson, M. A. 1933. Origin Apis of tissues in Equisetum scirpoides. Bot. Gaz. 94: 494, Botany DEPARTMENT, SutvaAsi University, KouHapur, INDIA. A New Genus of Salviniaceae and a New Species of Azolla from the late Cretaceous! Joon W. Hat The Salviniaceae are represented in the Cretaceous by two species of Azolla recently described from North America. Azolla cretacea Stanley is based on massulae; it has been reported from the Hell Creek Formation of South Dakota (Stanley, 1965), Montana (Norton and Hall, 1967) and the time-equivalent Edmon- ton Formation of Alberta (Srivastava, 1966). Azolla geneseana Hills and Weiner is known both as massulae and megaspores from the Edmonton Formation in Alberta (Hills and Weiner, 1965). These few reports are scarcely indicative of the place of the family in the Cretaceous, at least in the fluviatile deposits of eastern Montana that I have looked at. By using the simple sieving and sorting techniques customarily used for isolating mega- spores from sediments, large numbers of massulae and megaspores of Azolla have been found. They are so abundant and hetero- geneous as to suggest that the family was not only a conspicuous component of the aquatic vegetation, but was morphologically ‘Funds from the University of Minnesota Graduate School and NSF Grant GB-4090 helped support this publication AMERICAN FERN JOURNAL VotuME 58, Puate 10 Fic. 1. HoLorypr or Azouia crrcinata, X 325. Fies. 2 AND 3, MASSULAE FROM THE PARATYPE OF AZOLLOPSIS TOMENTOSA, 175. CRETACEOUS SALVINIACEAE 79 more diverse then than it has been subsequently. One new genus, with both megaspores and massulae, and a new species of Azolla representing what I consider to be a new section of the genus are described here. Among the extant species of Azolla, as well as the previously described fossils, there are two types of massulae. Section Azolla accomodates those species whose massulae have anchor-shaped glochidia; in sect. Rhizosperma the massulae lack glochidia. Section Azolla dates from the Cretaceous and sect. Rhizosperma from the Oligocene. The massulae of the species of Azolla described here possess glochidia that are filamentous and coiled. AZOLLA sect. Filifera J. W. Hall, sec and vegetative remains unknown. Type: Azolla circinata Oltz & Hall. Section Filifera includes massulae with glochidia which tend to resemble the perisporal hairs on the body of many Azolla megaspores, both living and fossil. Although megaspores with coiled perisporal hairs or with hairs having enlarged tips have not been reported, they do occur in some of the Cretaceous de- posits which I have examined; furthermore, there are massulae with straight, unbranched glochidia in some of these deposits. All of these await further, detailed study before their relation- ships and taxonomy can be clarified. My present opinion is that this section is primitive in Azolla because glochidia and megaspore ornamentation are only slightly, if at all, different. AzOLa circinata Oltz & Hall, sp. nov. or Massulae nearly circular to oval in outline, ca. 100-175 » in longest dimension, 75-150 » in shortest dimension, the surface often marked with wide, rugulo-reticulate thickenings, these aspect. Glochidia numerous, 1-2 u wide, up to ca. 15 y long, usually not extended greatly beyond the margin of the massula proper, variously coiled at the apex, some appearing merely in- flated at the tip, usually with 1 gyre, or sometimes with only a AMERICAN FERN JOURNAL VOLUME 58, PLATE 11 AZOLLOPSIS COCCOIDES CRETACEOUS SALVINIACEAE S1 partial gyre. Microspores small, often vps eae! obscured by the plasmodium of the massula, 20-24 » in diam., probably a and thin-walled. Megaspores and vegetative remains unknown. Type: Slide HH 16, coordinates 38.6 X 105.2, reference 48.0 pe Po fnaleobominien! collections, MIN), from the Hell Creek Formation (Maestrichtian) near the center of $16, T25N, R43E, McCone Country, Montana. Fig. 1. Azollopsis J. W. Hall, gen. nov. Megaspore apparatus with a conspicuous, tomentose perispore and a Hopes, trilete endospore. Numerous float-like structures imbedded in or attached to the hairs of the perispore. Massulae with few Sens baled and conspicuous multi-hooked or barbed glochidia. Vegetative remains unknown. Type: Azollopsis coccoides J. W. Hall. The long, tortuous but otherwise unstructured hairs of the peri- spore are like those on the megaspores of species of Azolla, but the mass of them is greater than in that genus. The floats are relatively small, very numerous, and scattered throughout the hairs of the perispore or attached to them at the surface of the spore. The number, size, and position of the floats serves to distinguish Azollopsis from Azolla, as do the barbed glochidia of the massulae. AZOLLOpsis coccoides J. W. Hall, sp. no Megaspore apparatus spherical to onl as in the genus, 560- 990 u (aver. 706 u) overall?; megaspore ‘walls finely foveolate, 6-9 u thick. Floats very numerous, 40-90 u (aver. 72 ») in diam., eirenlar | in outline, disk-shaped, sculptured as in A. tomentosa. : Massulae attached to the megaspore, Ca. 120 u in diam., with only 2 or 3 spores each, these circular, 44-50 (aver. 47 ») in diam., with laesurae extending to the equator, the surface ? Intact endospores could not be measured because all are obscured by the tomentose perispore ® The exact limits were mostly obscured because es massulae were deeply imbedded in cpa hairs and merged with the floa Fig. 4. HOLoTyPE IN REFLECTED LIGHT, X 90. Fi. 5. SAME, IN TRANSMITTED LIGHT, X 90. Fic. 6. GLOCHIDIA FROM A MASSULA OF THE PARATYPE, X 400. Fic. 7. MicRospoRE IN MASSULA FROM THE PARATYPE, X 400. AMERICAN FERN JOURNAL Fig. VoLuME 58, PLATE 12 