Eleven new species in the grammitid fern genus Melpomene (Polypodiaceae).

ABSTRACT.--Based on extensive studies of grammitid ferns (Polypodiaceae) in the field and as specimens in various herbaria, eleven new species in the genus Melpomene are recognized from the Neotropics: Melpomene albicans, M. caput-gorgonis, M. flagellate, M. huencabambensis, M. jimenezii, M. michaelis, M. occidentalis, M. paradoxa, M. personate, M. sklenarii, and M. 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.

KEY WORDS.--Andes, Caribbean, grammitid ferns, Melpomene, Mesoamerica

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The genus Melpomene belongs to the grammitid ferns, which form a monophyletic clade within the Polypodiaceae (Schneider et al., 2004) and comprise about 750 species (Parris, 2005) with a pantropical--southern temperate distribution (Parris, 1990, 2003). Melpomene was established as a genus solely on 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, 2003). Because of its young taxonomical age, the genus has never been revised as a whole. Partial revisions are available for Mexico (Mickel and Smith, 2004), Mesomamerica (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 of species and morphological emphasis caused by genetic drift may lead to an easier distinction of species. The majority of specimens come from the Andean region, where the genus occurs at 200-5200 m and displays the greatest morphological diversity.

In order to revise the whole genus, a total of 2100 collections from following herbaria were examined: AAU, B, BM, BHCB, C, COL, COLO, CUZ, GOET, K, LOJA, LPB, P, QCA, QCNE, RB, S, SP, TUR, UC, US, USM. Among them are 251 specimens collected by the author in Ecuador, Peru, Bolivia, and Argentina, giving him further insight about putative correlations between ecological conditions and morphological plasticity. As suspected from previous studies on grammitid ferns (Labiak, 2000; Rojas, 2001; Leon and Smith, 2004), several taxa in Melpomene were indentified as yet undescribed species.

Most of the following species have been collected prior to this study but were not recognized as something new. Instead, their morphological differences were dismissed as intraspecific variability or aberrations. In this study it was observed that some widespread species have a great morphological plasticity, reflecting a high potential to answer a wide array of ecological factors. Good examples are Melpomene moniliformis (Lagasca ex. Sw.) A. R. Sm. & R. C. Moran and M. flabelliformis (Poir.) A. R. Sin. & R. C. Moran, whose distributions cover almost the whole range of the genus, the latter species being the sole representative of the genus in the Paleotropics. They inhabit many different types of habitats and consequently vary in size enormously, which often is correlated with a change of the proportions of the segments and in the number of cells across the scale bases. Opposed 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 species descriptions presented here. The reasons for the respective recognition of a species are given in the comments following the descriptions.

Terminology follows that of Lellinger (2002). Grammitid ferns are beset with two types of laminar indument that are distinct and not part of a homologous series. The term "seta" has often been applied to the stiff, darkened hairs that 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 rachises in Melpomene. However, I find that the so-called "setae" are differently developed within Melpomene, sometimes representing species autapomorphies and sometimes reflecting different growth conditions. For example, the "setae" of M. firma (J. Sin.) A. R. Sin. & R. C. Moran are usually quite short and patent, for which the term "acicular hairs" would be more fitting (Lellinger, 2002). In M. flabelliformis and many related species, the "setae" are sometimes flaccid and twisted, often occurring with properly developed (i.e., straight and rigid) "setae". Such flaccid trichomes are better described as "ciliform hairs" (Lellinger, 2002). For the rigid "setae," the term "setiform hairs" is available (Lellinger, 2002). Furthermore, used in Latin diagnoses, the term "seta" would translate as "bristle" (Stearn, 2004), which is a notably different structure (Lellinger, 2002) that is not found in Melpomene or other grammitid genera. I will use the terms "ciliform," "setiform," and "acicular hairs" to describe the simple pluricellular trichomes that are usually dark red in color with acute tips, and "clavate hairs" for the minute, simple and branched, pluricellular hairs that are pale in color and bear clavate cells at their apices or along their length. This second type of hairs has been referred to as "glandular" (Parris, 2005) and they appear to be homologous to the secretive gland-bearing branched hairs of Zygophlebia (Bishop, 1989), but a secretive nature was not observed in the hairs of Melpomene so far. The same can be stated for the "glands" that occur on the scale apices of all Melpomene species. They are thin walled cells that are arranged singly or as pluricellular papillae or branched clavate hairs at the tips and 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 cells. Scales are only found on the rhizomes and sometimes on the petiolar bases.

The dimensions of the segments are measured with the midveins as cardinal points, which are assumed as medial lines in the central parts of the segments between the sori if they are obscure. Segment length is measured along the midvein from segment tip to rachis; segment width is taken as the orthogonal line at half of the length. The dimensions are more useful if the width is taken in the central part of the segments because deltate segments are usually strongly decurrent onto the rachises in their proximal half and width taken here may vary considerably.

Melpomene albicans Lehnert, sp. nov. TYPE.--BOLIVIA. Santa Cruz: Prov. Caballero, Comarapa, ca. 1 km de Siberia hacia Torecillos, 17[degrees]49'S, 64[degrees]40'W, 2650 m, 18 Mar 2003, Lehnert 714 (holotype GOET; isotypes LPB, UC). Figs. 1, 7A.

A Melpomene personata laminis abaxialiter albicantibus, a M. youngii (Stolze) B. Leon & A. R. Sm. pilis pluribus in soris confertis praestans.

Plants predominantly epiphytic or epilithic, growing in moss layers. Rhizomes moderately to short-creeping, horizontal (Fig. 1A), 0.8-1.2 mm diam., sometimes with short branches (5-10 mm) held at right angles. Fronds to 23 cm long, erect (Fig. 1A) or patent, inserted onto the rhizome at acute angles, but often held at nearly right angles to the rhizome, closely arranged (internodes 2-5 mm) but not caespitose (Fig. 1A). Scales 3.0-5.0(-7.5) x (0.3-) 0.6-0.8(-1.0) mm, (12-)16-20(-22) cells wide, lanceolate, clathrate (Fig. 1B), dark brown to brown, strongly iridescent, bases cordate to broadly cordate, tips acute to short-attenuate; apical cells 3-8, in a nodding cluster or palmate arrangement (Fig. 1C). Petioles (24-)35-85 mm long, 0.5-0.8 mm thick, marginate from the laminar bases, most parts terete, glabrous to glabrescent with dark brown setiform hairs 0.75-1.2 mm long (Fig. 1D); simple and branched clavate hairs (0.1-0.3 mm) of crosiers and young fronds often persist in older fronds. Laminae firm-chartaceous, to 120-75 x 18-35 mm (2/3 to 4/5 of frond length), narrow-elliptic to elliptic (widest in the middle), rarely obovate (widest above the middle), cuneate or somewhat tapering at bases, acute to attenuate at tips (Fig. 1A); surfaces abaxially whitish to white, often yellow or ochre when dried, usually eliminated if treated with alcohol. Rachises dark brown to black, planar and slightly sunken between segments adaxially (Fig. 1F), hemispherically protruding abaxially (Fig. 1E), with few to many branched clavate hairs like on petioles, otherwise glabrous or with scattered acicular hairs. Segments to 9.0-16.0 x 1.2-2.6(-3.0) mm (6-8 times longer than broad), flat, weakly ascending (80-70[degrees]), equilateral at bases or weakly decurrent basiscopically, fully adnate, linear-oblong to long-deltate, the tips obtuse or short-acute; midveins not visible, or obscurely so abaxially in dried specimens (Fig. 1E, 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. 1F). Sori 2-8(-9) pairs per segment, with 4-10 dark-castaneous setiform hairs 0.5-0.8 mm long (Fig. 1E).

[FIGURE 1 OMITTED]

The name refers to the white wax-like deposit on the abaxial laminae (Latin, albicans = being white).

Melpomene albicans grows in elfin forests, cloud forests, and moist montane forests at 1500-3400 m in Bolivia and eastern Brazil (Fig. 7A).

There are only a few other species of Melpomene with whitish abaxial laminar surfaces. Among these, Melpomene 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 the more strongly ascending segments; Melpomene sodiroi (Christ & Rosenst.) A. R. Sm. & R. C. Moran matches M. albicans in the hair distribution, but has gibbose segments with the sori slightly to deeply sunken (vs. laminae planar with superficial sori in M. albicans) and larger scales (on average 6.0-9.0 x 1.0-1.2 mm, 24-30 cells wide vs. 3.0-5.0 x 0.6-0.8 mm, 16-20 cells wide) with characteristically long tapering tips (vs. acute to short-attenuate). Melpomene erecta (C. V. Morton) A. R. Sm. & R. C. Moran, which only rarely has white laminar surfaces, has thicker rhizomes (to 2.4 mm diam. vs. to 1.6 mm diam. in M. albicans) and wider scales (on average 38-60 cells vs. 16-20 cells across base).

Melpomene albicans belongs to a complex comprising M. personata, M. peruviana (Desv.) A. R. Sm. & R. C. Moran, M. sodiroi, and M. youngii, which all have horizontally creeping rhizomes, relatively dark, lanceolate scales, and predominantly downward pointed fronds. Melpomene albicans is the only species in the complex that has patent fronds when growing epiphytically or even strongly ascending fronds when growning epilithically. All have relatively long setiform/ciliform hairs in the sori except for M. youngii, where the hairs are usually lacking or relatively short and sparse (up to 2 hairs per sorus). Melpomene personata and M. peruviana differ from M. 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 single hairs along the segment margins, which are always absent in M. albicans. Melpomene peruviana sometimes has whitish green laminae but lacks a wax-like layer. Additionally, M. peruviana is on average much smaller than M. albicans and M. personata (2.5-10.0(-17.5) cm vs. to 17.5-25.0 cm), tends to form dense tufts or cushions (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 of Melpomene 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 petioles. The Brazilian plants are more easily confused with M. xiphopteroides (Liebm.) A. R. Sm. & R. C. Moran than the Andean population because the white laminar layers are less developed and the scales are smaller, and thus closer to the morphological spectrum of M. xiphopteroides. However, even weakly hairy plants of M. xiphopteroides have more setiform/ciliform hairs on the rachises than M. albicans, in which 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[degrees]31'S, 41[degrees]53'W, 21 Mar 1999, Saline & Morals 4538 (UC). Parana: Curitiba, Serra Ibitiraque, Morro Cainapua (Mun. Campina Grande do Sul), 25[degrees]08'S, 49[degrees]04'W, 02 Mar 2000, Ribas et al. 3080 (UC). Rio de Janeiro: Theresopolis, ca. 22[degrees]25'S, ca. 42[degrees]58'W, Oct 1929, Brade 9094 (NY); boundary between municipio de Teresopolis and Municipio de Petropolis, Serra dos Orgaos National Park, ca. 5 km SW of city of Teresopolis, 22[degrees]27-28'S, 43[degrees]01-02'W, 30 Nov 1965, Eiten & Eiten 7168 (US).

BOLIVIA. Cochabamba: Prov. Ayopaya, San Cristobal, climbing along the trail that leads to San Miguel, 16[degrees]39'S, 66[degrees]43'W, 3100 m, 06 Jun 2002, Jimenez I. 1107B (GOET, LPB, UC); Prov. Carrasco, on the way from Comarapa to Siberia, 17[degrees]50'S, 64[degrees]42'W, 3000 m, 22 Jan 2000, Jimenez I. 283 (LPB, UC); 10 km from Siberia to Comarapa, 17[degrees]48'S, 64[degrees]42'W, 2600 m, 20 Oct 1996, Kessler et al. 9164 (LPB, UC); Prov. Carrasco, 10 km Cocapata-Cotacajes, 16[degrees]38'S, 66[degrees]41'W, 3000 m, 09 May 1997, Kessler et al. 9401 (GOET, LPB, UC); road Cochabamba-Villa Tunari, below Corani, 17[degrees]10.58'S, 65[degrees]53.67'W, 2700 m, 26 Nov 2002, Lehnert 512 (GOET, LPB, UC); road Cochabamba-Villa Tunari, below Corani, 17[degrees]10.58'S, 65[degrees]53.67'W, 2700 m, 26 Nov 2002, Lehnert 514 (GOET, LPB, UC); Prov. Chapare, Cochabamba 54 km hacia Villa Tunari, 2750 m, 30 Apr 1979, Beck 1424a (LPB). La Paz: Prov. Inquisivi, "Kinpaya," at the mouth of the Rio Jancha Kaihua where the Aquilani-Choquetanga trail crosses the Rio Ocsalla, 10 km N of Choquetanga, 16[degrees]45'S, 67[degrees]17'W, 3400 m, 07 Sep 1991, Lewis 39952 (LPB); comunidad Choquetanga-Wichupampa, Serranias de Lulini 13 km al N de Choquetanga, 2-3 km al NW del cerro Lulini, 16[degrees]45'S, 67[degrees]20'W, 3290 In, 17 Mar 1994, Salinas 2783 (US); Prov. Nor Yungas, Coscapa, on prehispanic trail Sillutinkara, 16[degrees]12'S, 67[degrees]53'W, 3100-3300 m, 07 Jan 2001, Jimenez I. & Vidaurre 526 (LPB, UC), 559 (GOET, LPB, UC); Unduavi, trench to the Valle de Coscapa, 16[degrees]17'S, 67[degrees]51'W, 3350 m, 04 Feb 2003, Lehnert 599, 601, 602 (GOET, LPB, UC). Santa Cruz: Prov. Caballero, Comarapa, between Torecillos and Siberia, 17[degrees]49.65'S, 64[degrees]40.14'W, 2600-2700 m, 18 Mar 2003, Lehnert 696, 707 (GOET, LPB, UC); from Siberia 4 km to the E, small laguna on the ridge (Laguna Tinque?), 2600 m, 18 Mar 2003, Lehnert 717 (GOET, LPB, UC); by small lake at summit of pass ca 4 km E of Siberia, 2800 m, 04 Jan 2000, Wood & Goyder 15792 (LPB).

