AMERICAN FERN JOURNAL Volume 104 January-March 2014 QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY A Pluviometric Fern Spore, Fungal Spore, and Pollen TVap Felipe Gdmez-Noguez, Blanca PSrez-Garcia, Aniceto Mendoza-Ruiz, and Alma Orozco-Segovia 1 Isoetes viridimontana: A Previously Unrecognized Quillwort from Vermont, USA Michael A. Rosenthal. Sharon R. Rosenthal, Gabriel Johnson, W. Carl Taylor, and Elizabeth A. Zimmer The Identity of Polypodium gyroflexum (■ Pleopeltis gyrojlexa, Comb. Nov, - Polypo- The American Fern Society Council for 2014 KATHLEEN FRYER, Dept, of Biology, Duke University, P.O. Box 90338, Durham, NC 27708. JAMES E. WATKINS, JR., Dept, of Biology, Colgate University, Hamilton, NY 13346. 1 MARY STENSVOLD, USDA Forest Service, 204 Siginaka Way, Sitka, AK 99835. JAMES D. CAPONE l I I, Div. of Biology, University of Tennessee, Knoxvil GEORGE YATSKIEVYCH, Missouri Botanical Garden, P.O. Box 299, St. WARREN D. HAUK, Dept, of Biology, Denison University, Granville, OH 43023. R. JAMES HICKEY, Dept, of Botany. Miami University, Oxford, OH 45056. r 127, 1 Membership Secretary MO 631660299. of Publications Bulletin Editors EDITOR WARREN D. HAUK Dept, of Biology, Denison University, Granville, OH 43023, ph. (740)587-5758, e-mail: hauk@denison.edu .Dept, of Botany, University of Haw; ASSOCIATE EDITORS Hobart 1 , Honolulu, HI 96822 R. JAMES HICKEY PAULO LABIAK Univen ROBBIN C. MORAN NATHALIE NAGALINGUM CARLROTHFELS . EMILY SESSA H. HAUn^ER Dept, of Ecology and Evolutionary Biology, Univt Lawrence, KS 66045-2106 Dept of Botany, Miami University, Oxford, OH 45056 Federal do Parana, Dept, de Botanica, 81531-980, Curitiba-PR, Brasil New York Botanical Garden, Bronx, NY 10458-5 126 — Royal Botanical Garden Sydney, Sydney, NSW, Australia Botany Dept, r ‘ f, Durham, NC 27708 153706 Herbanum The University of Vermont, Burlington, VT 05405 ..Dept of Biology, Colgate I ‘ y Dept.. Uni JAMES E. WATKINS, JR The “American Fern Journal” (ISSN 0002-8444) is an illustrated quarterly devoted to the general study of ferns. It is owned by the American Fern Society, and publi.shed at The American Fern Society, % Plant Resources Center, University of Texas at Austin, Main Bldg., Rm 127. 110 Inner Campus Dr. Stop F0404. Austin, TX 7871 2-1 7 1 1 . Periodicals postage paid at Austin, TX, and a&litional entry. ders for back issues should be addressed to Dr. Georee Yatskievych, Missouri Botanical Gankn, P.O. Box 299, Sl Lotus, MO 631664)299. Society Life Membership - S360 (add $12) Regular Memberdiip - USA, Canada, Mexh Regular Membership - All other countries ( i American Fern Journal 104(l):l-6 (2014) MISSOURI BOTANICAL A Pluviometric Fern Spore, Fungal S|lAlfe5 Pollen Trap GARDEN LIBRAF AMERICAN FERN JOURNAL: VOLUME 104 NUMBER volumetric trap (Hirst, 1952), which was designed to study airborne pollen. Both of these devices are commonly used in many types of research (e.g., Caulton et al., 2000; Hicks, 1999; Kasprzyk, 2004; Yang and Chen, 1998), as are specialized traps, such as the rotorod trap (Murray et al., 2007), the cyclone spore trap (Tate et al., 1980), and others reviewed by Gregory (1961) and Lacey and West (2006). Although these types of traps can be programmed to survey the air for different periods of time, they are expensive and require energy sources unavailable in some tropical environments (Gupta and Chanda, 1991; Potter and Rowley, 1960). Furthermore, they can be lost or stolen during prolonged field surveys. For these reasons, these traps are mostly used in urban areas, on the rooftops of available buildings (Estrella et al, 2006; Latorre and Caccavari, 2010; Ong et al., 2011; Ting et al., 2010). The use of simpler traps has been reported, such as moss clusters on trees (Limon, 1980), soil samplers (Anupama et al., 2002; Tovar-Gonzalez, 1987), and exposed petri dishes containing different nutritional media (Brown, 1971). However, simpler traps cannot be used to determine the palynomorph influx, and have low local flora representation (Tejero-Diez et al., 1988). Other traps, such as adhesive slices, have low uptake efficiency in prolonged surveys because of sampling area saturation and particle loss due to rain-washing (Melhem and Makino, 1978). Bush (1992) proposed an inexpensive, phenologically accurate gravimetric palynomorph trap composed of a funnel (sampling area) and a carafe; this trap was later improved by Gosling et al. (2003), but both of these traps have the disadvantage of not being able to record pluviometric values. Rainfall is the most important meteorological factor in particle deposition (Ramirez-Trejo, 2002; RamiTez-Trejo et al., 2004; Simabukuro et al., 1998, 2000), yet samplers are unable to measure rainfall. To preserve the pluviometric values associated with a spore-rain survey and to obtain more ecological inferences about rainfall and palynomorph depositions, we modified and simplified the Bush-Gosling trap. Our trap consisted of a funnel 9 cm in diameter attached to a two-liter carafe (Fig. 1 A and B). To prevent the evaporation of the rainfall deposited in the trap, a segment of PVC tube 15.5 cm (6.5 in) in diameter was added, surrounding the trap and serving as a base. A mosquito net or another type of mesh could be used to cover the funnel and thus prevent major detritus buildup. Moreover, the cylinder-base top edge should be beveled to avoid over representation of rainfall by splashing. The uptake area (top of the funnel) can be calculated as A=7cr^; 63.61725124 cm^ thus, the total rainfall can be expressed in millimeters by the following equation: mm = V/A, where V is the final volume of rain collected expressed in mm^ and A is the uptake area expressed in nun^. In addition, the particles may be expressed in density (particles/sampling area), which facilitates analysis and avoids the problems related in the percentage method used by Simabukuro et al. (2000). The advantage of this trap is that it allows better aeropalynological interpretations with the best airborne particle deposition factor (rain). To test GdMEZ-NOGUEZ ET AL.: FERN SPORE TRAP Fig. 1. Pluviometric spore/pollen trap. A. Pluviometric spore/pollen trap diagram: 1) funnel, 2) screen mesh, 3) beveled top edge PVC tube, 4} security seal to secure the mesh, 5) recapping with a hole for the funnel tube, 6) carafe with a 2 L capacity, 7) wall of the PVC tube. B. Trap mounted at 100 pm mesh, 2) 20 pm mesh, 3) graduated cylinder. D. Details of simple filter build up by a PVC tube segment, with a 100 pm mesh attached with a clamp. E. Particles captures with the device at 10 X magnification. F. Striate trilete fern spore aff. Alsophila firma (Baker) D. S. Conant (arrow), surrounded by diverse pollen grains at 100 X magnification. durability and efficiency of the trap, we used it to determine the spore rain in an area near the Malila River in the state of Hidalgo, Mexico. The trap was left in the field in its cylinder-base with monthly trap changes, which did not result in damage from environmental conditions. To vary the sampling period, one can change the carafe capacity and funnel size according to the total rainfall observed in previous years. Because we I (2014) Isoetes viridimontana: A Previously Unrecognized Quill wort from Vermont, USA Michael A. Rosenthal and Sharon R. Rosenthal Gabriel Johnson, W. Carl Taylor\ and Elizabeth j Department of Botany, National Support Center, i Haystack Pond, a small, acidic, oligotrophic h is distinguished from other Isoetes species in 1 high) and granulate to rugulate megaspore s. — Isoetaceae, Isoetes v Although plants in the genus Isoetes (Isoetaceae) are easily overlooked among other vegetation, members of this group have long had the attention of botanists in North America (Engelmann, 1882). This is especially true for several botanists working in New England in the late 1800’s (Dodge, 1897; Eaton, 1898; Underwood, 1888). By the 20^*^ century, Eaton (1900) was able to account for five species and at least five interspecific hybrids occurring in New England (although the hybrids were not originally recognized as such). New England is one of the best-botanized regions in North America and even though searches have been extensive, since 1900 only two additional Isoetes species have been found in this region (Britton and Goltz, 1991; Kott, 1981). Also, most of the possible interspecific hybrid combinations appear to have been reported, so it seems that Isoetes are fairly well documented in New England. Therefore, it is surprising that we are able to report the discovery of a previously unrecognized species of Isoetes from New England. While tracking historic occurrences of Isoetes in Vermont, one of us (M. A. Rosenthal) visited Haystack Pond in the summer of 2010 to verify a previously recorded occurrence of I. tuckennanii A. Br. Rosenthal collected the Isoetes in Haystack Pond, but when his specimens did not fit descriptions of /. tuckermanii, he asked one of us (Taylor) to examine the material. The small size of the mature plants and the granulate to rugulate texture of the megaspores clearly distinguished Rosenthal's collections from all other known species in northeastern North America. Analyses of recently collected voucher specimens from Haystack Pond confirm that that these specimens represent a previously unrecognized species. ^ Author for correspondence. Email: taylorwc@si.edu AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 1 (2014) nd Haystack Pond. A. I. viridimontana. Mature, fertile plant from ; Haystack Pond, Taylor 6743, 20 August 2012. B. Haystack Pond, Isoetes viridimontana M. Rosenthal and W. C. Taylor sp. nov. TYPE: U.S.A. Vermont: Haystack Pond (42.9170°N, 72.9170°W, elevation 2979 feet), 20 August 2012, C. Taylor 6743 with M. Rosenthal, S. Rosenthal, A. Gilman, and J. Taylor (holotype: US; isotypes: MO, NY, VT). Fig. lA Plants aquatic, submerged, not emergent, weakly rooted. Rootstock bilobed, less than 1 cm across. Leaves linear, light green toward apex, brownish to white toward base, gradually tapering to apex, spirally arranged, stiff, spreading, up to ca. 5 cm long, ca. 1 mm wide at mid-length, in tufts of ca. 5—17, semi-terete, with the adaxial surface flattened, abruptly dilated and spatulate at the base, with brownish membranous alae ca. 0.5 mm wide extending along lateral edges from base to ± one-third leaf length. Ligule triangular— cordate, ca. 1 mm long. Velum incomplete, covering ca. 50% of the adaxial surface of the sporangium. Sporangia basal, subglobose— reniform, ca. 2 mm across, adaxial walls spotted by small scattered patches of cells that are pigmented brown. Megasporangia containing up to ca. 40 megaspores. Megaspores globose, white, trilete, ca. 335^02 pm in diameter, averaging 364 pm in diameter, with granulate to rugulate texture. Microsporangia containing thousands of microspores. Microspores elliptic— reniform, echinate, light gray in mass, ca. 22.4—27.0 pm in length, averaging 25.0 pm in length. Chromosomes.— 2n = 22. Paratype.— U.S.A. Vermont: North end Haystack Pond, 11 September 2011, Michael Rosenthal 0021 (US); 27 June 2013, Taylor 6808 (US). Etymology, The specific epithet for this species viridimontana commem- orates the state of Vermont. Vermont translates from the French Wert mont’ , or Green Mountain and is often referred to as the Green Mountain State. Vermont I if? I! ROSENTHAL ET AL.: ISOETES VIRIDIMONTANA H2O, 0.2 lull Biolase® Taq enzyme (Bioline) 5 and 2.5 gl of sample DNA. PCR amplification of the LEAFY intron used the following thermal cycler program steps: Step 1 94°C 5 min., step 2 94°C 1 min., step 3 58"C 1 min., step 4 72°C Imin 10 sec., steps 2-4 for 9 cycles, step 5 94° 1 min., step 6 50° 1 min., step 7 72° 1 min., steps 5-7 for 31 cycles, step 8 72° 5 min. The LEAFY intron 2 region PCR products were size confirmed by electrophoresis for 30 min. at 100 volts in a 2% agarose gel in comparison with Hi-Lo® DNA Marker, (Minnesota Molecular, Minneapolis, Minnesota). PCR products were purified and concentrated by polyethylene glycol precipitation and then ligated into plasmid vector using the TOPO-TA® cloning kit (Life Technologies, Invitrogen, Grand Island, New York). Competent E. coli were heat-shocked, transformed with the vector and cultured on selective medium. Vector insert was PCR amplified for 16 colonies and cycle sequenced according to standard protocols; the sequencing products were analyzed with the ABI 3730x1 automated capillary sequencer (Life Technologies). Sequences were edited and a consensus produced for the clones of each species in Geneious 5.6 (Biomatters, Auckland, New Zealand). The consensus sequences were aligned in Geneious using the program MAFFT 7.0 and the alignment exported as a Nexus file for subsequent analyses. Parsimony analyses were carried out in PAUP* v.4.0bl0 (Swofford, 2003) using the Branch-and-Bound option; that option also was used for 1000 bootstrap replicates. Likelihood analyses were performed with the program GARLI 2.0 (Zwickl, 2006), using the HKY model produced via jModelTest v2.3 (Nylander, 2004). One hundred bootstrap replicates were run in GARLI. Posterior probabilities were produced in MrBayes 3.0 (Ronquist and Huelsenbeck, 2003) after running 10,000,000 generations with a burn-in of 25%. Results Spore size and texture.— Megaspore diameters ranged from 336 gm to 402 pm. The average megaspore diameter was 364 pm. Microspore lengths ranged from 22.4 pm to 27.0 pm. The average microspore length was 25.0 pm. The granulate to rugulate megaspore texture of 1. viridimontana is distinct from the megaspore textures of all other known, aquatic, basic diploid species in northeastern North America (Fig. 2A-G). The proximal surfaces of the megaspore are generally granulate to slightly verruculate whereas the distal surface is more verruculate to rugulate with the molded ridges occasionally anastomosing. A broad, granulate band or girdle is usually apparent along the distal side of the equatorial ridge. The microspores are echinate on all surfaces ^^'chr^llome counts.— Isomes viridimontana appears to be a basic diploid with a chromosome number of 2n=22 (Fig. 3). 1 Further evidence that I. viridimontana is a basic diploid comes from the cloned sequences used in the phylogenetic analysis. Eight cloned sequences of I. viridimontana yielded only a single haplotype. If I. viridimontana were an AMERICAN FERN JOURNAL: VOLUME 104 NUMBER ^ ot /. viridimontana to the six aquatic basic diploid species of Isoetes known to occur in northeastern North ROSENTHAL ET AL.: ISOETES VIRIDIMONTANA AMERICAN FERN JOURNAL: VOLUME 104 NUMBER : - Isoetes bolanderi Isoetes echinospora Isoetes prototypus 1 / 0 . 97 / 0.99 Fig. 4. Phylogenetic tree based on LEAFY intron 2 nucleotide sequences of the six known aquatic , basic diploid species of Isoetes occurring in northeastern North America. The tree is rooted with /. bolanderi, a western North American species. Values for analyses based on maximum parsimony bootstraps (lOOO reps, exhaustive search option), maximum likelihood bootstraps (1000 reps, HKY model) and Bayesian posterior probabilities (10 million generations, HKY model) are listed above the nodes. many useful changes that greatly improved this manuscript. The : individuals for their valuable assistance. Literature Cited BRiTT^^.jM^^d^J. 1991- Isoetes prototypus, a new diploid species from eastern Canada. Dodge, R. 1897. A new quillwort. Bot. Gaz. 23:32-39. Eaton, A. A. 1898. A new Isoetes. Fern Bull. 6:5-7. Eaton, A. A. 1900. The genus Isoetes in New England. Femwort Papers 2:1-16. Engelmann, G. 1882. The genus Isoetes in North America. Trans. Acad. Sci. St. Louis 4:358-390. Generous 5.6 created by Biomatters, Auckland. New Zealand. American Fem Journal 104(1):16-21 (2014) The Identity of Polypodium gyroflexum (= Pleopeltis gyroflexa. Comb. Nov. - Polypodiaceae) Pedro B, Schwartsburd Dept. Biologia Vegetal, Universidade Federal de Vigosa, Av. P.H. Rolfs s.n., Vigosa, MG, : Abstract.— P ofypodium gyroflexum has been a foigotten name since 1940. It is, in fact, an earlier lectotypification, a detailed characterization, and iUustrations for Pleopeltis gyroflexa, comb, nov., plus the synonymization of Pleopeltis repanda. Pleopeltis gyroflexa is known from two populations located ca. 2500-3000 km from each other, and they show some slight morphological differences. The gap between them comprises Amazon forest, the Amazon River, croplands, disturbed areas, and semi-desert vegetation (Caatinga). A vegetation map with their distribution data is also provided. Key Words.— Disjunct distribution, Ceard, north-eastern Brazil, Pleopeltis repanda, Venezuelan Guavana Among the first plants collected in the State of Ceara, northeastern Brazil, were those by J. Huber in 1897. Based on this, Christ (1898) published one of the first accounts of ferns and lycophytes from Ceara, citing 28 taxa, two of which he recognized as new: Pteris pedata var. huberi Christ and Polypodium gyroflexum Christ. A more comprehensive account for Ceara was published by Brade (1940), based on Huber’s collections and also on a more complete collection made by J. Eugenio in 1937. Brade (1940) recognized 67 taxa (including five new taxa), and regarded Pol. gyroflexum as endemic to Ceara. After Brade’s publication, Pol. gyroflexum has been forgotten, not listed in further accounts of Brazilian ferns (e.g., Barros, Lira, and Silva, 1988; Barros et al., 2004; Labiak and Hirai, 2010), nor in any synonymy. Smith (1990, 1995), when working on the ferns from Venezuela Guayana, noticed an undescribed Pleopeltis, that was unique in having repand lamina margins and lacking scaly paraphyses, and he described it as PI. repanda A.R. Sm. Later, Labiak and Prado (2007) expanded the known distribution of Pi. repanda into northwestern Brazil and Guyana. After analyzing types and recent collections, and performing field expeditions m northeastern Brazil, I have concluded that Polypodium ^oflexum and Pleopeltis repanda are conspecific. Polypodium gyroflexum has pnonty over PL repanda by almost 100 years; thus, the new combination for Pleopeltis gyroflexa in Pleopeltis is here provided, along with the synonymization of PI. repanda. ^ Materials and Methods Images of the type collections were analyzed through the websites of JStor Plant Science, and University of California and Jepson Herbaria. Materials SCHWARTSBURD: PLEOPELTIS GYROFLEXA from ALCB, HRB, and HUEFS herbaria were personally analyzed. Other herbaria from northeastern Brazil were visited, but they did not have specimens of Pleopeltis gyroflexa. The map was drawn using the software DIVA-GIS (Hijmans, 2013). For collections with no coordinate data, I estimated locations through searches in Google™ Earth. Since the record for Guyana is based on an informal communication (see Labiak and Prado, 2007; 119), I added it near the border between Venezuela and Brazil. Several localities in northeastern Brazil were visited during March-May 2011, especially in the States of Bahia, Pernambuco, and Ceard. Only one population of Pleopeltis gyroflexa was found, in Serra do Baturite, Ceard. Pleopeltis gyroflexa (Christ) Schwartsb., comb. nov. for Polypodium gyro- flexum Christ, Bull. Herb. Boissier 4:994. 1898. TYPE: BRAZIL. Ceara: Serra de Baturite, Guaramiranga, epiphyte sur les caf6iers, [auf cafestrauchem hinfig], Sep 1897, /. Huber s.n. [gl20] (lectotype, here designated: P- 00633206, image seen; isolectotype; MG-n.v.). Figs. lA-E. Pleopletis repanda A.R. Sm., Ann. Missouri Bot. Card. 77:259, Figs. 3A— D. 1990. TYPE: VENEZUELA. Bolivar: Altiplanicie Nuria, upper part of west- facing wooded slopes, E. of Miamo, 300-500 m, 8 Jan 1961, J.A. Steyermark 88177 (holotype; UC, image seen; isotype: US, image seen). Plants epiphytic. Stems creeping, 3^ mm diam., scaly; scales peltate, lanceolate, 1.5-2. 5 X 0.4-0.8 mm, centrally black, margins clathrate and erose- denticulate, commonly comose. Fronds subdimorphic, the fertile 25—40 cm long, with margins strongly repand, the sterile 20-25 cm long, with margins sinuous; petioles abaxially terete, adaxially grooved, 3-5 cm X 1-1.5 mm, light brown, glabrous; fertile laminae coriaceous, 20-35 x 1.3— 2.8 cm, linear- elliptic, broadest in the proximal 1/4 or in middle, the bases attenuate to cuneate, the apexes attenuate, the margins entire in the lower half, strongly repand in the distal half, abaxially with sparse lanceolate peltate scales, 0.3- 1 mm long, brown to blackish, with light brown margins, adaxially glabrous: sterile laminae coriaceous, 15-20 X 1.2-1.8 cm, linear-elliptic, broadest in the middle, the bases attenuate, the apexes cuneate to acute, the margins irregularly sinuate, the indument similar to fertile laminae; rachises abaxially and adaxially terete, stramineous with dark dots, essentially glabrous but with a few scales abaxially; veins irregularly anastomosing, adaxially some ending in dark hydatodes; sori submarginal, oblong to reniform, 3-5 X 2-3 mm, in one row on each side of midrib, not sunken, without scaly paraphyses; sporangia with long pedicels, 0.5-0. 7 mm long, persistent on receptacle after capsule fallen; spores reniform, monolete, whitish or yellowish. AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 19 Shorter Notes SHORT NOTES 2013. Collected under National Forest Service Forest Products Free Use Permit for Region 1, permit number 2011-7 to E. M. Sigel. The authors thank A. R. Smith and G. Yatskievych for their helpful comments and suggestions. — ^Erin M. Sigel and Anne K. Johnson, Department of Biology, Duke University, Box 90338, Durham, North Carolina, 27708, U.S.A., and Christopher H. Haufler, Department of Ecology and Evolutionary Biology, University of Kansas, 1200 Sunnyside Avenue, Haworth Hall, Lawrence, Kansas, 66045, U.S.A. PTERIDOLOGIA ISSUES IN PRINT The following issues of Pteridologia, the memoir series of the American Fern Society, are available for purchase: 1. Wagner, David H. 1979. Systematics of Polystichum in Western North America North of Mexico. 64 pp. $ 10.00 plus postage and handling. 2A. Lellinger, David B. 1989. The Ferns and Fem-allies of Costa Rica, Panama, and the Choco (Part 1: Psilotaceae through Dicksoniaceae). 364 pp. $32.(X) plus postage and handling. 3. Lellinger, David B. 2002. A Modem Multilingual Glossary for Taxonomic Pteri- dology. 263 pp. $28.(X) plus postage and handling. 4. Hirai, Regina Y., and Jefferson Prado. 2012. Monograph of Moranopteris (Polypo- diaceae). 1 13 pp. $28.00 plus postage and handling. For orders and more information, please contact our authorized agent for sales at: Missouri Botanical Garden Press, P.O. Box 299, St. Louis, MO 63166-0299, tel. 314-577- 9534 or 877-271-1930 (toll free). For online orders, visit: http://www.mbgpress.org. FIDDLEHEAD FORUM The editor of the Bulletin of the American Fem Society welcomes contributions from members and non-members, including miscellaneous notes, offers to exchange or purchase materials, personalia, horticultural notes, and reviews of non-technical books on ferns. SPORE EXCHANGE Mr. Brian S. Aikin, 3523 Federal Ave, Everett, WA 98201-4647 (spores.afs ©Comcast, net), is Director. Spores exchanged and lists of available spores sent on request, http:// amerfemsoc.oig/sporexy.html GIFTS AND BEQUESTS Gifts and bequests to the Society enable it to expand its services to members and to others interested in ferns. Back issues of the Journal and cash or other gifts are always welcomed and are tax-deductible. Inquiries should be addressed to the Membership Secretary, VISIT THE AMERICAN FERN SOCIETY’S WORLD WIDE WEB HOMEPAGE: http://amerfenisoc.org/ AMERICAN FERN JOURNAL aw Volume 104 Number 2 AprU-June 2014 QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY Terrestrial Clubm The Gametophytes and Young Sporophytes of Elaphoglossum decun ceae) in Costa Rica Tbereis Y. S. Choo, Fernando B. Matos, and Bobbin ate Endophyte (DSE) Fungal Association in Is, Eastern Ghats, Southern India iyan Sathiyadash, Rediscovery, Ecology, and Habitat o The American Fern Society Councii for 2014 KATHLEEN FRYER, Dept, of Biology, Duke University, P.O. Box 90338, Durham, NC 27708. President JAMES E. WATKINS, JR.. Dept, of Biology, Colgate University, Hamilton, NY 13346. President-Elect MARY STENSVOLD, USDA Forest Service, 204 Siginaka Way, Sitka, AK 99835. Secretary JAMES D. CAPONETTI, Div. of Biology, University of Tennessee, Knoxville, TN 37996-0830. BLANC LEOn, Plant Resources Center, University of Texas at Austin, Main Bldg., Rm 127, 110 Inner Campus Dr. Stop F0404. Austin, TX 78712-1711. GEORGE YATSKIEVYCH, Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 631660299. Curator of Publications WARREN D. HAUK, Dept, of Biology, Denison University, Granville, OH 43023. R. JAMES HICKEY, Dept, of Botany, Miami University, Oxford, OH 45056. Memoir Editor JOAN N. E. HUDSON. Dept, of Biological Science, Sam Houston State University, Huntsville, TX 77341-2116. DAVID SCHWARTZ, 9715 Christey Way, Bakersfield, CA 93312-5617. Bulletin Editors American Fern Journal EDITOR WARREN D. HAUK Dept, of Biology, Denison University, Granville, OH 43023, ph. (740)587-5758, e-mail: hauk@denison.edu MANAGING EDITOR TOM A. RANKER Dept, of Botany, University of Hawai‘i at Manoa, Honolulu, HI 96822 ASSOCIATE EDITORS ^CRYSTAL ARENS Dept of Geoscience, Hobart & William Smith Colleges, Geneva, NY 14456 JOSE MARIA GABRIEL Y GALAN Dept, de Biologfa Vegetal I, Universidad Complutense de Madrid, Madrid, Spain 28040 GERALD J. GASTONY Dept of Biology, Indiana University, Bloomington, IN 47405-6801 Biology Dept, Grand Valley State University, Allendale, MI 49401 CHRISTOPHER H. HAUFLER Dept of Ecology and Evolutionary Biology, University of Kansas, _ , „ Lawrence, KS 66045-2106 PAin V University, Oxford, OH 45056 Universidade Federal do Parana, Dept, de Botanica, 81531-980, Curitiba-PR, Brasil New York Botanical Garden. Bronx. NY 10458-5126 r Botanical Garden Sydney, Sydney, NSW, Australia FM^v - University, Durham, NC 27708 University of Wsconsin-Madison, Madison, WI 53706 Herbarium. The University of Vermont. Burlington. VT 05405 JAMES E. WATKINS JR Dept of Biology, Colgate University, Hamilton. NY 13346 f Journal” (ISSN 0002-8444) is an illustrated quarterly devoted to the general study of owned by to American Fern Society, and published at The American Fern Society, % Plant Resources TRjSl^rrpStS^JSt^^^d"^ A^” ^ 127, 110 Inner Campus Dr. Stop F(M04, Austin, TX fw missing issues, made 6 months (domestic) to 12 montMforeign) after the date of issue, and or- MOh^^S ^ Yatskievych, Missouri Botanical Garden. P.O. Box 299, inquiries concerning ferns should be addressed to^Secretai^^****^^^ ^ ^ Nlexico (includes Journal and Fiddlehead Forum) $30 So^Mrot^tap -All other countries (includes Journal and Fiddlehead Forum) $40 ^ (a 7.0) tap water. Under such situations, using sodium-softened or distilled water may be necessary to provide neutral to acidic conditions (pH < 7.0) (Husby, pers. comm ) Light.