Aspijial oO ® a ® ° c Pe Bearpaw Judith River 8. DRAWING OF THE HOLOTYPE OF AZOLLOPsISs coccoIDEs (cF. Fics. 4 ‘i 5 ot . ‘ AND 9), X 90. Fig. 9. GLocnipia oF AZOLLOPSIS TOMENTOSA (cr. FIG. 12), _ 450. Fig. 10. Giocuipia or A. COccoIpDEs (cF. Fig. 6), K900. Fic. il, IAGRAM OF STRATIGRAPHIC COLUMN IN EASTERN MONTANA SHOWING RELA- TIVE POSITION OF FORMATIONS IN WHICH AZOLLA CIRCINATA AND AZOLLOPSIS SPP, OCCUR CRETACEOUS SALVINIACEAE 83 probably psilate and the wall 1-2 » thick. Glochidia numerous, barbed, distinctly septate, 2-5 uw wide, with 6-15 barbs arranged in 2 rows, each glochidial ‘‘cell’’ with 2 opposite barbs. Type: Slide 223-3 (paleobotanical collections, MIN), from the Judith River Formation, Upper Cretaceous (Campanian), at the west end of the Fort Peck Reservoir at the “Robinson Bridge,” Philips County, Montana. Figs. 4, 5, 8, and 10. PARATYPE: Slide 223-1, coordinates 134.8 X 51, reference 131.8 X 72.3 (paleobotanical collections, MIN). Locality as for the holo- type. Figs. 6, 7, and 1 The holotype possesses both megaspores and massulae, but the glochidia of the latter are seen to better advantage in the paratype. Except for the slightly larger dimensions, which may not be significant because of the smaller number of specimens seen, the megaspores of this species are virtually identical to those of Azollopsis tomentosa; the differences between the two species are in the massulae. The only massulae seen were those attached to the megaspores, and every megaspore had several massulae attached. The apparent effectiveness of their attachment, to the extent that they are surrounded by floats and perisporal hairs, contrasts sharply with their lack of attachment in A. tomentosa. Compared with that species, A. coccoides has fewer spores per massula, laesurae extending to the spore equator, narrower, distinctly septate glochidia with numerous barbs, and smaller massulae. arranged granules and rods, sometimes vaguely reticulate and with the vacuolate aspect found in the floats of species of Azolla. f loats occurring only at the periphery of the perispore and extending ‘ The latter shape is due to the greater mass of floats and perisporal hairs at one end of the spore; it cannot be determined whether this is the distal or Proximal end because laesurae are not often seen. AMERICAN FERN JOURNAL VoLuME 58, PuatTe 13 HoLotyeE OF AZOLLOPSIS TOMENTOSA CRETACEOUS SALVINIACEAB 85 beyond its margin or completely surrounded by the tomentose airs. Massulae variable in size and shape, mostly oval to subcircular, occasionally elliptical or reniform, 146 & 215 ah to 245 X 630 p, mostly between 200 X 2504 and 200 x 300 u. Surface of the massulae with numerous, septate or non-septate glochidia ca. 30-90 » long, 6-10 » wide, each with 4-6 retrorse barbs, one of these terminal. Microspores usually 4-8 (sometimes 12) per massula, 25-50 u in diam., probably psilate, with short laesurae. Type: Slide 208-11, coordinates 59.6 X 139.1, reference 67.8 X 132.8 (paleobotanical collections, MIN), from the Colgate member, Fox Hills Sandstone, Upper Cretaceous (Maestrichtian), in $16, T13N, R55E, Dawson County, Montana. Figs. 12 and 13. ParatyPEs: Slide 208-1, coordinates 38.0 X 124.5, reference 12.1 X 119.5 (Fig. 2) and coordinates 35.8 X 125.4, reference 12.1 X 119.5 (Fig. 3) (both paleobotanical collections, MIN). Locality as for the holotype. The massulae of this species are extremely large; some of them approach the size of megaspores. None has been found attached to the megaspores, but this may be due to their being dislodged during preparation. The glochidia are stouter in this species than in A. coccoides, are less conspicuously septate, and have fewer arbs. DISCUSSION The term “float” is applied to the spherical objects in the perispore of the megaspore apparatus of Azollopsis in deference to its use in Azolla, although in that genus, as has been pointed out by many authors, the term is a misnomer, for the megaspores sink shortly after being shed. Whether this was also the case in Azollopsis will never be known, but, at least superficially, the so-called floats seem to increase the surface area. They are not hollow, and any supportive function would have come from their humerous vacuolar cavities. Those floats that cling to the peri- phery of the megaspore resemble small, glochidia-less massulae, and it was my first impression that this was actually what they Fig. 12. MEGASPORE IN REFLECTED LIGHT, X 100, Fie. 13. SAME, IN TRANS- MITTED LIGHT, 100. * AMERICAN FERN JOURNAL VoLuME 58, PLate 14 AZOLLOPSIS TOMENTOSA CRETACEOUS SALVINIACEAE 87 were. However, there are several reasons why they cannot be massulae, the most important being the lack of microspores; another is their presence deep among the perisporal hairs, where it is unlikely that a massula would be able to penetrate; and a third, of course, is the occurrence of massulae with barbed glochidia actually attached to the megaspore in A. coccoides. If one can attribute to Azollopsis the kind of ontogenetic development. of the megaspore and its swimming apparatus of Azolla, then the position of the floats, their large number, and their structure may be explained readily In the development of the megasporangium of Azolla there are 32 spores, separated by a periplasmodium derived from the tape- tum of the sporangium, and all but one spore aborts (Eames, 1936, p. 250). The periplasmodium and abortive spores fill the upper part of the sporangium, above the developing megaspore; three vacuoles appear in the plasmodial mass, the degenerating nuclei are distributed into each vacuolar region, and a float de- velops from each. The developing megaspore also lies in a