Melpomene caput-garganis Lehnert, sp. nov. TYPE.--BOLIVIA. La Paz: Prov. Nor Yungas, Cotapata, detras del gasolinero, 3200 m, 16[degrees]17'S, 67[degrees]51'W, 25 Sep 2002, Lehnert 367 (holotype: LPB; isotypes: GOET, UC). Figs. 2, 7B.

A Melpomene flabelliformi petiolis brevioribus, apicibus squamarum latioribus (3-4 vs. 1-2 cellulis latis), cellulis apicalibus creberioribus, absentia pilorum inter sores differt.

Plants predominantly epiphytic, growing in moss layers, rarely epilithic. Rhizomes moderately to short-creeping, horizontal (Fig. 2A), 1.0-1.5 mm diam. Fronds to 35-42 cm, arching to pendent, inserted onto the rhizomes at right angles (Fig. 2A), closely arranged (internodes 1-4 mm). Scales to 6.5 x 0.8-1.0 mm, (18-)20-26(-32) cells wide, clathrate, auburn to brown, strongly iridescent, broadly cordate to pseudopeltate, with obtuse to truncate tips ending in rows of 3-4 cells (Fig. 2B); apical cells numerous, sitting clustered on the wide tips (Fig. 2C). Petioles 15-50(-70) mm long, 0.6-0.8(-1.0) mm thick, decurrently marginate from the laminar bases, with brown flaccid ciliform hairs (1.0-2.0 mm) on both sides (Fig. 2D), simple and branched clavate hairs of crosiers and young fronds sometimes persistent in older fronds, but generally glabrescent. Laminae firm-chartaceous, rarely coriaceous (Lehnert 781), to 300(-350) x 14-22 mm, narrow-elliptic (broadest in the middle), decurrent at bases, acute to attenuate at tips, sometimes caudate (Fig. 2A). Rachises dark brown, flat, and slightly sunken between segments adaxially (Fig. 2E), weakly hemispherically protruding abaxially (Fig. 2F), glabrous except for the proximal part, with ciliform hairs as on petioles. Segments 4.5-9.0(-12.0) x 3.2-3.8(-5.0) mm (1.5-3 times longer than broad), weakly ascending (80-70[degrees]), oblong to lunate, equilateral at bases or weakly 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 pairs markedly smaller than the central segments (Fig. 2A), inequilateral at bases, basalmost auriculiform (e.g., Lehnert 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 receptacular and circumsoral ciliform hairs 1.0-1.5 mm long (Fig. 2F).

[FIGURE 2 OMITTED]

The name refers to the numerous apical cells on 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 in wet montane forests and elfin forests at 2680-3200 m in southern Peru and Bolivia (Fig. 7B).

The most distinguishing feature of this species is the broad scales with the abundant apical papillae. No other species of Melpomene has scale apices that provide a base for papillae several cells wide. Melpomene flabelliformis can be distinguished from M. caput-gorgonis by its distant fronds and glabrous or glabrescent petioles (vs. fronds close and petioles persistently hairy in M. caput-gorgonis). Melpomene flagellata is generally more slender in habit and most features, i.e., laminae to 9(-16) mm wide (vs. to 22 mm), rhizomes thinner and ascending, petiole scales narrower, and hairs in the sori longer.

Melpomene caput-gorgonis grows together with M. paradoxa, which can be distinguished by its longer, glabrescent petioles and shorter segments; the latter species also forms patches with the fronds erect to arching whereas M. caput-gorgonis grows solitary with arching to pendent fronds.

ADDITIONAL SPECIMENS EXAMINED.--PERU. Cuzco: Abra de Chaupimayo, Hacienda Pintubamba, 2700 m, Sep 1932, Bues 1945 (CUZ).

BOLIVIA. La Paz: Prov. Franz Tamayo, PN-ANMI Madidi, trail Keara-Mojos, below Chunkani, 2870 m, 14[degrees]38'S, 68[degrees]57'W, 08 Nov 2001, Jimenez I. & Gallegos 917 (LPB, UC); Prov. Larecaja, toll house above Mapiri, 2000 m, 12 Sep 1901, Williams 1154 (NY); Prov. Nor Yungas, Estacion Biologica Tunquini, Bajo Hornuni, senda del campo de Don Pedro al camino de lamina, 2550 m, 16[degrees]11'S, 67[degrees]53'W, 17 Aug 2000, Jimenez I. et al. 488 (LPB, UC); Coscapa, along the prehispanic trail Sillutinkara, 16[degrees]12'S, 67[degrees]53'W, 3100 m, 07 Jan 2001, Jimenez I. & Vidaurre 534 (LPB, UC); trench to the Coscapa valley, Parque Nacional Cotapata, 16[degrees]12'S, 67[degrees]33'W, 3000 m, 12 Dec 1997, Kessler et al. 1871 (LPB); 2 km from Chuspipata to Coroico, 16[degrees]22'S, 67[degrees]49'W, 2900 m, 19 Sep 1997, Kessler et al. 11921 (GOET, LPB, UC); Cotapata, behind the gas station, 16[degrees]17'S, 67[degrees]51'W, 3150-3200 m, 25 Sep 2002, Lehnert 367, 368, 369, 372, 373, 386, 392 (GOET, LPB, UC); Chuspipata Sacramento, 16[degrees]18'S, 67[degrees]49'W, 2680 m, 10 Nov 2002, Lehnert 496a (GOET, LPB, UC); 1.2 km E de Cotapata-Santa Barbara, trail to the Chuspipata electricity station, 16[degrees]17'S, 67[degrees]50'W, 3200 m, 02 Feb 2003, Lehnert 586 (GOET, LPB, UC); Chuspipata, old trail to Unduavi, 3200 in, 07 May 2003, Lehnert 781 (GOET, LPB, UC).

Melpomene flagellata Lehnert, sp. nov. TYPE.--BOLIVIA. La Paz: Prov. Nor Yungas, N side of Cerro Uchumachi above Coroico, 16[degrees]12'S, 67[degrees]45'W, 2350 m, 14 Jul 1989, Kessler & Kelschebach 107 (holotype LPB; isotype GOET). Figs. 3, 7C.

A Melpomene moniliformi pilis longioribus (1.0-2.0 mm vs. 0.5-1.0 mm), creberioribus in petiolis sorisque, rhizomatibus ascendentibus (vs. horizontaliter reptantibus) apicibusque segmentorum partim truncatis (vs. semper obtusis vel rotundis) differt.

Plants predominantly epiphytic, growing in moss layers, sometimes epilithic. Rhizomes erect or ascending, short to moderately long (Fig. 3A), 0.8-1.0(-1.5) mm diam. Fronds to 38 cm long, erect, inserted onto the rhizome at acute angles, closely arranged (internodes 1-5 mm), but not caespitose (Fig. 3A). Scales 3.0-5.0 x 0.5-0.6 mm, 12-16(-20) cells wide, clathrate (Fig. 3B), dark brown to brown, strongly iridescent, cordate to pseudopeltate, acute to attenuate at tips (Fig. 3C); apical cells 3-8, as furcate hairs or palmately arranged. Petioles 15-60(-75) mm long (Fig. 3A), (0.4-)0.5-0.6(-0.8) mm thick, decurrently marginate from the laminar bases, most parts terete (Fig. 3C), with many brown, setiform and ciliform hairs 1.0-2.0 mm long (Fig. 3C), simple and branched clavate hairs of crosiers and young fronds sometimes persistent in older fronds. Laminae firm-chartaceous, 150-320 x 4-9(-16) mm, linear to narrow-elliptic (broadest in the middle), long-decurrent at base, acute to attenuate at tip. Rachises dark brown to black, planar and slightly sunken between 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 and setiform hairs like on the petioles. Segments 1.8-3.0(-4.0) x 1.4-3.0 mm (1-2 times longer than broad), ascending (80-55[degrees]), fully adnate, inequilateral at bases, basiscopically decurrent, most segments lunate to deltate with the tips obtuse to truncate (Fig. 3E), sometimes distal segments short-oblong with 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 sori; margins without hairs; hydathodes present. Sori 2-4 pairs per segment, with (3-)12-20 hairs 1.2-1.8 mm long (Figs. 3D, E), rarely some sori of a frond without hairs.

The name refers to the narrow laminae with long curved tips, which are reminiscent of whips (Latin, flagellum).

Melpomene flagellata occurs in elfin forests and moist montane forests at 1950-3300 m in eastern Peru and Bolivia (Fig. 7C).

The segment shape of Melpomene flagellata varies strongly between trapezoid to rounded in small segments to deltate or short-oblong in larger ones. Melpomene flagellata replaces M. wolfii (Hieron.) A. R. Sm. & R. C. Moran, which at first sight is almost identical, in Bolivia and eastern Peru (Fig. 7C). Melpomene flagellata, however, has thinner rhizomes, thinner petioles with thinner alae, and narrower rhizome scales than M. wolfii; also, M. wolfii lacks hairs on the petioles. Pubescence of fertile laminae is similar and highly variable in both species, varying from glabrous to densely hairy in even one plant. In M. flagellata, the ciliform and setiform laminar hairs reach 1.2-1.8 mm length and are clustered in the sori; in M. wolfii, the laminar hairs are shorter (0.5-0.8 mm) and evenly distributed on the laminae.

[FIGURE 3 OMITTED]

Melpomene moniliformis differs from M. flagellata in the horizonally creeping rhizomes and shorter (0.5-1.0(-1.5) vs. 1.0-2.0 mm) setiform and ciliform hairs on the petioles. The soral hairs of M. flagellata are on average longer and more abundant than in M. moniliformis (12-20 hairs, 1.2-1.8 mm long vs. 1-5 hairs, 0.5-1.0 mm long) and thus more conspicuous. Melpomene moniliformis also lacks segments with truncate tips, which are characteristic of M. flagellata.

With the mentioned characters, Melpomene flagellata can be readily determined although it shows a considerable variation in size and appearance within its small range. Kessler 7320 has very narrow blades; Kessler 7318 from the same location is equal in frond length but has broader laminae; Kromer & Acebey 1368 is generally larger than average plants of M. flagellata but has characteristic short segments, fitting hair distribution, and small scales (16 cells wide across base). Molecular data indicate a stronger affinity of M. flagellata to M. moniliformis than to the similar M. wolfii (Lehnert et al., in press).

ADDITIONAL SPECIMENS EXAMINED.--PERU. Cuzco: Alturas de Sicre, La Convencion, 3000 m, Jun 1924, Bues 1574 (CUZ). Puno: Sachapata, Ostabhang der Cordilleren von Peru, Sep 1854, Lechler 2713 (B).

BOLIVIA. Cochabamba: Prov. Carrasco, 132 kin [on] old road Cochabamba-Villa Tunari, 17[degrees]06'S, 65[degrees]35'W, 1950 m, 15 Jul 1996, Kessler et al. 7318 (LPB, UC), 7320 (GOET, LPB, UC); 130 km [on] old road Cochabamba-Villa Tunari, 17[degrees]07'S, 65[degrees]36'W, 2000 m, 10 Jul 1996, Kessler et al. 7168 (GOET, LPB, UC); road Cochabamba-Villa Tunari, below Corani, 17[degrees]10.51'S, 65[degrees]54.02'W, 2750 m, 26 Nov 2002, Lehnert 517 (GOET, LPB, UC). La Paz: Prov. Franz Tamaya, PN-AMNI Madidi, trail Keara-Mojos, about half an hour from Tokuaque, 14[degrees]37'S, 68[degrees]57'W, 2420 m, 01 Jul 2001, Jimenez I. & Gallegos 527 (UC); Prov. Larecaja, Sorata, 15[degrees]44.04'S, 68[degrees]39.28'W, 3300 m, 06 Jan 2003, Lehnert 555 (GOET, LPB, UC); Prov. Nor Yungas, between Chuspipata and Yolosa, above Sacramento, 2760 m, 01 Jan 1994, Beck 21310 (LPB, UC); Parque Nacional Cotapata, surroundings of Estacion Biologica Tunquini, 16[degrees]11'S, 67[degrees]52'W, 2300 m, 26 Jul 2000, Kromer & Acebey 1368 (GOET, LPB, UC); Chuspipata-Sacramento, 16[degrees]18'S, 67[degrees]49'W, 2900 m, 10 Nov 2002, Lehnert 491 (GOET, LPB, UC).

Melpomene huancabambensis Lehnert, sp. nov. TYPE.--PERU. San Martin: Prov. Rioja, Buenos Aires, along road Pedro Ruiz-Rioja, 2000 m, 05[degrees]42'09"S, 77[degrees]53'06"W, van der Werff et al. 15352 (holotype: UC; isotype: MO). Figs. 4, 7D.