— Contrary to prior suggestions of Barrows, (1935), bright light is an important component of successful terrestrial clubmoss culture. Many tenesMa toa are considered stress-tolerating pioneers, forming robust colonies in frequently disturbed open habitats such as erosional scars, bunch grass p^amo. forest margins, talus slopes, road embankments, heaths, and ?nnQ 1979b; Rasmussen and Lawesson, 2002; Muller et al, 9m T T Marshfield, 2004; Wilfried Bennert et al., 2007; Horn et al., tw -f Lycopodiaceae has especially diverse representation, ( forest understories while all upland habitats (011gaard, 1979b). In understories of western North America, specimens of cons^enfi noticeably more diminutive in stature than their corn^nW." “nrgins (Benca, pers. obs.). Most hemisnhe^P understory taxa in the northern hrestfno on I that provide ample light- Zasada 19901 ^ q- deciduous canopies are absent (Nauertz and Spinulum Svensson et al., (1994) observed that high latitude runners and mn cuttings produced significantly longer horizontal evergreen Va7Z^- when grown under deciduous rather than such taxa in onen observations coupled with presence of clubmosses are more likelv^ latitudes and elevations suggest understory Additionally maintainin shade-tolerant than shade-dependent. iiy. maintaining open habitats via frequent disturbance regimes (e.g. BENCA: CULTIVATION TECHNIQUES FOR TERRESTRIAL LYCOPODIACEAE grazing, mowing, or controlled burning) has been a foundational practice in conservation efforts for dwindling populations of prostrate genera in Europe and North America (Byfield and Stewart, 2001; Rasmussen and Lawesson, 2002; Muller et ah, 2003; Gilman and Marshfield, 2004; Shackleford, 2004), Temperature. — Clubmosses are difficult to establish and maintain if the facility itself is not capable of providing moderate temperatures in conjunction with bright light and high humidity throughout the year. For example, at UCBG, high mortality rates occurred during the establishment phase in many taxa due to prolonged dormancy induced by persistently cold temperatures, low light availability, and short photoperiod in winter (House 1), and high tempera- tures/drying in summer (Propagation and Research houses). Such losses did not occur during the establishment phase in temperature and humidity- controlled research rooms of the UWBG or in growth chamber and laboratory facilities. Fer ttt.t7.e r. — In S. annotinum, extended periods of root growth and phosphate uptake along with efficient phosphorous utilization suggest adaptations for growing in oligotrophic substrates (Headley et al., 1985). This condition is likely widespread across the strobilate clades considering the habitats they associate with and the observation that Diphasiastrum digitatum cutting establishment is impaired by fertilizer application in the field (Railing and McCarty, 2000). In addition, premature rot of newly planted cuttings treated with fertilizer was observed across several genera at UWBG. However, cutting growth rate of most clubmosses observed at UWBG diminished if fertilizer solution was not provided within several months following establishment. Thus, previous accounts suggesting fertilizer has little (or even detrimental) effects on terrestrial clubmosses (Jones, 1987; Railing and McCarty, 2000) apply only to the rooting/ establishment stage. Moreover, increased productivity in established cuttings of all taxa following weak fertilizer application indicates that fertilizer is necessary for culture success following establishment. This response also suggests clubmoss sporophytes do not require mycorrhizal or cyanobacterial associates for successful culture, contrary to prior speculation (e.g. Mickel, 1994; Byfield and Stewart, 2007; McAuliffe, 2008). Medium. — ^Well-aerated inorganic substrates amended with mineral loams promote establishment in terrestrial clubmosses more readily than organic- based mixes possibly because these components take longer to degrade while increasing water use efficiency and nutrient buffering capacity (Owen et al., 2008). Mineral loams may also provide trace minerals necessary for growth. Furthermore, decomposing organic soil components can impede plant growth by decreasing aeration capacity of the medium through increased water retention and compaction (Bilderback et al., 2005; Calonje et al., 2010), while releasing harmful concentrations of CO2 in the rhizosphere (Whitcomb, 2003). Fermenting organic media components also release organic acids that can reduce root growth, increase root oxygen loss, and induce die-back in wetland plants (Nichols, 1981; Armstrong and Armstrong, 1999, 2001). AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 2 (2014) Prior accounts emphasizing the importance of using acidic peat- and sand- based substrates for clubmoss culture (Heim, 1988; Cullina, 2008) acknowl- edge such mixes are less productive for strobilate taxa than substrates obtained from their na ural habitats. Transplantation and greenhouse cultivation UnTted srr ° T** “derstory taxa of the Northeastern United States using well-aerated “native soils" (Barrows, 1935). Since substrates are prevalent across many productive znnl^r i ^atula, 1995; Page, 1997, fre likN!”” >>“nTu™e™*cM b”'*''h^^^ ™88est- three subfamilies of clubmosses in greenTome ®“ chamber facilities. Given the success o^e^a *^*»ratoty, and growth netic span of Lycopodiaceae, these techniqLsT™ T a ici^imiques can be used to increase BENCA: CULTIVATION TECHNIQUES FOR TERRESTRIAL LYCOPODIACEAE 45 accessibility of the entire clade to ex-situ conservation, experimental research, and horticulture. I thank Alan Smith, Chad Husby, Benjamin Ollgaard, George Yatskievych, Nathalie Nagalingum, and an anonymous reviewer for their feedback on drafts of the manuscript. Thanks also go to Doug Ewing, Holly Forbes, the staff of the University of Washington Botany Greenhouse, and the University of California Botanical Garden for generously providing facilities, equipment, care, and support for plants established during this work. I also thank Elbe Duffield, Phil Sheridan, Jason Larson, Chad Husby, Kapua Kawelo, Dawn Foglia, Richard Carter, Chris Carmichael, Martin Grantham, Charles Alford, Chad Wunderlich, and Tony Matthews for providing source material for many of the species covered in this work. Collection of plant material from Hakalau Forest National Wildlife Refuge, HI was conducted under a permit with Creighton Litton during fieldwork funded by National Geographic Young Explorer’s Grant #8985-11. Literature Cited Armstrong, J. and W. Armstrong. 1999. Phragmites die-back: toxic effects of propionic, butyric and caproic acids in relation to pH. New Phytol. 142:201-217. Armstrong, J. and W. Armstrong. 2001. Rice and Phragmites: Effects of organic acids on growth, root permeability, and radial oxygen loss to the rhizosphere. Am. J. Bot. 88:1359-1370. Atmane, N., a. S. Blervacq, N. Michaux-Ferriere and J. Vasseur. 2000. Histological analysis of indirect somatic embryogenesis in the marsh clubmoss Lycopodiella inundata (L.) Holub (Pteridophytes). Plant Sci. 156:159-167. Barrows, F. L. 1935. Propagation of Lycopodium; 1. Spores, cuttings, and bulbils. Contr. Boyce Thompson Inst. Pi. Res. 7:267-294. Bilderback, T. E., S. L. Warren, J. S. Owen Jr. and J. P. Albano. 2005. Healthy substrates need physicals too. HortTechnology 15:747-751. Brodribb, T. J. and S. A. Mcadam. 2011. Passive origins of stomatal control in vascular plants. Science 331:582-585. Byfield, A. and N. Stewart. 2007. Lycopodiella inundata (L.) Holub: Marsh Clubmoss, Lycopodiaceae. UK Biodiversity Action Plan: Plant Life, http://www.plantlife.org.uk/ uploads/documents/Lycopodiella_inundata_dossier.pdf Calonie, C., C. Husby and M. Calonje. 2010. Germination and early seedling growth of rare Zamia HortScience 45:679-683. Cheng, D. H., H. Ren and X. C. Tang. 1996. Huperzine A, a novel promising acetylcholinesterase Copeland, J. 2010. “Experiments in growing Lycopodium in pots.” American Fern Society, Fern Discussions, http://amerfemsoc.org/discus/messages/14/2719.html71289861660 CuLUNA, W. 2008. Native Ferns, Moss, and Grasses. New England Wildflower Society. Houghton Mifflin Company; New York. Field, A. R. and P. D. Bostock. 2013. New and existing combinations in Paleotropical Phlegmariurus (Lycopodiaceae) and lectotypification of the type species Phlegmariurus phlegmaria (L.) T. Sen & U. Sen. Phytokeys 20:33-51. Gilman, A. V. and V. Marshfield. 2004. Lycopodiella alopecuroides (L.) Cranfill Foxtail bog clubmoss New England Plant Conservation Program; Conservation and Research Plan for New England. New England Wildflower Society. Haines, A. 2003. The famiUes Huperziaceae and Lycopodiaceae in New England. VF Thomas Co. Bowdoin. Headley, A. D., T. V. Callaghan and J. A. Lee. 1985. The phosphorous economy of the evergreen timdra plant. Lycopodium annotinum. Oikos 45:235-245. AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 2 ™pe"81031 rhizoids on the thallus Elanh 7'* holbitidoid omm^osT'""’ “ '■®®®">hl®a ‘he sister genus Elaphogiossum decursivum MirvLi ^ (^^go^iarsino et al., 2012). Mexico Belize, Honduras, ^lontane forests of “d Smith, 2004), It is one of tie most^K fr""> >'b‘b^50 " (Mickel Cruces Biological Station to Sista1“ '»w-trunk fern epiphytes at pewmg on the trunks of the ^e ”"®‘ "omm™ly found (Moran et al., 2003). BlaphoXss!!^ f (^aker) D. S. Conant P oglossun, decuativum was not included in the AMERICAN FERN JOURNAL; VOLUME 104 NUMBER 2 (2014] H > a. .he Las . lower ttunks of Alsophila fima B^^SolitmT^ la^o numbers of sporophytes growing ophytes and young spomphidos on a ™ “8in»P«™- C- - - ^ uuiiK or /i. prma. D. Mature gametophyte in the root les om a gametophyte [upper left] to a young sporophyte CHOO ET AL.: GAM. AND YOUNG SPOR. OF E. DECURSIVUM Fig. 2. Gametophyte (A-B) and first sporophyte leaf (C-D) of Elaphoglossum decursivum. A. Ventral view of a mature gametophyte, with a cluster of archegonia at the center, long castaneous rhizoids, and tiny imicellular mar ginal hairs. Black circle indicates apical notch. B. Margin of the gametoph5rte showing secretory, papillate-glandular hairs at different stages of development, from a tiny bulge on the surface of the cell to a fully extended secretory hair with waxy cap. C. First leaf of the sporophyte showing numerous unicellular hairs along its margin. D. Detail of a papillate- surface of the thallus. Unicellular glandular hairs, often with a whitish waxy tip, were found on the thallus surface and margins. These glandular hairs often contained chloroplasts, especially when young, and measured about 50 pm long (Fig. 2B). We also observed minute, branched multicellular proscales (i.e., highly reduced, uniseriate scales; sensu Moran, 1986) on the surface of some of the gametophytes. These proscales often had swollen apical cells, as is typical of scales in many species of Elaphoglossum (Figs. 3A-B). Archegonia were located on the ventral surface near the apical notch (Fig. 3C). No antheridia were observed. Young sporophyte morphology . — The surface and margins of the first few sporoph5^e leaves were covered with minute papillate-glandular hairs like those found on the gametophytes (Fig. 2C— D). With each successive new leaf, the glandular hairs decreased in abundance and became increasingly restricted to the margins. Glandular hairs were absent by time the young sporophyte produced leaves that were about 2 cm long. This corresponded to the fourth or AMERICAN FERN JOURNAL: VOLUME 104 NUMBER : b«..„ .t J..I, .i"r. 0 Conclusions morphology oTmosTo^heff^SrfXgerur^^ being the elongate, strap-shaneH thnil ^ ^ prominent characters hai. on hre marginst‘’dtff:ce%t“ Stokey and Atkinson, 1957) Unlike ” = Elaphoglossum species grown in culture thp f many other decu^ivum lacked a prZiLt gametophytes of thalh. While there have been vp^/ f ^ marginal rhizoids, and branched between field collected and cultur^aro^^p^^'7 addressing the differences and Houston (2002) and Skelton (20^7) shfrihaTsoC^’A^ expected, especially with reoarrJ tL J “ * differences are to be y Wim regard to the expression of sexual systems and the CHOO ET AL.: GAM. AND YOUNG ; OF E. DECURSWUM dorsiventral presence of archegonia and rhizoids. The fact that several of the E. decursivum gametophytes were at least partially degraded seems to reflect the challenges of growing in a natural environment with limited resources. The continuous growth of these elongated thalli may also be an adaptation to growing in the dense root mantle of Alsophila firma, which is a highly heterogenous environment. Chiou et al. (1998) found that gametoph 5 ^es grown in culture from spores of five other Elaphoglossum species only produced archegonia after 8-24 months of growth. Taking this study as a guide, the presence of archegonia and the elongated thallus both suggest that the gametophytes observed were at least several months old, if not older. Perhaps spore germination and subsequent gametophyte growth started at the beginning of the wet season in early May. This observation is consistent with the idea that many fern epiphytes employ the strategy of having long-lived gametophytes (Chiou and Farrar, 1997; Farrar et al., 2008; Watkins et al., 2007). Chiou et al. (1998) did not find any evidence of an antheridiogen system in their study on Elaphoglossum gametophytes and hypothesized that intergametophytic mating was most likely for this group. As we did not find hermaphrodites or antheridia-producing gametophytes, our observations support this idea. However, locating the tiny, antheridia- producing gametophytes typical in a species with an antheridiogen system would be difficult in a field setting, so it is also possible that these individuals were present in our samples but were merely overlooked. One interesting observation from this study was the finding of proscales [sensu Moran, 1986) on the surface of the E. decursivum gametophytes. As is typical of larger “normal” scales in most species of Elaphoglossum (Moran, personal observation), the apical cells of these proscales were slightly swollen with darkened contents. Most important, the proscales are exactly like those found on the leaves, where complete serial homology can be demonstrated between these minute proscales and larger “typical” scales. This finding of proscales is the first observation of scales on an Elaphoglossum gametoph)^e. Without a comprehensive survey on the morphological characteristics of gametoph 5 des from the various sections of Elaphoglossum, it is not possible to confirm the position of E. decursivum within sect. Elaphoglossum based on gametophyte morphology alone. Nevertheless, the characters observed in our study do not present any compelling evidence to the contrary. For instance, the presence of glandular hairs with waxy tips in E. decursivum is consistent with its position outside sects. Squamipedia and Amygdalifolia, as these two sections have been reported as having no hairs or hairs without waxy tips on their gametophytes, respectively. Elaphoglossum decursivum differs from sect. Lepido^ossa by lacking scales bearing many marginal teeth, each consisting of a single, acicular cell, which is diagnostic for sect. Lepidoglossa (Vasco et al., 2009). The two types of indument (i.e., papillate hairs and proscales) found on the very young leaves of Elaphoglossum decursivum represent the first observa- tion of such indument in sect. Elaphoglossum. This observation raises questions on the evolution and development of glabrousness in this clade AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 2 (2014) and it would be interesting to see if the young sporophytes of other members of this clade have similar patterns of development. Our study has shown that the gametophytes and young sporophytes of Elapboglossum decursivum have additional morphological characters (i.e., papillate-glandular hairs) that are absent in the adult sporophytes. Studying these important stages in the fern life cycle not only provides a more complete picture of the biology of such organisms, but could also potentially be useful in a phylogenetic context, complementing the more traditional molecular and morphological studies. This study was conducted in January 2013 during the Tropical Ferns and Lycophytes specialty course sponsored by the Organization for Tropical Studies (OTS). The course was taught by Robbin Moran (The New York Botanical Garden) and James Eddie Watkins (Colgate University). We thank Miguel Chdves from OTS for his help with plant identification. We also thank two anonymous reviewers for their helpful comments and suggestions. This research was partially funded by grants to Matos by the Fulbright Scholarship Program and The CAPES Foundation, Ministry of Education of Brazil, to Choo by the Department of Plant Biology, Cornell University, and to Moran from the U.S. National Science Foundation (DEB1020443). Literature Cited Storey. 1964. Comparative morphology of etophyte of Cmou, W. L. and D. R. Farrar. 1997. Comparative gametophyte morphology of selected species of the family Polypodiaceae. Amer. Fern J. 87:77-86. Cmou, W. L., D. R. Farrar and T. A. Ranker. 1998. Gametophyte morphology and reproductive biology in Elaphoglossum. Canad. J. Bot. 76:1967-1977. Farrar, D. R., C. Dassler, J. E. Watkins, Jr. and C. Skelton. 2008. Gametophyte ecology. Pp. 222- 256, in T. A. Ranker and C. H. Haufler, eds. Biology and Evolution of Ferns and Lycophytes. Cambridge University Press, Cambridge. Gabriel Y GalAn, J. M. and C. Prada. 2012. Farina production by gametophytes of Argyrochosma nivea (Poir.) Windham (Pteridaceae) and its implication for Cheilanthoid phylogeny. Amer. Fem J. 102:191-197. Johnson, A. K., C. J. Rothfels, M. D. Windham and K. M. Pryer. 2012. Unique expression of a sporophytic character on the gametophytes of notholaenid ferns (Pteridaceae). Amer. J. Bot. 99:1118-1124. Lagomarsino, L. P., a. L. Grusz and R. C. Moran. 2012. Primary hemiepiphytism and gametophyte morphology in Elapboglossum amygdalifolium (Dryopteridaceae). Brittonia 64:226-235. LOriga, j., a. Vascx), L. Regalado, J. Heinrichs and R. C. Moran. 2014. Phylogeny and classification of the Cuban species of Elaphoglossum (Dryopteridaceae), with description of Elaphoglossum sect. Wrigbtiana sect. nov. PI. Syst. Evol. 300:937-951. Mickel, j. T. and A. R. Smith. 2004. The Pteridophytes of Mexico. Mem. New York Bot. Card. 88:1-1055. Mickel, J. T. and L. G. AtehortOa. 1980. Subdivision of the genus Elaphoglossum. Amer. Fem J. 70:47-68. Momose, S. 1967. Prothallia of Japanese ferns. Univ. Tokyo Press, Tokyo. Moran, R. C. 1986. The neotropical fem genus Olfersia. Amer. Fem J. 76:161-178. Moran, R. C., S. Kumas and M. Carlsen. 2003. Low-trunk epiphytic ferns on tree ferns versus angiosperms in Costa Rica. Biotropica 35:48-56. American Fem Journal 104(2):58-66 (2014) Effects of Seasonal Changes on Pigment Composition of Azolla filiculoides Lam. Taylan Kosesakal Abstract.— A zo77a is a floating fem, which contains a ■ azollae in the dorsal lohe cavitv nf cyanohacterium Anafenenn changes on the pigment composhion^f the effects of seasonal filicullides bega^bet^in Aptfl'^d May^^" ^^owed that the probable growth season of the A. Key Wvms.-Molla. chlorophyll, anthocyanin. carotenoids, seasonal variations co«;:rhelltt endosymbiCtc and Mayne, 1974). Azolln hL ^ . of the leaves (Peters temperate to tropical climates 5 ® distribution, ranging from sections are kno^ thrs^t in °t A^oUa in two taxonomic Lam., A. rubra R. Br A ^ includes the species Al. filiculoides micmphyUa Kaulf., while thrsection Bh ”***’ Willd and A. pinnala R. Br. and A. nilotica Mett (Vn^H includes the species A. Plant phenology is aff^oH , r°™ 2002)- (Lee et al„ 2009) It is well knnL"*th"!^ factors or by environmental factors and physiology. Fe^ Z setX com f P*'*"* 8^°'^'' seasonal acclimation of some physioloZd'di”.!”^ ^ tal conditions (Volkova et al 20111 It ' "“fp® *“ changing environmen- physiological responses during a pWs 1998). Green Azolla leaves n.rn Lo^ ® ? ®*®8®® (G™*ani el ^ temperature, and adverse environ I ,''°‘*®j ^‘Sh light intensity, low Photosynthetic pigments ■ 2004). the absorption of light (Lirhto th f ^ carotenoids), are responsible for fheuc pilnents aif of J photosyn- variations has been observed in tJ r ^ performance affected by seasonal and Landi, 2013). y^ttocvl “ ^ 7 '''““‘o™ ®' 20111 Paoli tissues. Anthocyanins are produred ^^lhe^®w ‘’T'’’” P‘™‘ the cytoplasm, and then transported KOSESAKAL: EFFECTS OF SEASONAL CHANGES ON AZOLLA FIUCULOIDES into the vacuole (Shirley, 1996). The synthesis of anthocyanins is induced by UV-B (Warren et ah, 2003), nutrient deficiency (Pinto et al, 1999), low temperatures (Rabino and Mancinelli, 1986), water deficit (Nogues et al., 1998), and heavy metal stress (Ling-Peng et al., 2006). The content of the most common plant pigments— chlorophylls, carotenoids, and anthocyanins— as well as their relative proportion determines both the color and appearance of plants (Abbott, 1999). , In Turkey Azolla filiculoides is found naturally only m the Thrace re^on (Unal and Uzen, 1996) and has been grown in Istanbul University Alfred Heilbronn Botany Garden for many years. The purpose of this study was to monitor and investigate the effects of seasonal changes on Azolla filiculoides Lam. For this reason, changes in the pigment content of A. filiculoides were investigated in 2007. Materials and Methods Plant material and sampIing.-The Azolla filiculoides that is grown in Istanbul University Alfred Heilbronn Botany Garden thrives (N 41“01'06", E 28°57'51") with Lemna spp. Pigment analysis of A. fihculoide is based particularly on the months of previous years where there were significant temperature increases and decreases. was collected from the botany garden in March Apn , November and December of 2007. Five replicates of ptots from the different parts of the pond for each month sampled. After washing three times with distilled water, plant fresh weight was measured and pigme contents of the leaves determined. nEanoPs in Seasonal temperature. -To investigate the seasonal Istanbul, maximum, minimum, and mean daily empera u ’ P . j the National Meteorology Service from the Florya meteorological station m carotenoid contents of the leaves in March, AP"1 May 2-4 A. plants totaling 700 mg leaves were used, and for chlorophyll, and carotenoid) were measme (Lichtenthaler and Spectrophotometer) and determined m pg g 8 Welbum, 1983). Mancinelli (1990) method was employed Anthocyanin determination. The Mane l i„avp<; According to for the determination of the arithocyanin content m the leav^.^to^^^^ this method, plant samples of 500 mg of fes weig methanol-Hcfliyo HCl. v/v) and und-mfrige^^^^^^^^^ C with occasional shakings. Then, the UV-Visible Spectropho- content in the filtrate was “ Pg g'' fre^h tometer). The anthocyanin content was expressea a fig g AMERICAN FERN JOURNAL: VOLUME ; NUMBER 2 (2014) ntemp.(»C) (2007) n temp.CO (1975-2006) max.temp.('’C) (2007) rnin.temp.(°C) (2007) -*• maxtemp.(“C) (1975-2006) -A> min.temp.(°C) (1975-2006) replicates of the chlorophyll, was performed using GraphPad Prism calculated. Statistical analysis «"raXL5;,srf.r^<“ Results mean and maximum tempTrato^^ot compared to the December. On the other hand ^ ^^^^"2006; not including December of 2007 were lower than temperatures in April and m Fig. 2, plant color 7ze ZT^ T ^^^8- D- October-November-December 2007 March-April-May and spring temperature) the plants were plants was very slow. At the besinnino r I ^ color, and the growth of red to green, plant growth increaspH ^ Z ^1"" ’ **'® plents’ color changed from gr wtlr mcreased, and vegetative multiplication accelemted. KOSESAKAL: EFFECTS OF SEASONAL CHANGES ON AZOLLA FIUCULOIDES AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 2 (2014) Table 1. Chlorophyll a, b (pg/gFW) and chi a and chi b ratio of A. filiculoides in March, April, May, October, November, December 2007. Chlorophyll a, b contents and chi a/chl b ratios results are means of 5 replicates in March-April-May and October-November-December, “±” indicates standard deviation. Significant differences determined by the Tukey’s Multiple Comparison test (p<0.05) are indicated by different letters (a-e). March May October Chlorophyll a Chlorophyll b 170.6 ± 10.8 d 66.9 ± 1.4 a 323.5 ± 23.8 a 63.4 ± 5.5 a Chi a/Chl b 5.1 ± 0.14 b 5.7 ± 0.07 a 5.6 ± 0.15 a In May, the plants had become fully green, were growing faster, and had completely covered the surface of the pond. During the October-November- December period, the color of the Azolla leaves changed from green to red, plant size decreased and plant growth also slowed down. In addition during the October-November-December period the Lemna spp. population density was higher than the A. filiculoides (Fig. 2). Although A, filiculoides was present in the small pond throughout the year 2007, it only reproduced vegetatively. While the average chlorophyll a/b ratio of the leaves was 2.6 for the March- April-May period, it was 5.5 for the October-November-December period. In April the chlorophyll a/b ratio (Table 1) and total chlorophyll (Fig. 3) were significantly (p<0.05) lower than in the other months, while total chlorophyll was significantly highest in October. The total chlorophyll amounts were 264, 157, and 237 pg g ^ during March, April, and May, respectively (Fig. 3). The carotenoid amount of the A. filiculoides leaves was lower in the March-April- May period when compared to the October-November-December period. The highest carotenoid content was determined in October (p<0.05), but values for November and December were not statistically different from each other (Fig. 3). While the highest level of anthocyanins was measured in March, the lowest was found in May, and these values are significantly different statistically from all other anthocyanin values measured. The anthocyanin content of the leaves was 0.6, 0.47, and 0.33 pg g ^FW throughout October, November, and December respectively (Fig. 3). Plants m their natural environment are exposed to different kinds of environmental stresses, and these conditions lead to both a reduction in growth and metabolism (Abraham, 2010). Azolla grows under different environmental conditions. Temperature is the most important factor affecting the growth and distribution of Azolla (Pabby ef al., 2004). The optimum growth KOSESAKAL: EFFECTS OF SEASONAL CHANGES ON AZOLLA AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 2 (2014) air temperature ranges from 18-28°C for Azolla spp. However, some species of Azolla are capable of tolerating a wide range of air temperatures from -5-35°C (Wagner, 1997). Janes (1998a), determined that under the most adverse winter conditions (January-March), Azolla filiculoides were very small and dark red, and in the spring/early summer, the plants’ color changed from red to green. This color change was also observed in reverse in the autumn in an outdoor culture. In our study, the color and morphological results obtained from A. filiculoides in March- April-May, and October-November-December showed a parallelism with the Janes (1998a) results. Bocchi and Malgioglio (2010) stated that the Azolla (Milan strain) biomass increased from March to April (under the range from 10.4 to 18°C mean temperature/day). In the present study, the A. filiculoides leaf dimensions and the biomass increased and leaves returned to green from April to May (under the range from 12.4 to 19.9°C mean temperature/day; Fig. 2). Thus, the growth season of A. filiculoides likely began between Aprd and May. This study also showed that under natural conditions, while A. filiculoides reproduced vegetatively, it did not reproduce sexually during 2007. The photosynthetic apparatus is quite crucial for communication between die plant and the environment (Mattoo etal., 1999). Chlorophyll concentration IS regulated by plants in order to balance their capacity to both absorb and utilize light energy. This regulation is considered as an adaptation of plants to seasonal fluctuations under environmental stress (Close et al, 2006). In May n growth temperature of plant), the amount of leaves higher than in April. On the other hand, the fwaTf "" ^ ^-2 in April. In addition. Stalling that the chi a/b ratio of the October-November-December . k compared to the ratio in the months of March- nho ‘^“ditional role in energy transdnction, the photosynthetic apparatns might also be an environmental sensor (Huner ef of., teriTneLure anthocyanins under high light intensity, low ha^e a rlddi h ^ environmental conditions, Azolla leaves generally w2r Sin population density was 0bS“ed Soc^ M™ March-April-May*^^ the opposite was iuSiSL Sm kT r hand, it was quite atom m 4) war^ 92^r”™S°'“^ hfarch (mean temper- temperature, 7 6°C) TW ^^s 0.33 pgg ^FW in December (mean response of the nlnntc • ’ may increase the antioxidant sene^ence ^ ^ consequence of exogenic stress or d.yifrculodfes likely begL between Iprili^d Mr;: 5 OF SEASONAL CHANGES C al 104(2):67-102 (2014) Arbuscular Mycorrhizal (AM) and Dark Septate Endophyte (DSE) Fungal Association in Lycophytes and Ferns of the Kolli Hills, Eastern Ghats, Southern India Raji Muthuraja, Thangavelu Muthukumar*. Kullaiyan Sathiyadash, Eswaranpillai Uma, and Perumalsamy Priyadharsini R oot and Soil Biology Laboratory, Department of Botany, Bharathiar University, Coimbatore 641046, Tamilnadu, India Abstract. — ^We examined the extent and type of arbuscular mycorrhizal (AM) and dark septate endophyte (DSE) fungal associations in three lycophyte and 44 fern species collected from three different sites in the Kolli Hills, Eastern Ghats, southern India. Of the 47 plant taxa (belonging to 21 families and 33 genera) examined, 46 had AM fungal and 33 had DSE fungal associations. But, fungal structures were absent in the aquatic fern Azolla pinnata (Azollaceae). This is the first report of AM and DSE fungal status for 16 and 28 species, respectively. Among terrestrial lycophytes and ferns, 26 species had dual association of both AM and DSE fungi, whereas 11 species had only AM fungal association. Vittaria elongata from epiphytic habitats had dual association of AM and DSE fungi. Likewise, Cheilanthes tenuifolia (saxicolous or terrestrial), Cheilanthes opposita, Lepisorus nudus, Pyrrosia lanceolata (terrestrial or epiphytic), and Asplenium lanceolatum (saxicolous or dual a 1 DSE fungi. Seventy two percent of the mycorrhizal lycoph)rtes and ferns had intermediate-type AM and 15 percent had both Paris- and intermediate-types at different sites. Significant variations in AM fungal structures were evident in 16 ferns occurring in two or more sites. Nine AM fungal spore morphotypes belonging to Acaulospora, Funneliformis, Glomus, Gigaspora, and Sclerocystis were found to be associated with lycophytes and ferns. Key Words. — arbuscular mycorrhiza, dark septate endophytic fungi, lycophytes, ferns, Paris-type, intermediate-type, Kolli Hills, Eastern Ghats A wide range of soil fungi colonize plant roots, of which the most common and widespread are the arbuscular mycorrhizal (AM) fungi belonging to the phylum Glomeromycota. These fungi facilitate the uptake of nutrients, especially phosphorus (P) from nutrient deficient soils in exchange for host photosynthates (Smith and Read, 2008). Other benefits for plants from the fungal association include improved water relations, and tolerance to various abiotic and biotic stresses. Surveys of AM associations in vascular plants for over a century have established their wide spread occurrence (Brundrett, 2009 and references therein). Nevertheless, many plant taxa from natmal ecosys- tems world-wide are yet to be examined for their mycorrhizal status. Despite their global distribution, the mycorrhizal status of lycophytes and ferns are scant. Kessler et al. (2010a) indicated that the 971 taxa of lycophytes and ferns whose mycorrhizal status was known to represent less than 10% of the global * Corresponding author; e-mail: tmkum@yahoo.com, tel.091 0422 2428306 AMERICAN FERN JOURNAL: VOLUME 104 NUMBER : lycophyte and fern diversity. Since 2010, gametoph5^es and sporophytes of several lycophytes and ferns from Malaysia and Indonesia (Kessler et al, 2010a), island of La Reunion (Kessler et al., 2010b), Argentina (Fernandez et al., 2010; 2012; Martinez et al., 2012), India (Muthukumar and Prabha, 2012, 2013; Sarwade et al., 2012), Honduras (Zubek et al., 2010) and Japan (Ogura- Tsujita et al., 2013) have been examined for their mycorrhizal status. The AM colonization patterns encountered within plant roots have been designated as Arum-, Paris- or intermediate-types based on the distribution of AM fungal structures. In Arum-type, the fungal hyphae spreads in the root cortex intercellulary forming arbuscules on the lateral intracellular hyphal branches (Dickson, 2004). In Paris-type, the spread of the fungus within the cortex is intracellular forming hyphal coils within cells. Sometimes these hyphal coils bear rudimentary arbuscules. Arum-type is presumed to be formed in roots with high growth rates, and when the root cortex possesses abundant intercellular spaces. In contrast, Paris-type is presumed to occur in slow Rowing roots with limited or no intercellular spaces (Brundrett and Kendrick, 1990). A range of intermediate types exist between typical Arum- and Paris-types exhibiting the characters of both the types (Dickson, 2004). Determining the morphological structures produced hy AM fungi is important because, the AM fungal structures like hyphae (inter-and intra-cellular), vesicles, arbuscules or arbusculate coils, and hyphal coils, have different roles m the symbiosis (Dickson et al., 2007). It has been adequately demonstrated that fungal structures such as arbuscules, hyphal as well as arbusculate coils ^e involved in nutrient transfers thereby indicating a functional association (Smith and Smith, 2011). In contrast, abundance of intercellular or intracellular linear hyphae and vesicles indicate a carbon cost to the host Sanders and Fitter, 1992). The AM morphology reported for a sample of limited lycophytes and ferns (10%), indicates the wide spread occurrence of Paris-type AM morphology (69%) compared to intermediate-type (28%) Dickson et a/. 