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New Grammitid Genus Eleven Species in the Fern Melpomene (Polypodiaceae) Marcus Lehnert ssenschaften, Abt. Systematische Botanik, Georg- D-37073 Germany Karspiile Gottingen, 3 2, — Abstract. Based on extensive studies of grammitid ferns (Polypodiaceae) in the field and as new specimens in various herbaria, eleven species in the genus Melpomene are recognized from Melpomene the Neotropics: albicans, M. caput-gorgonis, M. M. huancabambensis, M. flagellata, vulcanica. All are restricted to the Andes except for M. personata, which also extends to Mesoamerica and the Caribbean. The morphology and distribution of all species are illustrated. Words.— Andes, Caribbean, grammitid Melpomene, Mesoamerica ki-:\ ferns, The Melpomene genus belongs the grammitid which form to ferns, a monophyletic clade within the Polypodiaceae (Schneider et al, 2004) and - comprise about 750 species (Parris, 2005) with a pantropical southern temperate Melpomene was distribution (Parris, 1990, 2003). established as a genus on solely morphological characters (Smith and Moran, 1992) but was later supported as a natural monophyletic group in molecular analyses (Ranker et al, 2004; Lehnert et al, in print) that also confirmed some exclusions from and additions to the genus (Smith, 1995; Leon and Smith, Because 2003). of its young taxonomical genus age, the has never been revised whole. as a Partial revisions are available for Mexico (Mickel and Smith, Mesomamerica 2004), (Moran and Riba, 1995), and Brazil (Labiak and Prado, 2005), but these areas belong to the periphery of the total range of the genus, where a restricted assemblage and of species morphological emphasis caused by genetic drift may lead to an easier distinction of species. The majority specimens come of m from Andean the where region, the genus occurs 200-5200 and displays at the greatest morphological diversity. In order to revise the whole genus, a total of 2100 collections from following AAU, herbaria were examined: BM, BHCB, COL, COLO, CUZ, GOET, B, C, K, LOJA, LPB, P, QCA, QCNE, RB, SP, TUR, UC, US, USM. Among them S, are 251 specimens by collected the author in Ecuador, Peru, Bolivia, and him Argentina, giving further insight about putative between correlations and ecological conditions morphological As plasticity. suspected from previous studies on grammitid ferns (Labiak, 2000; Rojas, 2001; Leon and .owentor, Rosenstein NEW LEHNERT: OF MELPOMENE SPECIES 215 Melpomene Smith, were 2004), several taxa in indentified as yet undescribed species. Most of the following species have been collected prior to this study but were not recognized something new. as Instead, their morphological were differences dismissed as intraspecific variability or aberrations. In this study was observed some widespread have morpholo- that species a great it high answer wide gical plasticity, reflecting a potential to a array of ecological Good examples Melpomene factors. are moniliformis (Lagasca Sw.) A. ex. R. Sm. & Moran and M. A. Sm. & Moran, whose R. C. flabelliformis (Poir.) R. R. C. distributions cover almost the whole range of the genus, the species latter being the sole representative of the genus in the Paleotropics. They inhabit many and different types of habitats consequently vary in size enormously, which often correlated with a change of the proportions of the segments and is number Opposed in the of cells across the scale bases. to this, specialized species have to respond to a smaller amplitude of biotic and abiotic factors and consequently should display a lower morphological variability. In respect to the absolute ranges of the morphological variability, this leads to an apparent incongruence among The the species descriptions presented here. reasons for comments the respective recognition of a species are given in the following the descriptions. Terminology Grammitid