A Melpomene pilosissima (M. Martens & Galeotti) A. R. Sm. & R. C. Moran absentia vel praesentia irregulari hydathodorum, a M. michaeli, M. xiphopteroide et M. jimenezii pilis in paginis laminarum abaxialibus aequaliter distributis (vs. in soris confertis) differt.

[FIGURE 4 OMITTED]

Plants epiphytic in moss layers. Rhizomes stout, very short-creeping to erect, 1.0 mm diam. Fronds to 33.5 cm long, pendent (Fig. 4A), inserted onto the rhizomes at narrow angles, fasciculate (internodes 0.2-1.0 mm). Scales 2.0-3.2 x 0.3-0.5 mm, (10-)12-14(-16) cells wide, clathrate (Fig. 4B), dark brown, strongly iridescent, narrowly cordate, long-acute to attenuate at tips; apical cells 3-5, in a nodding cluster or linear arrangement. Petioles (12-)45-55 mm long, 0.6-1.0 mm thick, terete, densely hairy with setiform and ciliform hairs, usually rather lax, dark brown to castaneous, (1.8-)2.0-3.5 mm long (Fig. 4C); simple and branched clavate hairs absent or very sparse. Laminae papyraceous to chartaceous, to 140-290 x 29-36 mm, narrowly elliptic (broadest at or beyond the middle), cuneate to shortly decurrent at bases, long-acute at tips (Fig. 16A). Rachises deeply dark brown to black, planar adaxially (Fig. 4D), strongly hemispherically protruding abaxially (Fig. 4E), strongly hairy abaxially, moderately hairy adaxially, proximally more densely hairy than distally, setiform and ciliform hairs castaneous, 1.5-3.0 mm long. Segments 22-28 x 8 mm (3-4 times longer than broad), central segments patent or nearly so (90-80[degrees]), distal segments weakly ascending (75-65[degrees]), equilateral at bases, fully adnate, long-deltate to oblong, the tips obtuse; midveins obscurely visible in dried specimens (Fig. 4D, E); proximal segment pairs markedly smaller than the following segments (1/2 to 1/3 of longest segments) (Fig. 4A), sometimes the lowermost 2-5 segment pairs auriculiform; setiform hairs 1.0-2.0 mm long, evenly distributed on laminae abaxially, also in sori and along midveins but not clustered here, hairs present adaxially mainly along the midveins, always some on the margins (Fig. 4E); hydathodes lacking (Fig. 4D) or weakly developed in parts of a frond, small and inconspicuous. Sori 3-9 pairs per segment, (0-)1-3 setiform hairs to 1.5 mm long in them, more hairs around them on the laminae, often covering the sori (Fig. 4E).

The name refers to the geographic restriction of the species to the Amotape-Huancabamba region.

Melpomene huancabambensis grows in montane forests at 1900-2200 m in northeastern Peru and southern Ecuador (Fig. 7D).

This species is an ally of M. pilosissima, with which it matches in the densely hairy, terete petioles and the hairy laminae, which may have hairs along the segment margins and the adaxial surface. It is characterized by the lack or poor development of hydathodes (vs. always present and well-developed in M. pilosissima). The often large size of the plants, relatively short petioles, and the predominantly patent segments of M. huancabambensis create a habit that is quite different from that of M. pilosissima, where the petioles tend to be longer and the segments more ascending. Melpomene pilosissima further differs from M. huancabambensis in being usually less hairy on the segments adaxially; however, there are some exceptions. Some plants of M. pilosissima are equal to M. huancabambensis in the hairiness of the adaxial segment surfaces, but tend to have more acute segment tips, discontinuously visible midveins, and decurrently marginate petioles (all characters not present in M. huancabambensis).

Melpomene vernicosa (Copel.) A. R. Sm. & R. C. Moran and larger plants of Melpomene xiphopteroides are superficially similar to M. huancabambensis but they differ in having hairs clustered in the sori (vs. not clustered in sori in M. huancabambensis). Melpomene vernicosa also has glabrous segment margins (vs. hairy in M. huancabambensis) and more coriaceous laminae (vs. papyraceous to chartaceous). Other species with absent or weakly developed hydathodes (M. jimenezii, M. michaelis), which are characteristic of M. huancabambensis, differ in having ascending segments, more coriaceous laminae, and hairs clustered in the sori.

ADDITIONAL SPECIMENS EXAMINED.--ECUADOR. Zamora-Chinchipe: P.N. Podocarpus, park entrance "San Francisco," on new Loja-Zamora rd., 03[degrees]59.349'S, 79[degrees]05.713'W, 2151 m, 14 Dec 2006, Sundue 1123 (NY, UC). PERU. San Martin: Prov. Rioja, along road Pedro Ruiz-Rioja, 05[degrees]42'09"S, 77[degrees]53'06"W, 2000 m, 21 Mar 1998, van der Werff et al. 15353 (MO, UC); ibid., E1 Mirador, 1900 m, 26 Mar 1998, van der Werff et al. 15749 (MO, NY, UC).

Melpomene jimenezii Lehnert, sp. nov. TYPE.--BOLIVIA. La Paz: Prov. Franz Tamayo, Parque Nacional Madidi, trail Pelechuco-Mojos, locality Tambo Quemado (camping area), on the trail towards Qalla, crossing the fourth river and going up the trail which leads through the Polylepis forest, 14[degrees]41'S, 68[degrees]58'W, 3490 m, Jimenez I. 1859 (holotype: LPB; isotypes: GOET, UC). Figs. 5A-D, 7D.

A Melpomene pilosissima absentia hydathodorum (vel hydathodibus sparse et irregulariter praesentibus vs. semper paesentibus M. pilosissima), a M. michaeli squamis maioribus (usque 5.5-7.5 vs. 2.0-3.0 mm longis), pilis inter soros sparse distributis vel carentibus (vs. pilis crebris inter soros) praestans.

Plants epiphytic, growing in moss layers. Rhizomes short-erect (Fig. 5A), 1.2-1.8 mm diam. Fronds to 35 cm long, pendent, inserted onto rhizomes at acute angles, closely arranged (internodes 1-4 mm), caespitose (Fig. 5A). Scales 5.5-7.5 x 0.4-0.6 mm, (8-)14-18(-20) cells wide across base, clathrate (but usually many lumina occluded), dark brown to brown, weakly iridescent, subcordate to cordate at bases, long attenuate at tips (Fig. 5B); apical cells 5-12, palmately arranged or in a nodding cluster (Fig. 5C). Petioles 40-80 mm long, 0.8-1.0 mm diam., terete, with many dark brown setiform and ciliform hairs 1.2-3.0 mm long (Fig. 5A), simple and branched clavate hairs of crosiers and young fronds persistent on older fronds. Laminae 205-285 x 30-64 mm, coriaceous, adaxially matte, lanceolate or broadly elliptic (broadest at or below the middle), acute to attenuate at tips, bases rounded to cuneate (Fig. 5A). Rachises dark brown to black, planar to weakly protruding adaxially, hemispherically protruding abaxially (Fig. 5D), with few to many brown setiform and ciliform hairs on both sides (1.0-1.5 mm), usually hairier abaxially than adaxially; abaxially also with many simple and branched hairs. Segments 18.0-35.0 x 2.8-4.0 mm (6-12 times longer than broad), patent to weakly ascending (80-60[degrees]), fully adnate, equilateral at bases, or weakly decurrent towards the blade apices, tips obtuse (Fig. 5A); proximal segment pairs oblong to deltate, smaller than the following segments, but not auriculiform, often remote; margins entire or dentate basally in large segments (Fig. 5D), fertile segments often conduplicate; midveins not visible on both laminar sides, or obscurely so abaxially in dried specimens; stomata usually visible as dark brown to reddish dots (Fig. 5D); setiform/ciliform hairs to 2 mm long, clustered in sori, lacking or sparse along midveins or on laminae abaxially, margins and adaxial laminar surface glabrous (Fig. 5D); hydathodes usually absent, sometimes weakly developed in some parts of a frond. Sori 3-10 pairs per segment, each with 4-10 ciliform hairs (0.8-1.5 mm).

[FIGURE 5 OMITTED]

The species is named after Ivan Jimenez, Bolivian botanist and colleague, who collected the type specimen.

Melpomene jimenezii grows in elfin forest and moist montane forests at 2400-3800 m in Peru and Bolivia (Fig. 7D).

This species is remarkable because it lacks conspicuous hydathodes in most fronds. Occasionally, some segments have small hydathodes, while hydathodes are otherwise absent on the same frond. Other species with such irregular development of hydathodes are M. huancabambensis and M. michaelis. According to Parris (1997), absence and presence of hydathodes may change in many grammitid taxa in the Paleotropics, but until now it has been considered a reliably constant feature for the Neotropical genera.

The prevailing lack of hydathodes separates M. fimenezii from the superficially similar M. firma, which always has well developed hydathodes and also differs in having conspicuous black midveins (vs. midveins not visible or obscure in M. jimenezii) and fewer, shorter hairs on the petioles (vs. densely long-hairy).

From Melpomene pilosissima, M. jimenezii differs in having longer segments, ciliform hairs clustered in the sori (vs. evenly distributed on the laminae), and in lacking setiform or ciliform hairs along the margins and the adaxial laminae (vs. regularly hairy along the margins and the adaxial laminae). The smaller M. michaelis is similar, but differs from M. jimenezii by having setiform and ciliform hairs on the laminae between the sori (vs. restricted to the sori), lacking red stomata (vs. stomata often red), and having rhizome scales about half the size of those of M. jimenezii (2.0-3.0 vs. 5.5-7.5 mm long).

ADDITIONAL SPECIMENS EXAMINED.--PERU. Amazonas: Leimebamba, 2400 m, 30 Dec 1962, Woytkowski 7839 (UC). Pasco: Prov. Oxapampa, Huancabamba district, locality Lanturachi, sector Santa Barbara, 10[degrees]21'S, 75[degrees]39'W, 3800 m, 10 Oct 2003, Perera et al. 531 (MO, USM).

BOLIVIA. La Paz: Prov. Franz Tamayo, PN-ANMI Madidi, senda Pelechuco-Mojos, por el sendero que va hacia Qalla, cruzando el 4[degrees] rio y subiendo por el sendero que atravieza el bosque de Polylepis, 14[degrees]41'S, 68[degrees]58'W, 3360 m, 03 May 2003, Jimenez I. I842A (LPB, UC); ibid., localidad Tambo Quemado (lugar para acampar), 3250 m, 14[degrees]39'S, 68[degrees]57'W, 04 May 2003, Jimenez 1891 (LPB, UC).

Melpomene michaelis Lehnert, sp. nov. TYPE.--BOLIVIA. Cochabamba: Prov. Carrasco, carretera Cochabamba-Villa Tunari, debajo de Corani, 2750 m, 17[degrees]10.51'S, 65[degrees]54.02'W, Lehnert 519 (holotype: GOET; isotypes: LPB, UC). Figs. 5E-K, 7D.

A Melpomene pilosissima pilis in soris confertis, hydathodis partim reductis, a M. jimenezii squamis minoribus (usque 2.0-3.0 vs. 5.5-7.5 mm longis), pilis crebris inter soros (vs. pilis inter soros sparse distributis vel carentibus) differt.

Plants predominantly epilithic, also epiphytic. Rhizomes very short-creeping (Fig. 5E), ascending to erect, 0.8-1.4 mm diam., rarely branched. Fronds to 18 cm long, rather laxly pendent, inserted onto the rhizomes at acute angles, closely arranged (internodes 0.2-1.0 mm), caespitose (Fig. 5E). Scales to 2.0-3.0 x 0.4-0.6 mm, (8-)10-12(-16) cells wide across base, clathrate (Fig. 5F), dark brown to brown, weakly iridescent, narrowly lanceate, narrowly cordate basally, attenuate at tips; apical cells 1-8, linearly arranged (Fig. 5H); sometimes cells of scales unexpanded, then scales blackish and not iridescent (Fig. 5G). Petioles 26-55 mm long, 0.4-0.6 mm diam., terete, with many dark brown, setiform and ciliform hairs 1-3 mm long; simple and branched clavate hairs of crosiers and young fronds sometimes persistent on older fronds. Laminae to 115-125 x 11-20 mm (2/3 of frond length), firm-chartaceous, adaxially usually shiny, narrowly elliptic to linear-elliptic (widest at and/or below the middle), round to broadly cuneate at bases, short-attenuate to acute at tips (Fig. 5E). Rachises dark brown to black, planar and slightly sunken between segments adaxially, hemispherically protruding abaxially, densely hairy as on petioles abaxially and in the proximal half adaxially. Segments 4.5-7.2 x 1.4-2.2 mm (2.5-3.0 times longer than broad), weakly ascending (80-75[degrees]), inequilateral at bases, fully adnate, long-deltate to oblong, the tips obtuse (Figs. 5J, K); midveins not visible, or obscurely so abaxially in dried specimens; proximal 1-3 segment pairs markedly smaller than the following segments, but not auriculiform; few to many setiform and ciliform hairs 1-2 mm long on the abaxial laminar surface, clustered in and around sori, always some along the margins, rarely some adaxially; stomata not visible as dark spots; hydathodes small, inconspicuous and regularly lacking in some parts of the fronds (Fig. 5J). Sori 1-6 pairs per segment, with 1-3 long setiform and ciliform hairs 1.5-2.0 mm long (Fig. 5K).