2007). In a recent study, Zubek et al. (2010) demonstrated ^e wide spread occurrence of Paris-type AM in ferns and lycophytes from Honduras. However, intermediate-type AM morphology was reported in roots ot Lycopodium paniculatum and Equisetum bogotense from temperate forests (Fernandez et al, 2008). Muthukumar and Prabha (2013) showed that 93% of the lycophytes and ferns they examined from different habitats in the Eastern and Western Ghats, south India, had intermediate-type AM morphology. Plant roots including those of lycophytes and ferns are also colonized by melanised or hyaline, regularly septate hyphae, with or without mZ xu e “Us (Haselwandter and Read, 1982; Newsham, 1999). These fungi, commonly known as dark septate endophytes, appear to be ^ '“O™ to colonize over 600 plant species septate “'doph^e (DSE) fungal associations in lycophytes and ferns (Berch 2009^ rnh"l‘*’( “h. 2008, 2010; Hodson et al., 2009, Iqbal et al., 1981; Kessler et al., 2010a, 2010b; Lehnert et al , 2009; MUTHURAJA ET AL.: AM AND DSE FUNGAL ASSOCIATION IN LYCOPHYTES AND FERNS Muthukumar and Prabha, 2012; 2013). Recent studies suggest that DSE fungi could enhance plant growth and health under controlled conditions (News- ham, 2011). It has been speculated that DSE fungi could aid plants in the use ot organic nutrients (Cladwell and Jumpponen, 2003). Further, it has also been proposed that DSE-plant association need not be limited to nutrient acquisition, but could be multifunctional (Mandyam and Jumpponen, 2005). For example, DSE fungi could protect plants against pathogens and herbivores through minimizing the carbon availability in the rhizosphere or ^ngh die production of secondary metabolites (Mandyam and Jumpponen, 2005). It therefore essential to assess plants for DSE fongal associations. Approximately 900-1000 species of lycophytes ^d ferns ^ ?7n the Indian Himalayas and the Eastern and Western Ghats, Of these ™und species of lycophytes and ferns occur in south India ^ reports of AM fongal status and morphology in Indian lycophytes Jvy limited (see Muthukumar and Prabha, 2013 Eastern Ghats are isolated hill ranges occurring m ^ad? The over the three Indian states of Orissa. Andhra Pradesh, and Nad“Jh^ Kolli Hills are among the eight that occur in the f Ghats. Assessments of the florisUc diversity of the ‘ Hills ^ ment of farm lands and exotic plantations shifting ’ J’ 2oi| fire wood collection and tourism Sundaram and Parthasarathy, 2002). Hered the vegetation and large scale habitat destruction and substantially ^ Rere carbon stock (Jayakumar 2002; ^th the ethnobotan- Floristic analyses of Kolli Hills are m y meager for other ical or medicinal uses of angiosperms, an , 2011). Although, plant groups (Arokiyaraj et al, 2007; ^ro^nd 80 species of Gowrishankar et al. (2011) reported th P jg no lycophytes and ferns in their ®™^from this region. This prompted report on the root fungal assomations ferns of the Kolli us to assess the AM and DSE j J/jatiL^ttems and AM fungal Hills. Further, we This inforaation will improve our diversity associated with these distribution and abundance of root knowledge and understanding ^ diis fragile ecosystem, fungal associations in lycophytes and ferns in tms ir g Materials and Methods study sites and ranging from 200 78“ 30'E and a latitude of 11° 10 to 11 30 in wim ^L: VOLUME 104 1 I ■I i I i I I I I I =- I 1 L I I g I g 1 1 n } I ij i i MUTHURAJA ET AL.: AM AND DSE FUNGAL ASSOCIATION IN LYCOPHYTES AND FERNS < << << liiililii ii Iii 111 I I i'i I I ! }i i il Ii AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 2 (2014) MUTHURAJA ET AL.: AM AND DSE FUNGAL ASSOCIATION IN LYCOPHYTES AND FERNS li I i ] I s s s i s I II III 1 1 I I II I I 1 < < oa u < m < aa < m u 0.05). Life-forms differed significantly in average percentage root length with total AM colonization (H4 =78.261, p<0.001). The maximum average percentage root length with total AM colonization occurred in terrestrial taxa (78.81%) and the minimum occurred in ferns from marshy habitats (49.74%) (Fig. 2a). Although the variations in percentage root length with total AM colonization between species was significantly different, the differences between sites were not significant (Table 2, 3). The percentage root length with h)^hae varied among taxa and ranged from 0.67% (L. nudus) and 32.25% [Vittaria elongata, Vittariaceae). The variations in percentage root length with hyphae among sites were significant among species but not among sites (Table 2, 3). There were differences in percentage of root length with hyphal coils both among species and sites. The percentage of root length with hyphal coils ranged from 3.99% {Blechnum occidentale, Blechnaceae) to 51.66% (L. cernuum, Lycopodiaceae). The variation in percentage root length with hyphal coils among species, sites and species X site interactions were highly significant (p<0.001). Percentage of root length with arbusculate coils ranged between <1 [Drynaria quercifolia, Polypodiaceae) and 36.18% [Asplenium indicum, Aspleniaceae). The differ- ences in percentage root length with hyphal coils among species, sites and species X site interactions were significant (p<0.001). The percentage root length with vesicles ranged from <1% [Selaginella sp., Selaginellaceae) to 30.61% [Pseudocyclosonis xylodes, Thelypteridaceae) and the differences among species, sites and their interactions were highly (p<0.001) (Table 2). Life-forms AQ EP MS SX TE Life-forms AMERICAN FERN JOURNAL: I 1 ‘o I ■5 I 1 I -S I I I 1 0 1 i ‘iiiiiiiiiiiiiiiiiiiiiiiiiiiiiiii I'iiiiiiiiiiiiiiiiiiiiiiiiiiipiii 'iiPiiiiiiiiiiiiPiiiiiPiiiipi 'iiPiiiiiiiiiiiiiiiiiiiMiiyiii li 1 1 h I 1 m I f 111 ! MUTHURAJA ET AL.: . 'iiiiiiyiiiiiiiiiiiiiiiiiiiiii ^iiiiiiiiiiiiiiiiiiiiiiiiiiiiii j'iiiiiijiiiiPiiiiiyiiiiiiiiii *iiiiiiiijiiiiiiiiiiiiiiyiii» i<„u<-u< = u<.o< = u< = u75%) and low (<50%) incidence of AM in lycophytes and ferns (see Muthukumar and Prabha, 2013, and references therein). To our knowledge, AM association has been reported in 15 ferns and two lycophyte species for the first time here. The lack of AM association in the aquatic fern A. pinnata is consistent with previous observations of the aquatic ferns Azolla and Salvinia (Gemma et al, 1992; Lee et al, 2001; Muthukumar and Udaiyan, 2000; Ragupathy and Mahadevan, 1993). The non-mycotrophic nature of the free floating aquatic fern A. pinnata could be due to two causes. First, as the fern floats freely in water throughout the year, it has no chance of contacting mycorrhizal inocula of any type unless it drifts to the shores. Second, hydrophytes generally have a poorly developed root system as the necessary nutrients could be absorbed directly by roots and shoot surfaces that are in contact with water (Radhika and Rodrigues, 2007). In contrast to A. pinnata, M. quadrifolia and M. minuta examined firom marshy habitats in the present study were colonized by AM fungi, which corroborates previous findings (Bajwa et al, 2001; Bareen, 1990; Iqbal et al, 1988; Radhika and Rodrigues, 2007). As soil moisture levels in marshy habitats AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 2 (2014) Table 5. Extent of dark septate entophyte (USE) fungal colonization in lycophytes and ferns at different sites of the Kolli Hills. USE Colonization^^ Plant species Arachniodes amabilis Asplenium indicum Asplenium tenuifoliu Blechnum occidental Cheilanthes farinosa Cheilanthes tenuifolia t ± 1.38a 5.25 ± 0.95b MUTHURAJA ET AL.: AM AND DSE FUNGAL ASSOCIATION IN LYCOPHYTES AND FERNS 93 MUTHURAJA ET AL.: AM AND DSE FUNGAL ASSCXIATION IN LYCOPHYTES AND FERNS 95 Plant life-forms significantly affected the intensity of AM colonization. The average percentage root length with total AM fungal colonization of different life-forms was in the order of terrestrial > saxicolous > epiphytes > marshy plants. These results are in agreement with those of Fernandez et al. (2012) and Gemma and Koske (1995) where the incidence and intensity of AM was reported to be higher for terrestrial species compared to other life-forms. All epiphytic and saxicolous taxa observed in the present study were mycorrhizal as previously observed (Gemma and Koske, 1995; Muthukumar and Prabha, 2013; Muthukumar and Udaiyan, 2000). Nevertheless, epiphytic or saxicolous pteridoph 3 hes are often reported to be non-mycorrhizal or facultatively mycorrhizal in other studies (Berch and Kendrick, 1982; Fernandez et al., 2010, Zubek et al., 2010). Lycophytes and ferns growing on bare branches or rocks are frequently exposed to changes in water supply, as water holding capacities of these surfaces are very low (Hietz, 2010). Furthermore, in these extreme environments, high temperature along with strong wind currents may dry these surfaces quite rapidly resulting in vegetative desiccation (Oliver et al, 2000). Therefore, lycophytes and ferns existing on these habitats could depend more on AM fungi for water and nutrients under these stressful conditions as the association has been shown to ameliorate water stress (Smith and Smith, 2011). The lack of AM propagules has often been cited as a cause for the low incidence of AM in epiphytic and lithophytic habitats. Nevertheless, birds and animals could easily bring in the AM fungal propagules to these extreme environments (Gemma et al., 1992; Gemma and Koske, 1995). In addition, AM fungal propagules could reach rock surfaces and rock crevices through the movement of overhead dry soil, dispersal of mycorrhizal root fragments by wind activity, and surface runoffs carrying eroded soil (Berch and Kendrick, 1982). Root colonization directly through the rhizodermis and the presence of AM fungal hyphae within root hairs supports earlier observations (Berch and Kendrick, 1982; Cooper, 1976; Fernandez etal, 2012) where this phenomenon has been documented in lycoph 5 des and ferns. Likewise, the morphologically distinct types of intraradical AM fungal hyphae seen in roots have been reported in vascular plants including ferns (Bentivenga and Morton, 1995; Fernandez et al., 2012; Merryweather and Fitter, 1998). Arbuscule formation on the intraradical hyphae or hyphal coils varied from very limited (e.g., L. nudus, Diplazium sylvaticum, Woodsiaceae) to more elaborate (e.g., A. incisum, B. occidentale) forms. These observations suggests the colonization of pteridophyte roots by different AM fungal taxa as previously shown by both conventional (root squash) and molecular studies (Muthukumar et al., 2009; West et al., 2009). The consistent presence of mycorrhizae as evidenced by the presence of fungal structures in all the individuals of leptosporangiate ferns similar to the observations of Lee et al. (2001) and Fernandez et al. (2012), fails to support Boullard’s (1979) hypothesis that mycotrophy was inconsistent in the advanced leptosporangiate ferns and Zhao’s (2000) suggestion that the most recent common ancestor of pteridophytes was non-mycotrophic. In the present MUTHURAJA ET AL.; MUTHURAJA ET AL.: . MUTHURAJA ET AL.: AM AND DSE FUNGAL ASSOCIATION IN LYCOPHYTES AND FERNS . Mycorrhizal Martinez, A. E., V. Chicx:chio, L. T. Em, M. A. Rodriguez anc association in gametophytes and sporophytes of fem Ptei intraradices. Revista Biol. Trop. 60:857-865. McGonigle, T. P., M. H. Miller, D. G. Evans, G. L. Fairghild and J. A. Swan. 1990. A method which gives an objective measure of colonization of roots by vesicular- arbuscular mycorrhizal fungi. New Phytol. 115:495-501. Merryweather, J. and A. Fitter. 1998. The arbuscular mycorrhizal fungi of Hyacinthoides non- scripta. I. Diversity of fungal taxa. New Phytol. 138:117-129. Miller, S. P. and R. R. Sharttz. 2000. Manipulation of flooding and arbuscular mycorrhiza formation influences growth and nutrition of two semiaquatic grass species. Funct. Ecol. 14:738-748. Mishra, R. R., G. D. Sharma and A. R. Gathpon. 1980. Mycorrhizas in the ferns of North Eastern India. Proc. Indian Nat. Sci. Acad. 1346:546-551. Mohanraj, R., j. Saravanan and S. Dhanakumar. 2010. Carbon stock in Kolli forests. Eastern Ghats (India) with emphasis on aboveground biomass, litter, woody debris and soils. iForest Muthukumar, T. and K. Prabha. 2013. Arbuscular mycorrhizal and septate endophyte fungal associations in lycophytes and ferns of South India. S 5 nnbiosis 59:15-33. Muthukumar, T. and K. Udaiyan. 2000. Vesicular arbuscular mycorrhizae in pteridophytes of Western Ghats, South India. Phytomorphology 50:132-142. Muthukumar, T. and K. Prabha. 2012. Fungal associations in gametophytes and young sporophytic roots of the fem Nephrolepis exaltata. Acta Bot. Croat. 71:139-146. Muthukumar, T., K. Sathiyadash, E. Uma and V. Muniappan. 2009. Arbuscular mycorrhizal morphology in sporophyte of Psilotum nudum. Ph 3 d;omorphology 59:141-146. Muthukumar, T., M. Senthilkumar, M. Rajangam and K. Udaiyan. 2006. Arbuscular mycorrhizal morphology and dark septate fungal association in medicinal and aromatic plants of Western Ghats, Southern India. Mycorrhiza 17:11-24. Newsham, K. K. 1999. 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New Phytol. 120:517—524. Shorter Notes AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 2 (2014) O' +..» Maui Deparia /raa/aana Current Location \ Hawai i Deparia kaalaana Historic Locations Fig. 1. Distribution of Deparia kaalaana, Hawaiian Islands. made in a riparian site in fernsrlTh Wet Forest (Gagne & Cuddihy in Wagner, Sit Risf o Flowering Plants ofHawaPi. Univ. of Hawai'i ^rvLirit n“'T “ perennial stream Dool observaf^ t n™ “^'''taeging ledge on the western side of a small exh hi?^eT t the banks as well as in adjacent vegetation in strona floods capable of surviving submergence (Aspleniaceael adjacent are Asplenium excisum C. Presl Sel A SL rSr ? ^^otheca (Hillebr.) Copel. (Hymenophyila- (Sorene 1 Hvl rn Presl (Pteridaceae), Dryopteris wallichiana SKpELSST *• Hillebrand's and Kato's reports as weTla's thf^^ Kipahulu Valley. kaalaana is a rheophvte Katn f f specimen, suggest that D. roots and oblique pii^ae) suoDorted tb morphology (well-developed contrasting hiLriLnnf™ ^ assumption. With such scant and would be premature to state a single recent observation it reported a single geLma ^ rheophyte. Hillebrand gie gemma on the tip of one frond of Deparia kaalaana, however, SHORTER NOTES INFORMATION FOR AUTHO PTERIDOLOGIA ISSUES IN PRINT The following issues of Pteridologia, the memoir series of the American Fern Society, are available for purchase: 1. Wagner, David H. 1979. Systematics of Polystichum in Western North America North of Mexico. 64 pp. $10.00 plus postage and handling. 2A. Lellinger, David B. 1989. The Ferns and Fern-allies of Costa Rica, Panama, and the Choco (Part 1 : Psilotaceae through Dicksoniaceae). 364 pp. $32.00 plus postage and handling. 3. Lellinger, David B. 2002. A Modem Multilingual Glossary for Taxonomic Pteri- dology. 263 pp. $28.00 plus postage and handling. 4. Hirai, Regina Y., and Jefferson Prado. 2012. Monograph of Moranopteris (Polypo- diaceae). 1 13 pp. $28.00 plus postage and handling. 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Inquiries should be addressed to the Membership Secretary. VISIT THE AMERICAN FERN SOCIETY’S WORLD WIDE WEB HOMEPAGE: http ://amerfemsoc.org/ AMERICAN FERN JOURNAL Volume 104 Number 3 July-September 2014 QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY The American Fern Society Council for 2014 KATHLEEN FRYER, Dept, of Biology, Duke University, P.O. Box 90338, Durham, NC 27708. JAMES E. WATKINS. JR.. Dept, of Biology, Colgate University, Hamilton, NY 1 3346. MARY STENSVOLD, USDA Forest Service, 204 Siginaka Way, Sitka, AK 99835. Pre^i^ient Elect JAMES D. CAPONETTI, Div. of Biology, University of Tennessee, Knoxville, TN 37996-0830. ^ GEORGE VATSmvvcH,Mis»^B«a„i.a,G»fe„,P.O.B„,299.S,. Louis, MO WARREND.HAUK,Du,x„fBi„E,gy,Du„is„„U^„„„,,0,^,„„,0„,3^. Cu™u,.o/P».«cuuo». R- JAMES HICKEY. Dept of Bouu.,, Mi™i Ouiversify, Oxforf, OH 45056. Memoir Editor itsvme,TX 77341-2116. Bulletin Editors American Fern Journal WARREN D.HAUK Dept, of Biology, TOMA. RANKER.. managing editor -Dept, of Botany, Universit . A4.,.,ersity. Granville, OH 43023. (740)587-5758. e-mail: hauk@denison.edu 1. Honolulu, HI 96822 NANCRYSTALARENS Dent JOSE MARfA GABRIEL Y GAUn Hobart & William Smith Colleges. Geneva, NY 14456 ^ Biologfa Vegetal I, Universidad Complutense de Madrid, GERALD J. GASTONY . r. Madrid, Spain 28040 GARY K. GREER biology, Indiana University Bloomington IN 47405-6801 CHRISTOPHER H. HAUrIer ^P* ’ State University. Allendale, MI 49401 of Ecology and Evolutionary Biology, University of Kansas, R. JAMES HICKEY Lawrence, KS 66045-2 106 ^ Depf Botany University, Oxford, OH 45056 do Parana, Dept de Botanica. 815^1-qgn Cnritiha-PR. Brasil PALILOLABIAK universidarle F la KLAUS MEHLTRETER ™ Parana, Dept de Botanica, 81531-980, Curitiba-PR, Brasil i„„ odeEcologfa,A.C.,Xalapa 91070, Ver., Mexico York Botanical Garden, Bronx, NY 10458-5126 ROBBINC. MORAN NATHALIE NAGALINGUM CARI Rnmicci c «aihalieN.AGAL CARL ROTHFELS emilysessa MICHAEL SUNDUE .. New York Botanical Garden, Bronx, NY 10458-5l2f Botanical Garden Sydney, Sydney, NSW Australia “ _ Botany Etept., Duke University, Durham, NC 27708 ’ Hnivetsity of Wisconsin-Madison, Madison, W1 53706 ;;; He^anum. The University of Vermont, Burlington, VT 05405 ” ^ Biology, Colgate University, Hamilton, NY 13346 OCT 2 4 20H on 17 October 2014 GARDEN LIBRARY Megalastrum (Dryopteridaceae) in Andean South America, Part I MORAN ET AL.: ANDEAN MEGALASTRUM AMERICAN FERN JOURNAL; VOLUME 104 NUMBER 3 (2014) B (A-C) and echinulate (D-F) perispores. A, B, D, and E were taken with the SEM; C and F with the light microscope. A. M. falcatum. B. M. insigne. C. M. biseriale. D. M. fugaceum. E. M. nanum. F. M. fugaceum. A: Ceron 1483 (MO). B: Grubb et al. 1195 (NY). C: Navarrete & Ollgaard 3047 (AAU). D: Serrano et al. 5225 (GOET). E: Moran S' Robrbach 5380 (MO). F: Huaylla et al. 1554 (NY). In A, B, D, and E, scale bars = 10 micrometers. In C and F, scale bars = 50 micrometers. grooved or only shallowly so, scaly and pubescent abaxially, densely pubescent on the adaxial surfaces, the hairs whitish, spreading to antrorsely strigose, multicellular, if glands present, these ca, 0.1 mm wide, spherical, shiny, yellowish to orangish clear, sessile to stalked; basal basiscopic segment of pinnules becoming decurrent and adnate to the pinna rachises, the vein supplying the segment springing from the pinna rachis instead of the costule; hydathodes present or (more rarely) absent; indusia absent or (less commonly) present, circular, brown, firm, attached at the base of a sinus, in some species minute and fugacious; spores monolete, nongreen, the perispores cristate or spiny, non-fenestrate or rarely so (2 species); x=41 (Kramer and Green 1990). Etymology. — The name Megalastrum is taken from the Greek mega- for large, and -Lastrea, which refers to the genus Lastrea Bory named after Charles Jean Louis Delastre (1792-1859), a French botanist. When Ctenitis was first published (as a subgenus of Dryopteris), it included species now placed in Ctenitis (C. Chr.) C. Chr., Megalastrum, and Triplophyllum Holttum. Christensen later (1919 [1920]) recognized four informal species groups within the subgenus, one of which he called the “Group of D. subincisa.” This group is the same as what we are calling Megalastrum. AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 3 (2014) I MEGALASTRUM AMERICAN MORAN ET AL.: ANDEAN MEGALASTRUM MORAN ET AL.: ANDEAN MEGALASTRUM the scales like those abaxially; basal basiscopic pinnules of medial pinnae not enlarged or overlapping the lamina rachis; costules abaxially very sparsely provided with sessile glands, sparsely puberulent and sparsely scaly, the hairs like those of the pinna rachises, the scales ca. 1.5 X 0.3 mm, otherwise like those of the pinna rachises, adaxially eglandular, pubescent, scales absent, the hairs ca. 1 mm long, 5-7-celled, acicular, ascending, appressed; laminar tissue between veins abaxially sparsely provided with 0.1 mm long yellowish sessile glands, sparsely puberulent, the hairs 0.1-0.2(-3.0) mm long, 1- or 2-celled, erect, acicular, whitish, adaxially eglandular, densely pubescent, the hairs 0.3-0. 5 mm long, 2-3-celled, erect to appressed, acicular; ultimate veins visible on both surfaces, abaxially sparsely glandular and puberulent, the glands and hairs like those of the laminar tissue between the veins, adaxially eglandular, sparsely pubescent, the hairs 0.5- 0.7 mm long, 4-7-celled, acicular, erect to ascending; lamina margins eglandular, densely ciliate, the hairs 0.2-0.3 mm long, 2- or 3-celled, acicular, ascending; indusia present, conspicuous, circular, 0.5-0.7 mm wide, glandular, concolorous, brown, the margin and surface of the indusia glandular, pubescent to sparsely pubescent or sub-glabrous, the glands sessile, spherical, yellow to orange, the hairs 0. 1-0.2 mm long, acicular, spreading; spores cristate. Distribution and ecology.— Colombia, Ecuador, Peru; 1280-2150 m. Selected Speqmens Examined.-COLOMBIA. Huila: Cordillera Oriental, SE of Garzon, 1950 m, [9°8'7"N, 74°3'38"W], 7 Feb 1945, Uttle 9415 (US). ECUADOR. Napo: Road Baeza-Tena, 2 km N of Cosanga, 1900-2000 m, 0°34'S, 77°53'W, 19 Jan 1992, Ollgaard et al 99562 (AAU, QCA). Pichincha: Cordillera Occidental, casi 40 km por el camino oeste de Quito a Mono, 2150 m, 0°02'S, 78°38'W, 21 Jan 1984, Moran 3567 (NY). PERU. Junin; Chancamayo, Rio Rondayacu, 45 km from San Ramon, 1880- 1950 m, 11°20'S, 72°20'W, 15 Oct 1982, Smith et al. 2615 p.p. (AAU, F, MO, NY). San Martin: Huallaga, Saposoa, al sur de Anazco Pueblo, 2000 m, [6 56 S, 76°46'W], 3 Sep 2000, Quipuscoa et al. 2340 (F, NY, UC). Megalastrum acrosorum is distinguished by indusiate son and rachises and costules abaxially with pale flaccid appressed scales. The other indusiate species oi Megalastrum have pinna rachis scales firmer, spreading, denticulate (at least apically). The almost complete absence of glands on the pmna rachises and costules abaxially also helps to distinguish M. acrosorum from the other species. Most similar morphologically is M. andicola, which differs by the scale and pubescence characters given in the key. Megalastrum acrosorum was originally described as having son restncted toward the apex of the pinnules (thus the specific epithet). Although this is true for the original material, more recent specimens show that the son may occur throughout the length of the pinnules. The restriction of son tow^d the apex is diagnostic for M. apicale R. C. Moran & J. Prado, a species from Costa Rica and Pan^a long thought to be conspecific with M. acrosorum (e g., by Snu* and Moran 1995). Megalastrum apicale differs by pinna rachises abaxiaUy with dark brown scales and hairs 0.5-0.7 mm long (Moran and Prado 2010). AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 3 (2014) AMERICAN FERN JOURNAL: VOLUME 104 NUMBER : MORAN ET AL.: ANDEAN MEGALASTRUM 0.3 mm long, 2- or 3-celled, adaxially the hairs 0.3-0.5 mm long. 2^-celled; lamina margins eglandular, ciliate, the hairs 0.2-0. 3 mm long, 1-3-celled, ascending to spreading; indusia < 0.2 mm wide, circular, fugacious and usually seemingly absent, eglandular. pubescent by whitsh acicular hairs ca. 0.1 mm long; spores echinulate. AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 3 (2014) MORAN ET AL.: ANDEAN MEGALASTRUM 127 Distribution.— Peru, Bolivia, northern Argentina, southeastern Brazil; 750- 2100 m. Selected Specimens Examined. — PERU. Ayacucho: Aina, between Huanta and Rio Apurimac, 750-1000 m, [12°56'S, 74°15'W], 7-17 May 1929, Killip & Smith 22779 (US). Cusco: Distr. Urubamba, en el camino del proyecto arqueologico Mando-Paturusi, ca. 115 km de Cusco, en Machupicchu, 2100 m, 13°09'S, 72°31'W, 4-6 Apr 1988, Nunez V. 8929 (AAU, F, MO, NY, UC). Huanuco: Cochero, [9°35'S, 75°51'W], s.d., Dombey s.n. (P). BOLIVIA. Chuquisaca: Herando Siles, 1575 m, 20°20'16"S, 62°02'42"W, 23 Dec 2005, Serrano et al. 6873 (MO). La Paz: Prov. Nordyungas, Polo-Polo bei Coroico, [16°10'S, 67°43'W], Jun-Jul 1912, Buchtien 3598 (BM, F, S, US). Santa Cruz; Valle Grande, 12 km de Loma Larga a Masicuri, 1250 m, 18°47'S, 63°57'W, 21 May 1996, Kessler et al 5971 (GOET, UC). Tarija: Aniseto Arce Ruiz, Campamento de guardaparques Sidras, senda La Cascada, 890 m, 22°14'S, 64°32'W, 19 Jun 2004, Jimenez et al. 2468 (UC). ARGENTINA. Salta: Parque Nacional El Rey, [24°41'S, 64“36'W1, 21 Mar 1981, Brown 1339 (MO, NY). Tucuman: Depto. Tab, Quebrada de Cainzo, 900 m, [26°49'S, 65°37'W], 18 Nov 1951, Sleumer 2097 (BM). Megalastrum adenopteris has several distinctive characters that, in combi- nation, make it easy to identify. The species is characterized by large, highly divided leaves and laminae on both surfaces between the veins densely and evenly pubescent. The hairs are often gland-bpped with the terminal cell enlarged or globose, yellowish or orangish. The veins tips are slender, not enlarged and hydathodous as is typical in the genus. The indusia are distinchve by being minute (< 0.2 mm wide) and pubescent. After drying and contracting, they often appear as a tuft of whitish acicular hairs in the center of the sorus. This species is a member of the M. pulverulentum group. 3. Megalastrum alticola M. Kessler & A. R. Sm., Amer. Fern J. 96:36. 2006 Type.- BOLIVIA La Paz: Prov. Nor Yungas, 2 km de Chuspipata hacia Coroico, 16"22'S, 67°49'W, 2900 m, 17 Sep 1997, M. Kessler 11945 (holotype: UC! [barcode 1621224, 1621225); isotypes: GOETl, LPB-n.v.). Figs. 5C, 9F-K, lOD. Rhizomes erect, the scales 6.0-10,0 X 1.0-2.0 nm, ascendmg. Imear- lanceolate, brown, dull, Hat, sparsely denticulate; leaves 1. 0-2,5 m long; petiole base scales 10.0-15.0 X 1.0-1.5 mm, spreading to ascending Imein^ lanceolate, brown, dull, lustrous, flat, not tortuous denticulate, the teeth simple or sometimes bifid; laminae up to ca. 1.5 m long basally 3-pinnate- pinnatisect, medially 2-pinnate-pinnatisect; lamina rachises without hairs pinnule. M-O. M. decompositum. M. Adaxial surface of pimn Abaxial surface of pinna rachis, costule, and pinnules. aUetail A-G: from Brazil, isolectotype, Jurgens s.n. [Rosenst^k FiIims L: Isotype, Palacios 5844 (MO). M-O: Isotype, van der Werffet AMERICAN FERN JOURNAL: VOLUME NUMBER 3 (2014) MORAN ET AL.: ANDEAN MEGALASTRUM 129 abaxially; basal pinnae 20-30 cm long, inequilateral, longer basiscopically; pinna rachises abaxially eglandular, pubescent, scaly, the hairs 0.2-0.4 mm long, 3-5- celled, spreading to slightly ascending and tortuous, the scales denticulate, sometimes with bifid teeth, spreading to ascending or appressed. 2.5-7.0 X 0.5- 1.0 mm, lanceolate to linear-lanceolate, brown to light brown, lustrous, not clathrate, not bullate, adaxially very sparsely glandular, densely pubescent, sparsely scaly, the glands ca. 0.1 mm long, sessile, brownish, the hairs 0.6- 1.0 mm long, 4-7-celled, spreading to antrorsely strigose, scales like those on the abaxial surfaces; costules abaxially rarely glandular, pubescent, scaly, the glands ca. 0.1 mm, sessile, globose, yellowish, the hairs 0.2-0.4 mm long, 2- or 3-celled, spreading, scales 0.5-1.0 X 0.2-G.3 mm. spreading to loosely appressed, lanceolate, entire to denticulate, adaxially eglandular, sparsely to moderately pubescent, scales absent, the hairs 0.4-0.7 mm long, 2-5-celled, antrorsely strigose; laminar tissue between veins on both surfaces eglandular, glabrous; ultimate veins surfaces abaxially visible, obscure adaxially, eglandular, abaxially pubescent, the hairs 0.1- 0.2 mm long, 1- or 2-celled, spreading to ascending, adaxially the hairs absent or 0.2-0.5 mm long, 3-5-celled, spreading; lamina margins eglandular, ciliate, the hairs 0. 1-0.3 mm long, 1- or 2-celled, ascending; indusia absent; spores cristate. Distribution.— Ecuador, Peru, Bolivia; 1600-3400 m. Selected Specimens Examined.