with follows that of Lellinger (2002). ferns are beset two indument and homologous types of laminar that are distinct not part of a series. The term "seta" has often been applied to the darkened hairs that stiff, frequently occur in grammitid ferns and also in Melpomene (Smith and Moran, 1992; Labiak and Prado, 2005). The same treatments used the term "hairs" for the paler, often branched trichomes found mainly on young petioles and Melpomene. However, rachises in find that the so-called "setae" are I developed within Melpomene, sometimes representing species differently autapomorphies and sometimes growth conditions. For reflecting different & Moran example, the "setae" of M. firma Sm.) A. R. Sm. R. C. are usually (J. would more quite short and patent, for which the term "acicular hairs" be many M. and related species, the fitting (Lellinger, 2002). In flabelliformis "setae" are sometimes flaccid and twisted, often occurring with properly developed straight and rigid) "setae". Such flaccid trichomes are better (i.e., term described as "ciliform hairs" (Lellinger, 2002). For the rigid "setae," the used Furthermore, Latin "setiform hairs" available (Lellinger, 2002). in is which would (Steam, diagnoses, the term "seta" translate as "bristle" 2004), is Melpomene found a notably different structure (Lellinger, 2002) that not in is and or other grammitid genera. will use the terms "ciliform," "setiform," I "acicular hairs" describe the simple pluricellular trichomes that are usually to dark red in color with acute and "clavate hairs" for the minute, simple tips, and and branched, pluricellular hairs that are pale in color bear clavate cells at been their apices or along their length. This second type of hairs has referred to homologous as "glandular" 2005) and they appear to be to the secretive (Parris, gland-bearing branched hairs of Zygophlebia (Bishop, 1989), but a secretive Melpomene The same was so can be nature not observed in the hairs of far. AMERICAN VOLUME NUMBER FERN JOURNAL: 216 98 4 (2008) Melpomene stated for the "glands" that occur on the scale apices of species. all They are thin walled cells that are arranged singly or as pluricellular papillae branched and or clavate hairs at the tips rarely also along the margins of the rhizomes scales, strongly resembling the clavate hairs of the fronds. They are here simply referred to as apical and marginal Scales are only found on cells. the rhizomes and sometimes on the petiolar bases. The dimensions of the segments are measured with the midveins cardinal as which assumed points, are as medial lines in the central parts of the segments between the sori they are obscure. Segment length measured along the if is midvein from segment tip to rachis; segment width taken as the orthogonal is line at half of the length. The dimensions are more useful the width taken if is in the central part of the segments because segments deltate are usually strongly decurrent onto the rachises in their proximal half and width taken may here vary considerably. TYPE.— Lehnert, nov. BOLIVIA. Santa sp. Cruz: Prov. i km Comarapa, Caballero, de ca. 1 Siberia hacia Torecillos, 17 49'S, 64 40'W, 2650 m, 18 Mar 2003, Lehnert 714 (holotype GOET; isotypes LPB, UC). 7A. Figs. 1, A Melpomene personata laminis abaxialiter albicantibus, a M. youngii Leon & Sm. (Stolze) B. A. R. pilis pluribus in soris confertis praestans. Plants predominantly epiphytic growing moss or epilithic, in layers. mm Rhizomes moderately to short-creeping, horizontal 1A), 0.8-1.2 (Fig. diam., sometimes with mm) short branches (5-10 held Fronds at right angles. cm to 23 long, erect (Fig. 1A) or patent, inserted onto the rhizome acute at angles, but often held at nearly right angles to the rhizome, closely arranged mm) (internodes 2-5 but not caespitose X (Fig. 1A). Scales 3.0-5.0(-7.5) (0.3-) mm, 0.6-0.8(-1.0) (i2-)16-20(-22) wide, cells lanceolate, clathrate (Fig. IB), brown dark brown, to strongly iridescent, bases cordate broadly to cordate, tips acute to short-attenuate; apical cells 3-8, in a nodding cluster or palmate mm mm arrangement (Fig. 1C). Petioles (24-)35-85 long, 0.5-0.8 thick, marginate from the laminar most bases, parts terete, glabrous to glabrescent mm with brown dark setiform hairs 0.75-1.2 long simple and (Fig. ID); branched mm) clavate hairs (0.1-0.3 of and young crosiers fronds often persist mm in older fronds. Laminae firm-chartaceous, 120-75 X 18-35 to 4/5 (2/3 to of frond length), narrow-elliptic to elliptic (widest in the middle), rarely obovate (widest above the middle), cuneate somewhat or tapering bases, at acute to attenuate at tips (Fig. 1A); surfaces abaxially whitish white, often to yellow when or ochre dried, usually eliminated with treated alcohol. if Rachises dark brown to black, planar and slightly sunken between segments adaxially (Fig. IF), hemispherically protruding abaxially with few (Fig. IE), to many branched clavate on hairs like petioles, otherwise glabrous with or mm scattered acicular Segments X hairs. 9.0-16.0 to 1.2-2.6(-3.0) (6-8 times longer than broad), weakly ascending flat, (80-70 equilateral bases or at ), weakly decurrent basiscopically, fully adnate, linear-oblong to long-deltate, NEW LEHNERT: SPECIES OF A/// h\lH\h i't VOLUME NUMBER AMERICAN FERN JOURNAL: 218 98 4 (2008) the tips obtuse or short-acute; midveins not visible, or obscurely so abaxially in dried specimens (Fig. IE, F), without hairs; proximal 1-7 segment pairs markedly smaller than subsequent segments, sometimes the lowermost 1-4 segment pairs auriculiform; stomata sometimes visible as rusty red dots; segment margins without hairs; hydathodes present (Fig. IF). Sori 2-8(-9) mm pairs per segment, with 4-10 dark-castaneous setiform hairs 0.5-0.8 long (Fig. IE). The name refers to the white wax-like deposit on the abaxial laminae (Latin, = albicans being white). Melpomene albicans grows cloud and moist montane in elfin forests, forests, m 1500-3400 and forests at in Bolivia eastern Brazil (Fig. 7 A). There are only a few other species of Melpomene with whitish abaxial Among Melpomene laminar surfaces. these, youngii differs from M. albicans in being almost completely glabrous (rarely 1-2 hairs present in some sori vs. 4- 10 hairs in sori, petioles glabrous to glabrescent with few hairs in M. albicans) and more Melpomene & the strongly ascending segments; sodiroi (Christ Sm. & Moran Rosenst.) A. matches M. R. R. C. albicans in the hair distribution, but has gibbose segments with the deeply sunken laminae sori slightly to (vs. planar with M. and superficial sori in albicans) larger scales (on average 6.0- 9.0 X 1.0-1.2 mm, 24-30 wide 3.0-5.0 X 0.6-0.8 mm, 16-20 cells vs. cells wide) with characteristically long tapering acute tips (vs. to short-attenuate). Melpomene erecta V. Morton) A. Sm. & Moran, which only (C. R. R. C. rarely mm has white laminar surfaces, has thicker rhizomes diam. (to 2.4 vs. to mm 1.6 diam. in M. albicans) and wider scales (on average 38-60 16- cells vs. 20 cells across base). Melpomene albicans belongs complex comprising M. M. to a personata, peruviana (Desv.) A. R. Sm. & R. C. Moran, M. sodiroi, and M. youngii, which have horizontally creeping rhizomes, and all relatively dark, lanceolate scales, downward predominantly Melpomene pointed fronds. albicans the only is complex when species in the that has patent fronds growing epiphytically or even when strongly ascending fronds growning have epilithically. All relatively long setiform/ciliform hairs in the except M. where sori for youngii, the hairs are usually lacking and or relatively short sparse (up per to 2 hairs Melpomene sorus). personata and M. peruviana from M. differ albicans in their green laminae (vs. with a white wax-like layer abaxially), and they often have setiform/ciliform hairs along the midveins and