The name honors both Michael Kessler, my mentor at Gottingen University, and Michael Sundue, a colleague from New York Botanic Garden, who first recognized this species as being distinct.

Melpomene michaelis grows in wet montane forests at 2250-3450 m in Peru and Bolivia (Fig. 17A).

A noteworthy character of Melpomene michaelis is the inconspicuous, sometimes partially absent hydathodes. In gross morphology, M. pilosissima is virtually identical to M. michaelis, but has always well-developed hydathodes and evenly distributed setiform/ciliform hairs on the abaxial laminar surface; there may be few hairs present in the sori, but they are not clustered here like it is the case in M. michaelis. Melpomene pilosissima occurs in Mesoamerica and the northern Andes and shows no geographical overlap with M. michaelis, which is confined to the centrals Andes. Ecologically, Melpomene michaelis prefers the epilithic habitat whereas M. pilosissima is mainly found as an epiphyte.

The distribution of Melpomene michaelis overlaps with that of M. jimenezii, which--like M. michaelis--has setiform/ciliform hairs clustered in the sori. Apart from the larger size, M. jimenezii differs in having longer, more linear segments than M. michaelis and in lacking hydathodes, or having occasionally small hydathodes in some segments (vs. hydathodes inconspicuous but predominantly present in M. michaelis). It also lacks hairs between the sori on the abaxially laminar surface (vs. sparsely to densely hairy between the sori in M. michaelis), on the segments margins, and adaxial laminar surface (vs. sometimes with scattered hairs on margins and adaxial laminar surface).

A peculiar trend is observed in the rhizome scales of M. michaelis: already having scales with smaller cells than in most Melpomene species on average, the cells in the southern populations are often not expanded, with thick cell walls occluding the usually translucent lumina.

ADDITIONAL SPECIMENS EXAMINED.--PERU. Cuzco: Prov. Urubamba, Machu Picchu, along trail from Machu Picchu ruins to Winay Waina, 2300 m, 21 Jan 1976, Bishop 2513 (UC); entre San Luis y Abra Malaga, 13[degrees]06.22'S, 72[degrees]22.42'W, 3450 m, 16 Oct 2002, Lehnert 443 (GOET, UC, USM).

BOLIVIA. Cochabamba: Prov. Ayopaya, San Cristobal, subiendo por el sendero que va a San Miguel, 16[degrees]39'S, 66[degrees]43'W, 3250 m, 06 Jun 2002, Jimenez I. 1149 (LPB, UC); Prov. Carrasco, Sehuencas, encima cabanas del Country Club, 17[degrees]30'S, 65[degrees]17'W, 2250 m, 15 Dec 1993, Ibisch 93-1894 (LPB), 3 km de Siberia hacia Karahuasi, 17[degrees]48'S, 64[degrees]41'W, 2400 m, 18 Oct 1996, Kessler et al. 9141 (LPB, UC); Carretera Cochabamba-Villa Tunari, debajo de Corani, 17[degrees]10.51'S, 65[degrees]54.02'W, 2750 m, 26 Nov 2002, Lehnert 519 (GOET, LPB, UC). Santa Cruz: Prov. Caballero, 7.5 km (by air) N of Comarapa, vicinity of Tinque Laguna, 17[degrees]51'S, 64[degrees]32'W, 2625 m, 25 Nov 1999, Nee 50639 (LPB, NY, UC).

Melpomene occidentalis Lehnert, sp. nov. TYPE.--ECUADOR. Zamora-Chinchipe: New road Loja-Zamora, ca. 4 km E of pass "El Tiro," ridge from white cross on left road side (towards the valley), 03[degrees]59'S, 79[degrees]08'W, 2550 m, 25 Sep 2004, Lehnert 1343 (holotype: QCA; isotypes: GOET, UC). Figs. 6, 7E.

A Melpomene firma petiolis teretibus vel parce marginatis, glabris vel glabrescentibus (vs. valde marginatis adaxialiter pilosis), segmentis numquam deflexis (vs. basalibus deflexis), squamis latioribusque (0.8-1.6 vs. 0.4-0.6 mm) differt.

Plants predominantly epiphytic, rarely epilithic or terrestrial, growing in moss layers. Rhizomes moderately long to short, ascending to erect (Figs. 6A, B), (0.8-)1.2-1.8(-2.2) mm diam. Fronds to 22-28 cm long, erect (Figs. 6A, B), straight or weakly arching, inserted onto the rhizomes at acute angles, closely arranged (internodes 1-3(-5) mm), usually caespitose (Fig. 6B). Scales 6.0-9.0 x 0.8-1.6 mm, (18-)20-26 cells wide, clathrate, dark brown to brown, iridescent, bases cordate, tips long-acute to attenuate, margins with small hyaline retrorse papillae 1-3 cells long (Fig. 6C), apical cells 3-8, in a linear arrangement or nodding cluster (similar to Figs. 5C, H). Petioles (25-)30-95 (-140) mm long, 0.8-1.0 mm thick, terete (Fig. 6C) to weakly marginate throughout, without acicular or setiform/ciliform hairs, simple and branched clavate hairs of crosiers and young fronds sometimes persistent in older fronds. Laminae (110-)145-220(-330) x (22-)30-45(-80) mm, lanceate or broadly elliptic (broadest in the middle), acute to attenuate at tip; bases rounded to cuneate (Figs. 6A, B). Rachises dark brown to black, planar to weakly protruding adaxially, hemispherically protruding abaxially, with few to many short, brown, acicular hairs to 0.2-0.8 mm long on both sides, usually adaxially hairier than abaxially (Figs. 6D, E). Segments (8-)16-20(-42) x (1.5-) 1.8-2.2(-4.4) mm (5-8 times longer than wide) (Fig. 6D), patent or weakly ascending (90-75[degrees]) (Figs. 6A, B), fully adnate, inequilateral at bases, weakly surcurrent towards the blade apices (Figs. 6D, E), linear to long-deltate, tips obtuse; margins entire (Fig. 6E) or dentate proximally in large segments; midveins visible on both sides of the segments (Figs. 6E, F); proximal segments pairs smaller than the central segments, but not auriculiform, not remote, or if remote then connected by a thin strand of laminar tissue; stomata sometimes visible as dark brown to reddish dots; margins sometimes with ephemeral clavate hairs or cells; hydathodes conspicuous (Figs. 6E, F). Sori 5-12(-15) pairs per segment, with several stiff, castaneous acicular hairs 0.4-0.8 mm long mostly around them (Fig. 6E).

[FIGURE 6 OMITTED]

[FIGURE 7 OMITTED]

The name refers to the western centered distribution of the species in South America compared to the similar Melpomene firma, which extends E to the Guyana Highlands and central Bolivia.

Melpomene occidentalis grows in montane forests at 2100-3400 m in Venezuela, Colombia and Ecuador (Fig. 7E).

Molecular studies show that this species is sister to Melpomene firma (Lehnert et al., in press). The main differences of M. occidentalis to M. firma are the predominantly terete, completely glabrous petioles (except for clavate hairs vs. petioles always marginate, persistently hairy adaxially, or rarely glabrescent, with dark, acicular hairs in M. firma) and the patent to ascending segments (vs. patent to deflexed). Colombian plants may have sinuses wider than the segment width (Hagemann 1306, COL), but contrary to the remote segments of M. firma they are still connected by thin strands of laminar tissue. Plants from northern Ecuador have more deltate segments and may be confused with species of the M. pilosissima or M. personata complexes. Those species differ in lacking marginal cells on the scales (vs. scales with marginal cells in M. occidentalis). The M. pilosissima alliance (M. pilosissima, M. huancabambensis, M. jimenezii, M. michaelis) has more abundant, longer hairs (to 3 mm), especially on the petioles, and has hairs often occurring on the segments margins or the adaxial laminar surfaces. Melpomene personata and allied species (M. albicans, M. youngii) differ in their generally prostrate rhizomes and the adaxially not visible midveins from both M. occidentalis and M. firma.

ADDITIONAL SPECIMENS EXAMINED.--VENEZUELA. Tachirs: Paramo Tarma, near the Colombian-Venezuelan border, 2475-2550 m, 20-23 May 1967, Steyermark et al. 98600 (NY). Prov. unknown: [Merida?] Manzanos, 2800 m, Lindig 300 (B, P).

COLOMBIA. Antioquia: San Jose de la Montana, Alto El Cristo, 06[degrees]46'53"N, 75[degrees]41'45"W, 3420 m, 23 Jul 2002, Rodriguez W. et al. 3513 (COL). Huila: Cordillera Central, Cordillera del Buey, hike from Finca Loyola over the paramo down to San Antonio, 2100 m, 14 Dec 1975, Bishop 1993 (UC). Narino: La Botana (Pasto region), 2900 m, 29 Oct 1972, Hagemann & Leist 1306 (COL).

ECUADOR. Cotopaxi: Quevedo-Latacunga road, above Pilalo, 00[degrees]58'S, 78[degrees]58'W, 2850 m, 08 Apr 1973, Holm-Nielsen et al. 3251 (UC); along Quevedo-Latacunga road, between Pilalo and Pujili, 00[degrees]59'S, 78[degrees]58'W, 3400 m, 26 Nov 2004, Lehnert 1575 (GOET, QCA, UC). Loja: E of Nudo de Cajanuma, just N of "Centro de Informacion," sample plot site, 04[degrees]05'S, 79[degrees]10'W, 2900 m, 20 Sep 1989, Bosh 47959 (AAU); Parque Nacional Podocarpus, above Nudo de Cajanuma around "Centro de Informacion," 04[degrees]05'S, 79[degrees]10'W, 2800-3000 m, 16 Nov 1989, Bogh 86609 (AAU); Cerro Toledo, E of Yangana, between Loja and Valladolid, 04[degrees]23'S, 79[degrees]07'W, 3000-3100 m, 26 Oct 2004, Lehnert 1464a (GOET, QCA, UC); Cajanuma, SE of Loja, 04[degrees]07'S, 79[degrees]10'W, 2750 m, 03 Nov 2004, Lehnert 1507 (GOET, QCA, UC). Loja/Zamora-Chinchipe: limit of Parque Nacional Podocarpus, around pass on road Loja-Zamora, 03[degrees]58'S, 79[degrees]07'W, 2900 m, 08 Jan 1989, Madsen 85474 (AAU). Morona-Santiago: E of pass on Gualaceo-Limon road, 03[degrees]00.27'S, 78[degrees]39.10'W, 3000-3200 m, 16 Nov 2004, Lehnert 1558a (GOET, QCA, UC). Napo: Cartagena, km 25 from El Carmelo on road towards La Bonita, 00[degrees]37'N, 77[degrees]30'W, 2800 m, 13 Apr 1979, Lojtnont et al. 12334 (AAU); outskirts of Pifo, 2500 m, 08 May 1935, Mexia 7353a (UC). Pichincha: carretera Quito-Santo Domingo, 2500 m, 24 May 1987, van der Werff & Palacios 9600 (MO, UC). Zamora-Chinchipe: Estacion Cientifica San Francisco, above refuge, along trail "Antennenbergweg," just below junction with trail T1, study plot A 5, 03[degrees]59'36.4"S, 79[degrees]04'03.2"W, 2660 m, 26 Sep 2003, Lehnert 909a (GOET, QCA, UC); new road Loja-Zamora, ca. 4 km E of pass "El Tiro," 03[degrees]59'S, 79[degrees]08'W, 2650 m, 23 Sep 2004, Lehnert 1332 (GOET, QCA, UC).

[FIGURE 8 OMITTED]

Melpomene paradoxa Lehnert, sp. nov. TYPE.--BOLIVIA. La Paz: Prov. Nor Yungas, trocha al Valle Coscapa, Parque Nacional Cotapata, 16[degrees]12'S, 67[degrees]53'W, 3450 m, 09 Sep 1997, Kessler et al. 11717 (holotype: LPB; isotypes: GOET, UC). Figs. 8, 12A.

A Melpomene moniliformi squamis maioribus (squamis usque 2.0-2.5 mm longis cum 14-18(-20) cellulas supra basin vs. usque 3.5-4.0(-8.0) mm longis cum (20-)22-28(-32) cellulas), a M. flabelliformi absentia pilorum inter soros (vs. pilis inter soros dispersis) segmentisque brevioribus (rotundis, usque 4.5 mm longis vs. oblongis, usque 11.5 mm longis) differt.

Plants large, epiphytic in moss cushions. Rhizomes horizontal, long-creeping (Fig. 8E), (0.6-)0.8-1.2 mm diam. Fronds to 27-32 cm long, diffusely arranged (internodes 5-10 mm), not caespitose, mostly arching (Fig. 8E). Scales lanceate to ovate-lanceate, dark brown to brown, iridescent, 3.5-4.0 (-8.0) x 1.0-1.6 mm. (14-)22-28(-32) cells wide across base (Fig. 8C), broadly cordate to pseudopeltate at bases, acute to attenuate at tips; apical cells 3-8, bifurcately to palmately arranged, ultimate cells often elongate. Petioles 20-60 mm long, 0.5-0.8 mm diam., weakly marginate to rarely alate, sparsely hairy with hairs to 1.2 mm long. Laminae narrow, linear, glabrous except for the sori. Segments to 4.5 x 4.0 mm (only as long as broad), lunate to broadly deltate, patent to weakly ascending (90-80[degrees]), tips obtuse to round, midveins not visible or obscurely so in dried specimens (Fig. 8F); proximal segment pairs often gradually reduced to auricles. Sori single or 2(-3) pairs per segment, with 5-10 brown to castaneous, setiform or ciliform hairs 0.8-1.2 mm long (Fig. 8F).