-ECUADOR. Cotopaxi: Cantdn Sigchos, bosque en sucesion, dosel ca. 8 m, por fuente de agua, 3400 m, 0°35'40''S 78 49 54 W, 21 Jul 2003, Ramos et al 6371 (MO, QCNE). Imbabura: Carretera Otava o- Selva Alegre, 2800-3200 m. [0n4'N, 78 16'W1, 13 Feb 1989, van der Werff &■ Palacios 10570 (MO, MICH, QCNE). Morona-Santiago: road Plan de Mdagro- Gualaceo, Km 10.8, 2200-2250 m, 3°00'S, 78 32'W, 24-25 Nov 1997, 0llgaard & Navarrete 2755 (AAU, QCA). Pichincha: Valle de Lloa. 2800 m, [0 15 S, 78°34'W], 1919, Mille s.n. (GH, NY, P. S). ^ PERU. Cusco: Prov. La Convencion, Distr, 1750 m, 12”58'51"S, 72'03'31'W, 27 Jun 2003, Bomno et al. 746 (MO, NY UC). Huauuco: Cushi, 1600 m. [9»51'0'S, 75 41'0'Wl, 1^23 (un 1923, MocBn* 4855 (F US) Jum'n: Chanchamayo, Ghilpez ca. 26 km of San Ramon, 1720- 1850 m,’l0n6'k 75°22'W, 19 Oct 1982, Smith & Palacios 2639 (F. MO). Pasco: oxapampa, Distr. Huancabamba. Sector Quebrada Y^acta^ ‘^^9^0 .X nal Yanachaga-Chemillen), 2260 m. <5 S 75 28 55^ 18 2004^ Mellado Sr Becerra 1803 (MO); idem, 2410 m, 10 23 20 S. 75 28 263^, 20 Sep 2004, MeHado «• Becerra 183S (MO). ,, nQ'aQa"C BOLIVIA. Cochabamba: Prov. Tiraque, El Limbo, 2180 m. 17 09 39.4 S, 65»37-18'W 21 Aug 2003, Zabalaga 1282 (NY). La Paz: Coroico, Nor Yungas. Ltrada drCoroico para Caranavi, Prdximo a Chuspipata, 3000 m, 16 17 48-S. 67°48'46"W, 27 Feb 2008, Labiak 4730 (NY, UPCB). Among the large decompound species of the genus, Megalastrum alt, cola is characteLed by pinna rachises abaxially pubescent Md with l“cedmm biseriale Baker. Syn. Fil. W99 n™ biseriale (Baker) Diels, Nat. Pflanzenfam. 1(4):170. ImTibZa ^T 254- 1905. Ctenitis senaJjs (Baker) Lellinger, Fem Gaz. 11:108. 1975. Type.— ECUADOR. Ctenite bidecomta Lellinger, Proc. Biol. Soc. Wash 98-373 fie 5 1985 Mega nstrn^ , (Lellinger) A. R. Sm. & R. c. Moran, Imer. Fem Ld 9"47-N 83 Estrella-Sta. Maria -tpLto CRTefuS)'' " (holotype: US- "^CvISnTf «^344. tab. 7, fig. 17. 1907 TvPE.- Stubel 998 ’’8"25'W|, s.d., M. A. isolectotype: BM![bic;deMoraT‘^' ®’ ' 77:129. 1987 [publishp/q \a A. R. Sm., Amer. Fem J. Yungas, Simp^net ' Tvpe.-BOLIVIA. La Paz: Prov. Sud 1906, O. BuchJen 419 (lertr.r^'^^a 65°05'W1, 2000 m, 16 Nov P! [barcode 000600636); isolertot™:'uS-n 1 Megalastnim oequatoriensp A \/t' t-. B. 2008. Type.-ECUADOR 'S i- T^*-i Chico area of Awa Reserve’ 0°58 'n’ 7R°^'i'w^^' encampment, Gualpi S. Hoover, A. ArnuelTp r i ^^'W, 1350-1400 m, 21 Jan 1988, [barcode 3710680]; LotvnV ^609 (holotype: MO! Mephrodium subgla^Ts,^^' ECUADOR. Pichincha- “in c-i ’ 259. 1893. Type.— 1200 m, Oct 1891, A Sodiro^ 0''“’’® [0°19'S, 78»27'W1. 000568196); isolectotvnes- PI r^' '‘®?°*ype. here designated: P! [barcode Mzomes erect sc.,.. „ , Rhizomes erect, scales 7 0-17 QPLS?-n.v„ Q?-n.v,). linear-lanceolate, brown, lustrous 4”™ *“"®' ascending. Simple or bifid; leaves 0.6-1 2 m 1 denticulate, the teeth like those of the rhizomes but oftersnrr?”*^ ° ^ laminae 0.5-0.9 m long, basallv 2 “ more sparsely denticulate; to 2-pinnate-pinnatifid; lamina ^P^'^nate-pinnatifid, medially 1- 10.0-15.0(-23.0) cm long, equilaterart^^^T®°^- pinnae abaxially eglandular, pubescem scalv tb \ pinna rachises 3-celled strigose or oppressed to astndt' T mm long, 2- or brown lanceolate, lustrous, appressS m ^ °-2-0-5 sp^sely denticulate, not datSate nt t^ut not tortuous), pubescent, non-scaly, the hairs ^^axially eglandular, densely ascending to strigose, light brown- ’ ^ ^as»S€opic pinnules of medial pinnae AMERICAN FERN JOURNAL: VOLUME 104 NUMBER : MEGALASTRUM AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 3 (2014) MORAN ET AL.: ANDEAN MEGALASTRUM of the rhizomes; laminae 0.25-0.5 m long, basally 2-pinnate-pinnatifid, medially 1-pinnate-pinnatisect; lamina rachises pubescent abaxially; basal pinnae 3.5-12.0 cm long, equilateral or subequilateral; pinna rachises abaxially eglandular, sparsely pubescent, scaly, the hairs 0.5-1. 2 mm long, 4- or 5-celled, spreading, lax, the scales 0.5-1. 0 X 0.2-0.3 mm, dark brown, spreading, narrowly lanceolate, sparsely denticulate, not clathrate, not bullate, adaxially eglandular, sparsely pubescent, scales absent, the hairs 1.0-1.5 mm long, 5-9-celled, ascending, light brown to whitish; basal basiscopic pinnules of medial pinnae not enlarged or overlapping the lamina rachis; costules on both surfaces with indument like that of the pinna rachises; laminar tissue between veins abaxially eglandular, glabrous, adaxially glabrous; veins visible on both surfaces, abaxially subglabrous, with few hairs towards the apex, adaxially sparsely pubemlent, the hairs like those on the costules; lamina margins eglandular, ciliate, the hairs ca. 0.3 mm long, 2- or 1-celled, rarely on rocks; 600-2800 m. Selected Specimens Examined.-ECUADOR. Bolivar: Carretera f alinas-^cundo Vela, 2800 m. 1"15'S, 79°02'W, Sep 1994, Navarrete 725 (AAU). Mo™na- Santiago: along new road Mendez-Morona, Km 55-^2 MO ^ 40 S; 78“19'W], 23 Aug 1989, van der Werff S- Cudmo camino entre Baeza y Tena, 15 km al N de Tena, 1100 m, [0 59 S, 77 49 ^ 24 Jan 1984, Moran 3583 (AAU). Paslaza: Km 17 del '^“3 Uano-El Triunto. 600 m. OUSl'S, 7r30'W, 25 Feb 1994, Mamoa 12in (MO, QCNE). Zamora-Chinchipe: new road Loja to Zamora Pass, 2000 in, 4°00'S, 79^02'W, 14 Feb 1991, Moran 6r Rohrbach 5382 (AAU, “^PEmTLzonas: Bagua, 12 km E of La Peca. 1700 m, 23 67°52'3", 23 Aug 1998, Portugal et al. 246 (UCJ. Megalastrum bolivianum is distinguished by short laminae medially 1-pinnate-pinnatisect, md P‘““ uth diffembv minute long (0.5-1.2 mm) whitish hairs. Similar is M glandular pubescence between the “laminae 1-pinnate-pinnatisect eglandular). Also similar because of th differs from M. to/thnum by miform“"X and veins adaxially glabrous (vs. pubescent). AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 3 (2014) MORAN ET AL.: ANDEAN MEGALASTRUM One specimen we have identified as Megalastrum bolivianum (Qllgaard et al. 90906, AAU) is atypical by subdimorphic sterile and fertile leaves, veins glabrous adaxially, and pinna rachises abaxially with ovate scales. It might be new, but we refrain from describing it on the basis of this one specimen. 8. Megalastrum ciliatum M. Kessler & A. R. Sm., Amer. Fem J. 96:38. 2006. Type —BOLIVIA La Paz: Prov. Sud Yungas, camino Chulumani-Ocobaya, 2.5 km despues do Chulumani, 16"23'S. 67“31'W, 1600 m, 6 Jan 1992, k P. Schmit, T. Mione (r W, Zuazo 422 (holotype: UC! (barcode 1616956], isotype: LPB-n.v.). Figs. 14A, 15L-R, 18B. Rhizomes erect, scales 6.0-10.0 X 1.0-1.5 mm. spreading to ascending, linear-lanceolate, brown with margins black-bordered (at lustrous, twisted, strongly denticulate; leaves 0.5-0.8 m long; peho eb»e ^I« like those of the rhizomes but more spreading to loosely ascending, lam n . 0.7 m long, basally 3-pinnate-pinnaUfld, medially 2-pinnate-pinnatisect; lamma rachises ^thout hairs abaxially; basal pinnae 10.0-30.0 cm ‘“S; pinna rachises abaxially eglandular, long, 4-7-celled, spreading, acicular, the stales 1. 0-2.0 x 0. . . ,.®’ no“l“nZul”e%dSdly^^^^^^^^ hairs 0.7-1.0 mm long, 3-6-ceUed, spreading, the scales to ^ ^ ascending, brown, linear-lanceolate, lustrous, ^ ^ ^ j densely eglandular, ciliate, the hairs 0.4— 1.0 mm gi spreading; indusia absent; spores cnstate. Distribufion.— Bolivia, northern Argentina; 1600- o r. nniTVIA La Paz: Parque Nacional Madidi, Selected Specimens ExAM^° -®°^^^^-.4034ti2''S, 68 5 entre Carjata y Rio Yana Lomas, 1700 m, 14 iz Fuentes et al. 9013 (MO, UC). , aotail P Hair detail. Scale bars - 1 nun. A D: pinna rachis and segment. N. OUgaard et al 2219 (AAU). K-P: Isotype. Holotype. Kessler et al. 7630a (UC). E-J: Holotype, tauga AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 3 I MORAN ET AL.: ANDEAN MEGALASTRUM ARGENTINA. Jujuy: Depto. Ledesma, Parque Nacional Calilegua, Aguada del Tigre, 1600 m, 23°41'S, 64°53'W, 7 Apr 2010, Martinez Sr Prado 1893 (MCSN, NY, SP). Megalastrum ciliatum is characterized by relatively small leaves (up to 0.8 m long). Besides size, the species is further distinctive by veins adaxially with straightish hairs 0.5-1.0 mm long, laminar tissue abaxially pubescent with hairs 0.8-1.0 mm long, and pinna rachises and costules abaxially with strongly denticulate scales. o i u- This species most resembles Megalastrum nigromarginatum from Co ombia. Both have similar lamina size and cutting, and the rhizome scales are intermittently black-margined-a rare character in the genus. The species can be distinguished by characters given in the key. Megalastrum chaluw^ resembles M. marginatum and M. rupicola by lamina size and cutting but differe by the laminae between the veins abaxially pubescent. The densely P“buscent laminae may resemble that of M. pulverulentum but that ^ much larger leaves (to 4 m long), the presence of g ands pinna ™bis more strongly denticulate and often darkened apically, and echinulate spores. See Megalastrum nanum for comparison to that species. 9. Megalastrum clathratum R. C. Moran, J. Prado & *doud forest PERU. Pasco: Prov. Oxapampa, along trail Milpo ^ta. ^ qioo-3500 m 3 and puna vegetation, on rich soil, Nov 2009, H. von der Werff, R. Vdsquez M.. A Pena. L. Moteo R Avero 23175 (holotype: HOXA! [accession 0432311; isotype. MO. (barcode 6315956, 6315957). Figs. 8D, 14A, 19A-F. Rhizomes unknown; leaves up to 1.5 m long; petiole « ^nt“eL"o‘!frionrstiSy'i“^ sparsely glandular, densely and P“^®“X'hairro,2 J.6 mm long, 3-5- long, spherical, sessile, brownish or orMgis , denticulate to ciliate, celled, reddish, spreading, the scales , es, sometimes darker apically, non-bullate, ^ ’ lanceolate,’ spreading, the small ones ca, 0.2-0.6 X ca. 0.2 spreading, larger 5.0-10.0 X 0.2-0.5 mm basiscopic pinnules of ad^ially with indument like lamina rachis; costules medial pinnae not enlarged or ove pp g rachises abaxially sparsely glandular, the 3 o-5 0 X 0.2-0.3 mm long, but more sparsely scaly, the scales long, 3-5-celled, adaxially sparsely pubescent, ai • surfaces sparsely spreading, reddish; ^^^J^sile and spheri^^ glandular, glabrous, the glands ca. a ’ abaxially sparsely glandular orangish; ultimate veins on both surfaces obscure. AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 3 (2014) and sparsely pubescent, non-scaly, the hairs 0.3— 0.5 mm long, 3— 5-celled, adaxially eglandular, sparsely pubescent, the hairs 1.0-1. 8 mm long, 4-9- celled, spreading; lamina margins not glandular, ciliate, the hairs 0.2-0. 3 mm long, 2- or 3-celled, spreading; indusia absent; spores echinulate. Distribution. — ^Peru; 3100-3500 m. Selected Specimens Examined. — ^PERU. Pasco: Oxapampa, along tail Milpo to Sta. Barbara, 3100-3500 m, 10°22'53"S, 75°38'15"W, 3 Nov 2009, van der Werjf et al 23179 (MO). Megalastrum clathratum is distinguished by pinna rachis scales clathrate and reddish, and the pinna rachises and costules evenly and densely pubescent by reddish hairs 0.2-0. 6 mm long. The species has echinulate spores, a character that will separate it from all other species in the Andes except those in the “pulverulentum clade” (unpublished molecular results). The Andean members of this clade are M. adenopteris, M. fugaceum, M. pulverulentum, and M. nanum. The elevational range for this species is the highest in the genus. 10. Megalastrum crenulans (Fee) A. R. Sm. & R. C. Moran, Amer. Fern J. 77:127. 1987 [published 3 May 1988]. Aspidium crenulans Fee, Crypt. Vase. Bresil 1:139, t. 47, fig. 1. 1869. Ctenitis crenulata (Fee) Ching, Sunyatsenia 5:250. 1940. Dryopteris crenulans (Fee) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 8, 6:90. 1920. Type.— BRAZIL. Rio de Janiero: Rio de Janeiro, [22°56'S, 43°17'W], s.d., A. Glaziou 1781 (lectotype, designated by Christensen, 1920: C!, photos MICH!, MOI; isolectotypes: K! [barcode 000512192, 000512193], PI [barcode 00610618], P! [barcode 00610619], RBI-n.v.). Figs. 5C, 6C, llA-L. Dryopteris villosa (L.) Kuntze var. glandulosa Rosenst., Hedwigia 46:129. 1907. Dryopteris crenulans (Fee) C. Chr. f. glandulosa (Rosenst.) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 8, 6:91. 1920. Type. ^BRAZIL. Rio Grande do Sul: Mun. Rio Pardo, banks of Rio Cyriaco [29°59'22''S, 52°52'4m], 1906, L. C. Jurgens s.n. [Rosenstock 206] (lectotype, designated by Moran et al, BM. -000907729; S-n.v.; isolectotypes: B! [barcode 200053756], BM [b^code 000907728], GH!, HBI, Kl, MICH!, MO! [accession 1636088], t^arcode 00678938], NY! [barcode nofiiOR?^ ’ 006106231, P! (barcode 00610624], P! [barcode 00610625], UC! [barcode 441613], VT! [barcode 001519]). 0 2 ^ 1 * 0 ^™ “P ” '°"8! base scales 10.0-20.0 X "'““y ‘Oft’ ftoea'-, light brown, twisted, tortuous, sparsely denticulate; laminae ca. i m long basally 4-pinnate, ZnaXr^r'’’‘T“'*’ Smdulm mhe T "f inequilateral; pinna rachises abaxially glandular, pubescent, scaly, the glands ca. 0.1 nui long. 24:8lled, stalked. MORAN ET AL.1 ANDEAN MEGALASTRUM yellowish, the hairs 0.8-1.2 mm long. 4- or 5-celled. adcular. spreading, the scales ca. 1.0-1. 5 mm long, brown, lanceolate, subentire, not clathrate, bullate, adaxially densely pubescent, sparsely glandular, the hairs and glands similar to those of the abaxial side; medial pinnae not enlarged or overlapping the lamina rachis, costule abaxially glandular, pubescent, sparsely scaly, the glands ^ stalked, 2-celled, yellowish, the hairs 0.2-0.5 mm long, acicular, spreading, hyaline, the scales ca. 1 mm long, brown, lanceolate bullate, shiny, subentire, adaxially 'J' scales, the glands similar to those ahaxially, the . celled erect acicular; laminar tissue between veins abaxially densely glandular and puhescent, the glands stalked, 0.1 mm long, erect the hairs ca 0 1 mm long, 1-celled, erect, hyaline, acicular, adaxially moderately glandular, sparsely pubescent, ‘“^ument more dense towards the segment apex, the hairs ca. 0.2 mm long, 2-celled ^ glands similar to those abaxially; ultimate w abaxially very sparsely glandular, pubescent, the “f-" ! hairs, sometimes laminar hairs emerging through Jinus oi appear to be part of the indusium proper; spores ^ in Ltribution and ecology.-Veneznela. Brazil, and P^a^ay. m Venezuela, but to 725 m in Brazil and Paraguay (Moran el ai. 2009|. Selected Specimens Examined.— VENEZUE^. 'y^“o™^,.oro^'w of Serrania de peak. 8 km NW of-ttlement of Yuta e^ 4 to omo ^ Yutaje, ranging from slope below wet bl ^ bluff summit, 1500-1760 m, 5 41 S, 66 66 W, 4 Max * ’ . „ 21585 (MO, NY). Bolivar; Faldas del Cerro Apacar , [5“20'N, 62°12'W], 1 Jul 1946, Cardona 1590 (U6J. Megalastrum crenulans is toacto ^stinguishes the on the pinna rachises and costules. Th , ^08 (i.e., M. acrosorum. M. species from the other indusiate ones in ou . , ^ nhylogenetic results, andicola, and M. insignel. According to ““ Afrk^-Madagascan it forms a clade with the Brazilian M. f defined by blades M. lanuginosum (Willd. ex Kaulf.) Holttuin. fade >s d^ ^d costules abaxially with minute yellowish “4m from M. obundons abaxially with bullate scales. Megalastmm c ^ 20091. It differs from by the presence of an indusium (vs “ ^1^^ M. lanuginosum by petiole base scales . (vs. 0.5-0.7 mm long abaxially with hairs 0.2-0. 5 mm long an . , , ^ 5 mm; Rouhan and and 6-8-celled). and indusia 0.8-1.0 mm wide (vs. l-O-l- Moran 2011). MORAN ET AL.: ANDEAN MEGALASTRUM Megalastmm cteniloides is characterized by laminae and all pinnae, even the basal ones, equilateral. All otfte s^ments o ^ adnate for their entire width to the pinna rachises, and their ^ to truncate. The segment margins are entire, never lobed or Jto lamina mchises are pubescent and scaly, but the pinna ™*‘ses f “lally^ from glabrous to puLscent. Adaxially. the veins and “^nb °f segments are glatoous. The lamina margins are glabrous to sparsely and the less aivided biseJale. but that species differs by basal pinnae >n«q“dateral and genera ly more divided laminae. Also similar by ‘ammae med^ly 1 pmnate pinnatifisect are M. balivianum and M. Lstnles and veins on both surfaces pubescent with sfraight han^^ 1 mm long (vs. glabrous). The latter lobed, and slightly pinnae with the basal basiscopic segment enl^g , rarhises are 3.0- overlapping the lamina rachis. Also, its scales ^ bolivianum and M. 4.0 mm long and fibrillose (vs 2.0-3.0 ^ >“8). f * " fibrillosum occur on the eastern side of the Andes, wn known only from the western side. 12. Megalastrum decompositam K. C. Tambo, 12=10'S. Type.— ECUADOR. Bolivar: along road Chill b ^ ^ ^ 79"06'W1, 1700-2300 m. 19 lul M930696, 04930697), 12482 (holotype, QCNE!; isotypes, [tecod QCA-n.v., UcYlbarcode 15936661). Figs. 7M-0, 14D, 22B. Rhizomes erect; leaves up to 2.5 “ * teown, flat, dull, 1.0-2.0 mm spreading, linear to -- ^"pl,,,,atifid basally. 3- denticulate; laminae up to 1.5 m l g> P pubescent abaxially; basal pinnate-pinnatisect medially; lamina rac ^ ^ basiscopically; pinna pinnae ca, 75 cm long, wtlS Lsely scaly, the hairs rachises abaxially eglandular, densely pubem ^^^^j.^pj^jaaing, the scales 0,l-0.3 mm long, 2^-celled, sbgbtly not clathrate, 1.5-3,0 X 0.3-0.5 mm, brown, lanceolate, sp g 0.6-1.0 mm long, not bullate, adaxially densely 0 2-0.3 mm, lanceolate, 4-8-celled, spreading, the cpstules abaxially sparsely subclathrate, brown, appressed, den gpgsile, spherical, yellowish, glandular, pubescent, scaly, the glands c^ ‘ , ’i o-2 0 X 0.2-0.3 mm, linear- the hairs 0.1-0.4 mm long, 2-4-ceUed, ’ denticulate; laminar tissue lanceolate to linear, subclathrate, bro\^r P the hairs ca. 0.2 mm between veins abaxially eglandular, ^ ^^jmen, Harling & Andersson long, 2-celled, erect, adaxially puberulent or IP g^gHed, lax, tortuous, 22459) glabrous, the hairs abaxially eglandular, densely ascending; ultimate veins on both smfa 3-celled, ascending to erect, pubescent, non-scaly, the hairs ca. 0.2 • 5-0.8 mm long, 5-7-celled, adaxially eglandular, pubescent, the AMERICAN FERN JOURNAL: VOLUME 104 NUMBER ■ IT, ciliate, the hairs 0.1-0. 3 mm long, 2- or 3- celled, spreading; indusia fugacious, minute, consisting of a few hair-like scales, often covered by sporangia and apparently absent; spores cristate. Distribution. — Central and southern Ecuador, on the western side of the Andes; 1700-2300 m. Selected Specimens Examined. — ECUADOR. Bolivar: along road Chillanes-El Tambo, 1700-2300 m, [USS'S, 79°06'W], 19 Jul 1991, van der Werff et al. 12506 (MO, QCNE, UC). Loja: Celica-Zapotillo road, ca. Km 5, 2100-2200 m, [4 04'S, 80"03'W], 23 Feb 1985, Marling S' Andersson 22459 (NY). Tungur- agua: Banos, [0°23'S, 78°25'W], Jan 1892, Sodiro s.n. (NY). Megalastnim decompositum has large leaves up to 2.5 m long that are 4- pinnate-pinnatifid basally. It is one of the most dissected species in the genus, thus the specific epithet decompositum. The scales on the axes are distinctive y eing thin, subclathrate, and denticulate. Both surfaces of the laminae etween the veins are pubescent, except in one specimen [Marling & 22459). The hairs on the adaxial surfaces between the veins are S ig y ortuous and ascending, whereas those on the abaxial surfaces are mostly erect. This pubescence resembles that of M. mollicoma, a pecies 1 enng by pinna rachis scales entire, firmer, and opaque (vs. Tb 1^0 1 V. ’ subclathrate) and lamina margins with longer hairs (0.3- u.o vs. u.1-0.3 mm long). Ecol. Sist. 3(Supl. l):43-44, fig. 3A, on Rfn 7 ^ ECUADOR. Zamora-Chinchipe; along road between Zumbi 10 1 cordillera del Condor beyond Paquisha, 784^74"^mf Nangaritza bridge, 29.1 km E of Zumbi, 3^56'13"S, Katan Hannon, G. Walhert S' T. 17934751). FiJ,.2ri’oE,23A-^“4A" 7 9 _i 7 0 X ca. 1 mm long, appressed to ascending, simple or bifid- Ip twisted, sparsely denticulate, the teeth base sLles ca. 10.0 X 1.0 mm, laminae 0.5-0 q m i spreading and more sparsely denticulate, pinnate-pinnatifiH 2-pinnate to 2-pinnate-pinnatifid, medially 2- 25-0 cmLg ^baxially; basal pinnae 10.^ ular, pubescent seal v th i! ^^b^quilateral; pinna rachises abaxially egland or oppressed to Lc ndt. tT' ° ^^n long, 2- or 3-celled, strigose appressed to 2-5 X 0.2-0.5 mm, lanceolate, brown, sparsely denticulatJ ^“®^°us, twisted (but not tortuous), entire to pubescent, scales ahU^t bullate, adaxially eglandular, densely ascending to strigose habfu" b l°rig, 4- or 5-celled, acicular, not enlarged or overland basiscopic pinnules of medial pinnae ones soon becomine aHn^?^ lamina rachis, crenate to lobate, the proximal a e to the pinna rachis; costules abaxially eglandular. [ ET AL.: ANDEAN MEGALASTRUM AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 3 (2014) Rhizomes decumbent to erect, scales 15.0-20.0 X ca. 1 mm, appressed to ascending, linear-lanceolate, brown or golden, lustrous, flat to twisted, entire to sparsely denticulate; leaves up to 1.2 m long; petiole base scales ca 10.0 X 0.5 mm, spreading to loosely ascending, linear-lanceolate, brown lustrous, twisted, entire to sparsely denticulate; laminae to 0.8 m long basally 1-pinnate-pinnatisect or rarely 2-pinnate, medially 1-pinnate- pinnatisect; lamina rachises without hairs abaxially; basal pinnae 10.0- 20.0 cm long, equilateral, the segments adnate, often with the basal basiscopic pinnule enlarged and slightly more lobed on the basiscopic side; pinna rachises abaxially eglandular, sparsely pubescent (sometimes apparently without hairs), scaly, the hairs 0.1-0. 2 mm long, 1- or 2-celled, spreading, the scales 3. 0-5.0 X 0.2-0.3 mm, reddish-brown, linear- lanceolate, brown, lustrous, flat to twisted, spreading, entire to sparsely denticulate, not clathrate, not bullate, adaxially pubescent to glabrous, lacking scales, the hairs 0.3-1.0 mm long, 5-7-celled, antrorsely curved; basal basiscopic pinnules of medial pinnae often (especially in large leaves) enlarged and overlapping the lamina rachis; costules abaxially eglandular, without hairs, spmsely scaly, the scales 1.0-1. 5 X ca. 0.1 mm, otherwise like those of the pinna rachises, adaxially glabrous or sparsely pubescent, the hairs similar to those of the pinna rachises; laminar tissue between veins abaxially eglandular, glabrous, often with numerous appressed proscales, adaxially eglandular, glabrous; u mate veins abaxially glabrous, usually obscure, adaxially obscure or glabrous partially visible; lamina margins eglandular, sparsely ciliate, the hairs ca. . mm ong, 1- or 2-celled, acicular, ascending or sometimes appressed; indusia absent: spores cristate. ecotogy.-Ecuador, Peru. Bolivia, eastern side of Andes; 260-1700 m. Examined.-ECUADOR. Morona-Santiago: Morona Canton, fA ATI F Mnf M 78”08'W, 7 Aug 1993, Fay S- Fay 4014 Un-i ’ w Anangu, Rio Napo, 260-350 m, 0°31'S, 76°23'W, 8 Mar C^ntrtn Zamora-Chinchipe; Nangaritza Nang^iti^osn m Militar, mdrgen derecha de Rio MO, W^NE) ’ ^ PucXa 25 l™ NE of Tingo Maria on the road to Schunke Harie^’ ^ 4 Jul 1984, Moran 3707 (BM, NY), Jum'n: fun 1929 1400-1700 m, 11 07’S, 75=19'W, 8-12 Ang 1948 aZu US). Loreto: Fnndo chela, Sinchono, 3 Manu Rio Manu^Rf^ Parque Nacional forest on steeu mlv P’ Oxapampa, along road Chatarra-Cacazu, MORAN ET AL.: ANDEAN MEGALASTRUM BOLIVIA. Beni: Prov. Ballivian, 138 km N of Carmavi, 700 ^ 67n2'w, 20 Jun 1989. Fay Fay 2051 (MO) Cochabamba: Ch^are, El Palmar. 700 m 17°05'S. 65°32'W, 4 Sep 1996. Kessler et a!. 8126 (GOET. NY). La Paz. Prov Abel Iturralde. Parque Nacional Madidi. campamento de guardaparques sadlri “sadiri 900m. 14-10’S, 3 ,u 20 M,/— 6 Huaylla 2547 (NY). Santa Cruz: Prov. Ichilo, 4 km fW Jel C^p Macunucu, 17"44'S, 63"35'W, 27 Sep 1996. Kessler et d. 8680 (GOET). Megdastrum fibrillosum is characterized by laminae pinnatisect or 2-pinnate, pinnae equdateral, and (at least m ^ basal basiscopic segments often slighdy enlarged and o™riappmMho lam rachis. The overlapping segments are somewhat wrder "m lobed onjta basiscopic side. The species is further distinct by pinnae conspicuous dark spreading scales. The indnment on adaxLlly varies from glabrous to pubescent. Many specimens °f ^ were identified previously as M. honestun, (Baker) A. R. by Tryon and sfolze 1991). a naine here Several species resemble Megalastrum fibnll ^ ^ is M. platylobum. which differs by bolivianum and the scales sparse, ^conspicuous, and pp ^ straightish hairs differs by costules and veins adaxially wi g ‘,.11 g^^ly by wider (vs. glabLsl, and by the lamina and pinna rhizome lanceolate (not fibrillose) pinnae rachises scales smaller, 2. 5-4.5 mm long (vs. 15 ’margins elabrous or nearly so with lanceolate scales (vs. fibrillose), and (vs. ciliate). See M. molle for comparison with tha p 15. Megalastrum fimbriatum R. C. between La P^aya camp and PERU. San Martin: Prov. Mariscal Cacerej, trail » ^ Papayas camp, Rio Abiseo Nationa ^ ’ 1999770I; isotypes, AAU!, GH- Young & B. Leon 4958 (holotype, F! [b^code n.v.. USM-mv.). Figs. lOF, 14B, ^ Rhizomes erect, scales 1.5-1. 8 X ca. • sparsely denticulate, linear-lanceolate, light brown, lustrous, a ^5x2 mm, spreading to leaves up to 0.5 m long; petiole base scales up denticulate; ascending, lanceolate to linear-lanceo a , . A j gj. 3.piiinate, medially 2- laminae 20-30 cm long, basally 2:Piii^^t^P^”^f^biially; basal pinnae up to pinnate-pinnatisect; lamina rachises P^^esc , abaxially eglandular, 15 cm long, equilateral to subequilater^; P^ 4_6-celled, spreading, the densely pubescent, scaly, the hairs 0.5 • , lustrous, flat, spreading, scales up to 3 X 0.5 mm long, 1^"’ adSly eglandular, moderately entire to subentire, not clathrate, not bul a , ^ ^ ^ 5-8-celled, to sparsely pubescent, very sparsely sea y, g^tuies abaxially eglandulM, spreading, the scales like those ,i,ose abaxially, the scales like sparsely pubescent, sparsely scaly, the hairs lixe AMERICAN FERN JOURNAL: VOLUME 104 NUMBER : those of pinna rachises but shorter up to 2 mm long, adaxially sparsely pubescent, the hairs like those of the pinna rachises abaxially, scales absent; laminar tissue between veins abaxially glabrous, adaxially with scattered hairs toward the margins, the hairs 0.5-1. 0 mm long, 6-8-celled, appressed; ultimate veins on both surfaces visible, abaxially very sparsely glandular (often apparently eglandular), sparsely pubescent, the glands ca. 0.1 mm long, sessile, reddish, the hairs 0.2-0.5 mm long, 2^-celled, adaxially sparsely glandular, pubescent, scales absent, the glands ca. 0.1 mm long, sessile, reddish, the hairs ca. 1.0-1. 5 mm long, 5-8-celled, whitish, lax, slightly tortuous: laminar margins ciliate, eglandular, the hairs 0.5-1. 2 mm long, 5-9- celled, lax, spreading to ascending; indusia absent; spores cristate. Distribution. — Peru; 2280-2750 m. Specimens Examined. — PERU. Bagua: Amazonas, Third camp. Cordillera Golan SE of La Peca, 2280-2400 m, [5°38'S, 78“31'W], 30 Sep 1978, Barbour 3682 (MO). Megalastrum fimbriatum is distinctive by small laminae (20-30 cm long), pinnules mostly adnate for their entire width to the pinna rachises, pinna rachises abaxially densely pubescent and scaly, lamina margins fimbriate with long (0.8-1. 2 mm), lax cilia (thus the specific epithet fimbriata). It resembles M. peruvianum but differs by longer (0.5-1.5 vs. 0.2-0.5 mm) hairs on the veins adaxially and the tissue between the veins glabrous on both surfaces (vs. pubescent). Tryon and Stolze (1991) thought that the holotype represented a “depauperate specimen of high elevation variant” of M. pulvenilentum; however, this species does not have cristate spores and does not belong to the M. pulverulentum clade. It is amply distinct from M. pulverulentum by its small leaves (up to 0.5 m vs. up to 4 m long), less divided laminae (medially 2- pmnate-pmnatisect vs. medially 3-pinnate-pinnatisect), and pinna rachis scales not darkened distally. R. C. Moran, J. Prado & Sundue, sp. nov. Type.- BULIVIA. Santa Cruz: Prov. Florida, Canton Mairna, Parque Nacional Amboro, 7 km NNEby air from Mairana, 18°03'S, 65°55'W, 1900-2200 m, 23 f (holotype: LPB!; isotypes: AAU!, NY! [barcode 0^709511, UC! [barcode 1604363, 1604364]). Figs. 2D,F, 3G, 24B, 26 A-D, Rhizomes erect, scales 20.0-35.0 x 0.5-1.5 mm, linear, ascending, ^ 11 tortuous, sparsely to retrorsely »P>caUy w.th spreading or retrorse teeth; leaves up to 4 m long; mft- “nietimes forming a wool-like 3 ninnate ninn u ^ h^sally 3-pinnate-pinnatisect to 4-pinnate, medially MORAN ET AL.: ANDEAN MEGALASTRUM ovate to lanceolate, oppressed or spreading, light to dark brown oft™ d“ker distally, strongly denticulate, not clathrate, not bullate. adaxially ’ densely pubeLent, sparsely scaly, the hairs 0.3-0.5 mm 'o^ spreading, scales 0.5-3.0 X 0.1-0.4 mm. the apex, strongly denticulate; basal basiscopic pinnu es o j ^ent en4ed or overlapping the lamina rachis; oostules anally -thmdu^^^^^^ like that of the pinna rachises but scales larger, to 0.5 2.5 ^ ^ (smaller scales] to linear (longer scales), strongly denticulate, adaxially like the pinna rachises but hairs 2^ sSfaces celled, spreading; laminar tissue between ultimate eglandultn, glabrous or pubesce^ to hairs “5“®; mm"l4g."3^melled.®s;rigose m l»ly “ eglandular, ciliate, the hairs 0.1-1. 2 m g. g cluster of ascending; indusia minute, fugacious, Z the center of the sorus, not reddish proscales, ca. 0.1 mm long, protruding from the center easily observed in mature son; 1060-2300(-3500) m. Disfribution.— Ecuador, Peru, Bolivia, Argenui , i r m Hp Selected Specimens Examined.— ECUADOR. Carchu E^ejm ^ palacios Ceno Golondrina Hemhra, 2450 m, 0 5 7^ W’ [^059,5 78-57'Wl, s.d., 12645 (MO, QCNE). Chimborazo: in v^e Pdl^^^^g , I ..£1 Sodiro 113 (P). Loja: trails ca 5 ^ open trail banks in pastures Bosque” to Quebrada Romerillos and ;4t| /gno'W, 30 Nov 1994, and deep wet stream ravines, 2000-22 m, r-’ ^ton swampy forest on 0llgaard & Navarrete 105941 toward the river, 1850 m, NE side of Quijos, go N on the road ^ of the C g^^^ ^Y); road Baeza- 0°27'30"S, 77°53'W, 6-8 Aug 1992, Fay 7 77°53'W, 19 Jan 1992, Tena, 2 km N of Consaga, 2000-19 • Calacali-Nanegalito, 18 km 0IIgaard et al. 99556 (AAU). Pichmcha: jSO, tew disturbances, from Mitad del Mundo, inclination steep, or ‘ g. Kragelund 140 2040-2090 m, 0"01'N. 78=37'W. 20-25 Apr 199^ 13 km E of to (AAU, QCA). Zamora.Chmchipe: New road ^ ^ ^ ^g-orvl. 14 Pass, Wet forest along creek that crosses Feb 1991, Moran & Rohrbach 5380 p.p- ’ralouina, 1600 m, 12"52"01"S, PERU. Cusco; Distr. Huayopata, Huyr , 4^ Urubamba; Machu 72°32'46"W, 26 Jun 2003, Bonino et a . gayacmarca and Aobamba Picchu, on a slope, 5 km N of Ae union of ,,97 (MO). Rivers, 2160 m, [13^09'S, 72^32 W , 14 A una quebrada con aguas que BOLIVIA. Chuquisaca: Sud Cinti cercanms de^^ 64^.31'46"W, 11 Oct 2000, desembocan al Rio Albomiyoj, 230 antigua carretera Cochabam a Uiullyetal. 