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Luisma, a new genus of Grammitidaceae (Pteridophyta) from Colombia PDF

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Luisma, a New Genus of Grammitidaceae (Pteridophyta) from Colombia M. I Murillo Instituto de Ciencias Naturales, Herbario Nat ional Colombiano, Universidad Nacional de Colombia, Apartado 7495, Bogota D.C., Colombia. [email protected] Alan R. Smith University Herbarium, 1001 Valley Life Sciences Bldg. #2465, University of California, Berkeley, California, 94720-2465, U.S.A. arsmith@ut link4.berkeley.edu ABSTRACT. We describe and illustrate a monotyp- dium (Bishop, 1978), Enterosora (Bishop & Smith, ic new genus and species of fern, Luisma bivascu- 1992), Grammitis sensu stricto (Bishop, 1977). Lel- laris M. f. Murillo & A. B. Smith (Grammitida- lingeria (Smith et al., 1991), Melpomene (Smith & ceae), endemic from die Colombian Andes. Luisma Moran, 1992), Micropolypodium (Smith. 1992), is similar to narrow-leaved species of Campyloneu- Terpsichore (Smith. 1993), and Zygophlebia (Bish¬ rum (Polypodiaceae). However, it agrees with gram- op. 1989). Prior to these studies, Copeland (1952a, mitid ferns in having green, trilete spores (vs. hy¬ 1952b. 1955) dealt with various elements of Neo¬ aline or yellowish, monolete spores in nearly all tropical Grammitidaceae in a synoptical way, and Polypodiaceae) anti free veins (vs. anastomosing in Morton (1967) and Smith and Moran (in Moran & Campyloneurum). From all Grammitidaceae, Luis¬ Riba. 1995) treated grammitids from adjacent Ec¬ ma differs in having two petiolar bundles. Other uador and from Mesoameriea, respectively. distinguishing characters include suberect, radially Discovery of a new genus of ferns, based solely symmetrical rhizomes, sparse hyaline stipe hairs, on previously undescribed species, is an uncom¬ adaxial hydathodes, and clathrate, setulose rhizome mon event, notwithstanding several exciting discov¬ scales. eries in the last two years. Sun et al. (2001) recently described a new genus and species (Mankyua che- Rksumkn. Nosotros describimos e ilustramos un juense B.-Y. Sun et al.) of the eusporangiate fern nuevo geriero y especie tie helecho monotfpico, family Ophioglossaceae from Cheju Island. Korea. Luisma bivascularis M. T. Murillo & A. R. Smith Even more recently, Caohangia squamata (Poly¬ (Grammitidaceae), endemieo de los Andes colom- podiaceae) has been described as a new genus and liianos. Luisma es similar a especies tie Campylo¬ species from Vietnam (Smith & Zhang, 2002). In neurum (Polypodiaceae) con hojas angostas. Sin the Neotropics, the last such monotype named was embargo, concuerda con las gramitidaceas por te¬ Nephopteris maxonii Lellinger (1966), a poorly ller esporas verdes, triletes (vs. esporas hialinas o known plant (still known only from the type) from amarillentas en casi todas las Polypodiaceae) y Colombia, a country generally acknowledged to be venas libres (vs. anastomosadas en Campyloneu¬ among the richest for ferns (as well as vascular rum). De todas las Grammitidaceae Luisma tlifiere plants) in the Neotropics, and also among the rich¬ por tener dos haces peciolares. Otros caracteres est in the world (Cuatrecasas, 1958; Gentry. 