vacuole filled with a cytoplasmic fluid that becomes granulate. This ecyto- plasm develops into a foamy meshwork, and a perispore is formed, having a structure similar to that of the floats. Perisporal hairs then develop over the surface of the perispore; these seem homologous with glochidia. A float is the homologue of a massula. Only a slight departure from this pattern would produce the numerous floats of Azollopsis. If, instead of aggregating apically, the aborting spores remained scattered around the developing megaspore, and if each aborting spore developed in association with a small periplasmodial mass, the situation in Azollopsis would arise. Furthermore, if the development of floats occurred just prior to the formation of perisporal hairs (as in Azolla), many of the floats would then be surrounded by hairs. This presumed developmental pattern is somewhat like that for the development of the massulae, with microspores, in Azolla. Since massulae and floats are homologous in that genus, they probably are in Azollopsis G. 14. SEPTATE AND NON-SEPTATE GLOCHIDIA (CF. Figs. 1 AND 2), X 1045, Fa. es MIcROSPORES IN MASSULA, X 1045. SS AMERICAN FERN JOURNAL as well; the resemblance of floats to glochidia-less massulae is not difficult to imagine. The Judith River Formation, in Montana, is stratigraphically below any horizon from which remains of the Salviniaceae have been recovered until now. It lies in the Campanian stage of the European sequence; the Colgate Member, Fox Hills Sandstone is younger, and is in the Maestrichtian stage. Fig. 11 shows the stratigraphic relationships of the formations involved. The fluvial Judith River sediments were covered by a transgressing Bearpaw sea; when this regressed, the Fox Hills Sandstone was deposited; the Colgate Member is its uppermost, most shoreward phase. The Hell Creek Formation is again fluvial. Azollopsis must have occupied a non-marine refugium during Bearpaw and Fox Hills times, and then moved eastward, as the evolved species A. tomentosa, by Colgate times. The record of other pollen and spore types is not yet well enough known to substantiate this suggestion, but the small mammals apparently do. There are generic, but not specific, similarities in small mammals in the Judith River and Hell Creek formations (A. Sahni and R. E. Sloan, pers. comm.). Additional examples demonstrating species evolu- tion in the floras of these two formations can thus be expected. LITERATURE CITED Eames, A. J. 1936. Morphology of Vascular Plants. Lower Groups. IX + 4383 pp. McGraw-Hill, N.Y. Hitis, L. and N. Werner. 1965. Azolla geneseana n. sp. and revision of Azolla primaeva. Micropaleontology 11(2): 255-261. Norton, N. J. and J. W. Hay. 1967. Guide Sporomorphae in the Late Cretaceous—Early Tertiary of Eastern Montana (U.S.A.). Review of Palaeobotany and Palynology. 2: 99-110. Srivastava, S. K. 1966. Upper Cretaceous microflora (Maestrichtian) from Scollard, Alberta, Canada. Pollen et Spores 8(3): 497-552. : STanuey, E. A. 1965. Upper Cretaceous and Paleocene plant microfossils and Paleocene Dinoflagellates and Hystrichospherids from north- western South Dakota. Bull. Amer. Paleontol. 49(222): 175-384. DEPARTMENT or Botany, University or Minnesota, MINNE- APOLIS, Minn. 55455. SHORTER NOTES S9 Shorter Notes Notes oN WoopDWARDIA AREOLATA AND OTHER RaRE ILLINOIS Frerns.—On January 7, 1967 I discovered Woodwardia areolata (L.) Moore in a marshy seep spring nine miles northeast of Padu- eah, Kentucky, in southern Pope County (Schwegman 1089). Since this collection was of dead material, I returned to collect some live specimens on June 17, 1967 (Schwegman 1250). Both specimens are in the herbarium of Southern Illinois University (SIU). This station is about 40 miles north of stations for the Net- veined Chain Fern in Calloway County, Kentucky, and about 80 miles northeast of the stations in southeastern Missouri. The seep spring in which the specimens were found is a common habitat in the area of sandy, Cretaceous hills in southern Illinois. I have visited about 40 springs, but have found no other stands of Woodwardia. The Woodwardia dominates about a third of the spring, which measures about 40 feet long and 25 feet wide. The spring is in a dry oak-hickory woods, but Acer rubrum grows at its edge. Boehmeria cylindrica dominates another part of the spring, and Impatiens biflora, Polygonum punctatum, Carex incomperta, C. crinita, C. laevivaginata, and C. lurida are also present. The only other fern at this spring is Athyrium filix-femina. Ferns typical of adjacent springs include Athyrium filix-femina, Onoclea sen- sibilis, Osmunda regalis, O. cinnamomea, and Thelypteris palustris. Ophioglossum vulgatum was seen at one spring. Thelypteris noveboracensis (L.) Nieuwl. is rare in Illinois. The only previous specimen from southern Illinois was collected in Jackson County in 1880. I collected the New York Fern in damp, sandy soil of a small seep spring about 2.5 miles northwest of the Woodwardia station and 3.5 miles southwest of Bay City, Pope County, on June 27, 1967 (Schwegman 1268, SIU). The spring 1s on a north-facing wooded slope; the habitat is only slightly more moist than average for this hillside. The fern is in a circular stand about six feet in diameter. It is surrounded by Athyrium filix- femina, which it superficially resembles. 