101 (MO). Cochabamba. 130 Villa Tunari. 2000 m,17°08'S, 65 3® ^ Natural de Manejo Integrado UC). La Paz; Prov. Franz Tamayo. Area Na AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 3 (2014) Apolobamba, 2372 m, [14°39'S, 69°14'W], 10 Jun 2008, Huaylla et al. 2655 (MO). Santa Cruz: Manuel M. Caballero, bosque hiperhumedo de Ceja de Monte, Colecta en la comunidad El Locotal sobre el camino a San Mateo a 10 km del cruce El Empalme, 2364 m, 17°47'43"S, 64°43'15"W, 17 Jun 2003, Nunez & Huaylla 178 (NY, UC). Tarija; Arce, Mpio. Padyaca, Reserva Nacional de Flora y Fauna Tariquia, 1660 m, 22°00'S, 64°29'W, 14 Nov 2004, Serrano et al. 5225 (GOET, MO); idem, bajando de La Cumbre hacia Potreros, una sola planta, ladera con Rio permanente, 1706 m, 22°00'20.5"S, 64°31'59''W, 13 Dec 2004, Huaylla et al. 1554 (MO, NY). ARGENTINA. Jujuy; Depto. Ledesma, Parque Nacional Galilegua, RP 83, Aguada del Tigre, 1600 m, 23°41'S, 64°53'W, 7 Apr 2010, Martinez & Prado 1894 (MCSN, NY, SP). Tucuman: Monteros, Quebrada Pueblo Viejo, [27°10'S, 65°30'W], Jan 1965, de la Sota 4069 (US). Megalastrum fugaceum resembles M. pulverulentum by its large leaves (up to 4 m long) and scales of the piima rachises and costules usually darkened apically and strongly denticulate. Megalastrum pulverulentum differs by laminae with longer (1. 0-2.0 mm) and denser hairs; its lamina and pinna rachises are conspicuously pubescent to naked eye. In contrast, M. fugaceum is a less pubescent version. Its hairs are only 0.4-0.8 mm long and relatively sparse, not conspicuously pubescent to the naked eye. Whereas M. pulverulentum is pubescent between the veins abaxially, M. fugaceum is glabrous except in a few instances. In those instances, the hairs are much shorter (0.2-0.4 mm long) than those in M. pulverulentum. There is considerable variation in the occurrence of hairs in M. fugaceum. The type has sparse whitish hairs on both surfaces of the veins, whereas other plants are glabrous. 17. Megalastrum galapagense R. C. Moran, J. Prado & Sundue, sp. nov. Type.— ECUADOR. Galapagos Islands: Isla Isabela, SW slope of Volcano Cerro Azul, [0°38'S, 90°22'W1, 800 m, Aug 1975. H. H. van der Werff 2244 (holotype: NY! [barcode 010933221; isotypes: MO-n.v., U! [barcode 02495301). Figs. 14C, 20O-P, 27C. Rhizomes erect, the scales 20.0-30.0 X 0.6-1.1 mm, appressed to ascending, Imear-l^ceolate, golden brown, lustous or dull, flat to tortuous, the surfaces smooth (not setulose), denticulate, the teeth simple or bifid; leaves up to 2.5 m ong; petiole base scales like those of the rhizomes hut to 15.0 X 0.7 nun, spreading to loosely ascending; laminae up to 1.5 m long, basally and medially -pinnate-pmnatisect; lamina rachises pubescent abaxially; basal pinnae equilateral; pinna rachises abaxially eglandular, sparsely LcuW ™ 3-4-celled, spreading, lustrous ^ ^ ^ 0.1-0.3 mm, filiform to linear-lanceolate, brown, denticulate, not clathrate. not bullate. adaxially accular, tortuous, catenate; basal basiscopic pinnules of medial pinnae not MORAN ET AL.: ANDEAN MEGALASTRUM enlarged or overlapping the lamina rachis; coshdes ab^ally oglandukr. sparsely pubescent, sparsely scaly, the bans 0.3-O.5 mm long, ' eLt, aclLlar, hyaline, the scales ca. 0.5-1.0 X 0.1 mm, the pinna rachis, adaxially eglandnlar. f "\rJa^ absent, the hairs 0.3-0.5 mm long, 3- or 4-celled, edandular ascending, tortuous; laminar tissue between ^ acicular, erect, sparsely pubescent, the hairs 0.2-0.3 mm long, 2- ’iomie on both hyaline adaxially eglandnlar. glabrous; ultimate veins v site on tath surfaces, abaxially eglandnlar. sparsely pubescent, the ha. s s^mito to tose of the abaxial pinna costae; adaxially eglan u g acicular, eglandular, ciliate, the hairs 0.3 mm long, ascending; indusia absent; spores cristate. r-oianaont; Archinelaeo; Distribution and ecoiogy.-Isabella Island in the Galapagos Arcti.pe.ag Miconia belt vegetation; 800 m. S™s ExaMiNEn.-ECUADOR. Galapagos: I-bolla Island Volcano Cerro Aaul, 800 m, 10"01'S, 91“12'W1, Aug 1975. von dev w m Megalastrum galapagense and ^ Ihtpe,' bm M. galapagense Galapagos Islands. They are iaminae more divided differs by longer leaves, up to 2.5 m lo g I --notisect to (one specimen) (2-pinnate-pinnatisect medi^ly ^a^hises and costules abaxially 2-pinnate-pinnatifid medially), and P pnlaDasense the hairs on the sparsely pubescent (vs. densely ^ if M pleiosoros (0.4-0.5 mm abaxial surfaces of the laminae are shorter than m m p long, 3^-celled, vs. 0.6-1.0 mm Galapagos Islands. Megalas- The specific epithet „ia,S. whereas M. pleiosoros occurs trum galapagense occurs only on isaoeii on Isabella and others. 18. Megalastrum hirsutosetosuro f™urond pryopteris hirsatosetosa Fern J. 77:128. 1987 IP“bhshed 3 M y 1 (Hieron.) Hieron., Hedwigia 46;343, tab. 6. 16- “ t„e.-ECUADOR. Tungur- Lellinger, Proc. Biol. Soc. Wash. 89.709. . Ptntuc, s.d., ahiia?- nlatfvaii above AllpaV^cu between ^ i Q?n! B! [barcode 20 i?; plateau above Allpayacu r^stensen, 1920: B! [barcode 20 smbel 903 (lectotype, [barcode 0009077511). Figs. 0057934], photo BM; isolectotype: BM! (fragm.J t 24C, 28K-P, 29A. ^ lanceolate, golden brown. Rhizomes erect, scales ca. 10.0 X ca^ . _ Jong; petiole base scales ca. lustrous, tortuous, denticulate; brown lustrous, flat to twiste , 10.0 X ca. 1 mm. lanceolate, f ^^•i4“fiif2!™nnate to 2-plmiate-pinna^^ denticulate; laminae 0.5-0.8 m long, rachises pubescent abaxial y, sect, medially 1-pinnate-pinnatisect; ^ inequilateral, sometimes basal pinnae 8.M5^0 ‘“S’ ^dnto. pubescent, sparseiy equilateral; pinna rachises abaxially g AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 3 (2014) hairs ca. 1.5 mm long, 6-8-celled, acicular, spreading to erect, whitish, the scales ca. 1 X 0.1 mm, oppressed to spreading, filiform, brown, tortuous, denticulate, not clathrate, not bullate, adaxially eglandular, densely pilose, without scales, the hairs similar to those of the pinna rachises abaxially; basal basiscopic pinnules of medial pinnae not enlarged or overlapping the lamina rachis; costules abaxially eglandular, pubescent, without scales, the hairs ca. 1.5 mm long, 7-10-celled, oppressed to spreading, acicular, hyaline, adaxially eglandular, sparsely pilose, scales absent, the hairs like those of the adaxial pinna rachises; laminar tissue between veins both surfaces eglandular, pubescent, the hairs similar to those of the abaxial pinna rachises and costules: ultimate veins obscure and pilose on both surfaces, the hairs similar to those of the laminar tissue; lamina margins eglandular, ciliate, the hairs ca. 1.0 mm long, 5— 7-celled, acicular, spreading; indusia absent; spores cristate. Distribution and ecology. — Colombia, Ecuador, and Peru, eastern slopes of the Andes; 450-1200(-2250l m. Selected Specimens Examined.— COLOMBIA. Choc6-El Valle: at and on both sides of the principal ridge of the Serrania de Los Paraguas, along the trail from El Cairo to Rio Blanco, ca. 8 km SW of El Cairo, 2200-2250 m, 4°12'S, 75°48'W, 20 March 1971, Lellinger & de la Sota 840 (US). ECUADOR. Carchi: Border area between Prov. Carchi and Esmeraldas, about 7 km past Lita on road Lita-Alto Tambo, 550 m, [0'’50'N, 78 28'W], 27 Jun 1991, van der Wepfetal. 12094 (MO, QCA, QCNE). Esmeraldas: Pachicutza, at Escuela Fiscomisional Cardinal Dopfner”, Km 140 on road Loja-Gualaquiza, Tropical rain forest with cleared areas Rio Zamora and along the road, 900- 78“34'W, 26-27 Apr 1973, Holm-Nielsen et al 4502 (AAU, MU, NYJ. Napo: Holhn-Loreto road, 32 Km mark, 3-4 km S of road, path to pem^ent sampling plot, 1200 m, 0°35'S, 77°25'W, 25 Jan 1991, Moran Rohrbach 5160 (AAU, MO, NY). Pastaza: Rio Pastaza, between Banos and Mera, 1219 m [r23'S, 78°03'W], 4 Jul 1905, Tate 650 (US). Zamora- Chmchipe: Cordillera del Condor, Rio Wawaime watershed, tributary of Rio EcuaCorriente mine camp, 1300 m, 3°34'43"S, 26 07'YV, 26 Oct 2006, van der Werff et al. 21715 (MO, NY); Miazi, at S'w Chumbiriatza with Rio Nangaritza, 1200-950 m, 4°19'S, 0iigaard et al. 99268 (AAU, QCA). dp Ignacio, Distr. San Jose de Lourdes, localidad 5 42'04"S, 77“53'06"W, Feb 2002, & Smith 26037 fNY^mi Nicolas, 1100 m, 4-5 Jul 1929, Killip co^Tcl^vn r ‘“‘h densely and isUnnn^M bv llr “ f“dher rachis scales filfform cfl^^tf 2-pinate-pinnatisect and by » • 1 X 0.1 mm. Megalastnim platylobum is similar m MORAN ET AL.: ANDEAN MEGALASTRUM lamina dissection and size but differs by pinna rachises abaxidly densely pubescent by hairs of mixed lengths, all of which are shorter than those found in M. birsutosetosum. 19. Megalastrum insigne R. C. Moran, J. Prado & Sundue, »P- ”7- J™ - ECUADOR. Napo: Cantbn Quijos, south of Quijos |ust east of _ through pastures and up into the jungle, now logged tertestnd^ 0"28.5'S, 77»53.5'W, 2100 m, 4 Ang 1992, A. Fay S' ^ Fay 3827 (holot^e. NY! [barcode 010538651; isotypes: AAU! 3-sheets, MO [b^ 05083493], QCA-n.v., QCNE-n.v.). Figs. 2B, 4M-X, 6E, 24D. Rhizomes erect, the scales 10.0-20.0 X l ')-2.0mm appressed to ascending^ lanceolate, hrown to dark brown, lustrous, flat densely d®«dc'ikte>ajY P to 1.5 m long; petiole base scales like those of ^^^CSt loosely ascending; laminae 0.5-O.7 m abaxLlly; basal medially 2-pinnate-pinnatisect; lamina rachis p abaxiallv pinnae\5.cf-25.0 c^ long, strongly j-rl'^r^rng s"pt“ and' glandular, densely pilose, sparsely scaly, "“h _ ^ fcicular, spreading, sessile glands, the hairs 1.0-1,5 mm long, 5-8-ce led, acicu ^ hyaline, whitish to tei^ugineous or tan, t e sea e ^ ^ lanceolate, spreading, brown, not flaccid, shi y, opales densely pilose, not clathrate'^ no. bullate, adaxially eglanduto w.4om ^ the hairs 0.7-1.5 mm long. 5-12-celled, spreading the lamina basiscopic pinnules of medial pinnae not en arge glands rachis; costules abaxially sparsely glandular, pi ’ ^ 2.0 X 0.3 nun, like and hairs like those of the pinna rachises, e sc densely pilose, the those of the pinna rachises, adaxially sparse y g laminar tissue glands and hairs like those of the pinna J^3ely pubescent, between veins on both surfaces glandular, .g^sed, some of the the hairs 0.8-1.2 mm long, 5--8-celled,acicu ar ere ^ hairs gland-tipped, 0.2-0.4 mm l^girs and glands the 0.1 mm long, spherical, yellowish; ^^ins. ahaxially the veins same as those on the laminar tissue betwe gfilandular, ciliate, the partially visible, adaxially obscure; lamma mar^ 6^ ascending; indusia hairs 0.4-41.8 mm long. 3-7-celled. ^""'^^ “^^^^oncolorous, brown, glandular, present, ca. 1 mm wide, conspicuous, circul , ^ ^ long, pubescent, the glands sessile or short-sta e , ^ long, 3-7-celled, the stalked glands 0.2 mm long, the ai acicular, erect, hyaline; spores cristate. , Poni, both sides of the Distribution and ecology. — Colombia, Andes; 1300-2400 m. ^ Calderas, cerca de Selected Specimens Examined.— COLOMBIA. lUS). Caldas: Pueblo Cocorna,[5 58'N.75 06'Wl,Janl953,Dam^^ 1700-1900 m, Rico [Neiral, Cordillera Occidental, (US). Cauca: Montana [5 08'S. 75 3rWl, 17 Feb 1946, von Sneidem 5236 p P AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 3 (2014) del Oro, [5°08'N, 75°31'W], s.d., Lehmann 7416 (K). Valle: Finca la Pradera, 6 km SW of El Cairo on the trail to Rio Blanco, between El Brillante and Boqueron, 2150-2200 m, [4°12'N, 75°48'W], 27 Mar 1971, Lellinger 8r de la Sota 815 (US). ECUADOR. Loja: trails ca 5 km ENE of San Pedro de Vilcabamba, from “El Bosque” to Quebrada Romerillos and Banderilla, 2000-2200 m, 4°14'S, 79°10'W, 30 Nov 1994, QUgaard S' Navarrete 105938 (AAU). Napo: forest ridge just S of Baeza, 1950-2100 m, 0°28'S, 77°53'W, 31 Mar 1992, 011gaard 99904 (AAU, QCA, QCNE); road Baeza-Tena, 2 km of Cosaga, 1900-2000 m, 0°34'S, 77°53'W, 17 Jan 1992, QUgaard et al 99554 (AAU); Valley of Rio Oyacachi, ca 10 km along road W of El Chaco, bail along river to Rio San Juan Grande, 1760-1800 m, 0°17'S, 77°51'W, 12 Mar 1996, QUgaard & Navarrete 1630 (AAU). Pastaza: Montana de Canelos, 1600 m, [1°30'S, 78°03'Wl, Aug 1860, Spruce 5295 p.p. (BM, K, NY, P). Pichincha: Canton Quito, Rio Guajalito Reserve, 10 km W of Chiriboga, Km 59 of old road Quito-Santo Domingo, 1900 m, 0°14'S, 78°48'W, 10 Jul 1991, Fay & Fay 3370 (AAU, MO, NY, QCA, QCNE); Cerro Antisana, primary montane forest 2 mi. SE of Borja, 1737 m, 0°30'S, 78°W, 3 Aug 1960, Grubb et al. 1195 (BM, NY, US). Santiago-Zamora: high wooded slopes, on W side of Rio Valladolid, above Valladolid, 2100- 2400 m, [4°33'S, 79°03'W1, 15 Oct 1943, Steyermark 54713 (F, US). Tungurahua: San Antonio, eastern slope of Tungurahua Volcano, 1737 m, [1°28'1"S, 78°26'30"W1, 1924, Tate 573 (US). Zamora-Chinchipe: Parque Nacional Podocarpus, Quebrada Rio San Francisco, at new road Loja-Zamora, 2040-2250 m, 3“58'S, 79°05'W, 23 Jun 1988, QUgaard 74966 (AAU, QCA). PERU. Amazonas: Bongara, Hills 1-5 km. S-SE (150°) of Yambrasbamba, 2100-2400 m, [5°45'S, 77°54'W], 25 Jun 1962, Wurdack 1030 (GH, NY, US). Cajamarca: San Ignacio, San Jose de Lourdes, Gamana, 2000-2200 m, 5 00'00''S, 78°55'00"W, 20 Mar 1997, Campos & Corrales 3593 (GH, MO, NY). Cusco: La Convencion, Dist. Huayopata, Amaybamba, Quinsapun- m 12°59'18"S, 72°30'06"W, 22 Nov 2003, Bonino et al. 1104 p.p. [10°06'S, 75°09'W1, Aug 1947, Soukup M A oonn ^^iscal Caceres, alrededores de la m. 17“33'S, 76«47'W1, 25 Jun 1995, Quipuscoa et al. 11 r^,ah ' Abajo de La Morada, cerca del Ri'o " ^ characterized by the presence of conspicuous s^ai« Hen '')' *'“S 0,3-l.o mm long, and laminae with both (remarkable otstldi^^^^^^^^^ The specific epithet in«^« IpavA*: ^ deuse, conspicuous indument of the aminae -confused with M. andicola but differs by cXdtlHndustTa"? n ‘“T more consnimm ' t. * 3 7-celled). To the naked eye, the denser, acri:ZZ7M.!:^r distinguishes M. inst^e from the similar M, MORAN ET AL.: ANDEAN MEGALASTRUM 20. Megalastrum marginatum M. Kessler & A. R. Sm., Amer. Fern J. 96:40, fig. 2G-J. 2006. Type.— BOLIVIA. La Paz: Prov. Nor Yungas, Ceno 16”01'S, 67°52'W, 27 Nov 1998, A. Portugal 538 (holotype: UC. [barcode 1736806]; isotype: LPB-n.v.). Figs. 9A-E, lOA, 24A. Rhizomes erect, scales 5.0-12.0 X 0.2-0.6 nun, to ascending, brown to golden brown, lustrous, twisted simple or sometimes bifid, sometimes retrorse; leaves J" ' . base scales like those of the rhizomes but spreading to f; white, the scales 1.5-3.0 X 0.2-0.8 j’Xn’elv pubes- denUculate, not clathrate, Cg. 4-7-celled, spreading, of medial pinnae not enlarged or ^ the hairs like those of abaxially eglandular, sparsely pubescent, sp^s y J’, . ^ ^ smaller 0.5- the pinna rachises, the scales like those “f *e pinna m b e b 0.8 X 0.2-0.3 mm, adaxially eglandular. sparsely both the hairs 0.5-1.0 mm long, 4-6-celled; laminar surfaces, surfaces eglandular, glabrous; ultimate veins 0,2-0.4 mm abaxially eglandular, glabrous to adaxially pubescent, the long, 2- or 3-celled, spreading. ascending, straightish. hairs 0.7-1.0 mm long, 3- or ^ 0,3-0.5 mm long. 2- or 3- whitish; lamina margins eglandulm. ^ spores cristate, celled, straightish, whitish, ascending; indusia abse P Distribution. — Bolivia; 400-1900 m. , ann m r,rATTA7TA rarrasco: Cochabamba, 400 m, Selected Specimens Examined.— BOLIVIA. fTjn Cochabamba: Prov. 17°17'35"S, 64°52'87"W, 19 Jan 2000, fi5°29'35"W, 2 Sep 2003, Tiraque, Km 18 al Palmar, 900 m, 17 Varoue Nacional Madidi, Zabalaga 16 (NY). La Paz: Ab^l to^Jde. ^ gan Juan de campamento de guardaparques Sadir 1 Jimenez S' Huaylla Uchupiamonas), 950 m, 16°10'S, 67 5 - Ppcollo, Estacion Bioldgica 2518 (NY, UC); Prov. Nor Yungas, 24 Jul 2001, Bach 1234 Tunquiri, Homuni Bajo, 1900 m, 16°42 , ^ 15°32'S, 67"18'W, (UC); Prov. Sud YungL, Sapecho. Coloma Tupiza, 735 m. 29 Oct 1997, Kromer et al. 100 (GOET, Ubj. . r bv 0.5-17 mm long, straight, Megalastrum marginatum is bises costules, and veins. The whitish hairs on both surfaces of the lone straight hairs are always density of the hairs varies, but at le^t “ differs by the lack of present. The species most resembles M. P , and the pinna rachis scales hairs. In both species the lamina cutting ’ are similar by being lanceolate and shmy AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 3 I Moran and Prado (2010) cited Megalastrum longipilosum A. Rojas from Ecuador and Bolivia. The Bolivian report was based on a misidentified specimen of M. marginatum. The Ecuadorian report was based on two specimens {011gaard et al. 2247, Ankersen & Kragelund 211, both AAU, QCA) that represent a new species, M. obtusum, described below. Megalastrum longipilosum is endemic to Costa Rica and Panama. 21. Megalastrum martinicense (Spreng.) R. C. Moran, J. Prado & Labiak, Brittonia 61:284. 2009. Alsophila martinicensis Spreng., Neue Entdeck. Pflanzenk. 3:7. 1822. Phegopteris martinicensis (Spreng.) Fourn., Mex. PI. 1:90. 1872. Type.— MARTINIQUE, s.d., F. Kohaut s.n. (Sieber Syn. Fil. no. 162) (lectotype, designated by Proctor, 1989: L-n.v., photos MICH!, NYI, US!; isolectotypes: K!, P! [barcode 00600601]). Figs. 13F, 24A, 30A-D. Phegopteris epierioides Fee, Mem. Fong., 5. Gen. Filic.:248. 1850-52. Type. — CUBA, s.d., /. /. Linden “282 or 412” (holotype: P! [barcode 00610891]). Aspidium araguata Moritz ex Reichardt, Denkschr. Ak. Wien 17(2):38, tab. 2, figs. 28-31. 1859. Type.— VENEZUELA. Aragua: Colonia Tovar, [10°25'N, 67°16'W], 1844, /. W. K. Moritz 202, pro parte [mixed with M. subincisum] (lectotype, here designated: P! [barcode 610894], P! [barcode 610895]; isolectotypes: F!, GH!, P! [barcode 00600403, 00600404], US [barcode 00067054]). Rhizomes erect, scales 15.0-22.0 X 0.3-0.5 cm, linear-lanceolate, light brown, lustrous, denticulate; leaves 1.0-3. 0 m long; petiole base scales like those of the rhizomes but more spreading to loosely ascending; laminae up to 2 m long, basally 3-pinnate-pinnatiseGt, medially 2-pinnate-pinnisect; lamina rachises without hairs abaxially; basal pinnae ca. 0.5 m long, inequilateral; pinna rachises abaxially eglandular, without hairs, scaly, the scales 1.5-3.0 X 0. 1^.2 mm long, linear-filiform, tortuous, brown, denticulate, not clathrate, not bullate. adaxially eglandular, densely pubescent, sparsely scaly, the hairs 1.0-1. 2 mrn long 4-6-celled, the scales like those abaxially; basal basiscopic pinnules of medial pinnae not enlarged or overlapping the lamina rachis; costules abaxially eglandular, sparsely pubescent, sparsely scaly, the hairs .3-^.8 mm long 2-5-celled, acicular, erect to spreading, the scales like those on e pinna rachises abaxially but smaller, 1.5-2.5 X 0.1-0.2 mm, linear to mem-lanceolate, parallel sided, scarcely enlarged basally, adaxially densely pubescent, scalw absent, the hairs 0.5-l,o mm long, 4- or 5-celled, substrigose; InTtb r r.*™ glabrous; ultimate veins visible Ls ^ ogl^dular, sparsely pubescent, scales absent, the suholahmni:- 1 ^ ” 3-celled, adaxially eglandular, glabrous to 4 celled snm ciliate, the hairs 0.2-0.5 mm long, 2- 4-celled spreading to substrigose; indusia absent. PuertfmcrM‘’rt Jamaica, Haiti, Dominican Republic, Puerto Rrco, Martinique, St. Vincent, St. Lucia, Dominica, Saba. Trinidad, MORAN ET AL.: ANDEAN MEGALASTRUM northern Venezuela, Colombia; 1200-1860 m in Colombia and Venezuela; in the West Indies, 0-1600 m (Moran et al 2009b). Selected Specimens Examined.-VENEZUELA. faldas que miran al norte, a lo largo del camino 'N R7^15'W 16 lul hacia la represa. al snr de El Consejo, 1350-1400 m, 10 1 N 67 5 W^16 1979, Steyemmric ^ Stoddar 118183 (GH. Dmtn.o J to ^ 1852, Triana 615 (BM). Tolima; Mariquita, Combaima, 1200 m. 74°54'W], Jan 1843, Linden 1020 p.p. (B, P)- Megalastmm martinicense is “^^cales^^h^^^ al '«ast without hairs and with dark, ‘ort"™®;, _ “‘es ereatly resembles M. apically, are usually only 1-3 cells wide. P , costules golden subincLm, which differs by scales of the pinna "sjnto^m brown, lanceolate to linear-lanceolate, amguatam [Moritz 202] is The type number of the synonym Aspidium « P-006004021 mixed. Specimens of this number at K an represent M. subincisum. 22. Megalastmm microsorum (Kuntze) ^tolze, Field 1991. Nephrodium microsorum ^ook., bp. mi- ^ ^ Endl. 1833. Dryopteris microsora ^ leptosora C. Chr., Index nov. for Nephrodium microsorum Hook. Megalas- Fil. 274. 1905, nom. supetfl. for Nephrt^tummj^ ^ trum leptosorum (C. Chr.) A. R. Sm. ' fTuneurahua?]: at the foot 1987 [published 3 May 19881. Type.--EC^^^ (lectotype, of Mt. Chimborazo, [1°28 S, ^ ri ’de 000200224], photos R, designated by Tryon and Stolze, 199 . ^ ^537]^ pigs. 5A, 25A-D, 27B, GHlfisolectotype: BM! (fragm), [barcode 000921537J1 Rhizomes erect, scales 10.0-20,0 ‘’;^;l“ns'd”nsX''denticulate. the lanceolate, gold-brown, flat to tortuous, ^ uke those of the rhizomes teeth bifid; leaves 0,8-1.2 m long; pehole ^ basally 2-pinnate, but spreading to loosely ascending; lami^eO.^^ p„besce^ medially 1-pinnate-pinnatisect to 2 p equilateral; pinna rac abaxially; bLal pinnae 10.0-20 0 ™ evV pnbo™*®*’ „ abaxially eglandular, glabrous to densely and eve hairs (when present) 0.2-0.5 mm long, ^ guform, brown, ‘ortuous, acicular, hyaline, the scales ca. 3 X c . ^ bullate, adaxially egl ascending, lustrous, denticulate, not clathrate, n 168 AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 3 (2014) ular, densely pubescent, without scales, the hairs 0.5-0. 8 mm long, 4-6-celled, light brown, ascending, antrorsely strigose, acicular; basal basiscopic pinnules of medial pinnae not enlarged or overlapping the lamina rachis; costules abaxially eglandular, glabrous or sparsely pubescent, scales absent, the hairs (when present) 0.1-0. 5 mm long, 1-3-celled, like those of the pinna rachises, adaxially eglandular, glabrous to pubescent, scales absent, the hairs similar to those abaxially: laminar tissue between veins abaxially eglandular, glabrous or puberulent, the hairs (when present) 0.1-0. 2 mm long, 1- or 2-celled, acicular, erect, hyaline, adaxially eglandular, glabrous; ultimate veins on both surfaces visible, eglandular, glabrous; lamina margins eglandular, ciliate, the hairs 0.1- 0.2 mm long, 1- or 2-celled, acicular, appressed to ascending; indusia absent; spores cristate. Distribution. — Ecuador, Peru; humid secondary forests, fog forests; 600- 1300(-2800) m. Selected Specimens Examined.— ECUADOR. Cotopaxi: Tenefuerste, Rio Pilalo, km 52-53, Quevado, Latacunga, 750-1300 m, [l°02'S, 79°27'W], 7 Feb 1982, Dodson & Gentry 12280 (MO, QCNE). Guayas; Cordillera Chongon-Colonche, Bosque Protector Loma Alta, 600 m, 0°48'S, 80°47'W, 21 Dec 1996, Cornejo & Bonifaz 5459 (AAU). Loja: Vic. of Orianga, 1100 m, 3°52'S, 79°51'W, 5 Mar 1990, Madsen 86916 (AAU). Manabi: Cordillera de Congon, Parque Nacional Machalilla, S of San Sebastian, 500-600 m, [01°35'S, 80°41'W1 24 Mar 1993, Qllgaard et al 100773 (QCA). PERU. Cajamarca: Prov. Santa Cruz, ruta Chorro Blanco-Monteseco, 1750 m, [6^51 'S, 79°06'W], 21 Jan 1996, Leiva et al. 1759 (F). Junm: Paucartambo, 2800 m, [13 18'S, 71^35'Wl, 24 July 1961, Woytkowski 6742 (US). Megalastrum microsorum is characterized by laminae medially 1-pinnate- pinnatisect to 2-pinnate, basal pinnae equilateral, and pinna rachises with filiform tortuous scales. It is extremely variable as to the pubescence of the laminae abaxially, varying from subglabrous (e.g., Cornejo Sr Bonifaz 5459) to densely and evenly pubescent (type). Two species greatly resemble Megalastrum microsorum in division of the laminae. The first, M. platylobum, differs only by basal pinnae inequilateral (i.e prolonged basiscopically). The second, M. pleiosoros, is endemic to the Galapagos Islands. Megalastrum microsorum has similar filiform rachis scales as in M. pleiosoros, but differs by longer hairs (0.6-1.5 mm) on all parts of the niisee/ZuD, R. C, Moran. J. Prado & Sundae, sp. nov. Type.- ECUADOR. Zamora-Chinchipe: Canton Zamora, within 3 km of the town of lOOO 7 Sep 1994, Fay L. Fay 4382 mn*41912i [barcode 05034342], NY! [barcode 01053912], QCA?-n.v.). Figs. 22E, 31R-Z, 32A. mzomes unknoYm; leaves up to 2 m long; petiole base scales 15.0-25.0 X 1.0-1.2 mm, spreading to ascending, linear-lanceolate, brown, lustrous. MORAN ET . AL.; ANDEAN MEGALASTRUM twisted or flat, sparsely denticulate; laminae pinnatisect, medially 2-pinnate-pinna ^ rachises abaxially ally; basal pinnae 25.0-40.0 cm long, ^ 1 2-6-celled. eglandnlar, pubescent, spmsely scaly ae^ha^ 0.2^^. spreading, the scales 3.0-5. 0 • • sparsely denticulate spreading, linear-lanceolate, light tow , i„_j„iar densely pubescent, apically. not clathrate. not buUale. adaxially egtoduto, the haL 0.2-0.8 mm long, abaxially pinnae not enlarged or overlapping 0.1-11 mm long, 1-6- eglandular, densely 2 ’linear-lanceolate, brown, entire, celled, the scales 0.6-0.7 X gg scales absent; laminar adaxially with hairs like those of P^^^j^^^^gjately glandular, puberulent, tissue between veins abaxially sparse y onherical grading into gland- the glands ca. 0.1 mm long, sessile, emct. acicnlar, tipped hairs, the hairs ca. 0.1 mm long, ^Q^eon the abaxial surfaces, adaxially sparsely puberulent, the hau ^ sparsely glandular and ultimate veins on both surfaces with gland-tipped pubescent, the glands spherical, ye o ^ adaSally sparsely glandular to hairs, the hairs ca. 0.2 mm long, 1- or ’ iiowish the hairs 0.5-1.0 mm eglandular, pubescent, the glands eglandular, densely long, 4-6-celled, appressed to asLnding; indusia minute, ciliate, the hairs 0.2-0.3 mm long, 3 reddish proscales, ca. 0.1 mm fugacious, apparently consisting ® not easily observed in mature long, protruding from the center of t sori; spores cristate. Pnlivia on both sides of the Andes, DisWbubon.-Colombia. Ecuador, Peru, Bolivia, 550 — 2300 m. . (]]aldas Vereda Selected Specimens Examined.— COLOMBO. May 1984 Albert de La Cara. Finca La Oculta. 1950 m, 6 05 N, « M ^ 1 Escobare(al,4476(hni);Mpio. Salgar. ^ [5 54 N Regada, costado izquierdo de la ‘I'J® „ (i^yj Valle: Ansermanuevm 75"55'W1, 16 Jan 1988. ^'■b«f""'““„uevo. 1850-1875 m, [4 47 N. Josh del Palmar rd., ca. 30-37 km W 76°02'W), 13 May 1984, Lufeyn el al. 103 I Macas, 1100 m, 2 20 S, ECUADOR. Morona-Santiago: (a^U.MO, NY. UCJ. 2amo«- 78"08'W, 7 Nov 1993, Fay & ^us GuarderiaenRioBombu^ . Chinchipe: Zamora, Parque Nacion^ ^ 970 m, 103-46'S, 78-’37'Wl. 23 Jan 1995. P ' [6=27'S. 76 24 Wl. A g PERU. San Martin: Ml. Campana. near 1856, Spruce 4718 (PJ. ' ’ AMERICAN FERN JOURNAL: VOLUME 104 NUMBER : Megalastrum miscellum is characterized by pinna rachises and costules abaxially with linear-lanceolate mostly entire scales and hairs of decidedly mixed sizes (thus the specific epithet), lamina tissue between the veins pubescent, and veins adaxially with hairs 0.5-1.0 mm long. Megalastrum praetermissum is similar by narrow scales, minutely pubescent abaxial surfaces, and (often) long conspicuous hairs adaxially, but it differs from M. miscellum by pinna rachises abaxially without hairs and more strongly denticulate scales. 24. Megalastrum molle A. R. Sm., Novon 16:426, fig. 18. 2006, as “mollis. Type.— PERU. Amazonas: Distr. Bagua, along road from Chiriaco toward Bagua, [5 38'S, 78"32'W1, 750 m, 21 Mar 2001, H. van der Werjf, R. Vasquez Sr B. Gray 16300 (holotype: MO! [barcode 5675669]; isotypes: F! [accession 2251280], NY! [barcode 01093321], UC! [barcode 1777976]). Figs. 23M-Q, 24B, 29D. Rhizomes erect, scales 10.0-15.0 X ca. 0.5 mm, spreading to ascending, linear- lanceolate, brown, lustrous, twisted, subentire to sparsely denticulate; leaves 0.5-0. 7 m long; petiole base scales like those of the rhizomes but smaller, 5.0- 10.0 X ca. 0.5 mm, more spreading; laminae 25-50 cm long, basally 2-pinnate to 2-pinnate-pinnatifid, medially 1-pinnate-pinnatisect; lamina rachises glabrous abaxially; basal pinnae 7.0-9.0 cm long, slightly inequilateral; pinna rachises abaxially glandular, conspicuously pubescent, sparsely scaly, the glands short- stalked, ca. 0.1 mm long, 2-celled, yellowish, the hairs 1. 0-2.0 mm long, 5-7- celled, acicular, spreading, tortuous, whitish, the scales ca. 2. 5-3.0 X ca. 0.2 mm, appressed to spreading, linear-lanceolate, brown, lustrous, tortuous, entire to sparsely denticulate, not clathrate, not bullate, adaxially eglandular, densely pilose, scales absent, the hairs similar to those of the pinna rachises abaxially; basal basiscopic pinnules of medial pinnae often (especially in large leaves) enlarged and overlapping the lamina rachis; coshiles abaxially sparsely glandular, sparsely pubescent, the glands and hairs similar to those of the pinna rachises, adaxially eglandular, densely to sparsely pilose, scales absent, the hairs similar to those of the pinna rachis adaxially; laminar tissue between veins on both surfaces sparsely glandular, the glands ca. 0.1 mm long, stalked, 2- celled, yellowish, abaxially glabrous to sparsely puberulent, the hairs ca. 0.1 mm long, 1 -celled, erect, acicular, adaxially sparsely pilose near the lamina margins, the hairs ca. 0.5 mm long, 3- or 4-celled; ultimate veins visible on both surfaces, sparsely glandular on both surfaces, the glands similar to those of the lamina tissue, abaxially sparsely pilose, the hairs similar to those of the pinna rachises, adaxially pilose, the hairs 1. 0-2.0 mm long, 5-7-celled, whitish; lamina margins eglandular, ciliate, the hairs 0. 3-1.0 mm long, 2-6-celled, acicular, spreading; indusia absent; spores cristate. Distribution and ecology. — Ecuador, Peru; eastern side of Andes, on rocks or trail banks, premontane forest, 200-1200 m. Selected Specimens Examined.— ECUADOR. Napo: Km 86.4 along the road traversing Yasuni National Park, 200-300 m, 0°51'S, 76°16'W, 8 Mar 1998, 171 MORAN ET AL.: ANDEAN MEGALASTRUM ' PERU. Cusco: Quispicanchis, Hills around 3“,'*70"45'W, and Quince Mil Airport, forest 292 km o^ c_i K icm E of Shapaia on road to Mesofostmm mol/e is ch—d by 1— both surfaces glandular, pinna rachises a piliate bv hairs 0.3-1.0 mm 1,C^2.0 mm long, 5-7-celled, and lamina ^^tsal basiscopic long. On the type, which has .