The species is named for its puzzling morphology, which is intermediate between Melpomene flabelliformis and M. moniliformis.

Melpomene paradoxa is known from elfin forests and wet montane forests at 2800-3700 m in Peru and Bolivia (Fig. 12A).

This species matches M. flabelliformis in the rhizome and scale size, but is closer to M. moniliformis regarding the scale color, laminar shape, and hair distribution. The scales of M. paradoxa are not as strongly iridescent as in M. flabelliformis, and despite having often the same amount of hairs in the sori as that species, M. paradoxa lacks the hairs between the sori that are typical of M. flabelliformis.

Melpomene flagellata differs from M. paradoxa in having longer setiform/ ciliform hairs (1.2-1.8 mm) in a denser pubescence on the petioles; it also has most of the segment tips truncate (vs. always obtuse or rounded in M. paradoxa).

Melpomene caput-gorgonis has shorter petioles than M. paradoxa, often relatively wider, elliptic laminae (vs. laminae linear in M. paradoxa), and conspicuously multiple capitate hairs at the scale tips (vs. cells as a single branched clavate hair).

ADDITIONAL SPECIMENS EXAMINED.--PERU. Ayacucho: Condorcunca, 12 Oct 1964, Barrdn s.n. (USM). Cuzco: Prov. Urubamba, between San Luis and Abra Malaga, Km 154, 13[degrees]05.4'S, 72[degrees]22.2'W, 3300 m, 16 Oct 2002, Lehnert 439 (GOET, UC, USM); Machu Picchu, 4 km from the Runucuray-Sayacmarca pass in the Inca trail, above the lake between Sayacmarca and the pass (vegetation plot 166), 3665 m, 23 Aug 1982, Peyton & Tilney Peyton 1570 (UC).

BOLIVIA. Cochabamba: Prov. Carrasco, Colomi, along road, 62.2 km from Cochabamba (Rio Rocha), 17[degrees]12'84"S, 65[degrees]53'21"W, 3100 m, 30 Dec 1998, de Boer 1156 (LPB); 108 km antigua carretera Cochabamba-Villa Tunari, 17[degrees]09'S, 65"38'W, 2950 m, 22 Jun 1996, Kessler 6569 (LPB, UC); 94 km on old road Cochabamba-Villa Tunari, 17[degrees]12'S, 65[degrees]41'W, 3500 m, 28 Jun 1996, Kessler et al. 6773 (GOET, LPB); road Cochabamba-Villa Tunari, below Corani, near Km 71+00, trail up the mountain, 17[degrees]10.59'S, 65[degrees]53.67'W, 2800 m, 27 Nov 2002, Lehnert 525 (LPB). La Paz: Prov. Bautista Saavedra, Charazani, E of Chullina, 3500 m, 22 Dec 1993, Herzog H87 (LPB); Prov. Nor Yungas, Unduavi, 3300 m, Nov 1910, Buehtien 70 (P); roadside bank between Cotapata and Chuspipata, S-facing, La Paz-Caranavi road, 16[degrees]17'S, 67[degrees]50'W, 3200 m, 15 Aug 1990, Fay & Fay 3034 (LPB, MO); trench to the Valle Coscapa, Parque Nacional Cotapata, 16[degrees]12'S, 67[degrees]53'W, 3450 m, 09 Sep 1997, Kessler et al. 11717 (LPB, UC); Unduavi, trench to the Valle de Coscapa, 3400 m, 17 Dec 2002, Lehnert 536, 539, 542 (GOET, LPB, UC); 1 km W of Chuspipata, 16[degrees]17'S, 67[degrees]49'W, 3140 m, 24 Mar 1982, Solomon 7260 (LPB, UC).

Melpomene personata Lehnert, sp. nov. TYPE.--BOLIVIA. La Paz: Prov. Bautista Saavedra, 15 km de Charazani hacia Chullina, 15[degrees]10'S, 68[degrees]53'W, 3400 m, 05 Jul 1996, Kessler 10606 (holotype: UC; isotypes: GOET, LPB). Figs. 9, 12B.

A Melpomene pilosissima rhizomatibus longioribus horizontaliter reptantibus pilisque in soris confertis (vs. rhizomatibus brevibus erectis, pilis aequaliter distributis in pagina abaxiali), a M. youngii et M. albicante absentia strati albi in pagina abaxiali (vs. pagina abaxiali alba vel albicante) differt.

Plants predominantly epiphytic, rarely epilithic, growing in moss layers. Rhizomes horizontal, moderately to long-creeping, 0.8-1.2(-1.5) mm diam. (Fig. 9A). Fronds 14-33 cm long, stiffly pendent, inserted onto the rhizome at right angles, diffusely arranged (internodes 3-10 mm), not caespitose (Fig. 9A). Scales (2.0-)3.0-4.5(-6.0) x (0.3-)0.4-1.0 mm, (8-)10-14(-22) cells wide across base, clathrate (Fig. 9B), dark brown to brown, iridescent, narrowly lanceate, bases cordate, tips long-acute to attenuate; apical cells 1-3, linearly arranged (Fig. 9B) or 2-5 cells palmately arranged. Petioles 20-150 mm long, 0.6(-0.8) mm thick, terete or weakly marginate from the lowest segments, with few to many dark brown setiform/ciliform hairs 0.9-2.0 mm long (Fig. 9C); simple and branched clavate hairs of crosiers and young fronds sometimes persistent on older fronds. Laminae to 155-180(-220) x 26(-40) mm (2/3 to 3/4 of frond length, rarely less in single fronds of a plant), broadly linear to narrowly elliptic (broadest in the middle), bases cuneate to short-tapering, apices long-acute (Fig. 9A). Rachises dark brown to black, planar and slightly sunken between the segments adaxially (Fig. 9E), hemispherically protruding abaxially (Figs. 25D, F), sparsely hairy on both sides (Fig. 9E), usually more hairs abaxially, setiform/ciliform hairs to 1.2 mm long, brown. Segments 8.0-15.0(-22.0) x 1.2-2.5(-3.5) mm (6-8 times long than broad), ascending (70-60[degrees]), equilateral at base or weakly decurrent towards the bases, fully adnate, linear-oblong, the tips acute to obtuse (Fig. 9A); midveins visible abaxially (e.g. Jimenez I. 1773; Fig. 9D), at least discontinuously so (Fig. 9F); proximal 2-6 segment pairs markedly smaller than the subsequent segments (Fig. 9A), lowest ones usually auriculiform; setiform/ciliform hairs few (Fig. 9F) to many on the midveins, absent on margins and adaxial laminar surfaces; hydathodes present (Fig. 9E). Sori 2-10 pairs per segment, with 6-10 setiform or ciliform hairs 0.8-1.0 mm long (Fig. 9F).

The name alludes to the fact that the species has often been mistaken for M. pilosissima (Latin, personata = masked, disguised), and also alludes to the name of the genus: Greek actors wore masks (Latin, persona) to symbolize the different characters of the plays, including the tragedies that the muse Melpomene is representing.

Melpomene personata grows in shrubby paramos, elfin forests, and wet montane forests at (1850-)2700-4500 m in Mexico, Guatemala, Costa Rica, Panama, Dominican Republic, Venezuela, Colombia, Ecuador, Peru, and Bolivia (Fig. 12B) and is a common species in the Andes.

[FIGURE 9 OMITTED]

Many Central American plants differ from the Andean ones slightly in having more but smaller, isodiametric cells in the scales, more apical cells, and mostly weakly visible midveins. Apart from that, the characters are identical. Many specimens of M. personata have been erroneously determined as M. pilosissima, which has added greatly to the confusion within that species complex. Molecular data, however, indicate that M. personata is not closely related to M. pilosissima and allies, but in fact belongs to the M. moniliformis clade (Lehnert et al., in press). Morphologically, M. personata is characterized by horizontally creeping rhizomes (vs. usually erect in M. pilosissima); rhizome length can vary greatly, but usually is longer than in M. pilosissima and allies, with the fronds also placed further apart (3-10 mm vs. 0.5-2.0 mm in M. pilosissima). From M. pilosissima and M. huancabambensis, M. personata differs in having the hairs abaxially clustered in the sori and along the midveins (vs. hairs evenly distributed); from M. xiphopteroides, it differs in the terete petioles (vs. petioles marginate to alate); from M. huancabambensis, M. jimemezii, and M. michaelis in having shorter hairs (to 2 mm in M. personata vs. to 3 mm) and conspicuous hydathodes (vs. hydathodes reduced or lacking); from M. vernicosa and M. jimenezii in the thinner (0.4-0.8 mm vs. 0.8-1.5 mm), glabrous or glabrescent petioles (vs. petioles persistently hairy).

Other species for which Melpomene personata may be mistaken include M. sodiroi, M. albicans, and M. youngii. All differ from M. personata in having hidden midveins on the abaxial laminae (vs. midveins at least obscurely visible in M. personata). Melpomene sodiroi also has larger scales and usually gibbose segments; Melpomene albicans and M. youngii have a white wax-like deposit on the abaxial laminae, which is not present in M. personata; furthermore, M. youngii is completely glabrous and generally lacks hairs in the sori (sometimes 1-2 hairs in sori vs. hairy petioles, hairs clustered in sori in M. personata).

The Peruvian collection Philippi s.n. (B) belongs to Melpomene personata and was annotated as paralectotype of Polypodium firmum Klotzsch (= Melpomene firma). These two species can be easily distinguished, as M. firma (including the lectotype Schomburgk 1170, B) has an erect rhizome, patent segments, truncate laminar bases and small cells along the scale margins, while M. personata (including Philippi s.n., B) has horizontally creeping rhizomes, ascending segments, rounded to cuneate laminar bases, and lacks marginal cells on the scales.

ADDITIONAL SPECIMENS EXAMINED.--MEXICO. Chiapas: San Cristoba! Las Casas, E side of Zontehuitz near summit, 2800 m, 30 Nov 1971, Breedlove 22928 (NY); Union Juarez, SE side of the summit of Volcan Tacana, 3600 m, 03 Mar 1972, Breedlove 24310 (NY); ibid., 10 Nov 1972, Breedlove 29353 (NY); Mount Tacana, 2000-4038 m, Aug 38, Matuda 2379A (NY); N of San Cristobal las Casas on top of Cerro Zontehuitz, 3100 m, 27 Jul 1963, Mickel 1247B (NY).

GUATEMALA. Chimaltenango: Cerro de Tecpam, region of Santa Elena, 2400-2700 m, 26 Dec 1938, Standley 61075 (NY). Guatemala: Hyw. No. 1, 2900 m, 21 Feb 1947, Brenckle 47-61 (UC). Huehuetenago: Limestone region 3-15 km N of Chemal, Sierra de los Cuchumatanes, 3400 m, 02 Dec 1962, Molina-R. et al. 22138 (NY); Sierra Cuchumatanes between Km 136 to 150 to San Juan Ixcoy, 3000-3500 m, 12-23 Jan 1966, Molina-R. et al. 16541 (NY). Totonicapan: Maria Tecun, 3000-3600 m, 12-23 Jan 1966, Molina-R. et al. 16357 (NY).

COSTA RICA. Cartago: Cerro de la Muerte, 1 km NW of Villa Mills on Interamerican Highway, behind Hotel La Georgina, 2900 m, 08 Aug 1967, Mickel 3203 (NY); Km 89 Rt. 2 to paramo de la Muerte and San Isidro, 09[degrees]35'N, 83[degrees]42'W, 3300 m, 30 Oct 1993, Rivero 2488, 2494, 2495, 2497 (UC); Canton de Paraiso, R.F. Rio Macho, cuenca del Reventazon, carreterea interamericana, road Cartago-San Isidro, cerro de la Muerte, 3150-3300 m, 09[degrees]34'30"N, 83[degrees]45'W, 21 Apr 1999, Rojas A. 5082 (NY). San Jose: Cerro de la Muerte; 1 km NW of Villa Mills on the Interamerican Highway, cross from Hotel La Georgina, 2900 m, 08 Aug 1967, Bishop 869 (UC); along Inter-American Highway, Cerro de la Muerte, 09[degrees]35'N, 83[degrees]45'W, 3220 m, 18 Nov 1986, Hennipman et al. 7010 (UC); Cerro Sakira-paramo, 3300 m, Feb 2003, Kluge 1182 (GOET); Massiv of Cerro de La Muerte, 3200 m, 15 Jul 2003, Kluge 6841 (GOET); ibid., 3300 m, 16 Jul 2003, Kluge 6909, 6910 (GOET); ibid., 3400 m, 17 Jul 2003, Kluge 6946 (GOET); road from Cartago to San Isidro del General (Pan American Highway, Rt. 2), Km 96-97, ca. 1.5 km S of Villa Mills (near Siberia), 2900 m, 29 Jan 1986, Smith A. R. & Beliz 2061 (UC).