sometimes along single hairs the segment margins, which are always absent M. Melpomene in albicans. peruviana sometimes has whitish green laminae but lacks a wax-like layer. Additionally, M. peruviana on much average smaller than M. and is albicans M. cm personata (2.5-10.0(-17.5) 17.5-25.0 vs. to cm), tends form dense to cushions tufts or (vs. single plants or loose groups), and grows predominantly epilithic in drier or cooler habitats than the other two species (mostly epiphytic in elfin forests and wet montane forests). The specimens Melpomene of albicans from Brazil are less variable in size than the Bolivian specimens. They are smaller on average and have shorter petioles; Ribas et al. 3080 (UC) has also rather densely hairy The petioles. . NEW LEHNERT: OF MELPOMENE SPECIES 219 more Brazilian plants are easily confused with M. xiphopteroides (Liebm.) A. Sm. & Moran Andean than R. R. C. the population because the white laminar developed and layers are less the scales are smaller, and thus closer the to morphological spectrum M. of xiphopteroides. However, even weakly hairy plants of M. xiphopteroides have more on setiform/ciliform hairs the rachises than M. which albicans, in the rachises are usually glabrous. Additional specimens examined.—BRAZIL. Minas Gerais: Alto Caparao, Parque Nacional do & Caparao, along trail to Pico da Bandeira. 2600 m. ca. 20 31'S, 41 53'W, 21 Mar 1999, Salino Morals 4538 (UC). Parana: Curitiba, Serra Ibitiraque, Morro Camapua (Mun. Campina Grande do Mar Sul). 25 [)8'S, 49 04'W, 02 2000, Ribas et 3080 (UC). Rio de Janeiro: Theresopolis, al. ca. 22 25'S, ca. 42 58'W, Oct 1929, Brade 9094 (NY); boundary between municfpio de Teresopolis and Municipio de Petropolis, Serra dos Orgaos National Park. ca. 5 km S\V of city of Teresopolis, & Nov 22°27-28'S, 43°01-02'W, 30 1965, Eiten Eiten 7168 (US). BOLIVIA. Cochabamba: Prov. Ayopaya, San Cristobal, climbing along the trail that leads to San Miguel, 16 39'S, 6643'W, 3100 m, 06 Jun 2002, Jimenez 1107B (GOET, LPB, UC); Prov. Carrasco, I. on the way from Comarapa to Siberia, 17 50'S, 64 42'W, 3000 m, 22 Jan 2000, Jimenez 283 (LPB, I. km UC); 10 from Siberia to Comarapa, 17 48'S, 64 42'W, 2600 m, 20 Oct 1996, Kessler 9164 et al. km May (LPB, UC); Prov. Carrasco, 10 Cocapata-Cotacajes, 16 38'S, 66 41'W, 3000 m, 09 1997, Kessler et al. 9401 (GOET, LPB, UC); road Cochabamba-Villa Tunari, below Corani, 17 10.58'S, Nov 65 53.67'W, 2700 m, 26 2002, Lehnert 512 (GOET, LPB, UC); road Cochabamba-Villa Tunari, below Nov Corani, 17 10.58'S, 65 53.67'W, 2700 m, 26 2002, Lehnert 514 (GOET, LPB, UC); Prov. km Chapare, Cochabamba 54 hacia Villa Tunari, 2750 m, 30 Apr 1979, Beck 1424a (LPB). La Paz: km N Choquetanga crosses the Rio Ocsalla, 10 of Choquetanga, 16 45'S, 67 17'W, 3400 m, trail 07 Sep 1991, Lewis 39952 (LPB); comunidad Choquetanga-Wichupampa, Serranias de Lulini NW N km km Mar 13 al de Choquetanga, 2-3 al del cerro Lulini. L6 45'S, 67 20'W, 3290 m, 17 2 1994, Salinas 2783 (US); Prov. Nor Yungas, Coscapa, on prehispanic trail Sillutinkara. L6 S 1 ' & 67 53'W, 3100-3300 m, 07 Jan 2001, Jimenez Vidaurre 526 (LPB, UC), 559 (GOET, LPB, UC); I. Unduavi, trench to the Valle de Coscapa, 16 17'S, 67 51'W, 3350 m, 04 Feb 2003, Lehnert 599, 601, 602 (GOET, LPB, UC). Santa Cruz: Prov. Caballero, Comarapa, between Torecillos and Siberia, 17°49.65'S, 64°40.14'W, 2600-2700 m, 18 Mar 2003, Lehnert 696, 707 (GOET, LPB, UC); from km Mar Siberia 4 to the E, small laguna on the ridge (Laguna Tinque?), 2600 m, 18 2003, Lehnert km 717 (GOET, LPB, UC); by small lake summit of pass ca 4 E of Siberia, 2800 m, 04 Jan 2000, at Wood Goyder 15792 Sr (LPB). Melpomene TYPE.