® Lhis.’ln the paratypes, the pinnatisect and pinna rachises with glandular (vs. glabrous) and rachises and laminae between the veins 1.0-1-3 nun long (vs. 0.1 pinnae rachises conspicuously pubescen wi basiscopic pinnules 0.2 mm). Megalastrum bolivianum is ^ , • g M. molle differs by often enlarged and laminae adaxially wi ^ rachises of the lamina lamina tissue between the veins abaxially g , , lanceolate and non- and pinnae with linear-lanceolate, tortuous tortuous). ^ 25. Megalastrum mollicoma (C. molUcoma C. Chr., 77:128. 1987 [published 3 May f ^ath. Afd., ser. 8, 6:75_1920. Danske Vidensk. Selsk. Skr.. Natumdensk^ Mam Type.- Ctenitis mollicoma (C. Chr.) Ching, Qyacachi, Feb 1901, L So iro ECUADOR. Napo; “in silv. Stolze, (1991) and second s.n. (lectotype, firs, -step olectot^^^^ T'~49 step here designated:?! GH!, P! [barcode 00610849, 0009078241, photos F!, MICH!, S!t 006108511, US!), Figs. 20H-M, Megalastrum laxlpilosum A. Natural La Planada, 7 to 2008. TVPS.-COLOMBIA. Narmm -d Ricau^ X uo IN, / / a** v» , 780-800 [barcode 3912989)). appressed, lanceolate t< Rhizomes erect, the scales denticulate, flat to ^^^^^gpjading hnear-lanceolate. brown, entire to sp^ L-p£natifid, 0-5-2.5 m long; petiole base scale basally 3- to 4-pnm P to ascending ®latonae 0.5-1.5 -^a^^L pubescent ahaxtaUy. basal medially 2- to 3-pinnate-pinnatiset, la 172 AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 3 (2014) pinnae up to 30.0 cm long, inequilateral; pinna rachises abaxially sparsely to densely glandular, pubescent, scaly, the glands ca. 0.1 mm wide, short-stipitate to sessile and spherical, yellowish to orangish, the hairs 0.2-0.8 mm long, 3-6- celled, lax, spreading, the scales 2. 0-7.0 X 0.3-1.0 mm, dark brown to golden brown, lanceolate, flat (not twisted or tortuous), ascending to spreading, firm to thin and, lustrous, entire to sparsely denticulate, not clathrate, not bullate, adaxially eglandular, densely pubescent, scaly, the hairs 0. 6-2.0 mm long, 4-8- celled, spreading to ascending, the scales like those abaxially; basal basiscopic pinnules of medial pinnae not enlarged or overlapping the lamina rachis; costules with indument like that of the pinna rachises but with scales ca. 2.0 X 0.5 mm, adaxially like the pinna rachises but scales absent; laminar tissue between veins abaxially pubescent, sparsely glandular, the hairs 0.1-0. 2 mm long, 1- or 2-celled, erect, lax to acicular, the glands ca. 0.1 mm long, 2-celled, short-stipitate, yellowish, erect, adaxially pubescent (sometimes only so toward the margins), sparsely glandular, the hairs 0.1-1. 0 mm long, 2-8-celled, lax, slightly tortuous, ascending, appressed, the glands like those abaxially; ultimate veins on both surfaces visible to obscure, abaxially pubescent, the hairs 0.2— 0.6 mm long, 2-5-celled, spreading or appressed-ascending, lax, adaxially the hairs 0. 2-0.5 mm long, 2-4-celled, slightly tortuous, spreading to ascending; lamina margins eglandular, ciliate, the hairs 0.3-0. 8 mm long, 3-6-celled, lax, spreading; indusia absent; spores cristate. Distribution. — Colombia, Ecuador; (750-)1800-2450 m. Selected Specimens Examined.— COLOMBIA. Antioquia: Mpio. Fredonia, vereda Erbe, microcuenca La Chaparrala, 2150-2250 m, 5°59'01"N, 75°38'28.3"W, 20 Dec 2004, Rodriguez et al. 4805 (HUA, NY). Narino: Reserve Natural La Planada, sendero del Rondon, ca. 2 km de centro de investigacion, 1900 m, 1^09'55"N 77^58'44"W, 19 Jan 1997, Herrera 9130 (UC); trail to El Hondon, 5- (MO) 78°02'W, 6 Jan 1988, Gentry ECUADOR. Carchi: El Gualtal, faldas de Cerro Golondrina Hembra, 2450 m, 0°51'N, 78^072^, 21 Aug 1994, Palacios 8^ Clark 12644A (MO, QCNE, UC). rd from Lloa to the West, Km 20.6, 2450 m, 0°13'S, 78°38'W, 1 May 1991 0/7gaord 98891 (AAU). Tunguragua: without locality, [0°23'S, 78°25'W], ca. 1857, Spruce 5257 (P). Megalastmm moUicoma is distinctive by the pubescence on the adaxial surface of the laminae: the hairs are dense, slightly tortuous, appressed, and has pubescence like this. The erect Besidp pubescent but the hairs tend to be straighter and more erect. Besides these hairs, also distinctive are the pinna rachis scales -a the veins adaxially pubescent. ^ ““f“sed^ith If” vii^tut^'^hich has r P^boscent between the veins on both surfaces. Megala^rum vastuu, differs by hairs between the veins on both surfaces only MORAN ET AL.; ANDEAN MEGALASTRUM 0 lf-0 2) mm long, and scales of the lamina and pinna rachises usually dull brown. In M. vastum the scales of the lamina inconspicuous to the naked eye, whereas in M. mollicoma they conspicuous. 26. Megalastrum nanum R. C. Moran, J. Prado & Sundue, sp. , BOLI^A. Santa Cruz: Florida, Canton Mairana ^^94’ 7 km NNE by air from Mairana, 18°03'S, 65^55 J. m 008709451)’. R. C. Moran 7 (holotype, LPB!; isotypes, AAU., NY. [h Figs. 2E, 8A, 32B, 24A-E. cent abaxially; basal pinnae 15 . 0 - 20.0 cm long, (elongated basiscopically), the basal glandular, the pinna rachises abaxially densely pilose, sparsely scaly ® Tnt'cr o'ato long olate to lanceolate, mostly appressed, the similar indument but with denser ha , . tissue between veins indument similar to the pinna 1 4-6-celled, spreading abaxially pilose, glandular, the ^nes like those on the to erect, the glands both sessile md s ^ 2-CBlled, adaxially glabrous, pinna rachis, the stalked gl™ds c • „i,scure adaxially, abaxially eglandular; ulUmate veins visibl . , the hairs like those Distribution.-Ecuador, Peru, Bolivia; 1900-2100 m. f FPIJADOR Zamora-Chinchipe: New road Lo)a Selected Specimens Examined.— ECUAU - crosses the road, — PERU. Amazonas: San Martin, 1950 m, der Werffet al. 16720 (MO). ^arga a Valle Grande, BOUVIA. Santa Cruz: Vdle Gr^de, 5 ^ 2100 m, 18 = 43 'S, 63=54 W. » Jnn ^ y Fauna Tariquia, Rio Nogal, Aniceto Arce Ruiz, Reserve N 6 Oct 2004, /imenez Serrano campamento Alisos, 1700 m, 22 00 S, 64 33 vv, 2398 (GOET, NY. UC). AMERICAN FERN JOURNAL: VOLUME ,04 NUMBER 3 (20,4) by the scales on the pinna rachises and ■ f^goceum and M. pulverulentum echinulate spores and pinna rachis scales are darkened apicallv but nr«n the tips (not all character). The two cm be distinguish that basiscopic pinnule of the basal piLa- dissection of the basal with most of the segments ninnLfin ^ d ^tde wide with only one or twn h^ i • ^ ^p^t^ast, M. ciliatum is 1. 5-2.0 cm being entire. Other differences are segments pinnatifid, the rest 2.0 mm wide and yelST hrn 0.5 mn, wide end “■^- ••0-..5 .on. i„ M. nnnn. veLsns^-I^'etnTo™ j- r “ ^ “• - - ... . ,,,, «• - ascending, linear. *^old“il“broi^^^,h appressed to strongly intermittently), flat to twisted snar« f" ,'"a'*“e blaclt-bordered (at least petiole base scales like thot of * t denticulate) leaves 0.8-1.0 m long) ascending) laminae 0.4-0.6 m long tesallPr™®* spreading to pinnate-pinnatifid) lamina rachisM abav III ^ u" 'P^Usect, medially 2- long, strongly inequilateral; pinna rach^® P‘"“aa ™ pubescent, sparsely scaly, the hairs n n_n c “'’aiaally eglandular, densely spreading, the scales 2,5-4 5X0 3-0 n ^^'Celled, acicular, subentire to sparsely denticulatr'L?"’,"^®''*^' bi-own, flat, glandular, densely pubescent, scaks absent fh"*E’ “daxially 5-celled, ascending to spreading aciculm- br ' 1^^ ® *™B. 3- pmnae not enlarged or overkuknp ,E f eglandular, puberulent, sparsely a^calv k *“‘“1“ abaxially accular, spreading, the scales 1 5^x1^ ,1^T ® "™ *oog. 2-4-celled. lustrous, subentire, adaxially with inH, ' ‘*“®ar-lanceolate, brown, surfaces) laminar Ussue be^JTn ve^T abaxial “ eglandular, densely pubes- MORAN ET AL.: ANDEAN MEGALASTRUM 175 cent, the hairs 0.1-0.2 mm long, 1-3-celled, erect, acicular, adaxially glabrous; ultimate veins visible on both surfaces, abaxially eglandular, puberulent, the hairs like those on the lamina tissue, adeixially glabrous or only 1 or 2 hairs present distally, the hairs like those on the costules; lamina margins eglandular, ciliate, the hairs 0.1— 0.2 mm long, 1— 3-celled, ascending to spreading, acicular; indusia absent; spores cristate. Distribution. — Colombia; 2000—2700 m. Specimens Examined. — COLOMBIA. Cundinamarca; Cordillera Oriental, 2438 m, 3 Sep 1994, Little S' Little 8597 (GH). Santander: between Piedecuesta and Las Vegas, 2000-2500 m, [6°50'N, 73°02'W], 19-24 Dec 1926, Killip S' Smith 15555 (NY). Megalastrum nigromarginatum is distinctive by its black-bordered rhizome scales (thus the specific epithet). This border is often interrupted, not continuous. Also distinctive are the small laminae (0.4-0.6 m long). The indument is also distinctive. The laminae are pubescent between the veins abaxially by erect acicular hairs. Killipp S' Smith 15555 is slightly atypical by more glabrous axes, flaccid scales, and shorter pubescence between the veins abaxially. This species resembles Megalastrum ciliatum from Bolivia and northern Argentina. Both have small laminae and similar cutting, and their rhizome scales are intermittently black-margined — a rare character in the genus. The two species can be distinguished by characters given in the key. Also similar is Megalastrum pubescens, which also has small leaves and is pubescent abaxially between the veins. That species differs by ascending pinnules, laminae abaxially with some of the hairs gland-tipped, and concolorous rhizome scales. Also, its lamina tissue between the veins abaxially has slightly longer hairs (0.2-0. 3 vs. 0. 1-0.2 mm long) and longer marginal cilia (0.3-0.5 vs. 0.1-0.2 mm long). 28. Megalastrum obtusum R. C. Moran, J. Prado & Sundue, sp. nov. Type. ECUADOR. Pichincha: Reserva Biologica La Perla, 2 km from La Concordia, [0 O'N, 79 23'Wl, 400 m, s.d., H. Ankersen S' C. Kragelund 211 (holotype: QCA!; isotype: AAU! 2 sheets). Figs. 10B,C, 32C, 35A-E. Rhizomes erect, scales 5-15 X 0.3-0.5 mm, linear-lanceotae spreading to ascending, brown to golden brown, lustrous, twisted simple or sometimes bifid; leaves 0.4-1.0 m long; petiole base k hke those of the rhizomes but spreading to loosely ascending; ® basally 3-pinnate-pinnatisect. medially 2-pinnat^pmnatifif the “ lobes obtuse; lamina rachises pubescent abaxially; basal pinnae 6-^23^ ™ long, strongly inequilateral; pinna rachises abaxially eglandulM, pubescent, scaly, the hairs (Is-lO.lt-l.S nun long, 4-7-celled, X'ntfrTfo white, the scales 0.5-1.7 X 0.1-0.2 mm, linear, lustrous, “ sparsely denticulate, not clathrate, not bullate. ‘‘^axially «ly DiibPsrent snarselv scaly, the hairs 0.5-2.0 mm long, 4-7-celled, spreading, Lightish, white, L scLs like those abaxially; basal basiscopic pinnules of AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 3 (2014) medial pinnae not enlarged or overlapping the lamina rachis, obtuse; costules abaxially eglandular, sparsely pubescent, sparsely scaly, the hairs like those of the pinna rachises but smaller 0.2-0.5 mm long, 2^-celled, the scales like those of the pinna rachises but smaller 0.5-0. 8 X 0.2-0. 3 mm adaxially eglandular sparsely pubescent, the hairs like those on the pinnk rachises, without scales, the hairs 0.5-1.0 mm long, 4-6-celled; laminar tissue between vems on both surfaces eglandular, glabrous; ultimate veins visible on both surfaces, abaxially eglandular, pubescent, the hairs 0.5-1. 5 mm long 2- or 3- celled, spreading, straightish, white, adaxially pubescent, the hairs 0 7-\ 0 mm long, 3- or 4-celled, appressed, ascending, straightish, whitish; lamina margins eglandular, cil.ate, the hairs 0,3-0.5 mm long, 2- or 3-celled, straightish, whitish, ascending; indusia absent; spores cristate. Distribution . — Northwestern Ecuador; 250—700 m. Guapara, ca. 20 km OPA s? ?T' I o' ‘ 19-24 Jun 1967, Sparre 17168 (MO, Silanche NE of Pedro CAAU) ”■ 2 Dec 1996, Qllgaard et al. 2247 co^n runus o t 'P-^'Shtish white hairs. The hairs are also lo3 de™„,r„ " adaxially. This species has one of the lanceolate scales e„ ti. “ '”°^S’natum, a species that differs by Bolivia A nrevious re ^ P™® ibises abaxially and occurrence only in H Navarrete E Tp ^ 0iigaard, isotypes: QcAm.v., AAU! 5 sheets; 0.4-0.6 mm, sprerdina *tn*1 ™ **ase scales 30.0-40.0 X twisted and tortuous, sLnglydScuhtet”®’ pinnate-pinnatifid, medially 2-pinnate piinTt “ ^ hairs abaxially but with mLy scXiS^T f ' *** 50.0 cm long, strongly inequnateXlle ? a u P™'*® abaxially eglandular, glabrous, spailll s^riv P‘""“ narrowly lanceolate, spreading ascendine^’ ^ distally, strongly denticulate nnt rlatL 4°^’ brovm, sometimes darker densely pubescent, sparsely’ scalv the^h ’ adaxially eglandular, ascending, the scales 3.Q-5 0x0 ’2-^ . u ^ 4-6-celled, •^-0.5 mm, Imear-lanceolate, dark brown, MORAN ET AL.; ANDEAN MEGALASTRUM Strongly denticulate; basal basiscopic pinnules of medial pinnae not enlarged or overlapping the lamina rachis; costules abaxially eglandular, glabrous, sparsely scaly, scales like those of pinna rachises but smaller, 0.8-1.5 X 0.3- 0.4 mm, adaxially sparsely pubescent, the hairs like the pinna rachises; laminar tissue between veins on both surfaces eglandular, glabrous; ultimate veins abaxially visible, eglandular, glabrous, adaxially obscure, glabrous, eglandular; lamina margins eglandular, inconspicuously ciliate (often appar- ently eciliate), the hairs ca. 0.05 mm long, 1-celled, ascending; indusia absent, spores cristate. Distribution. — ^Ecuador; 1600-1750 m. Specimens Examined.-ECUADOR. Napo: road Baeza-Lago Agrio ca. 114 ^ from Lago Agrio, 1750 m, 0°16'S, 77°46'W, 8 Aug 1980, Qllgaard et al 35806 (AAU). Megalastrum oellgaardii is characterized by laminae on both surfaces between the veins glabrous, costular scales dark brown (often more so apical yi and strongly denticulate, and lamina margins inconspicuously ciliate, the ci ca. 0.05 mm long and often apparently absent. It resembles M. squomosissimum a species that differs by pinna rachis scales larger (2-10 X 0.M.5 mml, sptffs y denticulate, golden to pale brown, these often conspicuously ®PL"»oular, straightish, erect to spreading, the rntTcniL d V* oppressed dark brown, 0.8 mm od^ally e^andnlar, pubescent, scales abLnt, the hairs 0.5- brown- ®PP^®^®®^‘^sr:ending to antrorsely strigose, light o^an H ?(“"“•« of medial pinnae not enlarged or oXt'lT® r T"“ “‘’“^olly eglandular, subglabrous to ad^fan 1 ''’"T’ 00 Pmoo mchises abaxially, fai^o ^0**’“*!*^ pubescent with hairs like those on the pinna rachises; laminar hssue between veins abaxially eglandular, subglabrous to pubescent. MORAN ET AL.: ANDEAN MEGALASTRUM 183 rachis, and pinnules. B. Hair detail. C. Marginal hair. D. Clathrate scale. E. Abaxial surface of lamina rachis, pinna rachis, and pinnules. F. Hair detail. Scale bars = 1 mm. A-F: van der Werff et al. 23179 (MO). the hairs 0.1-0.4 mm long, 1-celled, erect, hyaline, acicular, adaxially glabrous; ultimate veins visible on both surfaces or sometimes obscure adaxially, abaxially glabrous to sparsely pubescent, eglandular, the hairs like those on the laminar tissue, adaxially glabrous; lamina margins eglandular, ciliate, the hairs 0. 1-0.3 mm long, 1-3-celled, acicular, ascending or appressed; indusia absent; spores cristate. Distribution— Feru; known only from the type; exact elevation unknown. Megalastnim platylobum is distinctive by hairs of mixed lengths along the pinna rachises abaxially. The smallest hairs are 0.1 mm long, and these intergrade with sparser hairs up to 0.5 mm long. Both types of hairs are erect and whitish. Also distinctive of this species are laminae medially 1-pinnate- pinnatisect to 2-pinnate-pinnatifid and basal pinnae inequilateral (i.e., prolonged basiscopically). The pinna rachis scales are sparse or seemingly absent. Megalastnim platylobum greatly resembles several other Andean species in size and cutting of the laminae, but none of these have the hairs of mixed lengths. The common and widespread M. biseriale differs by pinna rachises abaxially either glabrous or, if pubescent, the hairs sparser and more equal- sized. Megalastnim fibrillosum differs by pinna rachis scales more numerous, firmer, spreading, and if pubescent on the lamina rachises, then the hairs shorter (ca. 0.1-0.2 mm long). Megalastnim reductum is similar but differs by AMERICAN FERN JOURNAL: VOLUME NUMBER 4 (2014) MORAN ET AL.: ANDEAN MEGALASTRUM pinna rachises abaxially with smaller, evenly lengthened (0.1-0. 2 mm long), strigose hairs. Megalastnim bolivianum differs by long hairs on the veins adaxially. See Christensen (1912) for discussion of the synonymy and nomenclature of this species and the names proposed hy Balcer. 33. Megalastrum pleiosoros (Hook, f.) A. R. Sm. & R. C. Moran, Amer. Fern J. 77:129. 1987 [published 3 May 1988]. Polypodium pleiosoros Hook, f., Trans. Linn. Soc. London Botany 20:166. 1847. Dryopteris pleiosoros Svens., Bull. Torrey Bot. Club 65:316. 1938, as “pleiosora”. Ctenitis pleiosoros (Hook, f.) C. V. Morton, Leafl. W. Bot. 8:190. 1957, as “pleiosora”. Type.— ECUADOR. Galapagos Islands: James Island [San Salvador], [0°17'N, 90°42'W], without precise date, C. R. Darwin s.n. (holotype: CGE!). Figs. 24D, 27A, 28A-J. Rhizomes erect, scales 18.0-25.0 X 0.7-1. 3 mm, appressed, linear- lanceolate, gold-brown, lustrous, tortuous, the margins denticulate, the teeth simple or bifid; leaves up to 1 m long; petiole base scales like those of the rhizomes but shorter, to 12.0 mm long, and spreading: laminae to 0.8 m long, basally 2-pinnate, medially 1-pinnate-pinnatisect; lamina rachises pubescent abaxially; basal pinnae 8.0-15.0 cm long, equilateral, the segments adnate; pinna rachises abaxially eglandular, pubescent, and scaly, the hairs 0.6- 1.0 mm long, 6-8-celled, spreading, acicular, hyaline, the scales 2.0-3. 0 X 0.05-0.20 mm, spreading to loosely ascending, linear-lanceolate, tortuous, brown, lustrous, denticulate, the teeth simple or sometimes bifid, adaxially eglcmdular, densely pubescent, sparsely scaly, the hairs 1.0-1. 5 mm long, 7-9- celled, tan, acicular, spreading, the scales similar to those abaxially; basal basiscopic pinnules of medial pinnae not enlarged or overlapping the lamina rachis; costules abaxially eglandular, sparsely pubescent, the hairs like those of the pinna rachises, adaxially eglandular, pubescent, lacking scales, the hairs similar to those abaxially; laminar tissue between veins abaxially sparsely glandular, sparsely to densely pilose, the glands stalked, 0.3 mm long, the hairs 0.3 mm long, 2- or 3-celled, acicular, erect, hyaline, adaxially eglandular, subglabrous to sparsely pubescent, the hairs ca. 0.5 mm long, 3- or 4-celled, ascending, acicular; ultimate veins visible on both surfaces, abaxially eglandular, sparsely pubescent, hairs 0.3-0.5 mm long, 3- or 4-celled, spreading or erect, acicular, hyaline; adaxially eglandular, sparsely pubescent, the hairs 0.1 mm long, 4-7-celled, ascending to appressed, acicular, hyaline; lamina margins eglandular, ciliate, the hairs 0.5-0.7 mm long, 4-8-celled, erect, acicular, ascending; indusia absent; spores cristate. rachis, pinna rachis, and pinnule. M. Adaxial su of rachis lamina, pinna rachis, and pinnules. O. I of leaf rachis, pinna rachis, and pinnules. Q. Adi MacDougal et al. 3886 (MO); H-N: Herrera 9130 irface. N-Q. Af. galapagense. N. Ahaxial surface tachis scales; note hifid teeth. P. Ahaxial surface axial surface of pinna. Scale bars = 1 nun. A-G: (UC). N-Q: Isotype, van der Werff 2244 (U). AMERICAN FERN JOURNAL: VOLUME 104 NUMBER - MORAN ET AL.: ANDEAN MEGALASTRUM 187 Distribution. — Galapagos Islands; shaded moist gullies and potholes in lava (Wiggins and Porter, 1971); 400-800 m. Selected Specimens Examined. — ECUADOR. Galapagos: Fernandina Island, Green Crater, 600 m, [0°40'S, 90°33'W], 17 Sep 1977, Adsersen 887 (C); Idefatigable Island, Mt. Crocker, 9 May 1932, Howell 9230 (GH); Isabella Island, SW slope of Volcano Cerro Azul, 800 m, [0°01'S, 91°12'W1, Aug 1975, van der Werjf 2267 (NY, QCA, U); Pinta Island, in the old crater, central part of island, 590 m, [0°36'N, 90=45'W1, 25 Mar 1972, Hamann 915 (C); Santa Cruz Island, betvy^een Cerro Colorado II and Puntudo, 700 m, [0°38'S, 90 22'W], 18 Apr 1977, Adsersen 84 (C, QCA); idem, Miconia belt near Media Lunz, 800 m, [0°38'S, 90°22'W], May 1974, van der Werff 966 (NY, U); San Cristobal Island, Wrick Bay, 800 m, [0°48'S, 89°24'W1, 6 Jul 1905-1906, Stewart 914 (GH, MO, US); San Salvador Island, El Campamento Central, 570 m, [0"15'S, 90’42'W], 16 Aug 1972, Hamann 1985 (C). Megalastrum pleiosoros is characterized by laminae medially 1-pinnate- pinnatisect, pinnae equilateral, and pinna rachises and costules abaxially pubescent by hairs 0.6-1.0 mm long. This species and M. galapagense are the only Megalastrum in the Galapagos Islands. See M. galapagense for comparison. These two species greatly resemble M. microsorum from mainland Ecuador and Peru. That latter species differs by laminae abaxially either glabrous or shorter-pubescent with fewer-celled hairs (0.2-0.5 mm long, 2- or 3-celled). 34. Megalastrum polybotryoides R. C. Moran, J. Prado & Sundue, sp. nov. Type.— ECUADOR. Napo: proyecto de carretera Salcedo - Tena, 5 km desde donde termina el carretero desde Tena, camino a Verde- Yacu, 1 02'S, 77°52'W, 10 Aug 1994, H. Navarrete 538 (holotype: QCA! 2-sheets; isotjqies: AAU! 3-sheets, QCNE-n.v.). Figs. 5A, 21A-D, 29C. Rhizomes not seen; leaves 1.0-1.5 m long; petiole base scales ca. 10.0 x 1.0 mm, appressed to spreading, linear-lanceolate, light brown, lustrous, twisted, sparsely denticulate; laminae ca. 0.8 m long, basally 2-pmnate- pinnatifid, medially 2-pinnate; lamina rachises abaxially without hairs, glandular, the glands 0.1 mm long, globose, sessile; basal pinnae up to 22.0 cm long, equilateral, segments adnate; pinna rachises abaxially densely glandular, without hairs, scaly, the glands ca. 0.1 mm wide, sessile, spherical, brown, the scales 2.0-5.0 X 0.5 mm, appressed to loosely spreading, linear- lanceolate, brown, lustrous, tortuous, sparsely denticulate, adaxially densely glandular, sparsely pubescent, scaly, the glands and scales similar to those ■s and scales. All scale bars - 1 nun. A-D: Isotype, Navarrete 538 (AAU). E-H: Ollgaard ! (AAU). I-O: Isotype, Camp 3786 (K). Costal 1 AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 4 (2014) acicularf light brow *asraudi^h*°°f 5-celled, appressed to spreading, not enlarged or o^riapnino Z 1“ MORAN ET AL.: ANDEAN MEGALASTRUM 191 abaxially eglandular, sparsely and finely pubescent, the hairs 0.1-0. 3 mm long, 1- or 2-celled, acicular, erect, hyaline, adaxially glabrous to pubescent, the hairs (when present) 0. 3-2.0 mm long, 4-6-celled, ascending to spreading and slightly tortuous; lamina margins eglandular, ciliate, the hairs 0.1-0.4 mm long, 2- or 3-celled, ascending to spreading; indusia absent; spores cristate. Distribution. — Colombia, Ecuador, Peru, Bolivia, eastern side of Andes; 280— 2170 m. Selected Specimens Examined. — COLOMBIA. Putumayo: margenes del Rio Giiamues, San Antonio de Giiamues, 310 m, [0°34'N, 76"50'W], 19 Dec 1940, Cuatrecasas 11185 (F). ECUADOR. Azuay: Banos-Pintuc, (2"57'S, 79°06'W1, s.d., Stubel 943 (B). Morona-Santiago: Morona Canton, near city of Macas, 1100 m, 02'^20'S, 78°08'W, 22 Jul 1993, Fay 8^ Fay 4097 (AAU, MO, NY, QCNE); within 3 km of the town of Gualaquiza, 1000 m, 03°24'S, 78=34'W, 8 Mar 1993, Fay & Fay 4232 (AAU, MO, NY, QCNE). Napo: Santa Rosa at Rio Napo, [0°49'S, 75°30'W], 30 Apr 1972, Lugo 2064 (AAU, GH, MO, U). Pastaza: East of Puyo, along the road to Macas, and wet forest around the Municipal, 950 m, 01°30'S, 77°59'W, 14 Jul 1992, Fay & Fay 3662 (MO, NY, QCNE). Zamora-Chinchipe: Zamora, within 3 km of the town of Zamora, 1000 m, 4°03.5'S, 78°57.5'W, 7 Dec 1994, Fay & Fay 4403 (AAU, MO, QCNE). PERU. Bagua: valley of Rio Maranon above Cascades de Mayasi near Campamento Stte. Montenegro, Kms 280-284 of Maranon road, 450-600 m, [5“18'S, 78°25'W], 5 Sep 1962, Wurdack 1851 (US). Cajamarca: San Ignacio, Ricardo Palma, 1720 m, 5°07'29"S, 79°03'16"W, 19 May 1998, Campos & Lopez 4911 (MO, NY). Cusco: Paucartambo, Hacienda Villa Carmen, 540 m, [13"18'S, 71°35'W], 19 Jul 1963, Vargas 14685 (GH). Pasco: Oxapampa, Gran Pajonal, vicinity of Chequitavo, 1200 m, 10°45'S, 74°23'W, 26 Sep 1983, Smith 5260 (F, MO). BOLIVIA. Beni: Prov. Grab Ballivian, 16 km por el camino maderero, 12 km de Yucumo a Rurrenabaque, 450 m, 15=05'S. 67°07'W, 23 Jul 1997, Kessler et al 10774 (GOET, UC). Carrasco: Cochabamba, Localidad Dianpampa, 2160 m, 17°40'53"S, 64°40'55"W, 4 Sep 2003, Fernandez et al. 2366 (MO). Cocha- bamba: Prov. Jose Carrasco Torrico, 143 km antigua carretera Cochabamba- Villa Tunari, 1300 m, 17°07'S, 65°34'W, 23 Aug 1996, Kessler et al. 7631 (GOET, UC). La Paz: Prov. Abel Iturralde, Rio San Antonio, 46 km de Ixiamas a Alsto Madidi, 300 m, 13=38'S, 68°26'W, 13 Aug 1997, Kessler et al. 11131 (GOET, UC). Santa Cruz: Nuflo de Chavez, Serranfa de San Lorenzo, 2 km E of Est. Las Maras, 500-900 m, 16 15'S, 62°35'W, 3 Nov 1985, Killeen 1383 (F, MO, NY, UC). Abaxial surface of lamina and pinna rachises and pinnules. N. Rachis scale. O. Hairs on abaxial surface of lamina P Adaxial surface of pinnule. Q. Detail of hairs. Scale bars = 1 mm. A-E; holotype, Croat etai. 91165 (MO). F-L: Isotype, Killip & Smith 20453 (NY). M-Q: Isotype, van der Werffetal. 16300 [NY). AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 4 (2014) MORAN ET AL.: ANDEAN MEGALASTRUM 193 Megalastrum praetermissum is distinctive by the linear, brown denticulate scales along the pinna rachises. It varies considerably in pubescence on both surfaces of the lamina. In most specimens there are at least a few ca. 0.1 mm long, erect, acicular hairs between the veins abaxially. These hairs are easily overlooked, especially when viewed from above, and thus the specific epithet praetermissum meaning “overlooked,” or “passed by. These minute hairs are best seen in profile on folded portions of the lamina. Adaxially, the pubescence (when present) is quite different, with the veins and midribs of the ultimate segments bearing conspicuous whitish hairs 0. 3-2.0 mm long. These hairs vary from sparse and inconspicuous to dense and conspicuous. . . Megalastrum praetermissum resembles M. marginatum m lamina division, presence of filiform scales on the petiole bases and pinna rachises, and long hairs on the veins adaxially; however, M. praetermissum differs by pinna rachises abaxially glabrous and laminae between the veins minutely and inconspicuously pubescent. 36. Megalastrum pubescens A. Rojas, Metodos en Ecologi'a y Sistematica (MES) 3(Supl. 1):44, fig. 5. 2008. Type.— COLOMBIA. Norte de Santander. Eastern Cordillera, N of Toledo, Loso and vicinity, [7=02'N, 72^05 W), 220^ 2400 m, 6-7 Mar 1927, E. P. Killip & A. C. Smith 20453 (holotype: NY! [barcode 08715751; isotype: US-n.v.). Figs. 5A, 18D, 23F-L. Rhizomes erect, scales 5.0-13.0 x 0.3-0^ mm appressed to strongly ascending, linear, concolorous, flat, sparsely denticulate; leaves to 0.8-1 .0 m long; petiole base scales like those of the rhizomes but loosely spreading to ascending; laminae 0.4-O.6 m long, basally 3-pinnate-pinnatisect. medially 2- plnnate-pinnatiSd; lamina rachises pubescent abaxially; lamina rachises pubescent abaxially; basal pinnae ca. 30.0 cm long, strongly pinna rachises abaxially eglandular. glabrous to sparsely pubescent sparsely Laly, the hairs 0.3-0.5 mm long, 2-4-celled, acmular. spreading the scales 3 0-45 X 02-03 mm linear-lanceolate, brown, flat, subentire to denticulate, the teeth not bifid, admdally eglandular, densely pubescent, hairs 0 4-10 mm long, 4-7-celled, ascending to spreading, acicular, basal tLLscoptc ptaLules of medial pinnae not enlarged or overlapping the lamina rachis; costules abaxially eglandular, puberulent, sparsely scaly, ham M 0.5 mm long, 2^-celled, acicular, spreading, the scales up to 7.0 X 0.2^, linear-lanceolate, brown, lustrous, subentire, adaxially with indument simi ar to those of the abaxial surfaces; laminar tissue between veins ab^lly glandular, densely pubescent, the hairs 0 3 mm long 1-3-celled, erect, acicular, adaxially glabrous; ultimate veins visible on both surfaces, abaxially eglandular, on the lamina tissue, adaxially very sparsely pubescent, the haus like those on the costules; lamina margins eglandular, ciliate, the haus 0.3-0.5^ long, or 3-celled, ascending to spreading, acicular; mdtrsia absent; spores cristate. Distribution.— Colombia; 2000-2400 m. AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 4 (2014) MORAN ET AL.: ANDEAN MEGALASTRUM Specimens Examined. — COLOMBIA. Antioquia: Mpio. Angel6polis, Vereda Romeral, sector Bocatoma, acueducto multiveredal, Quebrada Las Animas, 2100-2200 m, 6°09'29"N, 75“42'05"W, 29 Oct 2005, Rodriguez et al. 5559 (HUA). Megalastrum pubescens is distinctive by laminae only 0.4-0. 6 m long, abaxially densely and conspicuously pubescent, and adaxially between the veins glabrous. The pinnules tend to be slightly ascending. Megalastrum pubescens resembles M. vastum but differs by laminae adaxially glabrous between the veins and longer hairs abaxially on the pinna rachises and costules (hairs 0.3— 0.5 mm long, 2— 4-celled vs. 0.1(-0.2) mm long, l(-2)-celled). Also their scales on the pinna rachises abaxially differ subtly, with those of M. pubescens being brown, linear-lanceolate, and firmer, whereas those of M. vastum are golden brown, lanceolate, and flaccid. 37. Megalastrum pubirhachis R. C. Moran, J. Prado & Sundue, sp. nov. Type. — ECUADOR. Napo: El Chaco Canton, just above N side of Rio Oyachachi, where the Lago Agrio Road crosses the river, 00 18'S, 77“21'W, 1500 m, 9 Aug 1992, A. Fay & L. Fay 3919 (holotype: QCNE; isotypes: AAU!, MO! [04670566, 04670570, 04670656,], NY! [barcode 00870997], QCA?-n.v.). Fig. 18E, 34F-L, 36A. Rhizomes unknown; leaves up to 2.0 m long; petiole base scales 10.0- 15.0 mm long, linear, brovra, shiny, smooth on both surfaces, spreading to loosely ascending, denticulate; laminae 0.4-1. 