90 AMERICAN FERN JOURNAL Dryopteris goldiana (Hook.) Gray is seldom collected in Illinois, and apparently is very rare in the southern half of the state; the only previous collection is from Jackson County. I discovered clumps of this fern growing with D. marginalis in rocky woods at the base of a north-facing hill along Lusk Creek, SW14, 521, T1285, R6E, Pope County, on August 18, 1967 (Schwegman 1367, SIU). I found Ophioglossum engelmannii Prantl, a fern of exposed lime- stone habitats, growing rather abundantly in limestone hill prairie on Millstone Knob, which is near Robbs, Pope County, on June 4, 1966 (Schwegman 454, SIU). I want to thank Professor Robert H. Mohlenbrock of Southern Illinois University, whose encouragement and interest in my collecting helped make this paper possible-—JoHN SCHWEGMAN, Mermet Wildlife Refuge, Belknap, Ill. 62908. A Sprny DENNSTAEDTIA FROM CENTRAL AMERICA.—During a ’” course in tropical pteridophytes for the Organization for Tropical — Studies last summer in Costa Rica, I found a new species of Dennstaedtia which is remarkable in having spiny stipes, rachises, and pinna axes, a condition found in no other New World species of the genus. DENNSTAEDTIA spinosa Mickel, sp. nov. Frondes fere 2 m longae, axibus spinulosis, spinulis usque ad 3 mm longis rectis vel paullo antrorsis; laminae quadripinnati- pinnatifidae; pinnae alternae, supra nitentes et fere glabrae, subtus hirsutae, pilis rectis hyalinis; venae apice tenues; indusia humilia lata plus minusve bivalvata, integra vel irregulariter dentata; sporae triletae, rugosae, rugis brevibus latis irregulariter coalescentibus. Rhizomes creeping, clothed with black hairs; fronds nearly 2m long; axes orange, spiny, the spines up to 3 mm long, straight or slightly antrorse; stipes ca. 1 m long, 1 em thick; blades approx: 1 m long, 1 m broad, deltoid, quadripinnate-pinnatifid, lacy; pinnae alternate, short-petiolulate (to 12 mm); basal pinnules opposite, not or only slightly reduced, the larger 10-13 em long, 2.5-3 em broad; laminae hirsute beneath with straight, hyaline SHORTER NOTES 91 hairs, shiny above and essentially glabrous; vein endings slender; sori small, ca. 0.6 mm broad; indusia entire to irregularly den- tate, low, broader than high, more or less bivalvate; spores trilete, with short, broad, irregularly coalescing ridges, 27.0-32.5 » in diameter. “Type: Costa Rica, Province of Puntarenas, Osa Peninsula, southwest of Rincon de Osa, in rain forest behind Tropical Science Center field station, ca. 50 m alt., Mickel 2730 (US; isotypes ISC, LP, USJ Paratype: Honduras, in forest on ridge above Lancetilla, Yuncker 4592 (US). This new species closely resembles D. cicutaria sensu lato but can be distinguished readily by the spiny axes and finer dissection. Dennstaedtia spinosa has distinctive internal anatomy and spore morphology as well. The petiole trace of D. cicutaria is extremely corrugated on the abaxial are and has much higher lateral bulges. The stipe bundles of D. spinosa, on the other hand, are smooth and uncorrugated on the abaxial side, and the lateral bulges are lower. The lateral grooves are so deep as to touch one another in the center of the stipe. In their rhizomes both species have di- cyclic solenosteles, but those of D. cicutaria are dorsiventral with the ventral side slightly corrugated and those of D. spinosa are terete and uncorrugated. The spores also differ; those of D. cicutaria are verrucate, whereas those of D. spinosa have low, irregularly anastomosing ridges. Furthermore, the indusia of the new species are more regularly dentate than are those of D. cicutaria. Although D. cicutaria occasionally has a few prickles on its axes and a few specimens, described as D. decomposita, are as dissected as D. spinosa, none has the regular and consistent spininess nor the anatomical and spore characteristics of D. spinosa. I have seen no Old World specimens of Dennstaedtia that closely approach the new species, and it seems clear that it belongs to the D. cicutaria complex. Joun T. Micxet, Department of Botany and Plant Pathology, Iowa State University, Ames, Iowa 50010. 92 AMERICAN FERN JOURNAL PoLYsTICHUM MUNITUM IN SouTH Daxora.—While collecting for the University of Kansas I discovered a small patch of Poly- stichum munitum (KKaulf.) Presl on a moist, grassy, shaded stream bank in the Black Hills of South Dakota three miles southwest of Rochford in Pennington County. This is the first record of the fern in South Dakota and extends the present range some 550 miles to the east from near Missoula, Montana. Elsewhere in North America it is known from Alaska to Montana and southward to California and Idaho. The numerous teeth of the long, narrow, sword-shaped pinnae are bristle tipped. This and the linear- lanceolate leaf distinguish the fern from other western species. The identification of the specimen, Ralph Brooks 427 (KANU), was verified by Dr. Ronald L. McGregor.—RautpxH Brooks, 5611 Maple St., Mission, Kansas 66202. TRICHOMANES PETERSII RevisITrED AT SaratoGa, MIssIs- stpp1.—During the summer of 1965, while teaching at the Univer- sity of Southern Mississippi, I learned from Mr. R. N. Dean of the existence of a colony of the rare T’richomanes petersii Gray at Saratoga, Simpson County, Mississippi, between Jackson and Hattiesburg. At that time Mr. Dean was a student at Valdosta State College. He, in turn, had been told about the colony by @ local high school teacher. In September 1965 I used Mr. Dean’s map to locate the colony of a few plants growing on a very damp outcrop of ferruginous sandstone. I collected a very small specimen which is deposited in the University of Mississippi Herbarium. Recently, I found that E. T. Wherry! had visited the site in 1936. Almost 30 years later I could still follow Wherry’s description of the area in order to locate the colony. The site, however, had been timbered and burned, apparently reducing the size of the colony from what it was in 1936—Samuet B. Jonus, Jr., Unt- versity of Georgia, Athens, Ga. 30601. ' Wherry, E. T. 1936. Trich i ee ee Fern J. 20: 141-142. richomanes petersii at Saratoga, Mississipp SHORTER NOTES 93 LYGODIUM MICROPHYLLUM, ANOTHER FERN ESCAPED IN FLormpaA.