PANAMA. Chiriqui: between Itamut and Bine Peaks, Fabrega massif, Bocas del Toro, 3200 m, 05-09 Mar 1984, Gomez L. D. et al. 22539 (UC); Volcan Baru, on road to towers at top, near towers at summit, 08[degrees]47'N, 82[degrees]32'W, 3300-3400 m, 13 May 1990, McPherson 15054 (UC).

DOMINICAN REPUBLIC. Peravia: 48 km S of Constanza (on road to San Jose de Ocoa), in area of La Nevera, 18[degrees]41'N, 70[degrees]35.5'W, 2070 m, 4 Mar 1981, Zanoui & Mejia 12209A (NY).

COLOMBIA. Antioquia: Guatape, Vereda Santa Rita, finca Montepinae, 1850 in, 06[degrees]15'N, 75[degrees]10'W, 06 Mar 1990, Contreras & Echeverri 217 (NY). Boyaca: Sierra Nevada de Cocuy, on steep S side of the Valle del Corallitos, at lower edge of this near river, 4000 m, 06 Sep 1957, Grubb & Guymer P102 (AAU); between Arcabuco and Villa de Leyva, trail Las Coloradas, above El Charizal, 3420 m, 31 Aug 1967, Jaramillo Mejia et al. 3166 (AAU). Cauca: Cordillera Central, Parque Nacional de Purace, trail from Pilimbala to the volcano of Purace, 3700 m, 10 Jul 1976, Jaramillo Mejia & van der Hammen 5219 (AAU). Cundinamarca: Laguna de Chisaca and surroundings, 3900-4200 m, 19 Oct 1958, Bishler 1517 (COL); Fomeque, Parque Nacional Natural Chingaza, surroundings of laguna de Chingaza, E shore of Rio Chuza, 2990 m, 05 Oct 1981, Franco P. et al. 495 (AAU); Cogua, Vereda Quebrada Honda, Reserva Forestal Protectora, 3200-3300 m, Aug-Oct 2003, Trujillo 13 (COL). Santander: Coromoro, laguna La Fiquera, quebrada de Coromoro, 3750 m, 29 Nov 1967, Jaramillo Mejia & van der Hammen 4380 (AAU); road between Bucaramanga and Pamplona, W slope of Paramo de Berlin, El Picacho, 3300 m, 11 Nov 1969, Murillo M. T. & Jaramillo Mejia 1342 (AAU). Prov. unknown: Excursion to the Llanos de San Martin, Toquisa area, Stubel 704 (B).

VENEZUELA. Merida: Dtto. Justo Briceno, Paramo Piedras Blancas, Laguna La Fea, SE de la Carretera Via Pinago, 3950-4500 m, 09 Jul 1982, Briceno et al. 635 (AAU); Dtto. Libertador, Parque Nacional Simon Bolivar, camino del teleferico (de la Aguada), Laguna La Fria, vertiente septentrional de la Sierra Nevada de Merida, 2700-3000 m, 21 Dec 1984, Pipoly et al. 6542 (UC); Dtto. Rangel, Las Escaleras-Laguna El Boqueron, paramo de Minugu, unos 10 kmal SE de San Rafael de Mucuchies, 3150-3400 m, 21 May 1972, Ruiz-Teran 7314 (UC). Trujillo: Dtto. Carache, via Paramo Cende sitio denominado "Las Cruces," 09[degrees]33'N, 70[degrees]08'W, 27 Nov 1987, Rivero & Diaz W. 1495 (UC).

ECUADOR. Azuay: Recreation Park Cajas, 4000-4100 m, 02 Sep 1984, Jaramillo J. 7198 (AAU); Cajas, N of Laguna Toreadora, 02[degrees]47.17'S, 79[degrees]13.10'W, 3850 m, 17 Nov 2003, Lehnert 1116 (GOET, QCA, UC). Carchi: Montufar, within 3 km of pueblo of Colonia Huaquena, 00[degrees]35.5'N, 77[degrees]42'W, 3500 m, 30 Jun 1994, Fay & Fay 4342 (AAU); base of Volcan Chiles, km 34-36 on road Tulcan-Maldonado, 3900-4050 m, 00[degrees]47'N, 77[degrees]57'W, 19 May 1973, Holm-Nielsen et al. 5922 (AAU, UC); El Angel-Tulcan main road, Km 1, turnoff towards W, ca. 8 km, 00[degrees]34'N, 77[degrees]54'W, 3460 m, 08 Aug 1990, Jorgensen et al. 92264 (AAU); road Tulcan-Maldondado, Km 32 de Tulcan, base del Volcan Chiles, Lehnert 145, 146 (GOET, QCA, UC). Chimborazo/Canar: W escarpment between Santa Rosa and Joyagshi, 2500-2700 m, 06-09 Jul 1945, Camp 4076 (NY). Imhahura: Laguna Grande de Mojanda, 15 km S of Otavalo, 3750 m, 00[degrees]08'N, 78[degrees]16'W, 14 May 1985, Eriksen 59365 (AAU); road Ibarra-Mariano Acosta, E of the pass, 00[degrees]20'N, 78[degrees]00'W, 3500-3600 m, 09 Aug 1976, Ollgaard & Balslev 8585, 8579 (AAU).

PERU. Cuzco: Prov. Paucartambo, Pillahuata, near Tres Cruces, + 130 km from Cuzco to Pilcopata, 13[degrees]05'S, 71[degrees]30'W, 2000 m, 13 Dec 1986, Nunez 7798 (CUZ, LPB); Achirani, Marcachea, 3000 m, 30 Jul 1939, Vargas C. 1573 (CUZ); Dtto. Marcachea; near Achirani, 2600 m, 30 Jul 1939, Vargas C. 11141 (CUZ, UC); Prov. Urubamba, Abra Malaga, 4330 m, 16 Oct 2002, Lehnert 423 (GOET, UC, USM); Machupicchu, at 88 kin and 112 km from Cuzco, Santuario Historico de Machu Picchu and along Inca trail, in Qorihuayrachina, llulluchayoc, Ronkuraky, Phuqupatamarca, Winayhuayna, and Intipunco, 13[degrees]09'10"S, 72[degrees]31'W, 4150 m, 14-22 Oct 1987, Nunez & Arque 8339 (UC); Altura Colca, Valle de Lares, 9000 ft, Mar 1932, BOles 1807 (CUZ). Junin: Prov. Satipo/La Convencion, Cordillera Vilcabamba, Rio Ene slope, near summit of divide, 3350-3400 m, 11[degrees]39'36"S, 73[degrees]40'02"W, Boyle et al. 4326 (UC, USM); Prov. Tarma, high region of second Cordillera, valley of Maranioch near Tarma, 1840, Philippi s.n. (B, isosyntype of Polypodium firmum Klotzsch). San Martin: Mariscal Caceres, Chochos, NW corner of Rio Abiseo National Park, 3500 m, 15 Jul 1987, Young & Leon 4716 (USM).

BOLIVIA. Cochabamba: Prov. Ayopaya, 2 km al SE de Saila Pata, 16[degrees]55'S, 66[degrees]55'W, 3550 m, 15 Nov 1997, Kessleret al. 12475 (GOET, LPB, UC); Prov. Carrasco, 100 km [on] old road Cochabamba-Villa Tunari, 17[degrees]12'S, 65[degrees]42'W, 3250 m, 26 Jun 1996, Kessler 6728 (GOET, LPB, UC); ibid., 115 km, 17[degrees]07'S, 65[degrees]38'W, 2700 m, 01 Jul 1996, Kessler et al. 6862 (GOET, LPB, UC); ibid., 63 km, 17[degrees]15'S, 65[degrees]43'W, 3750 m, 02 Jun 1996, Kessler et al. 6887 (GOET, LPB); ibid., 68 kin, 17[degrees]14'S, 65[degrees]13'W, 3600 m, 11 Jul 1996, Kessler et al. 7234 (GOET, LPB, UC); Prov. Chapare, ca. 8 km N [of] Maycamayu, ca. 70 km from Sacaba, 17[degrees]12S, 65[degrees]57'W, 3350 m, 12 Aug 1991, Kessler 2919 (AAU, LPB). La Paz: Prov. Franz Tamayo, PN-ANMI Madidi, trail Pelechuco-Mojos, locality Tambo Quemado (camping site), going down along the trail, a little past the second river, 14[degrees]41'S, 68[degrees]58'W, 3470 m, 29 Apr 2003, Jimenez I. 1773 (GOET, LPB, UC); Prov. Inquisivi, some 8 km from Quime towards Inquisivi, Camillaya arriba del pueblo, 16[degrees]58'S, 67[degrees]12'W, 3000 m, 29 Dec 1997, Beck 24364 (LPB); Prov. Murillo, arriba de la laguna de Viscachani al valle de Zongo, 16[degrees]13'S, 68[degrees]07'W, 4050 m, 10 Oct 1995, Kessler et al. 5885 (AAU, LPB); Prov. Nor Yungas, Estacion Biologica Tunquini, senda del Pantanon a Hornuni, 16[degrees]11'S, 67[degrees]53'W, 3350 m, 19 Sep 2000, Bach et al. 1080 (GOET, LPB, UC); pasando Unduavi antes de llegar a Cotapata, subiendo la senda antigua hacia Coroico, 3500 m, 22 Oct 1994, Beck & Ruthsatz 21492 (GOET, LPB, UC); Valle de Coscapa, 16[degrees]17'S, 67[degrees]51'W, 3400 m, 02 Oct 2002, Lehnert 396, 398, 404, 405 (GOET, LPB, UC); ibid., 17 Dec 2002, Lehnert 535, 537, 538, 541 (GOET, LPB, UC); Prov. Sud Yungas, Unduavi, near the mine Lourdes, 16[degrees]18'S, 67[degrees]52'W, 3450 m, 25 Nov 1995, Gonzales et al. 1557 (LPB, UC).

Melpomene sklenarii Lehnert, sp. nov. TYPE.--ECUADOR. Azuay: Cajas National Park, E flanks of Cerro Amarillo (4451 m), 02[degrees]45'S, 79[degrees]15'W, 13 Jul 1997, 4300-4400 m, Sklenar & Sklenarova 2592 (holotype: UC, isotype: PRC). Figs. 10, 12C.

A Melpomene peruviana squamis ovato-lanceolatis (vs. angusto-lanceolatis) frondibusque erectis distantibus (vs. frondibus pendentibus approximatis), a M. moniliformi et M. flabelliformi pilis aequaliter distributis in paginis abaxialibus (vs. pilis absentibus vel in soris restrictis) segmentisque basalibus interdum remotis (vs. segmentis semper confertis) differt.

Plants terrestrial or epilithic, growing in moss layers, rock crevices, or open soil. Rhizomes moderately to long-creeping, 0.6-1.0 mm diam. (Figs. 10A, B), regularly branching at wide to right angles (Fig. 10B). Fronds to 13 cm long, erect, inserted onto the rhizomes at right angles, diffusely arranged (internodes (5-)11-14 mm), not caespitose (Fig. 10A, B). Scales 2.8-4.0 x 0.6 mm, (10-) 14-18(-26) cells wide across their bases, clathrate (Fig. 10 C), dark brown to brown, weakly to rarely strongly iridescent, broadly cordate to pseudopeltate, acute to attenuate at tips; apical cells 3-8, palmately arranged; scales soon shed from rhizomes, especially between the fronds. Petioles (6-)12-55 mm long, 0.4-0.8(-1.0) mm thick, alate from the laminar bases, marginate towards the rhizomes, glabrescent or hairy, with dark brown, ciliform/setiform hairs 0.75-1.5(-2.0) mm long (Fig. 10D); simple and branched clavate hairs rarely present; petiolar bases with persistent scales. Laminae 40-80(-90) mm long, to 7-12 mm wide, firm-chartaceous to subcoriaceous, linear to narrowly lanceate (widest below the middle), round to weakly cuneate at bases, acute at tips (Figs. 10A, B). Rachises dark brown to black, planar and slightly sunken between the segments adaxially (Fig. 10E), weakly protruding to planar abaxially, with scattered brown acicular or setiform/ciliform hairs to 1.2 mm long abaxially (Fig. 10F). Segments to 4.2 x (2.0-)2.4-3.0 mm (ca. 1.5-2.5 times longer than broad), patent, inequilateral and decurrent towards the bases, fully adnate, deltate to oblong, the tips round to obtuse (Figs. 10E, F), slightly gibbose, abaxially pale green to whitish, but without wax-like deposit; midveins not visible, or obscurely so in dried specimens; basal pairs often remote, not notably smaller than the following segments, or only one pair weakly so (Fig. 10B); abaxially glabrous to densely hairy with evenly distributed, brown ciliform hairs 0.8-1.5 mm long (Fig. 10F); segment margins hyaline, one cell wide (2 cells wide in sinuses and at junctions with petioles), without clavate hairs (Fig. 10E, F); hydathodes present. Sori 1-3(-4) pairs per segment, surrounded by 5-8 dark brown setiform/ciliform hairs to 1.0 mm long (Fig. 10F).

[FIGURE 10 OMITTED]

The name honours Petr Sklenar from Charles University, Prague, who collected most of the known specimens during his studies of Ecuadorian paramos.

Melpomene sklenarii grows in paramos, punas (jalca), and elfin forests at 2900-4700 m in Colombia and Ecuador (Fig. 12C).

