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A New Microgramma Subgenus Solanopteris (Polypodiaceae) from Peru and a New Combination in the Subgenus PDF

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A New Micro gramma Subgenus Solanopteris (Polypodiaceae) from Peru and a New Combination in the Subgenus Blanca Leon Museo de Historia Natural tie la Universidad Nacional Mayor de San Marcos, Av. Arenales 1256, Apartado 14-0434, Lima 14, Peru, and Herbarium, Plant Resources Center, Bio Labs 311, University of Texas at Austin, Austin, Texas 78712, U.S.A. blanca. [email protected] Hamilton Beltran Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos, Av. Arenales 1256, Apartado 14-0434, Lima 14, Peru ABSTRACT. Based on a collection from north-cen¬ A Microgramma bismarckii et M. tuberosa squamis lam- inaribus inter soros positis et paraphysibus gracilibus, cel- tral Peru, a new tuberous species of Polypodiaceae lulis apicalibus elongatis luminibus hyalinis praeditis re- is recognized and described as Microgramma fas¬ cedit. ten. It differs from other coenosoric species by the presence of laminar scales intermixed among the Epiphyte (Fig. 1A). Rhizomes long-creeping, sori and slender paraphyses with elongate apical 1.5—2 mm wide, branched, glaucous to dark brown, cells possessing clear lumina and thin walls. A key apices and young parts covered with broadly ellip¬ for species identification in subgenus Solanopteris, tic non-clathrate peltate scales 0.5—0.75 mm long, to which the new species belongs, is provided. In white or brown with white margins (Fig. 1B-E); tu¬ addition, a new combination is made for a species berous rhizome hollow, 1.5—2 cm wide, covered previously recognized in the genus Solanopteris. with long acuminate scales from a round peltate Key words: Microgramma subg. Solanopteris. base, 1.4-1.5 mm long (Eig. IE), shiny dark-brown Neotropics, Peru, Polypodiaceae. in the center. Phyllopodia short, inconspicuous. Leaves dimorphic, 0.2—0.5 cm apart, subcoria- In the Neotropics, modified stems resembling tu¬ ceous, simple with entire margins, subsessile to bers generally inhabitated by ants are known only short-petiolate, petiole 1—2 mm long. Trophophylls in the genus Microgramma subg. Solanopteris (Co¬ (Fig. 1C) obovate to broadly elliptic, apex obtuse, peland) Lellinger (Wagner, 1972; Rauh, 1973; Go¬ 1.3—2 X 0.8—1.2 cm, scattered scales on both sur¬ mez, 1974; Lellinger, 1977; Moran, 1992). This faces; scales linear, 1.1-1.4 mm long (Eig. 1 H, 1); subgenus currently includes four species: M. bif- costa prominent abaxially % of leaf length, primary rons (Hooker) Lellinger, M. bismarckii (Rauh) B. veins slightly prominulous. Sporophylls (lig. 1J. K), Leon, comb, nov., M. brunei (Werckle ex Christ) oblong, apex retuse. 1.2—2 X 0.7—0.8 cm, adaxially Lellinger, and M. tuberosa (Maxon) Lellinger, all with scattered dark brown scales, persistent along with dimorphic fronds. In Peru, two species have margins, sporangia and paraphyses borne on a 2 been reported: M. bifrons from the Departments of mm broad receptacular line completely covering Amazonas, Cusco, Loreto, Madre de Dios, Pasco, the abaxial surface at maturity, with dark brown San Martin, and Ucayali, growing below 1000 m laminar scales (Eig. 1L, M) 1 X 0.5 mm, base pel¬ elevation; and M. bismarckii from Cusco, Pasco, tate with 0.1 mm long stalk, intermixed with spo¬ and San Martin from altitudes above 1000 m (Tryon rangia and paraphyses. Paraphyses hairlike, 0.8—1 & Stolze. 1993). Based on a collection made by the mm in length. 8—10 cells long. I cell wide, apical second author in the Department of Ucayali at 1220 cell hyaline, 40-60 pun long (Eig. 2A, B). Spores m, a new species is recognized and described here. 65—75 ^un long, with spines 8-13 /am long. Microgramma fosteri B. Leon & II. Beltran, sp. nov. TYPE: Peru. Ucayali: Cordillera Azul del The specific epithet honors Robin B. Foster for Biabo, cabeceras del no Pisqui, 8°28'45.6"S, his work in Peru and his efforts in neotropical con¬ 75°43'5.21”W. 1220 m, 12 Sep. 2000, //. Bel¬ servation. Microgramma fosteri is characterized by tran, R. B. Foster & W. Alverson 3643 (holo- its leaves with entire margins, scattered persistent type, USM). Kigures 1, 2A, B. abaxial scales, acrostichoid coenosori, I mm long Novon 12: 481-485. 