— BOLIVIA. La caput-gorgonis Lehnert, nov. Paz: Prov. sp. Nor Yungas, 3200 m, 16 67 51'W, 25 Cotapata, detras del gasolinero, 17'S, GOET, Sep 2002, Lehnert 367 (holotype: LPB; isotypes: UC). Figs. 7B. 2, A squamarum Melpomene apicibus brevioribus, flabelliformi petiolis latioribus (3-4 vs. 1-2 cellulis latis), cellulis apicalibus creberioribus, absentia pilorum inter soros differt. moss Plants predominantly epiphytic, growing in layers, rarely epilithic. mm Rhizomes moderately short-creeping, horizontal (Fig. 2A), 1.0-1.5 to diam. Fronds to 35-42 cm, arching to pendent, inserted onto the rhizomes at 1-4 mm). Scales X right angles 2A), closely arranged (internodes to 6.5 (Fig. mm, auburn brown, 0.8-1.0 (18-)20-26(-32) wide, clathrate, to strongly cells with obtuse truncate iridescent, broadly cordate to pseudopeltate, to tips numerous, ending in rows of 3-4 cells (Fig. 2B); apical cells sitting clustered mm mm on wide 15-50(-70) long, 0.6-0.8(-1.0) the Petioles tips (Fig. 2C). VOLUME NUMBER N 98 :.\L: 4 Melpomene Fig. 2. caput-g( habil [Kessler 1192, GOET); B. scale [Lehnert 868, GOET); C. detail of scale apices (//me LPB); D. petiole, upper part adaxially [Lehnert 868, GOET); , segment E. adaxially (Lehnei segment IET); F. fertile abaxially [Lehnert GOET). 868, NEW LEHNERT: OF MELPOMENE SPECIES 221 thick, decurrently marginate from the laminar with brown bases, flaccid mm) ciliform hairs (1.0-2.0 on both sides simple and branched (Fig. 2D), clavate hairs of crosiers and young fronds sometimes persistent in older fronds, but generally Laminae glabrescent. firm-chartaceous, rarely coriaceous X mm, [Lehnert 300(-350) 14-22 781), to narrow-elliptic (broadest in the middle), decurrent at bases, acute to attenuate sometimes caudate at tips, (Fig. 2A). Rachises dark brown, and slightly sunken between segments flat, weakly adaxially (Fig. 2E), hemispherically protruding abaxially (Fig. 2F), glabrous except the proximal with on for part, ciliform hairs as petioles. mm Segments X 4.5-9.0(-12.0) 3.2-3.8(-5.0) (1.5-3 times longer than broad), weakly ascending (80-70 oblong to lunate, equilateral bases or weakly at ), basiscopically decurrent, fully adnate, the tips obtuse to round (Fig. 2E, F); midveins not visible, or obscurely so abaxially (Fig. 2F), especially in dried specimens; proximal markedly pairs smaller than the central segments (Fig. 2A), inequilateral bases, basalmost auriculiform Lehnert at (e.g., 368), but never trapezoid; rarely few ciliform hairs scattered along the segment margins; hydathodes present (Fig. 2E). Sori 2-4 pairs per segment, with 3-10 mm and receptacular circumsoral ciliform hairs 1.0-1.5 long (Fig. 2F). The name numerous on refers to the apical cells the scale tips; these cells resemble the head of Medusa, one of the Gorgons in Greek mythology, which = had snakes instead of hair (Latin, caput head). Melpomene caput-gorgonis grows wet montane and in forests elfin forests at m 2680-3200 southern Peru and in Bolivia (Fig. 7B). The most distinguishing feature of this species the broad scales with the is abundant No Melpomene apical papillae. other species of has scale apices that Melpomene provide a base for papillae several cells wide. flabelliformis can be distinguished from M. caput-gorgonis by distant fronds and glabrous or its glabrescent petioles fronds close and petioles persistently hairy in M. (vs. Melpomene more and caput-gorgonis). generally slender in habit flagellata is mm most laminae wide 22 mm), rhizomes features, to 9(-16) (vs. to i.e., and and thinner ascending, petiole scales narrower, hairs in the sori longer. Melpomene which grows with M. paradoxa, can be caput-gorgonis together distinguished by glabrescent petioles and shorter segments; the longer, its whereas M. latter species also forms patches with the fronds erect to arching pendent caput-gorgonis grows with arching fronds. solitary to Additional examined.