—Two species of climbing ferns (Lygodium) have been reported from Florida. Lygodium palmatum (Bernh.) Swartz is a native of the eastern United States that has been recorded from Florida by several authors, including Chapman,! Small,? Correll, and Spurr.‘ For each report, however, some doubt has arisen, and its status in our flora has remained unsatisfactory. The other species, Lygodium japonicum (Thunb.) Swartz, is all too evident in many areas. It is an introduction from eastern Asia that has spread aggressively into moist woods and fields in most parts of Florida and adjacent states. It is largely lacking in southern Florida however. The University of Florida Herbarium has a March, 1932, collection from Dade County by the late Mary Diddell, “escaped on vacant lots, Miami,” but it has not persisted on any scale, if at all, in that area. More recent collections and field observations show the southern limits to be near Tampa Bay and in Highlands County. The above range data take on a special interest since the University of Florida Herbarium has recently acquired several collections of an escaped Lygodium from Martin County, in southeastern Florida: “(On mainland opposite Jupiter Island” and “Cabbage Palm Hammock on mainland, opposite Jupiter Island.” (both L. D. Ober, 15 June 1965) and “Jonathan Dickinson State Park. No fruiting plants.” (Carl W. Campbell, 12 Nov. 1966). These represent an apparent recent introduction that is established and spreading in the area, Lygodium microphyllum (Cav.) R Brown. This species is found throughout the Old World tropics from Africa to Australia, Asia, and Melanesia. It has long passed under the name Lygodium scandens (L.) Swartz, but Holttum?® ‘Chapman, A. W. 1897. Flora of the Southern py States, p. 63 - * Small, J. Kk. Flora of the Sere tga United States, 2nd Ed., Be 1913. ; * Correll, D. S. 1938. . net Check-List of Florida Ferns and Fern “s3 Amer. ha: a 28: Ree = the Distribution 0 Habits of the Ferns of OS ee Podinasar Florida. Proc. Fla. Acad. Sci. fig . ae _ ttum, R. E. 1959. Flora Malesiana, Series ir Vol. 1, part 1, pp. 44-41, 94 AMERICAN FERN JOURNAL has recently rejected this name. It is worth noting that Holttum refers to this plant as “‘sometimes a weed,” and indicates various disturbed habitats for it. A clue to the origin of these escaped plants is given by a collec- tion “in nursery, Delray Beach, Palm Beach Co.” (R. A. Long, 11 Feb. 1958). This is immediately south of the Martin County area. No doubt, Palm Beach County would yield colonies of the species if carefully searched. Since Holttum gives detailed descriptions and illustrations of Lygodium japonicum and L. microphyllum, there is little need to discuss their differences at length here. Lygodium japonicum has palmate to pinnately deeply lobed sterile leaflets which are not articulate at the bases. Lygodium microphyllum has articulate sterile leaflets that are unlobed or but infrequently lobed. The leaflets bear a superficial resemblance to those of Nephrolepis cordifolia. : It will be very interesting to watch the future spread and behavior of Lygodium microphyllum in Florida. It has presumably undergone several colder than average winters, so it should be able to maintain itself in this climate. Assuming it spreads out to overlap the range of L. japonicum to the northwest, we will be able to witness an experiment in competition between two closely allied and ecologically similar species.—JOHN BErcKNER, Depart- ment of Botany, University of F lorida, Gainesville, Florida 32601. Notes and News lgeRN Foray RESERVATIONS for September 1 and 2 should be made through Dr. Clara Frederick, Urbana College, Urbana, Ohio 43078. You may also write directly to the Holiday Inn Motel, Chillicothe, Ohio 45601. In either case, please write to Frederick, stating what reservations you have made or wish to make and whether you need transportation or can provide it for other participants. Rates at the motel are $9.00 for single rooms, $11.00 for double-bed rooms, and $14.00 for twin-bed rooms. ~~ . + aj . Society LuNcHEON RESERVATIONS should be made with Dr. RECENT FERN LITERATURE 95 Jane Decker, Department of Botany, Ohio State University, Yolumbus, Ohio 43210. The luncheon will be at noon on Tuesday, September 3. lERN WALK IN Wasuinaton, D. C.—Dr. James Johnston will lead a walk in Rock Creek Park at 2:00 pm on Sunday, July 14, 1968, starting at the foot trail at 16th Street N.W. opposite Whittier Street (not Whittier Place), which is two blocks south of Aspen Street, the southern boundary of Walter Reed Hospital. The walk is one of a series of guided nature walks on various topics held under the auspices of the National Park Service. A ranger naturalist from the Rock Creek Park Nature Center will also accompany the group. Details of seasonal growth and develop- ment of certain fern species will be featured.—D. B. L PoLYPODIUM DECUMANUM NEEDED.—I wish to obtain living ma- terial, either rhizomes or spores, of the large epiphytic fern of the Caribbean region Polypodium decumanum, sometimes known incorrectly as Polypodium leucatomos. Does any member know where it can be obtained?—EnGar T. Wuerry, Leidy Laboratory, University of Pennsylvania, Philadelphia, Pa. 19104 FERNS AVAILABLE.—I have a number of live ferns for exchange or as a gift. If the latter, please send postage. A list of species is available—Dr. I. KNosiocu, Department of Botany and Plant Pathology, Michigan State University, Kast Lansing, Mich. 48823. Recent Fern Literature An ANNOTATED BiBLioGRAPHY OF Mexican l'erns, by George Neville Jones. xxxiii + 297 pp. Univ. of Illinois Press, Urbana, 1966. $5.00.