1982; que lo distinguen incluyen un rizoma radialmente Moran, 1996; Rangel-Ch., 2000; Hammen, 1995). simetrico, con pelos hialinos esparcidos en el es- Below we describe another monotypic genus, Luis¬ tfpite. hidatodos adaxiales y eseamas del rizoma ma, with its sole species L. bivascularis. I he genus clatradas, setulosas. is known onlv from the type. Key words: Colombia, Grammitidaceae, Luis¬ Luisma bivascularis M. T. Murillo & A. B. Smith, ma. gen. et sp. nov. TYPE: Colombia. Risaralda: Municipio de Mistrato. entre los corregimien- In this paper we describe a new genus of Gram¬ tos de Geguadas y Puerto de Oro, selva de mitidaceae in Colombia. A number of recent papers pisones, 1550 m. zona de bosque pluvial pre- establish a generic framework for the family in the montano, bosque primario, 30 mar. 1992. ./. L. Neotropics. These include revisions or synoptical Fernandez 9613 (holotype, COL 468054). Fig¬ treatments of Ceradenia (Bishop, 1988). Cochli- ure 1. Novon 13: 313-317. 2003. 314 Novon Figure I. Luisma bivascularis M. T. Murillo & A. R. Smith, gen. et sp. nov., based on J. Fernandez 96/.i (COL). —A. Plant habit. —15. Rhizome scale. —C. Petiole and blade base. —I). Petiole, in cross section, showing two vascular bundles. —E. Venation of a portion of the blade. —F. Sporangium, with spores. Volume 13, Number 3 Murillo & Smith 315 2003 Luisma from Colombia Plantae epiphytical. Rhizomata radialiter symmetrica, within the Grammitidaceae remain intriguing but erecta vel suberecta, apice squamata, squamis fumeis, ni- unknown, at present. tentibus, clathratis, usque 2.2 X 0.25 mm. lanceolatis, basi peltatis ad marginem ciliatis, ciliis 0.05 rum longis, Habitat and distribution. The type locality is a liyalinis. 5ti[)ites teretes, basi cum duo fascibus vascular- subandean pluvial forest, where the environment is ibus in sectione transversali, 16-25 X 0.3-0.4 mm, fusci, favorable for the existence of diverse plant life, in¬ trichomatibus hyalinis simplibus, phyllopodiis carentibus. cluding numerous epiphytes, ferns, mosses, li¬ Laminae prasinae in statu sicco, anguste ellipticae, 9-20 X 0.3—0.4 cm, membranaceae, basi et apice longiattenu- chens. aroids, and orchids. In this area there have atae. margine integrae; costae venaeque glabrae utrinque; been found recently several new bird species, in nervi libri, 2—3-furcati; hydathodi adaxialiter prominentes Yireo (Salaman & Stiles, 1996) and Glaucidium ellipticae. Sori rotundati, ca. I mm diam., confinis costae, (Robbins & Stiles, 1999); a new lepidopteran in¬ uniseriales in lateribus ambabus costae, paraphysibus ab- sentibus, capsulis sporangiorum glabris. Sporae virides, sect, Antirrhea (Andrade, 1993); a new tree species globosae, 30-36 gm diam. (in acido laeteo), triletae. in the genus Matisia, Bombacaceae (Eernandez- Alonso, 2001); and a new Columnea, Gesneriaceae Plants epiphytic. Rhizomes radially symmetrical, (Amaya-Marquez, 2002). erect or suberect, scaly at the apex, scales smoky I he species is known from only a single collec¬ dark gray, shining, clathrate, up to 2.2 X 0.25 mm, tion. and is seemingly rare. However, the locality lanceolate, peltate at the base, margins ciliate, cilia is relatively inaccessible, and the great floristic di¬ 0.05 mm long, hyaline. Stipes terete, darkened versity of the region, as well as the paucity of col¬ (brownish), at the base with two vascular bundles lections from the area, may obscure the fact that it in cross section, 16—25 X 0.3—0.4 mm, trichomes could lx1 locally common. More likely, it is of very hyaline, simple; phyllopodia absent. Blades green¬ restricted occurrence. ish in the dried state, narrowly elliptic, 9—20 X At first glance, Luisma bivascularis looks like a 0.3-0.4 cm, membranaceous, at the base and apex species of Campyloneurum (Polypodiaeeae), but in long-attenuate, at the margins entire, midribs and studying the characters further, we conclude that it veins glabrous on