This species is easily confused with M. peruviana but can be distinguished by the evenly distributed hairs on the abaxial laminar surfaces (vs. clustered in sori in M. peruviana), more widely spaced fronds (internodes (5-)11-14 vs. 0.8-3.0 mm), and paler, more broadly lanceate rhizome scales (vs. dark brown to blackish and narrowly lanceate) which are often shed between the fronds (vs. usually persistent). Melpomene sklenarii grows in loose formations with the distant fronds held erect whereas M. peruviana tends to form dense mats and holds the fronds stiffly tip-downwards (or appressed to horizontal substrates).

The rather small range of Melpomene sklenarii matches that of M. vulcanica and both species apparently often grow closely together. Large specimens of M. vulcanica are easily separated (segments long-deltate with acute tips, midveins visible on both sides in M. vulcanica vs. segments oblong to deltate with round to obtuse tips, midveins not visible adaxially in M. sklenarii), but smaller plants may have these distinguishing characters more weakly developed and may be confused with M. sklenarii. Even if equal in size, Melpomene vulcanica has rhizome scales that are still larger (6.2 x 1.0 mm, 20-30 cells wide across bases vs. to 3.5 x 0.6 mm, 14-18(-26) cells wide across bases) and longer persisting than those of M. sklenarii. Both species have hyaline segment margins, but those of M. vulcanica are two cell rows wide and beset with ceils or clavate hairs whereas those of M. sklenarii are just one cell row wide in most parts and lack separate cells.

Small plants of M. flabelliformis can be distinguished from M. sklenarii by their setiform/ciliform hairs clustered in the sori and the proximally more strongly cuneate laminae with approximate segments (vs. hairs not clustered in sori and laminae proximally rounded to cuneate with often distant segments in M. sklenarii). The two species have not been found growing together so far, and especially the small forms of M. flabelliformis occur mainly outside the range of M. sklenarii (i.e., Mexico and Africa).

ADDITIONAL SPECIMENS EXAMINED.--COLOMBIA. Narino: Prov. Pasto, Volcan Galeras, [ca. 01[degrees]12'N, ca. 77[degrees]28'W,] 3950 m, 06 Aug 1977, Pinto et al. 1828 (COL). Boyaca: Cordillera Oriental, Sierra Nevada del Cocuy, surroundings of Salto de Correlitos, Sta. 13 above and E of Salto, E of Laguna San Paulito, ca. 05[degrees]34'N, ca. 72[degrees]37'W, 4200 m, 14 Apr 1959, Barclay & Juajibioy 7370 (COL); Sierra Nevada del Cocuy, paramo Cocavo, Cuchilla Puentepiedra ca. 2 km to the NE of Laguna Pintada, 4510 m, 30 Sep 1972, Cleef 5668 (COL). Caldas: Paramo del Ruiz, [ca. 05[degrees]28'N, ca. 75[degrees]39'W,] 4200-4630 m, 29 Aug 1957, Barclay 5240 (COL); Nevado del Ruiz, [ca. 05[degrees]28'N, ca. 75[degrees]39'W,] 4700 m, 06 Aug 1958, Bishler 1482 (COL); ibid., sandy area 2 km SW of the refuge with many outcrops of volcanic rock, [ca. 05[degrees]28'N, ca. 75[degrees]39'W,] 4520 m, 18 Mar 1972, Cleef & van't Hart 2446 (COL); road from Manizales to Nevado, above the "termales," 3500 m, 07 Jun 1966, Murillo M. T. et al. 861, 874 (COL); road from Manizales to Nevado, 4230 m, 05 Jul 1959, Pinto 437 (COL); Cordillera Central, surroundings of the refuge of Ruiz, road to El Silencio, [ca. 05[degrees]28'N, ca. 75[degrees]39'W,] 4310 m, 07 Oct 1978, Rangel et al. 1735-A (COL).

ECUADOR. Carchi: road Tulcan-Maldondado, Km 32 from Tulcan, base of Volcan Chiles, 06 Jul 2002, Lehnert 155a (GOET, QCA, UC); road Tulcan-Maldonado, Km 34 from Tulcan, 06 Jul 2002, Lehnert 156 (GOET, QCA, UC); S slopes of volcan Chiles, 02[degrees]49'N, 77[degrees]57'W, 4100 m, 21 Oct 1987, Ramsay & Merrow-Smith 872 (AAU). Chimborazo: El Altar, N side of the volcano, on the ridge below the Canoningo peak, 01[degrees]41'S, 78[degrees]24'W, 4200-4400 m, 19 Aug 1995, Sklenar & Kosteckovar 967 (UC); El Altar, N side of the volcano, on the ridge below the Canoningo peak, 01[degrees]41'S, 78[degrees]24'W, 4500 m, 19 Aug 1995, Sklenar & Kosteckovar 88_7 (UC); Chimborazo volcano, base of the terminal moraine on the E side of the mountain, 01[degrees]28'S, 78[degrees]46'W, 4500 m, 03 Jul 1997, Sklenar & Sklenarova 2198 (UC); Chimborazo volcano, on the E slope of the mountain, 01[degrees]28'S, 78[degrees]46'W, 4200-4250 m, 02 Jul 1997, Sklenar & Sklenarova 2308 (UC). Chimborazo/Morona-Santiago: Cerro Yanaurcu, N ridge of the mountain, 02[degrees]14'S, 78[degrees]30'W, 4200-4300 m, 29 Oct 1995, Sklenar & Kosteckovar 1499 (AAU); Cerros Yuibug-Pailacajas (4730 m), E side of the mountain ridge, 01[degrees]45'S, 78[degrees]27'W, 4300-4350 m, 31 Jul 1997, Sklenar & Sklenarova 3025 (UC). Imbabura: SW slopes of the volcano Cotacachi, 00[degrees]22'N, 78[degrees]21'W, 4100-4320 m, 09 Nov 1983, Boysen Larsen et al. (AAU); slopes of Volcan Cotocachi, 00[degrees]35'N, 78[degrees]20'W, 4150 m, 11 Oct 1987, Ramsay & Merrow-Smith 796 (AAU); Cerro Imbabura, in a gully on the S side of the volcano, 00[degrees]15'N, 78[degrees]10'W, 4400 m, 05 Jun 1995, Sklenar & Kosteckova 520 (AAU); Nevado Cotacachi, SE ridge of the volcano, 00[degrees]21'N, 78[degrees]21'W, 4200-4400 m, 09 Sep 1995, Sklenar & Kosteckovar 1237 (UC); Cerro Imbabura, 00[degrees]15'S, 78[degrees]10'W, 4300 m, 05 Jun 1995, Sklenar & Kosteckovar 31-16 (QCA), 31-17 (AAU). Loja: Cerro Toledo, E of Yangana, between Loja and Valladolid, 04[degrees]23'S, 79[degrees]07'W, 3000-3100 m, 26 Oct 2004, Lehnert 1465 (GOET, QCA, UC). Napo: Laguna Yuragcocha, 3 km E of Cerro Quilindana, 00[degrees]47'S, 78[degrees]21'W, 4050 m, 31 Mar 1979, Hohn-Nielsen 16375 (AAU); the SW slope 1.5 km from Cerro Quilindana, 00[degrees]47'S, 78[degrees]21'W, 4100 m, 01 Apr 1979, Holm-Nielsen 16416 (AAU); Cordillera de los Llanganatis, NE side of Laguna Encantada, 01[degrees]11'S, 78[degrees]12'W, 3430 m, 16 Mar 1983, Holm-Nielsen et al. 41858 (AAU); around Laguna Yuragcocha, 3 km E of the peak of Cerro Quilindana, Cordillera Oriental, 00[degrees]47'S, 78[degrees]21'W, 4100 m, 31 Mar 1979, Lojtnant & Molau 11567 (AAU); Volcan Antisana, rocky slopes on the W side of the mountain, 00[degrees]30'S, 78[degrees]10'W, 4500-4550 m, 21 Jul 1997, Sklenar & Sklenarova 2803 (UC); Volcan Antisana, rocky slopes on the W side of the mountain, 00[degrees]30'S, 78[degrees]10'W, 4500-4550 m, 21 Jul 1997, Sklenar & Sklenarova 2806 (UC). Pichincha: heading down W-SW from the highest point of Sincholagua, 00[degrees]35'S, 78[degrees]21'W, 4600 m, 02 Jun 1985, Bosco Nowak 171 (QCA); NE side of Cayambe mountain, 4420 m, 10 Dec 1961, Cazalet & Pennington 5750 (B, UC); Volcan Atacazo, SW slope, Km 19 from San Juan, 00[degrees]21'S, 78[degrees]39'W, 2900 m, 25 Aug 1980, Holm-Nielsen & Azanza 25180 (AAU); Nevado Cayambe, ladera S, 00[degrees]00.5'N, 78[degrees]00.95'W, 3700-3800 m, 29 Jun 2002, Lehnert 108 (GOET, QCA, UC); "In Monte Pichincha," 3500 m, 1921, Mille s.n. (P). Pichincha/Cotopaxi: NE slope of Illiniza Sur, 00[degrees]40'S, 78[degrees]42'W, 4400 m, 28 May 1995, Sklenar & Sklenarova 19_2 (UC). Pichincha/Napo: W side of a mountain ridge, ca. 2 km to the W from cerro Sara Urcu, 00[degrees]06'S, 77 57'W, 4400 m, 29 Aug 1995, Sklenar & Kosteckovar 100-9 (AAU). Tungurahua: Volcan Tungurahua, N side of the mountain, steep slope to the right of the summit, 01[degrees]27'S, 78[degrees]27'W, 4100 m, 08 Aug 1997, Sklenar & Sklenarova 3207 (UC).

Melpomene vulcanica Lehnert, sp. nov. TYPE.--ECUADOR. Nape: road Olmedo-Laguna San Marcos, E of the pass, 00[degrees]07'N, 77[degrees]59'W, 3640 m, 08-09 Jul 1980, Ollgaard et al. 34159 (holotype: AAU; isotype: QCA). Figs. 11, 12D.

A Melpomene pseudonutante (Christ & Rosenst.) A. R. Sin. & R. C. Moran frondibus lanceolatis basin versus truncatis vel breve cuneatis (vs. frondibus longe obovatis basin versus decurrentibus) rhizomatibusque tenuioribus (0.8-1.5 mm vs. 1.8-2.5 mm), a M. personata pilis laminarum abaxialium aequaliter dispersis (vs. pilis in soris confertis) frondibusque erectis (vs. pendentibus) differt.

Plants epiphytic or terrestrial, in mosslayers on trunks of trees or the caudices of Blechnum, in ditches, sometimes directly rooting in soil. Rhizomes horizontal, moderately to long-creeping, regularly branching at wide to right angles, (0.8-)1.2-1.5 mm diam. (Fig. 11A, B). Fronds 10-25 cm long, erect, inserted onto the rhizomes at right angles, or at narrow angles and strongly ascending, diffusely placed (internodes (2-)10-20 mm) (Fig. 11A). Scales 3.5-6.2 x 0.6-1.0 ram, (18-)20-30(-46) cells wide, clathrate, cell walls thick and dark brown to thin and brown, weakly iridescent, broadly lanceolate to ovate-lanceolate, cordate to pseudopeltate at bases, acute to attenuate at the flat to flaring tips (Fig. 11C); apical cells 3-8 (rarely more), palmately or linearly arranged, relatively small and ephemeral. Petioles (20-)35-65(-80) mm long, 0.6-1.0 mm thick, marginate from the laminar bases to semiterete, glabrescent to hairy, with brown, acicular or ciliform hairs 1.0-1.5 mm long (Fig. 11D); simple and branched clavate hairs to 0.2 mm long rarely present; petiolar bases with scales. Laminae (45-)90-220 x (10-)16-30(-34) mm, lanceate (broadest below the middle, normally after the second segment pair), rather abruptly ending at bases, truncate or widely cuneate, acute to attenuate at tips (Fig. 11A). Rachises dark brown to black, planar and slightly sunken adaxially (Fig. 11E), hemispherically protruding abaxially (Fig. 11F), with scattered, brown, setiform/ciliform hairs 1.0-1.2 mm long. Segments (9.0-)10.0-14.5 (-17.0) x (2.8-)3.2-3.8 mm (ca. 3-4 times longer than broad), patent to ascending (80-60[degrees]), inequilateral, decurrent towards the bases, fully adnate, long-deltate to linear-oblong, the tips acute (Figs. 35E, F), in smaller fronds also obtuse; segments abaxially with scattered hairs like those on the rachises; midveins black, visible at least abaxially, in large fronds usually well visible from both sides with some scattered hairs like those on the rachis (Fig. 11F); segment margins hyaline, (1-)2 cells wide, with few clavate hairs; proximal 1-3 pairs markedly smaller than the subsequent segments, but not auriculiform. Sori 2-4 pairs per segment, surrounded by (0-)2-8 brown setiform/ciliform hairs to 1.2 mm long (Fig. 11F).

[FIGURE 11 OMITTED]

[FIGURE 12 OMITTED]

The name refers to the fact that this species has been found mainly on the slopes of volcanoes.

Melpomene vulcanica grows in paramos and elfin forests at (2600-)3200-4500 m in Colombia and Ecuador (Fig. 12D).