—In this useful volume Professor Jones has amply fulfilled his aim of providing “a useful bibliography of the principal literature pertaining to Mexican ferns.”’ The extensive cross indices (about 3,000 entries) lead to more than 1,200 annotated references arranged by author. Bibliographic citations run only through 1962, which is understandable considering the lengthy and diverse cross indices, which must have taken oe to prepare. A few examples among the ateg: re general indices to 96 AMERICAN FERN JOURNAL annotated catalogues, bibliography, ecology, ethnology, explora- tion and collection, floras and revisions, herbaria, horticulture, and nomenclature. There are also special biographic, geographic, and taxonomic indices. The title of the book is exceedingly modest, for the range of material far surpasses the boundaries of Mexican ferns. The book will therefore be useful to all pteridologists, even those whose interest in Mexican ferns is slight —D. B. L. Two Cuinese FERN Booxs.—Two books by Shu-hsia Fu have recently come to my attention. One is called (transliterated) “Chung-kuo chiieh lei chih wu chih shu,” which may be trans- lated “Genera of Chinese Ferns and Fern-allies.” It is in the Oriental Section of the Library of Congress, no. QK 529 F8 1954. It contains descriptions and keys to the genera, but unfortunately these are not usable, being wholly in Chinese, and also many line drawings. A number of combinations, mostly or all attributed to Ching, are indicated as new, but these are not validly published, since the place of publication and dates are not cited, as required by the Code for works published since J anuary 1, 1953. The second book, ‘‘Chung-kuo chu yao chih wu t’u shuo,”’ i.e., ‘Illustrations of Important Chinese Plants: Pteridophyta” (no. QK 529 F78 1957), is similar except that the illustrations are larger and better. Again the book is wholly in Chinese. A validly published new combination is Gymnopteris vestita var. auriculata (Franch.) Ching (p. 85), which is cited as based on Gymnogramme vestila var. auriculata Franch. Nouv. Arch. Mus. Paris 10: 123. 1887. Gymnocarpium continentale (p. 129) also meets the requirements for valid publication, but the combination is superfluous, having been made by Pojarkova in 1950. The combination Dryoathyrium viridifrons var. okuboanum (Makino) Ching (p. 125) is not validly published because the basionym cited, Aspidium okuboanum Bot. Mag. Tokyo 6: 47. 1892, is a nomen nudum at the place cited. These two publications indicate that some good work in botany is being done in China, but that it is unusable to western botanists, both because the books are so rare in western institu- tions and because they are written wholly in Chinese, which most botanists are unable to read.—C. V. M. Exotic and Hardy Ferns Begonias BOLDUC’S GREENHILL NURSERY 2131 Vallejo Street St. Helena, California 94574 Open Saturdays and Sundays from 10 A.M. to 4 P.M. and by appointment Phone 963-2998—Area Code 707 Mall orders accepted UNUSUAL AND RARE FERNS SHIPPED DIRECTLY TO YOU © List Available e LEATHERMAN’S GARDENS 2637 N. Lee Avenue South El Monte, Calif. 91733 Vou. 58 JULY-SEPTEMBER, 1968 No. 3 American Fern Fournal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY 0. V. MORTON BOLLA M. TEYON CONTENTS A Culture Chamber for Tropical Rain Forest Plants Donatp R. Farrar 97 A Presumable Asplenium Hybrid from Kentucky... Tuomas N. McCoy 103 The Correct Name of a Common Tropical American Oleandra C. V. Morton 105 Vegetative ————— in the Ferns I. Leaf = of Grammitis tenella. Ricuarp A. Wuite 108 ‘Prien: Revision of the Polypodiaceous Genera of India-III. Pleopeltis.......... ae _.S. 8. Brr anp Cuanver K. TrIkHA 119 Trends of = reseae in the Stipe Anatomy of Dennstaedtia and Relate Gaels. es Ricnarp C. Keatine 126 Shorter Note: A Note on Aspidotis........-..-++0+eeeseserrees omnes 140 Notes and News Se eee oe eae 141 Recent Fern Literature ba oe ee be en on 68 ee es 142 American Fern Society... oc. cc cecscsscecscrcsstenererccvceters 144 Mreeou RI Botanica. OCT 2- 1968 GARDEN - imosbv Che American Fern Hocietp Council for 1968 Irvine W. Knosiocu J sap toner of Botany & Plant Pathology, par State BS amet "East Lansin ichigan Warren H. WacNneR, JR., Teniceeiak of Botany, University of Michigna, Ann Arbor, Michigan 48 48104. ice-President LenettTe R. ATKINSON, 415 S. Pleasant Street, Amherst, ‘deccoeea 01002. Secretary y K. Henry, Division of Plants, Carnegie Museum, Pittsburgh, Penn- sylvania 15213. re er Davip B. Le.iincer, Smithsonian Institution, Washington, D. C. 20560. Editor-in-Chief Hational Society Representatives Warren H. Waoner, Je., Dept. of Botany, University of Michigan, Ann Peace ae 48104. Representative to A. A. A. 8. GNER, ~ Dept. of Botany, atvening of Michigan, Ann arco ks "Michigus’s 8104. Eepresentative to A. I. B. 8. American Fern Journal EDITORS Davin B. LELLINGER____Smithsonian Institution, Washington, D. O. 20560. C. V. Mortow __._______Smithsonian Institution, Washington, D. C. 20560, Rota M. Tryon | Gray Herbarium, Harvard Universi bridge, Mass. 02138. ' ty, Cam Im L 1. Wieoms_—_Dudley ey Herbarium, Stanford University, Stanford, Calif. An illustrated — devoted to the general study of ferns, rigid a4 the American Fern Society, and p ublished at 3110 Elm Ave., Baltim Md. 21211. haope chnas postage sald &: eels. The pages of the Jour. nal are open to members who wish arrange ex exchanges ; & mem p list is published at intervals, to assist those interested in obtaining