2002. 482 Novon laminar scales intermixed with the sporangia, 1 (fig. 2A. 8). The paraphyses in this new species cell-wide paraphyses with elongate apical cells do not resemble those found in the discrete-sori having clear lumina, and spores with long spines species (fig. 2C, I)), where the most distal cells are 8—13 pun. It differs from 17. bifroris and M. Brunei broader than long and the conformed apex has one by its trophophylls with entire margins, acrosti- or two blunt cells. They differ from those other coe¬ ehoid coenosori, and paraphyses with elongate api¬ nosoric species (Fig. 2F), in which the distal cells cal cells. It can he distinguished from M. bismarckii are also broader than long, and the terminal cell and M. tuberosa, both of which also have coenosori, has the lumina dark yellow-brown filled with tannin by its trophophyll with a retuse apex, completely at maturity. This is similar to the situation in Po¬ covered abaxially with sporangia when mature (vs. lypodium virginianum L. (Peterson & Kott, 1974) having a nonsoriferous expanded apex), the pres¬ in which the marginal cells are similarly darkened. ence of shiny dark brown laminar scales among the Affinities between subgenus Solanopteris and sporangia (vs. none or inconspicuous), and the pa¬ some species of subgenus Microgramma are man¬ raphyses with elongate cells, clear lumina, and thin ifest in the paraphyses (e.g., Sota, 1986). For M. apical cell walls (vs. having blunt apical cells with baldwinii Brade and M. ulei (Ule) Stolze, both also dark lumina). Microgramma bismarckii and M. tub¬ found in Peru, paraphyses are one cell wide. In M. erosa, which are known from few collections, are baldwinii, the three to four of the most distal cells poorly understood taxonomieally and may represent are broader than long. In some cases, the third row different phases of a single species. is bicellular, as in M. bifroris and M. Brunei, and Wagner (1986) studied soral patterns in pleopel- the apical cell is hooked, with the lumina filled with tid ferns and commented on the parallel situation tannin. In M. ulei, the apical cell is blunt and dark, in Solanopteris and Microgramma with regard to more like those seen in M. bismarckii. Further re¬ soral series ranging from discrete to confluent sori search on paraphyses development and character¬ to coenosori. The soral pattern in this newly de¬ istics for these and other species of Microgramma scribed species links coenosoric development with may contribute to a better understanding of these a reduction of the fertile blade. The presumed clos¬ affinities. est relative. M. bismarckii, better represented in herbaria, presents laminar expansions beyond the Nkw Combination proximal and distal border of the sori but no con¬ current frond reduction. In the overall distribution Microgramma bismarckii (Rauh) B. Leon. comb, ol Microgramma subg. Solanopteris, coenosoric nov. Basionym: Solanopteris bismarckii Rauh. species occupy altitudes above 1000 m. Abh. Akad. W iss. Abh. Lit. Mainz, Math.-Na- turwiss. Kl. 5: 232. 1973. TYPE: Peru. Pasco: Iaxonomic characters used for species recogni¬ Oxapampa, 1800-2000 m, Ceja de la Mon¬ tion in subgenus Solanopteris have included leaf morphology, venation patterns, and soral shape. tana. July 1971, Werner Rauh 25790 (holotype, HEID). However, with the exception of Wagner (1972). lit¬ tle attention has been paid to other characters such The following key distinguishes all currently rec¬ as those found in paraphyses. Paraphyses in Solan¬ ognized species of Microgramma subg. Solanopter¬ opteris can be described as one cell w ide for most is. of their length. Differences from species to species appear in the distal portions, where variation occurs kiT to i ni. Si’kciks oi Microgramma si no. Soiaaorteris in cell number, dimensions, shape, and coloration of the apical cell lumina (Fig. 2A-E). The distinc¬ 1. Sporophylls with discrete sori; leaves charta- tive acrostichoid coenosori in M.fosteri mostly con¬ ceous to subeoriaceous; paraphyses witli blunt apical cells . 