The size of the laminae and the thickness of the cell walls in the scales vary considerably in this species, and every extreme of one character can be found within the whole range of the other without apparent correlation to the life form. However, plants with large fronds seem to grow in sheltered spots, i.e., they are present in nearly all epiphytes, but also in terrestrial plants from rock gullies. Smaller scales with thick cell walls are found in terrestrial plants growing directly on soil; large scales with wide, iridescent lumina are produced if the rhizomes grow in thick moss layers, i.e., in all epiphytes and some terrestrial plants from paramos. However, the scales are always larger than in M. sklenarii, which is similar to the small forms of M. vulcanica.

Rhizomes of Melpomene vulcanica are usually long-creeping and do not bear any developed fronds over a length of the apices. This clearly separates this species from M. pseudonutans, which has a rather compact growth (although the posterior parts of the rhizome persist and contribute greatly to the total length). Melpomene pseudonutans also has thicker rhizomes (usually more than 2.0 mm diam. vs. usually less than 1.5 mm in M. vulcanica), thicker, less hairy petioles (1.0-1.2(-1.8) mm vs. 0.6-1.0 mm), and longobovate blades (vs. lanceolate). Other species with long-creeping rhizomes similar to those of M. vulcanica have hairs clustered in the sori, like M. personata, and/or the midveins not or only partially visible on the abaxial laminae, like M. flabelliformis and M. moniliformis.

Ollgaard et al. 1194 (AAU) and Sklenar & Sklenarova 3019 (UC), both from Ecuador, are good examples of plants with strongly branching rhizomes. Bishop 1986 (UC), from Colombia, has exceptionally thick but nevertheless long-creeping rhizomes.

ADDITIONAL SPECIMENS EXAMINED.--COLOMBIA. Cauca: Cordillera Central, Parque Nacional del Purace, camino de Pimabala al volcan del Purace, 3700 m, 19 Jul 1976, Jaramillo Mejia & van der Hammen 5218 (COL); left side of road Toez-Tacueyo, 3640 m, Sep 1980, Rangel 2396 (COL). Huila: Cordillera Central, Cordillera del Buey, hike from Finca Loyola over the paramo down to San Antonio (2100 m according to residents), 2600 m, 14 Dec 1975, Bishop 1986 (UC). Huila-Cauca: Macizo Colombiano, paramo Las Papas, cerros y alrededores de la laguna La Magdalena, 3530 m, 16 Oct 1958, Idrobo et al. 2954 (AAU). Narino: Paramo El Tabano, Alto de la Cordillera, entre Pasto y El Encano, vertiente occidental, 3200 m, 11 Jan 1941, Cuatrecasas 11920 (COL); Pasto, Volcan Galeras, paramo al S de la cima, 3700 m, 24 Dec 1972, Hagemann & Leist 1783 (COL); vertientes de Cumbal, 3400-4300 m, 22 Jan 1973, Hagemann & Leist 1974 (COL). Risaralda: Santa Rosa, Cordillera Central, entre la hacienda La Sierra y Termales de Santa Rosa, Quebrada La Sierra, 3525 m, 26 Jan 1980, Jaramillo Mejia et al. 5931 (COL).

ECUADOR. Carchi: Paramo El Angel, in the pass on road El Angel-Tulcan, 00[degrees]41'N, 77[degrees]54'W, 3750-3850 m, 15 May 1973, Holm-Nielsen et al. 5476 (AAU); carretera Tufino-Maldonado-Lagunas Verde, 01[degrees]28'S, 79[degrees]13'W, 3900-4000 m, 01 Oct 1994, Navarrete 756 (AAU); Volcan Chiles, along gully on the SW side of the volcano, 00[degrees]48'N, 77[degrees]57'W, 4150-4200 m, 23 Jun 1995, Sklenar & Sklenarova 637 (UC). Chimborazo: El Altar, N side of the volcano, on the ridge below the Canoningo peak, 01[degrees]41'S, 78[degrees]24'W, 4200-4400 m, 19 Aug 1995, Sklenar & Kosteckovar 950, 93-15 (UC). Imbabura: E slopes of Cayambe peak, 3200 m, 16 Jul 1944, Wiggins 10407 (NY). Morona-Santiago: trail Alao-Huamboya, around the pass, between Cuspipaccha and alt. 3700 m on E slope, 01[degrees]47'S, 78[degrees]25'W, 3550-3950 m, 07 May 1982, Ollgaard et al. 38233 (AAU); trail Alao-Huamboya, around the pass, 01[degrees]47'S, 78[degrees]25'W, 3550-3950 m, 07 May 1982, Ollgaard et al. 38290 (AAU); Cerros Yuibug-Pailacajas, E side of the mountain ridge, 01[degrees]45'S, 78[degrees]27'W, 4300 m, 31 Jul 1997, Sklenar & Sklenarova 2968 (UC). Napo: Cordillera de los Llanganates, loma between Rio Topo and Rio Verde Grande, 3 km WNW of Cerro Hermoso, 01[degrees]13'S, 78[degrees]18'W, 4000 m, 10 Nov 1980, Holm-Nielsen & Jaramillo J. 28329 (AAU); Cordillera de los Llanganatis, NE side of Laguna Encantada, 01[degrees]11'S, 78[degrees]12'W, 3430 m, 16 Mar 1983, Holm-Nielsen et al. 41769, 41802, 41811, 41842 (AAU); S side of the crater of Cerro Sumaco, 00[degrees]34'S, 77[degrees]43'W, 3780-3820 m, 26 Apr 1979, Holm-Nielsen et al. 17334 (AAU); E side of Cerro Sumaco, 00[degrees]34'S, 77[degrees]43'W, 3750 m, 30 Apr 1979, Holm-Nielsen et al. 17469 (AAU); SE side of Cerro Sumaco, 00[degrees]34'S, 77[degrees]43'W, 3750 m, 30 Apr 1979, Holm-Nielsen et al. 17489 (AAU); ibid., 3750-3800 m, 01 May 1979, Holm-Nielsen et al. 17550, 17577, 17590 (AAU); ibid., 3200 m, 02 May 1979, Holm-Nielsen et al. 17699 (AAU); ibid., 3350 m, 05 May 1979, Holm-Nielsen et al. 17969 (AAU); carretera Quito-Baeza, Quijos, 4300 m, 08 Jul 2002, Lehnert 176 (GOET, QCA, UC); N-facing slopes at the W side of Laguna Parcacocha, 00[degrees]16'S, 78[degrees]09'W, 4100 m, 18 Mar 1979, Lojtnant & Molau 11164 (AAU); NE-facing ridge on the N side of Cerro Sumaco, 00[degrees]35'S, 77[degrees]39'W, 3600-3700 m, 24 Apr 1979, Lojtnant & Molau 12664 (AAU); S side of Cerro Sumaco, 100-200 m S of the main crater, 00[degrees]35'S, 77[degrees]39'W, 3700-3800 m, 29 Apr 1979, Lojtnant & Molau 12954B, 12954C (AAU); Oyacachi, 5 km despues del paso, bosques en los margenes del carretero, 00[degrees]12'S, 78[degrees]06'W, 3500 m, 28 Dec 1996, Navarrete 1367 (AAU, QCA); Oyacachi, Yarupaccha, 00[degrees]12'S, 78[degrees]07'W, 3620-3680 m, 16 Jan 1996, Navarrete 1416 (AAU), road Quito-Baeza, 7-8 km NW of the Laguna Papallacta (Paramo de Guamani), 00[degrees]19'S, 78[degrees]08'W, 3800 m, 20 Jul 1976, Ollgaard & Balslev 8162 (AAU); Paramo de Soguillas, near Las Torres de Llanganati, 01[degrees]08-09'S, 78[degrees]15-16'W, 3850-4000 m, 16-17 May 1982, Ollgaard & Holm-Nielsen 38752 (AAU); Llanganati, paramo SE of Chosa Aucacocha, between Aucacocha and Pan de Azucar, 01[degrees]09'S, 78[degrees]18'W, 3800-3900 m, 15 May 1982, Ollgaard et al. 38496 (AAU); Volcan Antisana, rocky gully on the W side of the mountain, 00[degrees]30'S, 78[degrees]10'W, 4400-4500 m, 22 Jul 1997, Sklenar & Sklenarova 2784 (AAU, UC); NE side of Volcan Antisana, 00[degrees]27'S, 78[degrees]08'W, 4300 m, 17 Aug 1997, Sklenar & Sklenarova 3402 (AAU, UC); ibid., 00[degrees]27'S, 78[degrees]08'W, 4200 m, 18 Aug 1997, Sklenar & Sklenarova 3438 (AAU, UC); Hacienda Yanahurco, 3800 m, 28 Aug 2001, Smith A. R. et al. 2875 (UC). Pichincha: Guamani pass, E of Pifo, Cordillera Oriental, 4000 m, 10 Nov 1944, Ewan 16447 (UC); 2 km S of Paso de la Virgen on road Quito-Baeza, 00[degrees]20'S, 78[degrees]13'W, 4000-4200 m, 19-20 May 1984, Laegaard 52148 (AAU); Carretera Quito-Baeza, 00[degrees]20.2'S, 78[degrees]13.2'W, 4200 m, 08 Jul 2002, Lehnert 168 (GOET, QCA, UC); ibid., Km 256, 4300 m, 08 Jul 2002, Lehnert 174 (GOET, QCA, UC); Paramo de Guamani, Carretera Pifo-Papallacta, 00[degrees]19'S, 78[degrees]12'W, 3960 m, 13 Jan 1990-26 May 1991, Leon S. 1164 (AAU); Volcan Cayambe, N slopes, 00[degrees]03-05'N, 77[degrees]59'W, 3750-3850 m, 09 Jul 1980, Ollgaard et al. 34235A (AAU); road and trail from Chaupi-paramo of Volcan Corazon, 00[degrees]34'S, 78[degrees]41'W, 4140 m, 15 Mar 1995, Ollgaard et al. 1194 (AAU); W side of a mountain ridge ca. 2 km to the west from Cerro Sara Urcu, 00[degrees]06'S, 77[degrees]57'W, 4100 m, 30 Aug 1995, Sklenar & Sklenarova 108-2 (AAU); Pichincha/Napo border, edge of Antisana reserve, 3800 m, 28 Aug 2001, Smith A. R. et al. 2871 (UC); along road Quito-Pallapacta, 2800-3900 m, 27 Feb 1994, van der Werff & Gray 13356 (UC); at pass of road Quito-Pallapacta, 4000 m, 14 Jul 1991, van der Werff & Palacios 12357 (AAU). Pichincha/Napo: Volcan Cayambe, N slopes, 00[degrees]03-05'N, 77[degrees]59'W, 3750-3850 m, 09 Jul 1980, Ollgaard et al. 34290 (AAU); road Olmedo-Laguna San Marcos, E of the pass, 00[degrees]07'N, 77[degrees]59'W, 3620-3800 m, 10-11 Jul 1980, Ollgaard et al. 34462 (AAU); W side of a mountain ridge, ca. 2 km to the west from cerro Sara Urcu, 00[degrees]06'S, 77[degrees]57'W, 4400 m, 29 Aug 1995, Sklenar & Kosteckovar 1118 (AAU). Tungurahua: Santiago de Pillaro Canton, paramos de Pisayambo, surroundings of laguna de Pisayambo, 01[degrees]05'S, 78[degrees]23'W, 3600-3900 m, 11 Oct 1998, Cueva 249 (UC); Santiago de Pillaro, Parque Nacional Llanganates, W of Cerro Hermoso, near saddle between headwaters of Rio Verde and Rio Topo, 01[degrees]11'40"S, 78[degrees]19'34"W, 3950 m, 12 Nov 1999, Neill et al. 12005 (UC); Cerro Hermoso, SW ridge of the mountain, 01[degrees]14'S, 78[degrees]18'W, 4100 m, 06 Sep 1997, Sklenar & Sklenarova 3625 (AAU); Patate Canton, Parque Nacional Llanganates, slopes of Cerro Pan de Azucar, on transect Paramo de Soguillas-Cerro Pan de Azucar, 01[degrees]09'S, 78[degrees]17'W, 3800 m, 13 Oct 1998, Vargas H. et al. 2820 (UC).

ACKNOWLEDGMENTS

I thank my colleagues Michael Kessler, Alexander Schmidt-Lebuhn, Alan Smith, Michael Sundue, Robbin Moran, Nicki Mandl, Maarten Christenhusz, Maria Teresa Murillo, Julio Betancur, Jorge Jacome, Thomas Janssen, and S. Robbert Gradstein for their help and support in this study. I am indebted to the curators and director of the following herbaria for making their collections accessable to me: AAU, B, BM, COL, COLO, F, GH, HUT, LOJA, MO, NY, P, Q, QCNE, QPLS, RB, S, SP, UC, US, and especially Stephan Beck (LPB), Hugo Navarrete (QCA), and Asuncion Cane (USM). I furthermore thank Blanca Leon and an anonymous reviewer for their helpful comments on the manuscript.

The financial support by the DAAD (German Academic Exchange Service) and the DFG (German Research Foundation) is greatly acknowledged.

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MARCUS LEHNERT

Albrecht-von-Haller-Institut fur Pflanzenwissenschaften, Abt. Systematische Botanik, GeorgAugust-Universitat Gottingen, Untere Karspule 2, D-37073 Gottingen, Germany

Current address: Staatliches Museum fur Naturkunde Stuttgart, Am Lowentor, Rosenstein 1, D70191 Stuttgart, Germany. [e-mail]