specimens ere Matter for publication should be addressed to Dr. David B. Lellinger, sg cme —— Washington, D. C. 20560. Change address, applications for membership, subscriptions, orders for co oan and other business communications should be addressed to the Treasurer. eg Es $4.50, exclusive of agency handling fees; ens free to — bers o American Fern Society (annual dues. , $4.00; su boral ney “iy membership, $80.00). Extracted repeinta, if ordered i furnish will thors at plus postage ck volumes — to $6.25 each; single ‘inch cuibers of 64 pages oF less, $1.25; 65-80 pages, $2.00 each; over 80 pages, $2.50 each; Cumulative Index to Velulng 125, 50 cents. Ten p ercent discount on orders of six more. volumes or Librarp and Herbarium Dr. W. H. Wagner, Jr., Department of Botany, University of Michigan, Ann Arbor, —— 48104, is librarian and curator. Members may borrow moda and spec at any time, the borrower paying all postal or express Spore Exchange Mr. Neill D. Hall, 1225 Northeast 95th Street, Seattle, Washington 98115, is Director of the Spore orang Viable spores are received and dis patehed, and lists of the collecti ms are sen American Fern Fournal VoL. 58 JULY-SEPTEMBER, 1968 No. 3 A Culture Chamber for Tropical Rain Forest Plants Donatp R. Farrar! A typical greenhouse, although it may function as a tropical house or humid house, usually fails to meet the requirements for growing certain types of tropical plants, notably such rain forest epiphytes as filmy ferns, grammitid ferns, and lycopods. Humidity appears to be the most critical factor for these plants. Temperature and light must also be correct, but they are easier to control. The great success of workers at Kew Gardens and elsewhere in Great Britain in growing these difficult plants has been attributed largely to a climate which enhances efforts to maintain conditions of high humidity, moderate temperature, and indirect light. My research at the University of Michigan involves the study of fern gametophytes which propagate themselves vegetatively by means of gemmae. Self-propagating gametophytes are found in three primarily tropical fern families: Hymenophyllaceae, Vittari- aceae, and Grammitidaceae (Farrar, 1967). The culture of these gametophytes apart from their sporophytes has not been difficult. Most of the variety of techniques commonly used for gametophyte culture are suitable (cf. Stokey, 1930; Steeves et al., 1955; Klein- schmidt, 1957; Miller and oo bates provided that spores or clean, healthy gametop lating material. For my own work I sabe the oui described by Basile (1964) for the culture of bryophytes. A problem arises, however, when ‘I wish to acknowledge the help of Forrest Cochran, W. F. Kleinschmidt, M. F. Tessene, and W. H. Wagner, Jr. in designing and constructing this chamber. Volume 58, No. 2 of the JourNat, pp. 49-96, was issued June 24, 1968. CTION Fig. 1. DIAGRAM OF CHAMBER AME. Fig. 2. DIAGRAM OF CONST OF WET WALL. CULTURE CHAMBER FOR TROPICAL PLANTS 99 it becomes desirable to observe in culture the entire life cycle of these ferns, including their sporophytes. Although earlier experi- ence had shown that filmy fern sporophytes could be kept alive in small terraria with a water-saturated atmosphere, the growth rate under these conditions was slow, and the plants could not be examined conveniently. With a knowledge of the British method of growing filmy ferns and with some first-hand knowledge of the natural habitats of these ferns, I constructed in one of the green- houses at the University of Michigan Botanical Gardens a chamber that would simultaneously maintain condit’ons of 100% relative humidity, 65°-75°F temperature, and indirect natural light at an intensity not greater than 300 foot candles. The basic components of the rain forest chamber are (1) an enclosure constructed of redwood and heavy, clear polyethylene plastic film and (2) a wet wall within, all of which is supported by (3) an elevated table that allows for drainage. The chamber frame (Fig. 1) is constructed of 2” X 2” redwood with doors spaced at convenient intervals, and is covered on the inside aad outside by 4 mil (0.004 inch) clear polyethylene film. This produces nearly a two inch wall of insulating air around the chamber and helps to insulate the inside of the chamber from the remainder of the greenhouse. The wet wall (Fig. 2) is constructed from 6” X 8” X 16” hollow cinder blocks which are stacked (without mortar) in an interlocking position. The holes run vertically and are filled with sphagnum. The entire wall is kept saturated by water dripping onto the top of the wall from a galvanized trough. The trough has 146” perforations spaced six inches apar: and is fed from above by a pipe with mist nozzles spaced three feet apart. Mist nozzles are used because they maintain equal pressure throughout the pipe and thus equalize flow through all the nozzles This method of maintaining high humidity is favored over d'rect spraying of the plants because of possible damage from mineral salts in the water. Water flow is controlled either manually, allowing for a slow continuous dripping, or electrically, using a timer which turns the water on periodically to fill the trough. In either case the water drains slowly from the trough into the wall. The latter control 100 AMERICAN FERN JOURNAL method is preferred because of the possibility of a drip flow either stopping or increasing to an undesirable rate due to gradual change in the controlling valves over a period of time. The table supporting the wall is filled with sphagnum to a depth of three inches to increase the evaporating surface and to serve as a large reservoir of moisture in case of failure