2 tain thin paraphyses with elongate cells, and the L. Sporophylls only with coenosori, occasionally most distal cells have a clear lumina and thin walls intermixed discrete sori present; leaves eoria- Figure 1. Microgramma fosteri 8. Lertn & H. Beltrdn. —A. Habit. —B-E. Rhizome scales. —I). Central basal portion of rhizome scale. —F. Tuber scale. —G. Abaxial view of trophophyll. —If. Adaxial trophophyll scale. —1. Abaxial trophophyll scale. J. Abaxial view ol sporophyll. —k. Adaxial view of sporophyll. —L. Adaxial sporophyll scale. — M. Abaxial sporophyll scale. Drawn from Beltran. Foster & Alverson 5824 (USM). Stippled areas represent dark lumina cells; striped areas represent insertion. The 0.2 mm bar applies to B, C, E, F, H, I, L, and M. The 0.8 cm bar applies to G, J, and k. Volume 12, Number 4 Leon & Beltran 483 2002 Microgramma fosteri from Peru 0.8 cm B 200 iim 130 fim Volume 12, Number 4 Leon & Beltran 485 2002 Microgramma fosteri from Peru ceous; paraphyses with elongate apieal cells or and Alan R. Smith for providing early comments broader than long apical cells with lumina filled and photocopies of Solanopteris at UC. We also with tannin . 3 thank Robbin C. Moran and an anonymous review¬ 2(1). Trophophyll margins lobed; leaves chartaceous .M. hi from er for their comments. 2'. Trophophyll margins entire or rarely slightly re- pand; leaves subcoriaceous.M. brunei Literature Cited 3(1). Sporophylls with retuse apex; sori acrostichoid; Gomez, L. 1). 1974. Biology of the potato-fern Solanopteris dark brown laminar scales dispersed on both brunei. Brenesia 4: 37—61. surfaces; paraphyses with elongate apical cells Lellinger, D. B. 1977. Nomenclatural notes on some ferns and terminal cell without tannin .M. fosteri of Costa Rica, Panama, and Colombia. Amer. Fern J. 3'. Sporophylls with mucronate to acute apex; sori 67: 58-60. elongate to discrete; laminar scales absent or Moran. R. C. 1992. The potato fern. Fiddlehead Forum inconspicuous; paraphyses with distal cells 19; 18-20. broader than long; terminal cell brown filled Peterson, R. L. & L. S. Kotl. 1974. The sorus of Polypo¬ with tannin . 4 dium virginianum: Some aspects of the development 4(3). Trophophyll apex mucronate.M. bismarckii and structure of paraphyses and sporangia. Canad. J. 4'. Trophophyll apex acute or obtuse . . M. tuberosa Rot. 52: 2283-2288. Ranh. W. 1973. Solanopteris bismarckii Rauh. Abb. Akad. Acknowledgments. We thank the Environmental Wiss. Abb. Lit. Mainz, Matli.-Naturwiss. Kl. 5: 223- Conservation Programs of The Field Museum, Chi¬ 256. cago, for inviting the second author to participate Sota, F. 1986. Sobre la posicion sistematica tie Polypo¬ in the field expedition to Cordillera Azul. We also dium fuscopunetatum Hook, y Polypodium percussum thank Peter Jprgensen, John M. Mendenhall, Car¬ Cav. (Polypodiaceae s. str., Pteridophyta). Physis (Buen¬ os Aires) 44: 19-28. men Ulloa Ulloa, Tyana Wachter, and Kenneth Tryon, R. M. & R. G. Stolze. 1993. Pteridophyta of Peru. Young for their help and support during the prep¬ Part V. Solanopteris. Fieldiana Bot. n.s. 32: 178-183. aration of the manuscript. For providing insightful Wagner, W. H., Jr. 1972. Solanopteris brunei, a little- comments, suggestions, and literature we thank known fern epiphyte with dimorphic stems. Amer. Fern J. 62: 33-43. George Yatskievych. We thank llenk van dec Werff -. 1986. The New World fern genus Marginariopsis for his comments and help with the Latin diagnosis. (Polypodiaceae): An example of leaf dimorphy and coe- We thank David Dellinger for providing information nosory in generic delimitation. Bull. Torrey Bot. Club on paraphyses characters of Microgramma tuberosa 113: 159-167. Figure 2. Paraphyses. A, B. Paraphyses of Microgramma fosteri. Drawn from Beltran, hosier & Alverson 3824 (U5M). —A. Paraphyses. —B. Detail of distal portion. —C. Paraphyses of Solanopteris bifrons. Drawn from Dick 270 (MO). _I). Paraphyses of Solanopteris brunei. drawn from Forero et al. 6022 (MO). —F. Paraphyses of Solanopteris bismarckii, drawn from Rauh 35685 (LISM). —F. Paraphyses of Solanopteris bismarckii. two left figures drawn from Ceron et al. 5790 (MO), two right figures from Gentry & Smith 45184 (MO). The 200 pm bar applies to A, C, I), and E. The 130 gm bar applies to B and F.

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