both sides; nerves free, forked; has no close affinity with that genus. Clearly, the hydathodes adaxially prominent, elliptical. Sori species is a member of the Grammitidaceae be¬ round, ca. 1 mm diam., adjacent to (touching) the cause of its chlorophyllous (green) trilete spores midrib, uniseriate on each side of the costa, lacking and free veins (although a few Polypodiaeeae sensu paraphyses; sporangial capsules glabrous. Spores stricto also have free veins, e.g., some species of green, globose, 30-36 /cm diam. (in lactic acid), Polypodium and also Pecluma, and a few grammi- trilete. tids have anastomosing veins, e.g., Zygophlebia). Also the rhizome scales, which are strongly clath¬ Etymology. The genus is dedicated to Luis Ma¬ rate and with hyaline marginal setulae, are remi¬ ria Murillo (father of the first author), pioneer in niscent of Lellingeria (Grammitidaceae) and unlike entomology in Colombia and author of many sci¬ anything known in Polypodiaeeae. The rhizomes entific works. He was one of the founders in 1929 appear to be radially symmetrical and suberect, of the Academia Colombiana de Ciencias Exactas, like some grammitid ferns but unlike almost all Po- Ftsicas y Naturales, and editor of the journal for 10 lypodiaceae. We are unable to place the specimen years (1952—1961). He belonged to the Soeiedad in any known genus of Grammitidaceae in either Colombiana de Geografta. was an honorary member the Neotropics or Paleotropies. From all known of the Soeiedad Real Entomologica de Belgiea, and genera and species of Grammitidaceae, Luisma bi¬ founded tlie Instituto de Altos Estudios del Go- vascularis differs in having two vascular bundles in bierno Naeional. After more than 40 years of con¬ the petioles. Many species of grammitids have been tinuous service in the Ministry of Agriculture as checked for this character (Parris, 1990; A. R. "Jefe de Sanidad Vegetal”, Colombia’s President Smith & B. S. Parris, unpublished data), including Alberto Lleras Camargo conferred the Cruz de Boy- some that are larger than Luisma. In addition, Luis¬ aca in 1962, in recognition of his outstanding work ma differs from the largely Neotropical genera Cer- and exceptional scientific merits. We cite some of adenia. Enterosora, and Zygophlebia in having his many publications (Murillo. 1937. 1944, 1951. veins ending in prominent hydathodes adaxially. 1956). Luisma further differs from Ceradenia in having rile species epithet refers to the petioles, which clathrate rhizome scales and from Zygophlebia in in cross section have two vascular bundles; this is lacking distinct phyllopodia. the only genus known in Grammitidaceae that has from Terpsichore, Melpomene, and Micropolypo¬ thi s vascular configuration. The implications of this dium, other Neotropical grammitid genera, Luisma finding for the phylogenetic position of Luisma differs in lacking long reddish or red-brown setae 316 Novon on the blades and petioles; in fact, die fronds of black sclerenchymatous margins and lustrous, con¬ Luisma are nearly glabrous or with only sparse eolorous, orange to tan. entire rhizome scales; and Iximaphlehia (Greater Antilles), with anastomosing whitish hairs on the stipes. Melpomene further dif¬ veins and rhizome scales similar to those of Coch¬ fers in having entire marginal rhizome scales and lidium and Grammitis. American species of Gram¬ pinnatifid to pinnatiseet leaves. Like Micropolypo¬ mitis sensu stricto lacking a black margin occur in dium and some species of Terpsichore, Luisma has southern South America, but these have orangish, hyaline marginal setae