of the water supply. A number of problems arose in the development of the chamber. One of these involved the question of how well the chamber would maintain its humidity if the outside water supply were shut off or if the doors were opened repeatedly. When the doors are opened, the humidity immediately drops to near that of the greenhouse. However, the plants have never shown any ill effects from routine use of the doors and, in fact, have shown evidence of drying only when the doors were left open for an hour or more. Humid ty rises to nearly 100% within a few minutes after the doors are closed. When the external water supply was shut off, chamber humidity remained near 100% for as long as three days, which was the most extended period tested. However, plants on the wall did begin to dry out in this time. Similarly, plants mounted on fern fiber that was separated from the wet wall by sections of cork oak bark dried out completely within a few weeks, although the relative humidity was maintained near 100%. Considerable water must pass from the wall to the plants via capillarity, which perhaps is similar to “stem flow” in nature, and blockage of this flow causes desiceation. A second problem was the effect of humidity on metals and wood used in the chamber and in mount'ng the plants. Redwood was used to make the chamber because of its known decay resistance under humid conditions. The pipe and trough were galvanized steel. The plants were initially potted in six inch plastie pots on a medium of fir bark chips and peat. After becoming established, some were transferred to blocks of fern fiber or sections of tree fern trunk which were fastened against the wet wall. Copper wire was used by mistake in fastening some of the plants to fern fiber sections. Within a month the wire had corroded badly and plants up to one inch from the wire had been killed. The wire was re- CULTURE CHAMBER FOR TROPICAL PLANTS 101 placed by nylon cord, which has proved to be quite satisfactory. During the winter months the temperature in the chamber remained near that of the greenhouse (70°-75°F). With increasing solar radiation in the spring it became apparent that the chamber acted as a heat trap; the temperature rose above 90°F on bright, Fig. 3. END VIEW WITHIN CHAMBER SHOWING WET WALL, PLASTIC POTS IN SPHAGNUM, AND AIR CONDITIONER AT FAR END. sunny days. After several exposures to temperatures above 80°F some of the plants developed necrotic patchés : and appeared to have ceased growing. To cireumv ent this problem a 5000 BTU air conditioner was installed. in one end of the chamber. When the air conditioner begins a cooling cycle, relative humidity drops to about 90% because of condensation. on the evaporator coils, but rises again to 100% as the air temperature is low ered. Although 102 AMERICAN FERN JOURNAL an optimum temperature has not yet been estab-.ished, the plants show excellent growth at 65°-75°F. Measurements of light intensity that I have made in the field in Costa Rica indicate that rain forest trunk epiphytes (particularly the Hymenophyllaceae and Grammitidaceae) thrive in less than 500 ft-c, and often in less than 100 ft-c. To maintain this low light intensity the greenhouse roof was whitewashed heavily so that no direct sunlight fell upon the chamber. To minimize excess solar light and heat, the chamber should be placed against a north- facing wall. The chamber is now working well and is highly useful for research with tropical rain forest plants. Such a chamber would also maintain specimens for teaching and display purposes. For the latter purpose a suitable portion of the polyethylene cover- ing could be replaced with glass or with clear vinyl plastic. The chamber has proved to be an effective means of maintaining a habitat not usually found in temperate North American green- houses. LITERATURE CITED Basite, D. V. 1964. New procedures of bryophyte culture which t alteration of the culture medium during the life cycle. The Bryologist 7: 141-146. Farrar, D. R. 1967. Gametophytes of four tropical fern genera reproducing independently of pee sporophytes in the southern Appalachians. Science 155: 1266-126 Kierscumipt, W. F. 1957. A ocd of preparing spores for fern cultures. Amer. Fern J. 47: 95-98. Miter, J. H. and P. M. Miner. 1961. The effects of different light conditions and sucrose on the growth and development of the gametopnyte of the fern Onoclea sensibilis. Amer. J. Bot. 48: 154-159. Sterves, T. A., I. M. Sussex and C. R. ParTanen. 1955. In vitro studies on abnormal growth of prothalli of the bracken fern. Amer. J. Bot. 2. Stoxey, Atma G. 1930. Prothallia of the Cyatheaceae. Bot. Gaz. 90: 1-45. Boranicau Garpens, University or Micuigan, ANN ARBOR, Micu. 48105 ASPLENIUM Hysrip 103 A Presumable Asplenium Hybrid from Kentucky Tuomas N. McCoy In June 1961 when I was scouting the Kentucky countryside in preparation for the Society’s annual fern foray, I found a sterile hybrid Asplentum in a small hole on a moist sandstone cliff growing in a clump with four other Asplenium species. The mass of roots, about one centimeter in diameter, was so matted that the fronds seemed to have a single source. The clump was sent to Dr. Edgar T. Wherry, who realized that some of the fronds were an apparent new hybrid and identified the others as A. pinnatifidum, A. montanum, A. trudellii, and A. trichomanes. The clump was predominantly A. trudellii. Dr. Wherry suggested that Dr. Warren H. Wagner, Jr. see the new hybrid. Dr. Wagner wrote that the plant might be ‘derived from the gametes of Asplenium X trudellii (3X ‘apogamous’) and A. pinnatifidum (2X sexual) and therefore, a pentaploid of the constitution MMR