5 m long, basally 3-pinnate- pinnatisect, medially 2-pinnate-pinnatisect; basal pinnae 30.0-45.0 cm long, inequilateral; pinna rachises abaxially eglandular, densely pubescent, scaly, the hairs (when present) 0.1-0.2 mm long, 2 or 3-celled, mostly retrorsely strigose, the scales 3.0-4.0 X ca. 0.1-0.2 mm, dull brown, oppressed (mostly) to loosely spreading and ascending, filiform to linear, slightly tortuous, denticulate, adaxially pubescent, sparsely scaly or lacking scales, hairs like those abaxially, the scales like those abaxially; basal basiscopic pinnules of medial pinnae not enlarged or overlapping the lamina rachis; costules abaxially eglandular, pubescent, without scales or only sparsely scaly, the hairs ca. 0.1 mm long, 1- or 2-celled, acicular, ascending to strigose, the scales (when present) like those of the pinna rachises, adaxially pubescent, the hairs like those on the pinnae rachises adaxially; laminar tissue between veins abaxially eglandular, glabrous to sparsely puberulent, the hairs ca. 0.1 mm long, easily overlooked, erect, adaxially eglandular, glabrous; ultimate veins abaxially visible, glabrous to sparsely puberulent, the hairs ca. 0.1 mm long. Rachis scale. Scale bars = 1 mm. A-D; Conn 2547 (UC). L-O: Lectotype, Spruce 4656 (K). MORAN ET AL.: ANDEAN MEGALASTRUM acicular, adaxially obscure, glabrous; lamina margins eglandular, ciliate, the hairs 0.1-0. 2 mm long, 1- or 2-celled, acicular, ascending to appressed; indusia absent; spores cristate. Distribution. — Ecuador, Peru, eastern side of the Andes; 1300—2100 m. Selected Specimens Examined. — ECUADOR. Napo: 65 km along rd. NE of Baeza, Cordillera Oriental, 1300 m, [0=05'S, 77°30'W1, 26 Jan 1984, Moran 3594 (B, MO, NY). PERU. Amazonas: Bagua, 12 km E of La Peca, 1700 m, [5°36'S, 78"26'W1. 20 Jun 1978, Barbour 2394 (MO, NY). Megalastrum pubirhachis is distinctive by pinna rachises abaxially with linear denticulate scales and dense, even, retrorsely strigose hairs. By these characters it resembles M. reductum, a species that differs by filiform scales and smaller (0.4-0.8 m long vs. 0.4-1. 5 m in M. pubirhachis), less divided laminae (basally 2-pinnate-pinnatisect, medially 1-pinnate-pinnatisect to 2- pinnate-pinnatifid vs. basally 3-pinnate-pinnatisect, medially 2-pinnate- pinnatisect in M. pubirhachis). Megalastrum pubirhachis occurs only on the eastern side of the Andes, whereas M. reductum is only on the western side. The specific epithet refers to the dense hairs on the abaxial surfaces of the pinna rachises. 38. Megalastrum pulvenilentum (Poir.) A. R. Sm. & R. C. Moran, Amer. Fern J. 77:129. 1987 [published 3 May 1988]. Polypodium pulverulentum Poir., Encycl. Meth. Bot. 5:555. 1804. Aspidium lutescens Willd., Sp. PI. 5:272. 1810, nomen superfl. Dryopteris pulverulenta (Poir.) C. Chr. in Urb., Symb. Antill. 9:305. 1925. Ctenitis pulverulenta (Poir.) Copel., Gen. Fil. 124. 1947. Lectotype.— designated by Proctor (1985): Plunder, Traite Foug. Amer. 27, pi. 34, 1707, illustrating a plant from Hispaniola. Fig. 8C,E, 26H-L, 36B. Polypodium barbatum Desv., Mem. Soc. Linn. Paris 6:242. 1827. Type.— HISPANIOLA. Without exact locality, s.d., A. N. Desvaux s.n. (holotype: P! [barcode 00600637], photo US!). Polypodium karstenianum Klotzsch, Linnaea 20:390. 1847. Phegopteris karsteniana (Klotzsch) Mett., Abh. Senck. Nat. Ges. Frankfurt 2:314 1858. Nephrodium villosum (L.) Kuntze var. karstenianum (Klotzsch) Jenman, Bull. Bot. Dept. Jamaica n.s. 3:114. 1896. Dryopteris subincisa (Willd.) Urb. var. karsteniana (Klotzsch) C. Chr., Index Filic. 295. 1905. Dryopteris karsteniana (Klotzsch) Hieron., Hedwigia 46:348. 1907. Ctenitis karsteniana (Poir.) Vareschi, FI. Venez. 1(1):405. 1969. Scale bars = 1 mm. A-D: Isotype, Moran 5911 L: Fay S' Fay 3845 {NY). (UC); E-G: Isotype, Qllgaard et at. 99556 (AAU). H- 198 AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 4 (2014) Megalastrum karstenianum (Klotzsch) A. Rojas, Rev. Biol. Trop. 49:483. 2001. Type.— VENEZUELA [“as Colombien”]. Dec 1848, H. Karsten II, 3 (lectotype, designated by Moran and Prado, 2010: B! [barcode 200058400], photos B!, US!; isolectotypes: H-n.v.). MORAN ET AL.: ANDEAN MEGALASTRUM Phegopteris hirsuta Fee, Mem. Foug. 5:248. 1852. Type.— VENEZUELA [“Mexico”]. Merida: Aug 1846, H. Funck & L /. Schlim 975 (lectotype, designated by Moran and Prado, 2010: P [barcode 00600642); isolecto types, BM! [barcode 000907809], P! [barcode 00600643, 00600644), P [barcode 583962, 00583963]). Aspidium erythrostemma Christ, Bull. Herb. Boiss., ser. 2, 4:961. 1904. Dryopteris erythrostemma (Christ) C. Chr., Index Filic. 263. 1905. Type.— COSTA RICA: in 1903, C. Werckle s.n. (lectotype, designated by Moran and Prado, 2010: P! [barcode 00600398]). Rhizomes erect, scales 20.0-40.0 X 0.3-0.5 cm, appressed to strongly ascending, linear-lanceolate, light brown to golden, twisted, lustrous, the margins denticulate; leaves up to 4.0 m long; petiole base scales like those of the rhizomes but spreading to loosely ascending; laminae 1. 2-3.0 m long, basally 4-pinnate-pinnatisect, medially 3-pinnate-pinnatisect; lamina rachises pubescent abaxially; basal pinnae to 1.4 m long, strongly inequilateral; pinna rachises on both surfaces densely pubescent, scaly, glandular, the hairs 1.0- 2.0 mm long, 5-12-celled, spreading, the scales denticulate to ciliate (the cilia often darker than the body of the scale, especially towards the apex), non- bullate, of two types, small ones ca. 0.2-0.6 X ca. 0.2 mm long, ovate to lanceolate, mostly appressed, the larger 3. 0-8.0 x 0.2-0.5 mm long, linear to linear-lanceolate, spreading, the glands ca. 0.1 mm long, spherical, sessile, yellowish, adaxially with indument like that abaxially; basal basiscopic pinnules of medial pinnae not enlarged or overlapping the lamina rachis; costules abaxially glandular, densely pubescent, sparsely scaly, the indument like that on the pinna rachises but more sparsely scaly adaxially; laminar tissue between veins glandular and pubescent on both surfaces or only abaxially, the glands short-stalked to sessile and spherical, yellowish, the hairs 1. 0-2.0 mm long, 5-8-celled, spreading to erect; ultimate veins on both surfaces visible to obscure, abaxially glandular, sparsely pubescent, non-scaly, the hairs ca. 0.5-2.0 mm long, 4-8-celled, adaxially eglandular, sparsely pubescent, the hairs 1.0-1. 8 mm long, 4-9-celled, spreading; lamina margins glandular, ciliate, the hairs 0.4-1.0 mm long, 4- or 5-celled, spreading, the glands few, stalked, yellowish; indusia absent; spores echinulate. Distribution— Mexico, Guatemala, Honduras (Stolze, 1981), Costa Rica, Panama, Jamaica, Haiti, Dominican Republic, Guadeloupe, Martinique, St. Vincent, Dominica, Saba; Venezuela, Colombia, Ecuador, Peru; 1200—2500 (-3200) m (in West Indies from 0-1600 m; Moran et al., 2009b). Selected Specimens Examined. — ^VENEZUELA. Aragua: Colonia Tovar and vicinity, 2000-2100 m, [10"22'S, 67°16'W], 28 Dec 1921, Pittier 10029 (GH, NY, US). Distrito Federal: Carretera El Junquito-Colonia Tovar, [10 28'N, 67"05'W], 24 Oct 1963, Agostini 207 (NY). Lara: Moran, 8.7 mi SE of Sanare, in Parque Nacional Yacambu, 1500 m, 9=41'N, 69"35'W, 13 Nov 1982, Smith et al. 1249 (MO). Merida: Dpto. Sucre, en las Selvas Nubladas situadas al norte de la localidad conocida como La Trampa, 2000 m, 9 34' N, 71 26' W, 2 Feb 1987, Ortega et al. 3030 (MO). Trujillo: Mpio. Bocond, entre las antenas de relevo en AMERICAN FERN JOURNAL: VOLUME NUMBER 4 (2014) MORAN ET AL.; ANDEAN MEGALASTRUM el Paramo de Guaramacal y la Vega de Guaramacal, 1200—3200 m, [9 11 N, 70 12'W], 23-24 Jul 1984, Ortega et al. 2013 (MO, NY). GOLOMBIA. Antioquia: Jardin, Vereda Quebrada Bonita, sector Bocatoma. quebrada La Mendoza, 2110-2150 m, 5 35'04.3"N, 75 46'36.1'"W, 7 Jan 2005. Rodriguez et al. 4915 (HUA, NY). Cauca: near Popayan, 1800 m. [2 26'N, 76=37'W], 14 Apr 1939, Alston 8026 (MO). Cundinamarca: Fusagasugd. m. [4 20'N, 74=22'W], s.d., Andre 91 (K, NY). Magdalena: Cordillera Oriental, Sierra de Perija, 10 km ENE of Manaure, 46 km E of Valledupar, 3 km from Venezuela border, 2300 m, [10 28'N, 73 15'W]. 4 Feb 1945, Grant 10810 (NY, US). Norte de Santander: Eastern Cordillera, Loso and vicinity. N of Toledo, 2200-2400 m, [7°17'N, 72°25'W], 6-7 Mar 1927, Killip Er Smith 2041 1 (GH, NY, US). Santander: Rio Surata valley, above Suratd, 2000-2300 m, (7 30'N. 72°58'W], 5-6 Jan 1927, Smith 16659 (GR, NY, P, S, US). ECUADOR. Napo: Quijos Canton, within 5 km of the town of Baeza. 1900 m. 0°27'5"S, 77"53'00"W, 20 Jun 1992, Fay & Fay 3845 (AAU, MO. NY (^NE)_ Pichincha: camino Santo Domingo de los Colorados a Alagoa, 2.5 km E ot Cornejo Astorga, bosque primario por un no, Cordillwa Central 1200 m, [0=26'S, 78 47^], 8 Jan 1984, Moran 3546 (F, MO, NY). Without locality: 1857-1859, Spruce 5257 A (GH, K, NY, P, S). , o i PERU. Cusco: La Convencidn, Distr. Santa Ana, Madre Selva, 2100 m, 12°53'49"S, 72°45'02"W, 19 Mar 2004, Valenzuela et al. 3019 (MO). Megalastnim pulverulentum is one of the largest species in the genus, with leaves to 4 m long and basal pinnae to 1.4 m long. Its lammae are densely pubescent on both surfaces by long (1-2 mm) whitish hairs The pinna rachm Lales are prominently toothed and darker apically. Yellowish gl^ds ^ present on both surfaces and may be either sessile or short-stalked. It ^eatly resembles M. fugaceum (see discussion of M. fugaceum for comparison). 30. Megalastrum reductum A. Rojas, Met. Ecol. Sist. 3(Supl. ^ 2008 TYPE.-ECUADOR. Guayas/Canar/Chimborimo/Bd.var: onhe W cordillem near the village of Bucay, (2 40'S, 79 40 W|, ^ MS lun 1945 W H Camp 3786 (holotype: US-n.v.: isotypes: F. (areession 1409933),’ K!, MO! (barcode 1634138), NY! [barcode 0871594), P. (barcode 00568190]). Fig. 5B, 21J-0, 29B. Megalastrum dorsiglabrum A- M^^E^VaUe^ C. 2008, as -dorsoglabrum’ . Type.— PAN fhnlotvnP- [8=37'N, 80 07'W], 1000 m, 4 Jun 1934, A. H. G. Alston 8730 (holotype. CR!; isotypes: BM-n.v., US-n.v.)- costule. Q-X. ^ scale. T. Costu surface of rach n Q. Abaxial surface of rachis and pi hairs and scale. U. Detail of abaxial 1 W Rachis hairs. X. Hairs along cc ^ Rohrbach 5160 (MO). Q-X: Leiva asta. Scale bars = etal. 1759 (F). AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 4 (2014) Fig. 29. Leaf dissection in four species of Andean Megalastrum. A. M. hirsutosetosum. B. M. reductum. C. M. polybotryoides. D. M. molle. A: Ollgaard et al. 99268 (AAU). B: Panama, Moran 5060 (MO); C; Navarrete 539 (AAU). D: Isotype, van der Werffet al. 16300 (F). Rhizomes erect to decumbent, up to 10.0 cm long, scales 10.0-20.0 X 1.0- 2.0 mm, ascending, linear, golden brown, dull, firm, setulose on both the margins and surfaces; leaves up to 1.2 m long; petiole base scales like those of the rhizomes but spreading to loosely ascending; laminae 0.4-0.8 m long, basally 2-pinnate-pinnatisect, medially 1-pinnate-pinnatisect to 2-pinnate- pinnatifid; lamina rachises glabrous abaxially; basal pinnae 12.0-30.0(-40.0) cm long, equilateral or nearly so; pinna rachises abaxially eglandular, densely puberulent, scaly, the hairs (when present) 0.1-0.2 mm long, 1-3-celled, spreading to retrorsely strigose, the scales 3.0-4.0 X ca. 0.1 mm, dull brown, loosely spreading to ascending, filiform to linear, tortuous, entire, adaxially glabrous or pubescent, sparsely scaly or lacking scales, hairs like those abaxially, the scales ca. 1.0 X 0.1 mm, dull, brown, filiform, entire, appressed; basal basiscopic pinnules of medial pinnae not enlarged or overlapping the lamina rachis; costules abaxially eglandular, sparsely puberulent or rarely glabrous, sparsely scaly or scales absent, the hairs ca. 0.1 mm long, 1- or 2- celled, acicular, ascending to strigose, the scales ca. 0.5 mm long, filiform, dull brown, entire, adaxially glabrous or subglabrous to sparsely pubescent, the MORAN ET AL.: ANDEAN MEGALASTRUM hairs like those on the pinnae rachises adaxially; laminar tissue between veins ahaxially eglandular, glabrous to puberulent, the hairs ca. 0.1 mm long, 1- or 2- celled, acicular, erect, adaxially eglandular, glabrous: ultimate veins ahaxially visible, sparsely puberulent with hairs like those on the costules, adaxially obscure, glabrous; lamina margins eglandular, ciliate, the hairs 0.1-0.2 mm long, 1- or 2-celled, acicular, ascending to substrigose; indusia absent; spores cristate. Distribution. — Eastern Panama, Colombia and Ecuador, western side of Andes; 300-1000 m. Selected Specimens Examined. — COLOMBIA. Antioquia: Mpio. Frontino, Parque Nacional Natural Las Orquideas, vereda Venados Abajo, camino de la cabana de Venados a la vereda Alto Bonito, 900-920 m, 6°32'10"N, 76°19'15"W, 24 Jul 2011, Samn et al. 5176 (NY). Caldas: Santa Cecilia, Cordillera Occidental, Fatama, vertiente occidental, 800 m, 5°03'S, 75°39'W, 25 Nov 1945, von Sneidern 5024 (F). ECUADOR. Bolivar: San Miguel, within 3 km of Balzapamba, 800 m, 1°47'S, 79°10'W, 10 Jul 1995, Fay & Fay 4626 (MO, QCNE). Cotopaxi: Cacaoal, 300 m, [1°01'S, 79°12'W], 6 May 1951, Bell 920 (BM, S). El Oro: 10.6 millas E de Saracay hacia Pinas, 700 m, [3°37'S, 79°55'W], 23 Nov-18 Dec 1978, Albert de Escobar 943 (NY). Esmeraldas: Quininde, Bilsa Biological Station, Montanas de Mache, 35 km W of Quininde, 5 km W of Santa Isabela, 400-600 m, 0°21'N, 79°44'W, 10 Dec 1994, Pitman Bass 1073 (MO, QCNE). Los Rios: Rio Palenque Estacion Biologica, Km 56 al norte de Quevado, 150—220 m, [1°22'S, 79°42'W], 29 Jan 1984, Moran 3609 (F, NY, QCA, US). Manabi: Canton Pedernales, Cerro Pata de Pajaro a 10 km al este de Pedernales, a 5 km del rancho de la familia Arroyo, 800 m, 00°01'N, 079°57'W, 10 Mar 1997, Vargas 1295 (MO, QCNE). Pichincha: Hotel Tinalandia, casi 25 km al este desde Sto. Domingo de los Colorados, Cordillera Occidental, 1000 m, [0'’17'23"S, 79°04'06"W], 8 Jan 1984, Moran 3543 (MO, NY, QCA). Megalastrum reductum is distinctive by the combination of leaf dissection and indument. The leaves are medium-sized, up to only 1.2 m long, with the basal pinnae equilateral or nearly so, not prolonged on the basiscopic side as in many other species of Megalastrum. The first two proximal pinnules are often slightly reduced (thus the specific epithet), and apices of the pinnules are typically obtuse or broadly acute. The pinnules are mostly adnate and appear short and stubby. The abaxial surfaces of the pinna rachises are evenly puberulent with hairs 0. 1-0.2 mm long, 1- or 2-celled, and spreading to retrorsely strigose. The scales on the pinna rachises ahaxially are scattered, dark, filiform to linear, and slightly tortuous. Shorter hairs (ca. 0.1 mm long) occur ahaxially on the costules and sometimes along the veins, and in some specimens short erect acicular hairs (these easily overlooked) occur on the lamina tissue between the veins. The rhizome scales appear setulose on both surfaces, this apparently from the upturned and projecting ends of the cells. Some specimens have pinna rachises adaxially glabrous, and these have been named Megalastrum dorsiglabrum A. Rojas (Rojas 2008). We place M. AMERICAN FERN JOURNAL; VOLUME 104 NUMBER 4 (2014) MORAN ET AL.: ANDEAN MEGALASTRUM dorsiglabnim in synon5niiy because no other character appears to correlate with the lack of hairs adaxially on the pinna rachises. Megalastnim vastum is similar by its short even pubescence on the laminae abaxially; however, it differs by larger, more divided laminae (up to 3-pinnate- pinnatisect), linear lanceolate (vs. filiform) scales on the pinna rachises abaxially, and laminar tissue between the veins pubescent adaxially especially toward the margins. Megalastnim biseriale is similar hut differs by inequilateral basal pinnae (at least in large leaves), scalier leaf axes abaxially, and pinna rachises abaxially with longer hairs (0.4-0. 6 mm long, 3- or 4-celled) and wider, more lanceolate scales. See Megalastnim pubirhachis for comparison with that species. 40. Megalastnim rhachisquamatum R. C. Moran, J. Prado & Sundue, sp. nov. Type.— ECUADOR. Napo: Canton El Chaco, Proyecto Hidroelectrico Coca, Punto ST3, margen derecha del Rio Quijos, ca. 10 km S de Reventador, 00°39'S, 77°39'W, 1500 m, 3-5 Oct 1990, W. Palacios 5844 (holotype: QCNE!; isotypes: MO! [barcode 4062147, 4062148], NY! [barcode 0871596], QCA?-n.v., UC! [barcode 1593127]). Figs. 7H-L, 22D, 36C. Rhizomes erect; leaves 1.0-1. 2 m ]ong; petiole base scales ca. 15.0 x 2.0 mm, brown, flat, ascending to spreading, sparsely denticulate, sometimes with denticulae on the surface; laminae 0.7-0. 8 m long, 3-pinnate-pinnatisect at base, 2-pinnate-pinnatisect medially; lamina rachises pubescent abaxially, densely and conspicuously scaly; basal pinnae up to 27.0 cm long, inequilateral, longer basiscopically; pinna rachises on both surfaces very sparsely glandular, densely pubescent, densely and conspicuously scaly, the hairs 0.4-0. 5 min long, 4-6-celled, lax, spreading, the scales 5. 0-7.0 X 1.0- 2.0 mm, brown, lanceolate with conspicuously narrowed base (the point of attachment thickened), spreading, flat, lustrous, denticulate, adaxially egland- ular, densely pubescent, scaly, the hairs 1.0— 1.5 mm long, 5— 8-celled, spreading, lax, scales like those on the abaxial surfaces; costules abaxially glandular, pubescent, scaly, the glands stalked, orangish, the hairs and scales like those of the pinna rachises abaxially, adaxially eglandular, pubescent, sparsely scaly, the hairs 1.0-1.5 mm long, 5-8-celled, spreading, lax, the scales like those abaxially; laminar tissue between veins abaxially glandular, pubescent, the glands 0.1-0.2 mm long, 2-celled, stipitate, yellowish, erect, the hairs ca. 0.1 mm long, 1- or 2-celled, erect, acicular, adaxially sparsely glandular, pubescent, the glands like those abaxially, the hairs 0.3-0.4 mm long, 2- or 3-celled, lax, spreading to appressed; ultimate veins not visible on Segment apex. M. Hairs along costule. N. Abaxial surface of pinna rachi: tail of adaxial sinface. Scale bars = Moritz 202 p.p. (F).*E-G: Isotype, Steyermark 85913 (US). H-P: from Ja 1 pinnule. O. Detail of un. A-D: Isolectotype, ;a, Clute 132 (US). 206 AMERICAN FERN JOURNAL: VOLUME 104 NUMBER - MORAN ET AL.: ANDEAN MEGALASTRUM 207 both surfaces, abaxially sparsely glandular, pubescent, the glands like those from the laminar tissue, the hairs 0.2-0. 1 mm long, 2- or 3-celled, lax, spreading, adaxially like the abaxial side but hairs 0.7-1. 2 mm long, 3-7- celled, spreading to appressed; lamina margins eglandular, densely ciliate, the hairs 0.3-0.5 mm long, 3-5-celled, ascending to spreading; indusia absent; spores cristate. Distribution. — Ecuador, Peru; 1350-2300 m. Selected Specimens Examined.— ECUADOR. Zamora-Chinchipe: Cordillera del Condor, Chinapintza, trail leading to Destacamento Mayaycu Alto, 135^ 1480 m 4°03'S, 78°35'W, 6 Dec 1990, 0Ugaard 98413 (AAU); vie. of the mining camp at the Rio Tundaime. along road to miliary base El Condor, 1500 m, 3°38'02"S, 78°25'32'W, 6 Nov 2004, van der Werff 19383 (QCNE, UQ. PERU. Cusco: La Convencidn, above Qillouno, 2300 m, 12°28 35 S, 72°29'15"W, 29 Apr 2006, van der Werff 21116 (MO, NY). Megalastrum rhachisquamatum has petioles and rachises densely and conspicuously scaly (thus the specific epithet), the scales 1-2 mm wide and spreading. Also helpful in identification are the laminae 3-pinnate-pinnatisect at base and tissue between the veins pubescent on both surfaces. The pubescence adaxially is distinctive by the hairs between the veins being shorter (0.3-0.4 mm long) than those along the veins and midnbs of the ultimate segments (0.7-1.2 mm long). This species resembles M. ^o^coma a species that differs by shorter, slightly tortuous, and appressed-ascendmg hairs adaxially on and between the veins. 41 Meaalastrum nipicola M. Kessler & A. R. Sm., Amer. Fern J. 96:41 fig. 2Xi- F 2006 Type -BOLIVIA, La Paz: Prov. Nor Yungas, Estacion Biologica de TunqutnLTajo Homuni, senda del campo de Don Pedro *1 cammo de la mina, 16”12'S, 67°53'W, 21 Jul 2000, D. [barcode 1749582); isotypes: GOET!, LPB-n.v.). Figs. 12A-C, 13E, 36B. Rhizomes erect, scales 5.6-10.0 X 0.5-0.8 r^, iTm 'rnr;^oir;i:: rx'i loosely ascending; j rachises AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 4 (2014) 209 MORAN ET AL.; ANDEAN MEGALASTRUM 14°35'52"S, 68°55'46"W, 29 Jun 2005, Fuentes et al 9034 (MO, NY); Prov. J. Bautista Saavedra M., Pauji-Yuyo, entre Apolo y Charazani, 1450 m, 15 02'S, 68“29'W, 8 Jun 1997, Kessler 9900 (UC). Megalastrum rupicola is characterized by pinna rachises abaxially without hairs, pinna rachis scales lanceolate, brown to golden brown, shiny, and veins adaxially glabrous or subglabrous. The species most resembles M. marginatum, which differs by laminae pubescent with long, straightish, white hairs. 42. Megalastrum squamosissimum (Sodiro) A. R. Sm. & R. C. Moran, Amer. Fern J. 77:129. 1987 [published 3 May 19881. Nephrodium squamosissimum Sodiro, Crypt. Vase. Quit. 256. 1893. Dryopteris squamosissima (Sodiro) C. Chr., Index Filic. 294. 1905. Ctenitis squamosissima (Sodiro) Copel., Gen. Fil. 125. 1947. Type.— ECUADOR. Chimborazo: Mt. Tungurahua, [1 28'S, 78°26'W], Dec. 1857, R. Spruce 5262 (neotype, here designated: BM! [barcode 000907806]; isoneotypes: BM! [barcode 000777142, (fragm.) 000890191], CHI, K! [barcode 000200216], NY! [barcode 00871582], P! [barcode 00600375, 00610642]). Figs. 12 D-F, 18A, 36C. Dryopteris subincisa (Willd.) Urb. var. bogotensis Hieron Hedwigia 46:349 1907. Dryopteris squamosissima (Sodiro) C. Chr. var. bogotensis (Hieron.) C. Chr.. Koneel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math Afd_., segments. Q. Adaxial surface oi cosiuie. : Moran 3594 (MO). U-Q: Holotype, Tryon & Tryon 6103 (GHJ. F-L: AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 4 (2014) MORAN ET AL.: ANDEAN MEG/U^SraUM lone, 4-6-celled, antrorsely strigose, scales like those on the abaxial surfaces; basal basiscopic pinnules of medial pinnae not enlarged or overlapping the lamina rachis; costules abaxially eglandular. glabrous or pubescent sparsely scaly, hairs (when present) ca. 0.2 mm long, 2-celled, acicular, scales O.fr-3 5 X 0.3-0.5, appressed, ascending, lanceolate, entire, adaxially eglandul^ sparsely to moderately pubescent, scales generally absenb the hairs 0.4- 10 mm long, 4-6-celled, antrorsely strigose; laminar tissue between veins on both surfaces eglandular. glabrous, or rarely with scattered ^ ca. 0.1 mm long; ulUmate veins on both surfaces visible or adaxially, glabrous; lamina margins eglandu ar ciliate, the “ ^ ■ long, 1- or 2-celled, ascending to spreading; mdusia absent; spores cristate. Distribution.— Venezuela. Colombia, Ecuador; 1400-2800 m. Selected Specimens Examined.-VENEZUELA. Yaracuy; Las Parchitas, Tierra Fria y Ojo de Agua, 1500-1600 m, 10 20 N, 68 57 W, 18 xESSStoSs por San Pedro de la f'’™’ g “t^der; Mpio. Tone, Corregimiento 1983, Santo S- Escobar 642 (HUA). Ltween Pamplona and Corcova, Vereda Las ^“‘ana alo^ ^y^^ Bucaramanga, 2220 m, 7 07 N, 73 0 • 78°37'W1, 24-26 Aug ECUADOR. Tungurahua: vie. of Ambato, u 1918, Rose 22379 (NY, US). Among the largedecompoundspeciesoWo^^^^^^^ is characterized by pinna rachises ^ ,o„spicuous to the large (5-12 mm long) laminae and pinnae. Sometimes naked eye, especially along ' msLt along the rachises. The smaller scales resembling scurf-like this scurf-like appearance, long cilia from the bases of large scales acce atypical by having Two specimens [Daniel 1898, Ortega 6^ ^^^fff'ribaxially acicular, erect, 0.1 mm long hairs ^y pinna rachises abaxially Similar is Megalastrum abaxially. pubescent and smaller scales along P ^ , j r -1 n R Hair details. S. Abaxial surface his. costule. and veins. O. Hair de^l. S pinnule. U. Hair detail. Scale bars rachis, costule, and veins. T. Detail 4250 [VC). O-U: Isotype, Moran et al. A-E: Sparre 1 7168 (MO). F-M: Isotype. Fay S' Foy AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 4 (2014) AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 4 (2014) SEi™ Specd^ns Exammed.-VENEZUELA. Aragua: Colonia Tov^, 110”22'N, 67“1B'W1, 1846. Funck » Schlim 412 (BM, NY. P). Barinas: Dislr. Boh™. Sjm T.ifiro ca 5 mi NW of La Soledad along Barinas-Sto. Domingo (Edo. Menda) road 1000 m 8°50'N, 70“35'W, 19 Nov 1982, Smith et al. 1398 (MO). Carabobo: Limite Distr. Bejuma Distr. Montalban, Cordillera de la Costa, vertiente norte, al este de la carretera Bejuma-Canoabo, al este de Las Rosas, faldas siiperiores y fila de la montana, bosque secundario y cafetales, 950- 1100 m, 10°16'N, 68°15'W, 26 Dec 2001, Meier 8768 (NY). Distrito Federal: Upper Catuche wood (above 1400 meters), near Caracas, above 1400 m, 110“30'N, 66°55'W], 8 Jun 1921, Pittier 9595 (GH, NY). Miranda: Distr. Guaicaipuro, Cordillera de la Costa, Serrania del Interior, vertiente norte, 6 km distancia aerea al S del pueblo Altagracia de la Montana, cuenca de la Quebrada Agua Fria, 800 m, 10°04'30"N, 67°03'W, 3 Mar 2002, Meier S' Nehbn 8939 (NY). Falcon: Sierra San Luis, cerca del Hotel Parador, 1500 m, [11°12'N, 69°43'W], 25 Aug 1978, Wingfield & van der Werff 6562 (U, UC). Lara: Moran, carretera Villanueva y Ojo de Agua, SE de Guarico, 1500 m, [9°37'N, 69°47'W], 6 Mar 1983, Ortega et al. 1635 (NY). Merida: Dpto. Sucre, Estanquez-Paramo, Las Coloradas, 1600-1200 m, [8°27'N, 71^30'W], 16 Sep 1984, Ortega 6 Berti 2120 (NY). Nueva Esparta: Isla de Magarita, Juan Griego trail, 450 m, [11°00'N, 64°10'W], 29 Jul 1903, Johnston 187 (NY, S). Portuguesa: Guanare, 2300 m, 8°39'N, 71°21'W, 24 Nov 1985, Ortega & van der Werff 2938 (MO, NY, UC). Sucre: Distr. Arismendi, Peninsula de Paria, SE de Cariipano, NE de Maturincito, Cerro La Cerbatana a lo largo del camino de La Sierra al caseno Guarico y subiendo la fila, 650-850 m, 10°38'N, 63nO'W, 9 Jan 2005, Meier et al. 11193 (NY). Trujillo: entre las antenas de relevo en el Paramo de Guaramacal y la Vega de Guaramacal, 9°13'N, 70 lO'W, 23-24 Jul 1984, Ortega et al. 2025 (MO). Vargas: Monumento Natural Pico Codazzi, Carretera Colonia Tovar-La Victoria, Sector Matapalo, 1950 m, 10°26'N, 67"17'W, 23 Jun 1999, Mostacero & Howorth 209 (VEN). Yaracuy: Distr. Bolivar, Cordillera de la Costa, Serrania de Aroa, S del pueblo Aroa, 1300-1400 m, 10 23'N, 68°54'W, 25 Aug 2002, Meier &■ Flauger 9107 (VEN). COLOMBIA. Antioquia: Mpio. Angelopolis, vereda Romeral, Hacienda La Argentina, quebrada Las Animas, 2050-2130 m, 6"08'22.9"N, 75’41'58'W, 20 Nov 2005, Rodriguez 5629 (HUA, NY); Mpio. Betania, alrededores de parcela permanente (UN), 1900-2100 m, 5"44'10"N, 76 OO'll'YV, 5 Nov 2004, Rodriguez et al. 4589 (HUA, NY). Arauca: Sarare, Santa Librada, 1300^ 1600 m, [6°29'N, 7ro2'W], 24-25 Mar 1959, Bischler 1951 (BM, MICH • aoco-Antioquia, 2100-2200 m, 4 Apr 1971, Lellinger 8^ de la Sota 917 (US). Cundin^arca: Mpio. de Venecia, Vereda Las Mercedes, alrededores de la Quebrada La Chorrera, 2200-2300 m, [4 22'N, 78 28'W], 24 Jul 1981, Diaz Santa Marta, 1219 m, (10 55'N, 73 38'W], Mar 1898- 1901, Smith 1047 [B, BM, F, GH, K, MICH, MO, NY, P, S, U, US, VT). Nanno: caime, sendero hacia Hondon, 1800 m, [1 07'N, 70 14'Wl, 25 Jun 1995. Sanchez-Baracaldo 64 (MO, UC). Norte de Santander: road from Pamplona to ^ and Rio Mesme, 2800-3000 m, N, 72 39 W], 27-28 Feb 1927, Killip & Smith 19998 (NY). Santander: vic. MORAN ET AL.: ANDEAN MEGALASTRUM 217 of Las Vegas, 2600-3000 m, [6°33'N, 73°08'W], 21-23 Dec 1926, Killip & Smith 16136 (NY). Tolima: Mariquita, Combaima, 1200 m, [5°12'N, 74"54'W1, Jan 1843, Linden 1020 p.p. (B, P). Megalastnim subincisum is characterized by pinna rachises abaxially without hairs but with linear-lanceolate, golden brown scales. The veins adaxially are glabrous to subglabrous (only one or two hairs present), and sparse hairs are often present on the costules abaxially, especially distally. In South America this species greatly resembles M. aureisquamum from Peru and Bolivia. They can be distinguished morphologically only by the characters of the rhizome scales as given in the key. Christensen (1920) construed Megalastnim subincisum in the wide sense, occurring from Mexico to Bolivia. He admitted much variation /he type ot laminar scales and the presence versus absence of hairs and glands. We tind, however, that indument characteristics can be used to divide M subincisum into smaller, less variable species. In this restricted sense, M. subincisum is more narrowly distributed than conceived by Christensen, occurring only m toe Momct and Honduras, toe West Indies, and Colombia and northern Venezuela. M»t of the Central American specimens identified by Christensen^ M. ^ referahle to M. galeottii (M. Martens) R C. Moran & J. Prado (Moran & Prado, 2010). 44. Megalastnim subtile R. C. Moran. J. Prado & Sun due, ECUADOR. Morona-Santiago: small ravine ca. ^ of Li ■ ’ 78”24’03'W. 960 m, 3 Mar 2005. R. C. Moran D. Aviles 7808 (holotype: QCA!; isotypes: NY! [barcode 1053900), QCNE.). Figs. 13D, 14A, 350-U. Rhizomes erect, scales 10.0-20.0 X 1-2 nun. ascending, linem. bro^. dull, firm, the margins denticulate, the surfaces smooth (not setu J’ ^ ^ 1.0 m long; petiole base scales ^ loosely ascending; laminae 0.5-0. 6 hLs abaxially; basal medially 2-pinnate-pinnatifid; l^ma rac^ pinnae ca. 20 cm long, equilateral or yearly , P ^ ^ ^ ^ eglandular, without hairs, sparsely scaly, e giiform to linear, brown, appressed to loosely spreading gcaly or lacking scales, slightly tortuous, entire, adaxially pubesce , p ^ ^ ^ q ^ j^m, filiform, hairs ca. 0.3 mm long, antrorsely strigose, medial pinnae not entire, appressed, dull, brown; basal basisc p P abaxially eglandular, enlarged'^or overlapping the lamina raclus; costul^ 3-celled, sparsely pubescent and scaly, ^ prown, entire, adaxially ascending to strigose, the scales ' rachises adaxially; laminar pubescent, the hairs like those on th p glabrous; ultimate tissue between veins ab^ially obLure, glabrous; lamina margins veins abaxially visible, glabrous, adaxial y , ^ or 2-celled, acicular, eglandular, ciliate, the hairs 0.2-0.3 mm ascending to substrigose; indusia absent; spor 218 AMERICAN FERN JOURNAL: VOLUME 104 NUMBER - Distribution. — Ecuador, eastern side of the Andes; 900-1180 m. Specimens Examined. — ECUADOR. Morona-Santiago: trail along Rio Inimkis, ca 3 km NE of village of Inimkis (San Luis) toward foothills of Cord. Cucutu, 900- 1000 m, 02°23'S, 78°06'W, 14 Nov 1995, Ollgaard & Navarrete 1426 (AAU). Napo; Archidona, Reserva de Biosfera Sumaco, cordilla de Galeras, bosque protector de la comunidad de Mushullacta, 1180 m, 0°45'50"S, 77°33'28"W, 27 Mar 2003, Huaylla 831 (MO, QCNE, UC). Among the decompound species of Megalastrum, M. subtile is distinctive by relatively small leaves (up to 1 m long) and the pinna rachises abaxially hairless with filiform to linear entire scales. It resembles M. martinicense by lack of hairs and filiform scales, but that species differs by large leaves, strongly denticulate scales, and occurrence in northern Venezuela and Colombia. Also similar is M. reductum, which differs most conspicuously by pinna rachises abaxially densely and uniformly pubescent by minute hairs. The specific epithet refers to subtile differences between this species and M. reductum. 