on the rhizome scales, but coneolorous, entire rhizome scales and are very it differs from all species in those genera in having small-bladed species, generally less than ca. 5 cm entire leaves. From Lellingeria, which lias similar long. We do not believe that any American genus rhizome scales. Luisma differs in its simple, entire of Grammitidaceae is closely related to Luisma. nor blades, lack of forked hairs on the stipes and ra- can we demonstrate that any Old World genus of ehises, and generally quite glabrous blades. Grammitidaceae has close relationships. Without Other simple-bladed species of Grammitidaceae molecular data, the closest affinities of Luisma will in the Neotropics include Cochlidium, the species probably remain obscure, and we believe it likely of which have dull, coneolorous, orange to tan, en¬ that Luisma is one of the most isolated species and tire rhizome scales; Grammitis, with species having genera in the family, phylogeneticallv. Artikiciai Key to Genera of Grammitidaceae in Colombia la. Blade s entire, deeply serrate, crenate, or broadly and shallowly lobed. 2a. Petioles each with two vascular bundles; rhizome scales clathrate, with ciliate margins. Luisma 21). Petioles each with a single vascular bundle; rhizome scales non-clathrate, entire or at least not ciliate. .5a. Blade margins sclerified, dark brown or black. Grammitis 3b. Blade margins not sclerified, greenish or yellowish. 4a. Rhizomes radially symmetrical; blades entire to deeply serrate, usually loss than 5 mm wide; hydathodes evident or not adaxiallv.Cochlidium 4b. Rhizomes dorsiventral; blades entire, crenate, or broadly lobed, usually more than 5 mm wide; hydathodes absent adaxially. Enterosora Blade (cid:9632)s pinnatifid, pinnatiseet, or 1-pinnate. 5a. Hydathodes absent adaxially. 6a. Veins free; opaque whitish stalked glands borne in soil and sometimes also on blades . . Ceradenia 6b. Veins areolate; opaque whitish glands lacking in sori and on blades.Zygophlebia 5b. Hydathodes present adaxially. 7a. Rhizome scales not clathrate. 8a. Blades less than 1 cm wide, with one sorus per segment; main veins in segments (pinnae) simple or with a single acroscopic branch; rhizomes radially symmetric .... Micropolypodium 8b. Blades usually more than 1 cm wide, with more than one sorus per segment; main veins in segments (pinnae) pinnately branched; rhizomes dorsiventral to radial.Terpsichore 7b. Rhizome scales usually strongly clathrate. 9a. Setae lacking on margins of scales, but sometimes with several minute papillae at scale tips .Melpomene 9b. Setae on margins of scales hyaline to reddish. 10a. Cell luminae clear; blades lacking black elavate fungal fruiting bodies.Lellingeria I Ob. Cell luminae darkened; blades often with black elavate fungal fruiting bodies, espe¬ cially along rachises, costae, and among sporangia. Terpsichore Acknowledgments. We express our gratitude to I .iterature Cited the Working Group on Grammitidaceae for their comments: Tom Ranker, Barbara Parris, Chris Amaya-Marquez, M. 2002. Una nueva especie de Col¬ umned seccion Collandra (Gesneriaceae). Caldasia 24: Haulier, and Harald Schneider. We also thank 225-228. members of the Instituto de Ciencias Naturales de Andrade, G. 1993. Una nueva especie de Antirrhea (Lep- la Universidad Nacional de Colombia for facilities idoptera: Nymphalidae: Morphinae) de la Cordillera Oc¬ ami suggestions on the manuscript. Lastly, we cidental de Colombia. Revista Acad. Colornb. Ci. Exact. thank Juan Carlos Pinzon for executing the illus¬ 18(71): 555-558. trations. Bishop, L. E. 1977. The American species of Grammitis Volume 13, Number 3 Murillo & Smith 317 2003 Luisma from Colombia sect. Grammitis. Amer. Fern J. 67: 101—106. tanical Garden, St. Louis; The Natural History Museum, -. 