—PERU. Cuzco: Abra de Chaupimayo, Haci< specimens 2700 m, Sep 1932, Sties 1945 (CUZ). BOLIVIA. La Paz: Prov. Franz Tamayo, PN-ANMI Madidi, trail Keara-Mojos, below Chunkani, & 2870 m, 14 6857'W, 08 Nov 2001, Jimenez Gallegos 917 (LPB, UC); Prov. Larecaja, toll '38'S, I. Williams 1154 (NY); Prov. Nor Yungas, Estacion , « m, Dec to the Coscap.i :*U00 12 1997, Kesslei et al. 1871 (LPB); 2 km from Chuspipata to Coroico, 16 22'S, 67 49'W, 2900 m, 19 Sep 1997 Kessleret 11921 (GOET, LPB, UC); Cotapata, behind the gas station. 16 17'S. 67 51 \Y. 3150- al. 3200 m, 25 Sep 2002, Lehnert 367, 368, 369, 372, 373, 386, 392 (GOET, LPB, UC); Chuspipata- VOLUME NUMBER AMERICAN FERN JOURNAL: 222 98 4 (2008) km Sacramento, 16 18'S, 67 49'W, 2680 m, 10 Nov 2002, Lehnert 496a (GOET, LPB, UC); 1.2 E de Cotapata-Santa Barbara, to the Chuspipata electricity station, 16 17'S, 67 50'W, 3200 m, 02 trail May Feb 2003, Lrluurt 586 (GOET. LPB. UC): Chuspipata. old trail to l.'nduavi, 3200 m, 07 2003, Lehnert 781 (GOET, LPB, UC). TYPE— Melpomene BOLIVIA. La Nor flagellata Lehnert, nov. Paz: Prov. sp. N Uchumachi Yungas, side of Cerro above Coroico, 16 12'S, 67°45'W, & 2350 m, 14 1989, Kessler Kelschebach 107 (holotype LPB; isotype Jul GOET). Figs. 7C. 3, mm A Melpomene mm), moniliformi longioribus (1.0-2.0 0.5-1.0 pilis vs. creberioribus in petiolis sorisque, rhizomatibus ascendentibus horizonta- (vs. reptantibus) apicibusque segmentorum partim semper liter truncatis (vs. obtusis vel rotundis) differt. Plants predominantly growing moss sometimes epiphytic, in layers, Rhizomes epilithic. erect or ascending, short to moderately long (Fig. 3A), mm cm 0.8-1. 0(-l. diam. Fronds to 38 long, erect, inserted onto the rhizome 5) at acute angles, closely arranged (internodes 1-5 mm), but not caespitose mm, X (Fig. 3A). Scales 3.0-5.0 0.5-0.6 12-16(-20) cells wide, clathrate brown dark brown, (Fig. 3B), to strongly iridescent, cordate pseudopeltate, to acute attenuate to at tips (Fig. 3C); apical cells 3-8, as furcate hairs or mm palmately arranged. Petioles 15-60(-75) long 3A), (0.4-)0.5-0.6(-0.8) (Fig. mm thick, decurrently marginate from the laminar most bases, parts terete mm many with brown, and (Fig. 3C), setiform ciliform hairs 1.0-2.0 long (Fig. 3C), simple and branched clavate hairs of crosiers and young fronds sometimes persistent in older fronds. Laminae firm-chartaceous, 150-320 X 4- mm, 9(-16) linear to narrow-elliptic (broadest in the middle), long-decurrent at base, acute to attenuate Rachises dark brown planar and at tip. to black, sunken between slightly segments adaxially, hemispherically protruding abaxially (Fig. 3D), glabrous or glabrescent with branched clavate hairs distally, usually sparsely to densely hairy basally on both sides with ciliform mm and setiform hairs like on the petioles. Segments 1.8-3.0(4.0) X 1.4-3.0 (1-2 times longer than broad), ascending (80-55), fully adnate, inequilateral at bases, basiscopically decurrent, most segments lunate to deltate with the tips obtuse truncate sometimes to segments with (Fig. 3E), distal short-oblong round tips (Fig. 3D); proximal segment pairs markedly smaller than the central segments, trapezoid with truncate tips, often auriculiform; midveins not visible (Figs. 3D, E), or obscurely so abaxially in dried specimens; segments glabrous except for hairs in margins without hydathodes sori; hairs; present. mm 2-4 Sori pairs per segment, with (3-)12-20 hairs 1.2-1.8 long 3D, (Figs. E), some rarely sori of a frond without hairs. The name narrow refers to the laminae with long curved which are tips, reminiscent whips of (Latin, flagellum). Melpomene flagellata occurs in elfin forests and moist montane forests at m 1950-3300 in eastern Peru and Bolivia (Fig. 7C). The segment shape Melpomene of between flagellata varies strongly rounded trapezoid to in small segments to deltate or short-oblong in larger & GOET]; Kelschebach Melpomene [Kessler 107, B. A. flagellata: t & GOET); Kelschebach GOET); apices [Kessler 107, D. petiole, C. detail ;ale GOET); oblong segments abaxially. sporangia upper part nert 517, E. part, right truncate segments abaxially, sporanj d ,PB); F. il {