45. Megalastrum tepuiense R. C. Moran, J. Prado & Sundue, sp. nov. Type.— VENEZUELA. Bolivar: Sierra de Lema, Cabeceras del Rio Chicanan, 80 km (en linea recta) al SW de El Dorado, at base of high NNE-facing sandstone bluffs, 6°05'N, 62°W[sic], 500 m, 27 Aug 1961, /. A. Steyermark 89513 (holotype: VEN! [barcode 52682]; isotypes: NY! [barcode 000870956], US! [barcode 00798656, 00798657, 00798658]). Figs. 13B, 30E-G, 32D. Rhizomes not seen; leaves up to ca. l.l m long; petiole base scales 12.0-20.0 X 0.3-4).5 mm, ascending to spreading, filiform, golden brown, lustrous, flat or twisted denhculate; laminae 0.4-0.7 m long, basally 3-pinnate-pinnatisect, medially 2-pinnate-pinnatisect; lamina rachises without hairs abaxially; basal pinnae 25.0-35.0 cm long, inequilateral; pinna rachises abaxially eglandular, glabrous sparsely scaly, scales up to 3.0 X 0.2 mm, light brown, linear- anceolate, lustrous, spreading to slightly appressed, flat or twisted, subentire adaxially eglandular, densely pubescent, scaly, the ^-0 mm long, 3-6-celled (the contents of the cells brown), spreading, sometimes with bifid teeth; basal rarbiQ r ^ of medial pinnae not enlarged or overlapping the lamina 1 0 U n“l '’8'“dular, sparsely pubescent, scaly the hairs 0,5- rachisBs ahalieii u . u ’ a«cular, scales like those of the pinna • “P to 2 mm long, adaxially eglandular, densely tissue between ve^ * u pinna rachises, scales absent; laminar bo7 srf^rvwr ™ eglandular, glabrous; ultimate veins on to^Dse rf *e cn ^ ? eglendular, sparsely pubescent, the hairs like Wng,thm^ the^airs 0.5-1.0 mm long. MORAN ET AL.; ANDEAN MEGALASTRUM 219 Megalastrum tepuiense is characterized by decompound laminae, petiole bases with linear-lanceolate scales, pinna rachises abaxially without hairs but with long, linear-lanceolate scales, and costules and veins on both surfaces pubescent by whitish hairs 0.5-1.0 mm long, 3-6-celled. This species and M. adenopteris are the only species of Megalastrum with acicular hairs on fugacious indusia (the indusia often apparently absent in mature sori). Only two other species of Megalastrum occur in the tepuis region of southern Venezuela: M. biseriale and M. crenulans. The first species can readily distinguished by 1-pinnate-pinnatifid laminae, and the second by large, persistent indusia. , i • Similar is Megalastrum praetermissum, which differs by the laminae abaxially between the veins minutely and finely pubescent, rachis scales strongly denticulate, and sori without acicular hairs. , Megalastrum tepuiense resembles M. subincisum by lamina ® pinna rachises abaxially glabrous. It differs by costules an mi ^ ^ , ultimate segments abaxially pubescent by long (0.5-1 0 hairs (vs. glabrous to sparsely and inconspicuously Pubescent wiAhatrs on y 0.2-O.5 mm long). Megalastrum tepuiense occurs only m and does not overlap geographically with M. subincisum, w ic northern Venezuela, Colombia, and the Antilles. 46. Megalastrum vastum (Kunze) A. R. Sm. & R- C. Mor^, t iJjj^gai 129. 1987 [published 3 May 19881. Polypodium ^ ^359 9:50. 1834. Phegopferis vasfa (Kunze) Mett Fil Lechl„ ^ Nephrodium vastum (Kunze) Hieron., Bot. ^ Hedwieia 46:347. ifieg., non Baker (1867). Pryopferisvasta (Kunze) Hieron Hed^^ 1907. Type.-PERU. Huanuco: “inter Pampayaco by Ventanilla de Cassapi,” 1829, E. Poeppig 217 ’ 20^69277))^ Tryon and Stolze, 1991: B! [barcode 200069274, 200069276, 200069277Ji Figs. 9L-Q, 33A, 36D. Polypodium extensum C. Presl. Rel. HaeiA. Forster (1786). Tvpe.-PERU. s.d., Haenke s.n. (holotype. PK BM!,NY!). n V n 7_1 n nun appressed to spreading. Rhizomes erect, scales 10.0-20.0 X 0.7 1. ’sgiy denticulate; leaves linear-lanceolate, brown, dull or lustrous, twis ’ P, . g b^t spreading to 1.0-2.2 m long; petiole base scales like those of ^""[^^a.Jpinnatifid, loosely ascending; laminae 1.0-1.5 m nubescent abaxially; basal medially 2-pinnate-pinnatisect; lamina rac « p^ abaxially eglandular or pinnae 20-25 cm long, puberulent, sparsely scaly, the rarely sparsely glandular, densely and even Y P 0. 1-0.2 mm long, glands (when present) sessile, spherical, yellowis ’ 3 3 X 0.5 nun, 1-3-celled, acicular or lax, erect to strigose, e , ^ adaxially densely entire or with a few teeth distally, flaccid, p ^ „pU Id- basal basiscopic pubescent, eglandular, the hairs 0.6-1.0 nun long. AMERICAN FERN JOURNAL: VOLUME 104 NUMBER ■ pinnules of medial pinnae not enlarged or overlapping the lamina rachis; costules abaxially eglandular or nearly so, densely pubescent, sparsely scaly, the hairs like those of the pinna rachises, the scales ca. 1 X 0.2 mm, like those of the pinna rachises, adaxially with indument like that of the pinna rachises but scales absent; laminar tissue between veins abaxially eglandular or rarely glandular, densely pubescent, the hairs ca. 0.1 mm long, l(-2)-celled, erect to curved-ascending or oppressed, acicular, glands ca. 0.1 mm long, 2-celled, stalked or sessile, yellowish to orangish; adaxially puberulent, the hairs like those on the abaxial surfaces, often more abundant toward the segment margins; ultimate veins visible on both surfaces, pubescent on both surfaces or sometimes glabrous adaxially, eglandular or very sparsely glandular, the hairs and glands like those on the laminar tissue between the veins, or hairs sometimes longer, being 0.5-1. 0 mm long, 3-5-celled; lamina margins eglandular, ciliate, the hairs 0.2-0.3 mm long, 1-3-celled, acicular, substrigose, ascending; indusia consisting of a tuft of minute reddish fugaceous hairs (often apparently absent); spores cristate. Distribution. — ^Venezuela, Colombia, Ecuador, Peru, Bolivia; 400-2200 m. Selected Specimens Examined.— VENEZUELA. Yaracuy; Distr. Nirgua, Cordillera de la Costa, Macizo de Nirgua, SW de Nigua, entre el Picacho de Nirgua y San Antonio-Buenos Aires, 1200-1400 m, 10°08'N, 68"38'W, 5 Mar 2004, Meier & Nehlin 10084 (UC). COLOMBIA. Antioquia: Mpio. Liborina, Vereda Barrio Nuevo, quebrada La Honda, limites con el municipio de Sabanalarga, 1910 m, 6 46'54.3'TJ, 75H7'34.2"W, 26 Sep 2002, Rodriguez et al. 3636 (HUA, NY). Caldas: Pueblo Rico [Neira], Cordillera Occidental, Vertiente Occidental, 1700—1900 m, [5^08'N, 75=31'W1, 17 Feb 1946, von Sneidern 5236 p.p. (F, US). Norte de Santander: Cordillera Oriental, region del Sarare, Hoya del Rio Cubugon entre El Caranoy El Indio, 600-470 m, l7°01'N, 72"ll'Wl, 12 Nov 1941, Cuatrecasas 13021 (GH). Risaralda: El Cedral, trocha a El Silento, 20 km de Pereira, 2140 m, [5=18'N, 7542'W], 15 Oct 1989, Rangel et al 5762 (MO). ECUADOR. Bolivar: Chazo Juan, 600 m, t0=23'S, 79 05'W1, 1943, Acosta 6517(F). Carchi: near Maldonado, 1400 m, [0°54'N, 78"06'W1, 29 Jul 1989, van der Werff 8- Gudino 10711 (AAU, MO, NY). Chimborazo: El Limdn, Mt. Chimborazo, 1000 m, [0°28'S, 78 49'Wl, Jul 1860, Spruce s.n. (K). Cotopaw: anton La Mana, Reserva Ecologica Los Ilinizas, sector del Brasil, acceso desde Occidental, 1480 m, 0 40'37"S, 79 05'09"W, 13 Jul Rnl’ 9088 (MO, QCNE). El Oro: Road from Pinas to Sta. Es^; ^ 79'44'W1, 7 Oct 1979, Dodson et al. 8958 (US)^ Biologica Station, Mache mountains. 35 km W of CfaTfr w ^ ^ “o-eoo 0 21'N. 79 44'W, 14 May 1995, SWW of r Rios: Bosque protector Cerro Sam^- 2^1^ ^ La Clementina. 600-730 m, 0’40'S. 79’20'W. CuaialitoR.™ PI Pichincha: CantOn Quito. W" ' 'to R^urve. 10 km W of Chiriboga, Km 59 of old road Quito-SanM MORAN ET AL. : ANDEAN MEGALASTRUM 221 Domingo, 1900 m, 0°14'S, 78°48'W. 10 Jul 1991, Fay & Fay 3352 (AAU, MO, QCA, QCNE); New road Sto. Domingo-Quito 6-8 km E of Tandajapa. 1150 m, [OHIO'S, 78=46'W], 23 Jan 1981, Balslev et al 1693 (AAU, MO, NY). PERU. Huanuco; Leoncio Prado, km 478 on Lima-Tip Maria road. Thicket along road , following stream to waterfall, 1400 m, [9'05'S, 76 20 W], 6 Jun 1981, Young Sullivan 852 (MO). BOLIVIA. Cochabamba: Chapare, Territorio Indigena Parque Nacional Islboro-Secure, Cordillera de Mosetenez, Laguna Carachupa. 1300 m. 16°14'S, 66"25'W, 29 Aug 2003, Kessler 6^ Jimenez 12995 (UC). Among the large decompound species of the genus, characterized by pinna rachises evenly pubescent ab».ally ^ ^ 0.2 mm long, pinna rachis scales entire or nearly so, the bssne veins on both Lfaces pubescent by short hairs ca 0.1 mm '“"B- and mdusu absent. In many specimens, the hairs on the adaxial surface of the “ more frequent toward the margins. Glands are often lackmg on lamma^ tissue and veins abaxially. except for 0%oard 1559 and Bakiev 1693 wh.ch have spherical, sessile, orange glands abaxially along e vei ' ... j MegLstrum vastun, resembles M. nL^en the lamina size and cutting but differs from by the ‘^'nae be we“ adaxially with hairs ca. 0.1 mm long, acicular, hyatae, (vs. longer and lax). Also, its scales differ, being are and often brown; in contrast, those of M. am firmer, shiny, and yellowish brown. The sc ® ^ aureisquama and inconspicuous in M. vastum, but ^es M. owophilum, a M. mollicoma. Megalastrum vastum also great y denticulate species that differs by petiole and rachis scales golden brown and (vs. dark brown and entire or nearly so). cnecies. Some of the Christensen (1920) had a mixed concep different species. The specimens he cited are considered by us to P represent either M. specimens he cited of ^^Salastnim vastum o Colombia [Lehmann skutchii or M. palmense (Moran and Prado 20 L Paraguay [Hassler 8924) represents M. reductum; the e ci ^ ^ j^oran (Moran et al 10421) represents M. umbrinum (C. Chr.) A. . _ ^senis M. miscellum. 2009a); and the specimen from Peru [Spruce 4718) P ^ Photos distributed by US (Morton negative polypodium vastum; ■ llVLcimen and therefore it is on their labels that the plant depicted however, there is no locality information uncertain whether it is a type. with the specimei Names OF Uncertain Appucation OR7 1905 nom. nov. tor Jyepmuuiu. Dtyopteris fusca C. Chr., Index Fihcum 2 • pichincha: “Crece polylepis Sodiro, non Baker (1891). Type. 222 AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 4 (2014) en los bosques de los Colorados entre 400—500 m,” s.d., A. Sodiro s.n. (P?- We did not find a type of this name. Sodiro did not designate types, and a specimen of his with sufficient locality information and date was not found by either us or Christensen (1920). A specimen at S (seen on-line) was annotated as the type of Nephrodium polylepis Sodiro, but its collection information does not agree with the protologue. The specimen was collected by Sodiro near Oyacachi in January 1901, but has the wrong locality, and the annotation was made after the name Nephrodium polylepis Sodiro had been published. The specimen was annotated by Christensen as “D. vasta” {=Megalastrum vastum). Polypodium honestum Kunze, Linnaea 9:49. 1834. Phegopteris honesta (Kunze) Mett., Pheg. u Aspid. Nr. 59. 1858. Dryopteris honesta (Kunze) C. Chr., Index Fil. 271. 1905. Ctenitis honesta (Kunze) R. Tryon & A. Tryon, Rhodora 84:127. 1982. Megalastrum honestum (Kunze) A. R. Sm. & R. C. Moran, Amer. Fern. J. 77:128. 1987 [published 3 May 1988]. Type.— PERU. Huanuco: Pampayacu, 1829, E. Poeppig 22 (LZ, destroyed). The type was destroyed during WWII, and apparently no duplicates exist. The description is inadequate to apply the name. Morton negative 5268 (MICH, UC, US) shows a photo of a Poeppig collection at H identified as this species, but it is doubtful that this specimen is the type. The photo depicts a plant that is 2-piimate-pinnatisect, whereas according to the original descriptions the lamina was only 1-pinnate-piimatifid. Unusual Specimens We have not been able to identify the specimens cited below with any of the species recognized above. These specimens might represent new species, but more material is needed to confirm this. Bonino et al. 1186 (MO). Bolivia. This specimen resembles Megalastrum aureisquama but differs by dark appressed scales along the lamina rachises (not golden brown scales). The cutting of the laminae also differs, being only 2- pmnate pinnatifid basally (vs. 3-pinnate-pinnatifid) and with the proximal pinnae soon becoming adnate the pinna rachises distally. The pinnules are obtuse to acute (vs attenuate or acuminate) higher elevation (2700 ml. Widely adnate to the pinna rachis. MORAN ET AL.: ANDEAN MEGALASTRUM MORAN ET AL.: ANDEAN MEGALASTRUM \ Copel. (1) AMERICAN FERN JOURNAL: VOLUME 104 NUMBER Ctenitis hirsutosetosa (Hieron.) Lellinger (18) Ctenitis honesta (Kunze) R. M. Tryon & A. F. Tryon (see Uncertain Applicatioi Ctenitis karsteniana (Poir.) Vareschi (38) Ctenitis mollicoma (C. Chr.) Ching (25) Ctenitis pleiosoros (Hoolc. f.) C. V. Morton (33) Ctenitis pulverulenta (Poir.) Cope). (38) Ctenitis subincisa (Willd.) Ching (43) Ctenitis villosula (C. Chr.) Ching (4) Dryopteris acrosora (Hieron.) C. Chr. (1) Dryopteris adenopteris C. Chr. (2) Dryopteris andicola C. Chr. (4) Dryopteris andicola C. Chr. f. lehmanniana (Hieron.) C. Chr. (4) Dryopteris andicola C. Chr. f. spruceana (Hieron.) C. Chr. (4) Dryopteris biserialis (Baker) C. Chr. (6) Dryopteris crenulans (F§e) C. Chr. (10) Dryopteris crenulans (F6e) C. Chr. f. glandulosa (Rosenst.) C. Chr. (10) Dryopteris erythrostemma (Christ) C. Chr. (38) Dryopteris fusca C. Chr. (see Uncertain Application section) Dryopteris hirsutosetosa Hieron. (18) Dryopteris karsteniana (Klotzsch) Hieron. (38) Dryopteris leptosora C. Chr. (22) Dryopteris macrotheca (F6e) C. Chr. (6) Dryopteris microsora Kuntze (22) Dryopteris mollicoma C. Chr. (25) na (Klotzsch) C. ( Dryopteris platyloba (Baker) C. Chr. (32) Dryopteris pleiosoros Svens. (33) Dryopteris pulverulenta (Poir.) C. Chr. (38) Dryopteris squanrosissima (Sodiro) C. Chr. (42) Dryopteris squamosissima (Sodiro) C. Chr. var. bogotensis I Dryopteris subincisa (Willd.) Urb. (43) Dryopteris subincisa (Willd.) Urb. var. bogotensis Hieron. ( Dryopteris subincisa (Willd.) Urb. i Dryopteris tarapotense (Baker) C. Chr. (32) Dryopteris vasta (Kunze) Hieron. (46) Dryopteris villosa (L.) Kuntze var. tomentosa Rosenst. (2) Dryopteris villosa (L.) Kuntze var. glandulosa Rosenst. (10) Dryopteris villosa (L.) Kuntze var. inaequalis Gilbert (4) Dryopteris yungensis Christ & Rosenst. (6) Megalastrum acrosonim (Hieron.) A. R. Sm. & R. C. Moran (1) Megalastrum adenopteris (C. Chr.) A. R. Sm. & R. C. Moran (2) Megalastrum aequatoriense A. Rojas (6) alticola M. Kessler & A. R. Sm. (3) . Sm. & R. G Moran (4) 7 andicola (C. Chr.) A. R. Sm. & R. C. Moran f. lehmai MORAN ET AL.: ANDEAN MEGALASTRUM . Prado & Sundue (9) & R. C. Moran (10) indue (14) o & Sundue (15) er & A. R. Sm. (5) Megalastmm bidecoratum (Lellinger) A. R. Sm. & R. C. K Megalastnun biseriale (Baker) A. R. Sm. & R. C. Moran Megalastrum bolivianum M. Kessler & A. R. Sm. (7) Megalastrum ciliatum M. Kessler & A. « ffll Megalastrum clathratum R. C. Moran, • ■ , crenu/ans (Fee) A. R. Sm I ctenitoides A. Rojas (11) n R. C. Moran, Megalastrum dorsiglabrum A. Rojas (30) Megalastrum falcatum A. Rojas (13) Megalastrum fibrillosum (Baker) R. C. Mo Megalastrum fimbriatum R. C. Moran, J. I Megalastrum fugaceum R. C. Moran, J. Prado & Sundue linj Megalastrum galapagense R. C. Moran, ). Prado & Sun ue Megalastrum hirsutosetosum (Hieron.) A. R. Sm. & R- C. Motm Megalastrum honestum (Kunze) A. R. Sm. & R- 0. Moran Megalastrum insigne R. C. Moran, J. Prado & Sundue ( Megalastrum karstenianum (Klotzsch) A. Rojas (38) Megalastrum laxipilosum A. Rojas (25) uUsirun, ,apL™m (C. Chr.) A. R. Sm. . R C. M Megalastrum marginatum M. Kessler & A. R. Sm. |20 Megalastrum martinicense (Spreng.) R- C. Moran, J. Megalastrum microsorum (Kuntze) Stolze (22) Uegalastrva. n,iscellun. R. C. Morm. J. Prado & Sundua (23) Megalastrum molle A. R. Sm. (24) coma (C Chr.) A. R Sm. & R C. Moran (25) an R. C. Moran, J. Prado & Sundue (26) yiegaiastrum mgromarginatum R. C. Moran, ^ J™ ° degalastrum obtusum R. C. Moran, J. Prado & 'i® Megalastrum oellgaardii R. C. Moran, J. Prado & ^ogalaslram oreopMua, R C Mo^l ^ Megalastrum peruvianum R. C. Moran, J. Moran (32) Megalastrum platylobum (Baker) A. R^Sm. & R^ ^ Megalastrum pleiosoros (Hook, f.) A. R- ^ gundue (34) Megalastrum polybotryoides R. C. Moran, )• a (35) Megalastrum praetermissum R. C. Moran, J- egatastrum pubescens A. Koja^ ^ egalastrum pubirhachis R. C. Moran, J. Moran (38) ’egalastrum pulverulentum (Poir.) A. R. m. Megalastrum reductum A. Rojas (39^ ^ ^ ^ ^ Sundue (40) „ M. Kessler » „ c. Moran (42) "‘^‘TiRSrArcMoranrar.lv^-- AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 4 (2014) Megalastrum ubincisum (Willd.) A. R. Sm. & R. C. Moran (43) mbtile R. C. Moran, J. Prado & Sundue (44) epuiense R. C. Moran, J. Prado & Sundue (45) rastum (Kunze) A. R. Sm. & R. C. Moran (46) Megalastrum villosulum (C. Chr.) A. R. Sm. & R. C. Moran (4) Megalastrum yungense (Christ & Rosenst.) A. R. Sm. (6) Nephrodium acrosorum Hieron. (1) Nephrodium biseriale (Baker) Diels (6) Nephrodium microsorum Hook. (22) Nephrodium polylepis Sodiro (see Uncertain Application section^ Nephrodium squamosissimum Sodiro (42) Nephrodium subglabrum Sodiro (32) Nephrodium vastum (Kunze) Hieron. (46) Nephrodium villosum L. var. karstenianum (Klotzsch) Jenman (31 im Mett. ex Hieron. f. opacum Nephrodium villc Nephrodium villc Nephrodium villc ubincisum (Willd.) Hook, ex Sodir ubincisum (Willd.) Jenman (43) Phegopteris epierioides F6e (21) Phegopteris hirsute F6e (38) Phegopteris honesta (Kunze) Mett. (see Uncertain Application sei Phegopteris karsteniana (Klotzsch) Mett. (38) Phegopteris martinicensis (Spreng.) Foum. (21) Phegopteris subincisa (Willd.) Fee (43) Polypodium barbatum Desv. (38) Polypodium biseriale Baker (6) Polypodium extensum C. Presl (46) Polypodium honestum Kunze (see Uncertain Polypodium karstenianum Klotzsch (38) Polypodium platylobum Baker (32) Polypodium pleiosoros Hook. f. (33) Polypodium pulverulentum Poir. (38) Polypodium subinch Polypodium tarapotc sum Willd. (43) ense Baker (32) Kunze (46) Application section) Appendix 3: List of Exsiccatae and listed in Appendix 1. Collection numbers in boldface are ty^s. Abbott, J. R.: 15112 (39) Acebey, A.: 595 (14); 730 (20) Acosta, S. M.: 6517 (46) Adsersen, A.: 9, 84, 774, 887, 1248, 1659, 1819 (33) MORAN ET AL.: ANDEAN MEGALASTRUM 229 Agostini. G.; 207 (38) Aguilar, P.: s.n. (14) Albert de Escobar, L.: 943 (39) Albert de Escobar, L. et al.: 3900 (43); 4476 (23) Alston, A. H. G.: 8026 (38); 8730 (39) Altamirano, S. et al.; 826 (3) Alvarez, R.: 18 (43) Andre, E.: 91 (38) Ankersen, H.: 27 (19); 140 (16); 168 (35) Ankersen, H. & Kragelund. C.: 211 (28) Arbelaez, A. et al.: 87, 119 (43); 276 (231; 287 (21) Arroyo, L.: et al. 1912 (4) Asplund, E.: 16407 (39) Atehortua, L.: 107 (46) 15 (19) Brbom, 'ps^^OdW); 2499 (7); 2504 (3); 2587 (19); 3682 (15) Barfod, A. et al.: 48891 (18) Beck, G.: 1705 (35) Bell, P. R.; 252 (46); 920 (39) Bittne ,1 753 (6); 761, 1113 (5); 1116 (31): ' ;n): 1104, p.p. (16): 1104. p.p. (19) (18); 258, 279 (19); Bonino, E. et al.: 734 (16): 746 (3): 750 (5); 1178 (16); 1186 (unusual spec Bonino, R.: 66 (7); 117 (19): 143 (7);: Borgtoft, P. et al.: 104324 (18) Boyle, B. et al.: s.n. (3) Bredemeyer, F.: s.n. (43) s .... ..... .« ■“ “ Si;';: Calleias, R. et al.; 311a (H) Camp, W. H.; 3786 (39) Campos. I. &Corrales, S.: 3593(19) Campos. J.& Garcia, Z.: 4049 (19) Campos, J. & Lopez, P.: 4911 (35) Campos, J. & Vargas, W.: 3892 (19) Cardenas, M.: 2849 (2) Cardona, F.; 1590 (10) 3. 1137a (16): 1160 AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 4 (2014) Castilldn, L.: 40958 (16) Cer6n, E. C.: 1483, 1489 (13) Clark, J. L. & Watt, C.: 862 (46) Cook, F. O. & Gilbert, G. B.: 1128 (16) Cornejo, X.: 8165 (46) Cornejo, X. & Bonifaz, C.: 5459 (22) Croat, T. B.: 56541 (42); 71469 (25); 72158 (46) Croat, T. B. & Menke, M.: 89543 (14) Croat, T. B. et al.: 91009, 91135A, 91165 (13) Cuatrecasas, J.: 11185 (35); 13021 (46); 23889 (19) Daniel, Bro.: 2013 (23); 4473 (19) Daniel, H.: 1898 (42) Darwin, C.: s.n. (33) de la Sota, R. E.: 4069, 4298 (16) Diaz, S.: 3130 (43) Dodson, C. H. et al.: 7215 (46); 7844 (39); 8958 (46) Dodson, C. H. & Gentry, A.: 10088 (39); 12280 (22) Dombey, J.: s.n. (2) Dorr, L. J. & Stergios, B: 8720 (38) Dudley, R. T.: 11512 (35) Eberhardt, D. et al.: 137, 142 (41); 204 (6); 402 (20); 513 (6) Eliasson, U.: 379, 382, 1354, 1851 (33) Escobar, L. et al.: 3854 (43) Evoy, J.: 78 (39) Fagerlind, F.: 3036 (33) Fay, A. & Fay, L.: 2051 (14); 2205 (6); 3287 (19); 3316, 3335. 3349, 3350. 3351, 3352 (46); 3370, : (19); 3662, 3663 (35); 3827 (19); 3845 (38); 3870, 3876 (16); 3910, 3919 (37); 4014 (14); 4 4046. p.p. (23); 4046, p.p., 4097, 4232, 4250 (35); 4381 (14); 4382 (23); 4403 (35); 4595, 4626 Fendler, A.: 156, 202 (43); 447 (38) Fernandez, E. et al.: 2362 (4); 2366 (35) Foster, M. S.: 85-144 (1) Foster, R. & Wachter, T: 7423 (14) Foster, R. et al.: 11598 (14) Freire, E. et al.: 1480 (16) Fuentes, A. et al.: 9013 (8); 9034 (41); 11098 (20); 14359 (14) Funck. H. & Schlim. L. 282 (21); 407, 412 (43); 975 (38); 1575 (43) Gentry, A. et al.: 22867 (19); 60443 (25); 60492 (25) Giraldo, F. & Mejia. I: 2098 (46) Gonzales, N. et al.: 7 (43) MORAN ET AL.: ANDEAN MEGALASTRUM Marling, G. & Andersson, N. J.: 22459 (12 Marling, G. et al.: 7396 (35) Merrera, G.: 9130 (25) Merrera, G. & Chacon, A. 2794 (11) Moldridge, L. R.: 1622 (39) Molm-Nielsen, L. et al.; 4502 (18); 4507 (: 24853 (6); 26458 (37); 26896 (14); 26£ Moover, S. W. & Wormley, S.; 1620 (46) I; 4547 (23); 4618 (14); 6999 (16); / Muamantupa, I. & Quispe, W: 3836, 3837 (16) Huaylla, M.: 792 (18); 831 (44) iz! I. & Huaylla, M.: 2518 (20); 2547 (14); 2551, 2583 (20); 2589, 2589 (14); 2590 (20) !Z, I. & Moguel, A.; 1428 (4); 1489 (16); 1641 (3) ■z, I. & Serrano, M.: 2398 (26) >z, I. et al.: 2468 (2) on, J. R.: 187 (43) (26); 6379 (4); 62 7189 (16); 7379 (5); 7426 (4 Z 7630a p’);' 7631 (35); 7644 (5); 7751 (41); 7789 (5); 7907 (41); 79 (14)- 9088 9105, 9488 (16); 9900 (41); 10019B (14); 10774, 10897 [c U4)riTl3T (35711604 (14);’ 11621 (35); 11944 (6); 11945 (3); 12948 (14); 12950 (. J(14) 7522 (5); 7554 C 8126 (14); 8256 Su:L^%"c"ir5'S; 16136, 19998, 43);304n^ Killip, (6); 24659 (14); 26021 (6); 26037 Killip, E. P. & Varela, J. D.: 34671 (t Knapp, S. & Mallet, Kromer, T. et al.: It Labiak, P. M.: 4730 (19); 7369 (4); 7416 (19): 8924, 8925 (39) 1759 (22) 232 AMERICAN FERN JOURNAL: VOLUME 104 NUMBER - Lellinger, D. B. & de la Sota, E. R.; 815 (19); 840 (18); 917 (43) Leon, B. et al.: 3047 (31) Liesner, R. & Holst, B.: 21585 (10) Liesner, R. et al.: 12829 (43) Limminghe, A. M. A.: s.n. (38) Linden, M.: 184 (21); 243, 843 (4); 844 (38); 1020, p.p. (21); 1020, p.p. (43); 1033 (46); s.n. (21) Lindig, A.: 159 (4); 331 (42) Little, E. L. & Little, R. R.; 8597 (27); 9415 (1) Lliully, A. et al.: 101 (16) Lugo, H.: 2064 (35) Luteyn, L. J. et al.: 6590 (13) MacDougal, J. M. et al.: 3886 (11) Madrinan, S. & Barbosa, C. E.: 187 Madsen, J.: 86916 (22); 87002 (6) Madsen, J. & Rosales, C.: 8162 (23) Maguire, B. et al.: 36871 (30) Martinez, O. & Prado, J.: 1893 (8); 1894 (16) Meier, Meier, W.: 2862, 8768 (43) W. & Flauger, N.: 9107 (43) W. & Gutierrez, R.: 12333 (38) W. & Molina, [?]; 9226 (43) Mejia, R. Mellado, jt al.: 11193 (43) et al.: 3312, 3517 (4) . F.: 1686 (19) . F. & Becerra, E.: 1754, 1803, 1835 (3) al.: 11879 (unusual specimen) ^oran, R. C.: 3510 (6); 3522 (16); 3531 (6); 3543 (39); 3546 (6); 3546B (38); 3546A (46); 3567 (l); 3583A, p.p. (7); 3583, p.p. (13); 3591 (6); 3594 (37); 3609 (39); 3689 (13); 3707 (14); 5911 (16); L C. & Rohrbach, C.: 5133, 5160 (: , 5220 (19); 5229 (6); 5380, p.p. (16); 5380, p.p. Moran, R. C. et al.: 5949 (46); 6925 (39); 7443 (6); 7542 (19); 7545 (6); 7608 (44); 7634 (14) Moritz, J. W. K.: 202, p.p. (21); 202, p.p., 385, p.p., 389, 435 (43); 459 (38); s.n. (43) Morrone, O. et al.: 2863 (16) Mostacero, J. & Howorth, R.: 209 (43) Navarrete, H.: 538, 539, 540 (34); 725 (7); 789 (6); 798, 847, 896 (19); 1684 (6) Navarrete, H. & Asimbaya, P.: 1646 (6) Neill, D. et al.: 5956 (6) MORAN ET AL.: ANDEAN MEGALASTRUM Nunez, R.: 760 (35) Nunez, R. & Huaylla, H.: 178 (16); 199 (23); 279 (4) 74966 (19); 98257 (6); 98379(35); 98410 Ufl: 98413 (40); 98471 (23); 98888 (6); 98891 (25); 98919 (3); 98930 (16); 98942. 99593 (6); 99738 (37) 9 ^ (46); 99763 (13); 99890 (37); 99904 (19); 99993 (6); 100629 (46); 100631 (19); 100743 (14) 011gaard, B. & Borchsenius, F: 100674 (39) 37844 (46): 57852 (13); 74815 (24): 90906 (7), 90909 (16 . yu 99562 (1); 98468, 98694, 99792 (18): 99923 (14); 100773 (22) Ortega, F.; 2934 (43) & Berti, L.: 2120 (43) & Smith, A. R.: 2476 (42) , (43); : Paniagua C. N. Pedersen, H. B. et al.: Peyton, B.: 1497 (16) Peyton, B. & Peyton, S Stergios, B: 1502 (43) .an derWerff.H.: 2938, 2951 (43) al.: 1635, 1825 (43); 2013 (38); 202 ' !o 3?4070 (18); 5844 (40); 10986 (18); 12113 (7); 12645 (16) & Cen, [?]: 13883 (22) & Clark, J.: 12644A (25) & Tirado, M.: 13214 (23) & van der Werff, H.: 3600 (19) et al.: 8764 (14): 8766 (18); 9569 (37); 9724 (6) N. et al.; 4216 (35) ,2 (35) 3 (39) Pittier, H.; 9498, 9595 (43); 10029 (38) Plowman, T. et al.: 3942 (37) Poeppig, E.: 34 (46) Portuga), A. 8 )im6ne2 I.; 452 (41) 53, (20) Portugal, A. et al.: 223 (41): 246 (7), 250, i Quijada, C.; 25 (43) Qoipuscoa, V. » Baniales, I - „ ^described species); 2340 (1); 2404 (19, Quipuscoa. V. et al.; 1083 (1); 1088 (probably an AMERICAN FERN JOURNAL: VOLUME 104 NUMBER 4 (2014) Rangel, O. et al.: 5762 (46) Rodriguez. W. et al.: 3209, 3598 (43); 3636 (46); 4589 (43); 4591 (46); 4805 (25); 4905 (43); 4915 (38); 4934 (6); 5559 (36); 5575 (43); 5578 (46); 5594, 5629 (43) Rose, J. N.: 22379 (42) Rothfels, C. J. et al.: 3714 (19) Rusby, H. H.: 425, p.p. (8); 425, p.p. (31); 426, p.p. (31); 426, p.p. (8) Sanchez-Baracaldo, P.: 42, 60 (25); 64 (43) Sanin, D. et al.: 5176 (39) Santa, J. & Escobar, R: 642 (42) Schunke, C.: 45, 46, 47, 66, 87, 225A, 453, 510, 704, 841, 862, 1555 (6); s.n. (18) Seidel, R. et al.: 9035 (35) Serrano, M. et al.: 5225 (16); 6873 (2) Silverstone-Sopkin, P.: 9088 (46) Silverstone-Sopkin, P. et al.: 2933 (11) Sleumer, H.: 2097 (2) Smith, A. R. et al.: 906 (43); 1180 (43); 1249 (38); 1342 (21); 1398 (43) Smith, D.: 5260 (35) Smith, D. et al.: 2615, p.p. (l); 2615, p.p. (5) Smith, D. & Palacios, W.: 2639 (3) Smith, D. N. & Vdsquez, R.: 4884 (19) Smith, H. H.: 1023, 1024 (30); 1047 (43) Sodiro, L.: 112 (25); 113 (16); 23827, 23/36 (19); 23/36 (46); 44/145a (25); s.n. (1); s.n. (3); s.n. (6); s.n. (12); s.n. (16); s.n. (19); s.n. (25); s.n. (38); s.n. (39); s.n. (42); s.n. (46) Soukup, J.; 296 (26); 1032 (6) Soukup, S. S.: 3354, 5261 (19) Spane, B.: 13840, 14759 (39); 14898 (46); 16754, 16761 (38); 17168 (28) Spmca. R.; 4340 (35); 4656 (32); 4718 (23); 4742 (14); 5257 (25); 5257A (38); 5262 (42); 5295. p.p. (4); 5295, p.p. (19); s.n. (6); s.n. (22); s.n. (46) Stewart, A.: 914 (33) Steyermark, J. A. & Nilsson, S.: 762 (6) Steyermark, J. A. & Stoddard, A.: 118183 (21) Steyermark, J. A.: 53768 (6); 54713 (19); 62438, p.p. (unusual spt Steyermark, J. A. et al.: 121505 (43) Stu)*). M. A.; 896 (23); 142 (1); 903 (18); 943 (35); 852 (19); 998 Suarez, O. et al.: 4887 (6) Sundue, M. A. & Vasco, A.: 1260 (46) Sundue, M. A. et al.: 3155 (46); 3265 (11); 3265 (46) Tate, G. H. H.: 573 (19); 627 (4); 650 (18); 653 (6) Triana, J. J.: s.n. (42); s.n. (4); 615 (21); 639 (42) Trujillo, D. et al.: 7608 (44) (6); 418 (42); 770 (6); s.n. (1) Tryon, M. R. & Tryon, A. F.: 5412 (16); 6103 (27) Tuomisto, H. & Ruokalainen, K.: 11724 (24) MORAN ET AL.: ANDEAN MEGALASTRUM 235 Uribe, L.: 3286, 3362 (4) Valenzuela, L. et al.: 3019 (38); 3164 (31) van der Werff, H.: 906 (43); 966 (33); 2244, 2256, 2257 (17); 2267 (33) van der Werff, H. & Gudino, E;10711 (46); 11122 (18); 11295 (35); 11369 (7) van der Werff, H. & Palacios, W.: 10570 (3) van der Werff, H. et al.: 12094 (18); 12324 (19); 12482, 12506 (12); 13143 (18); 16300 (. (26); 18275, 18305 (14); 19150 (18); 19383 (40); 21125 (5): 21140. 21371 (31); 21715, 21884 (18); : Vargas C. C.: 14685, 11284 (35); 12938, 16837 (16); 17373 (35) Vargas. H.: 1295 (39) Vargas, I. et al.: 7195 (2) Vasquez, R. & Flores, S: 26345 (19) Vasquez, R. et al.: 26569 (19) Verleysen, J.: 145, s.n. (35) Vogl, C.: 869 (43) von Sneidem, K.: 5024 (39); 5236, p.p. (19); 5236, p.p. (46) Werner, F. A. & Sundue, M. A.: 2151 (23) Wiggins, I.: 661, 18535 (33) Williams, L. & Alston, A. H. G.: 10781 (43) Wingfield, R. & van der Werff, H.: 6562. 6567 (43) Wood, J. R. I.: 14903 (20) Woytowski, F.: 6742 (22) Wurdack, J. J.: 1030 (19); 1851 (35) Young, K. & Le6n, B.: 4958 (15) Young. K. & Sullivan, G. A.: 852 (46) Zabalaga, M. A.: 16 (20); 1124, 1282 (3) Zak, V. & Jaramillo, ).: 3606 (13) Zarate. M. & Muriel, E.: 779 (41); 831 (20) Zuloaga, O. F.: 6166, 6168 (16) American Fem Journal 104(4):237 (2014) Published on 24 October 2014 Referees for 2014 All manuscripts submitted to the journal are peer reviews on a the American Fem Society, as well as quali^of articles in the American Fem voluntary basis. It is their work that contnbutes to the hi^ ^ ^ M. we also smcately dtank Sudi. Kaar for excallea. volunteer editing of manuscripts. Helena FernAndez Aubthur Gilman Christopher Haufler Nathalie Nagalingum Carrie Olson-Manning W. 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