1978. Revision of the genus Cochlidium (Gram- London. mitidaceae). Amer. Fern J. 68: 76-94. Morton, C. V. 1967. The genus Grammitis in Ecuador. -. 1988. Ceradenia, a new genus of Grammitida- Contr. U.S. Natl. Herb. 38: 85-123. ceae. Amer. Fern J. 78: 1—5. Murillo, L. M. 1937. El sentido de una lueha biologica. -. 1989. Zygophlebia, a new genus of Gramm i I i- Revista Acad. Colomb. Ci. Exact. I: 376—410. daceae. Amer. Fern J. 79: 103-118. -. 1944. Los insectos y el clima en Colombia. Cal¬ -& A. R. Smith. 1992. Revision of the fern genus dasia 10: 465—471. Enterosora (Grammitidaceae) in the New World. Sysl. -. 1951. El amor y la sabidurfa de Francisco Josd Rot. 17: 343-362. de Caldas. Revista Arad. Colomb. Ci. Exact. 8(30): Copeland. F. R. 1952a. Grammitis. Philipp. J. Sci. 80: 149-153. 93-271. -. 1956. Colombia, un arehipielago hioldgico. Re¬ -. 19521). The American species of Xiphopteris. vista Acad. Colomb. Ci. Exact. 9(36,37): I-XX. Amer. Fern J. 42: 41—52; 92—100. Parris, B. S. 1990. Grammitidaceae. Pp. 153—157 in K. -. 1955. Ctenopteris in America. Philipp. J. Sci. 84: Kubitzki (editor). The Families and Genera of Vascular 381-475. Plants, Vol. I. Pteridophytes and Gymnosperms. Spring¬ Cuatrecasas, J. 1958. Aspectos de la vegetacion natural er-Verlag, Berlin. de Colombia. Revista Acad. Colomb. Ci. Exact. 10: Rangel-Ch., 0. 2000. La megadiversidad biologica de Co¬ 221-268. lombia: (iRealidad o ilusionV In J. Aguirre (editor), Me- Fernandez-Alonso, .1. E. 2001. Bombacaceae Neotropicae morias del Primer Congreso Colombiano de Botanica. Institute de Ciencias Naturales, Universidad Nacional, novae vel minus cognitae—II. Novedades taxondmicas y eorolbgieas en Matisia y Quararibea del norte de Su- Bogota. [CD-ROM version.| Robbins, M. B. & F. G. Stiles. 1999. A new species of dameriea. Caldasia 23: 351-382. pygmy-owl (Strigidae: Glaucidium) from the Pacific Gentry, A. H. 1982. Phytogeographic patterns as evidence lor a Chord reluge. Pp. 1 12—136 in G. T. Prance (edi¬ slope of the northern Andes. Auk I 16: 305—315. Salaman, P. G. W. & F. G. Stiles. 1996. A distinctive new tor), Biological Diversification in the Tropics. Columbia species of vireo (Passeriformes: Vireonidae) from the Univ. Press, New York. western Andes of Colombia. Ibis 138: 610-619. Hammer), T. van der. 1995. El estudio de la vegetarian Smith, A. R. 1992. A review of the fern genus .1hcro- en Colombia y su relacion con ecosistemas y biodiver- polypodium (Grammitidaceae). Novon 2 : 419—425. sidad. Pf>. 27—40 in Fondo FEN Colombia, Thomas van -. 1993. Terpsichore, a new genus of Grammitida¬ der Hammen premio a la vida y obra 1995. Grupo ed¬ ceae (Pteridophyta). Novon 3: 478— 489. itorial 87. Bogota. -& R. C. Moran. 1992. Melpomene, a new genus Lellinger, 1). B. 1966. Nephopteris, a new genus of fern of Grammitidaceae (Pteridophyta). Novon 2: 426—432. from Colombia. Amer. Fern J. 56: 180—182. -& X.-C. Zhang. 2002. Caobangia, a new genus Moran, R. C. 1996. The importance of the Andes as a and species of Polypodiaceae from Vietnam. Novon 12: barrier to migration, as illustrated by the pteridophytes 546—550. of the Chord phytogeographic region. P. 75 in .1. M. -, R. C. Moran & E. E. Bishop. 1991. Lellingeria, Camus, M. Gibby & R. J. Johns (editors), Pteridology a new genus of Grammitidaceae. Amer. Fern J. 81: 76— in Perspective. 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