Synopsis of Mickelia, a newly recognized genus of bolbitidoid ferns (Dryopteridaceae) ROBBIN C. MORAN1, PAULO H. LABIAK2, AND MICHAEL SUNDUE1 1 The New York Botanical Garden, Bronx, NY 10458-5126, USA; e-mail: rmoran@nybg.org; e-mail: msundue@nybg.org 2 Depto. de Botânica, Universidade Federal do Paraná, C.P. 19031, 81531-980, Curitiba, PR, Brazil; e-mail: plabiak@ufpr.br Abstract. Our recent molecular phylogenetic study revealed a previously unrecognized clade of six species that is sister to Elaphoglossum. Within this clade, four species are currently classified in Bolbitis, one in Lomagramma, and one in Acrostichum. For this clade, we propose the name Mickelia, with M. nicotianifolia as the type species. We also make new combinations for the species in our phylogenetic study shown to belong to Mickelia (M. bernoullii, M. guianensis, M. hemiotis, M. nicotianifolia, M. oligarchica, and M. scandens) and two other species believed to belong to the clade based on morphology (M. lindigii, M. pergamentacea). A new hybrid and two new species are also described (M. ×atrans, M. furcata, and M. pradoi). In total, Mickelia consists of ten species and one hybrid. It is entirely neotropical. We provide a key to the genera of bolbitidoid ferns and a synopsis of Mickelia that gives for each species a complete synonymy, geographical distribution, comparative discussion, and illustration. Key Words: Bolbitis, taxonomy, neotropics. Resumo. Análises filogenéticas recentes revelaram a existência de uma clado composto por seis espécies, irmão de Elaphoglossum. Das espécies incluídas nestas análises, quatro são atualmente tratadas em Bolbitis, uma em Lomagramma, e uma em Acrostichum. A este clado propomos o nome Mickelia, com M. nicotianifolia sendo sua espécie-tipo. Neste trabalho são apresentadas as novas combinações para as espécies de Mickelia, tanto as que foram tratadas nas análises filogenéticas (M. bernoullii, M. guianensis, M. hemiotis, M. nicotianifolia, M. oligarchica e M. scandens), quanto para duas outras que são aqui incluídas com base em evidências morfológicas (M. lindigii e M. pergamentacea). Um híbrido novo e duas espécies novas são também descritos (M. ×atrans, M. furcata e M. pradoi). No total, Mickelia compreende dez espécies e um híbrido. O gênero é inteiramente neotropical, e é caracterizado pelo hábito hemiepifítico, nervuras anastomosadas, e pela ausência de gemas (essas, se presentes, surgindo próximas à junção da pina com a raque). São apresentadas uma chave para os gêneros das samambaias bolbitidoides, e uma sinopse para Mickelia contendo uma lista completa de sinônimos, dados sobre distribuição geográfica, discussões comparativas e ilustrações. A recent phylogenetic analysis (Moran et al., 2010) of the bolbitidoid ferns (equivalent to “the former lomariopsid ferns” of Schuettpelz & Pryer, 2007) has shown that Bolbitis is diphyletic. The first clade, to which the name Bolbitis Schott applies, is pantrop- ical and consists of about 50 species. It is sister to the rest of the bolbitidoid ferns (i.e., Arthrobotrya, Elaphoglossum, Lomagramma, and Teratophyllum). The second clade, which we called the “Bolbitis nicotianifolia clade” (Moran et al., 2010), is entirely neotropical Brittonia, 62(4), 2010, pp. 337–356 © 2010, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A. ISSUED: 15 December 2010 338 [VOL 62 BRITTONIA and consists of ten species. It is sister to Elaphoglossum. We propose herein to name this second clade as Mickelia. We believe it is best to recognize the clade as a distinct genus, instead of including it in Elaphoglossum, for two reasons. First, its species have never been thought to be closely related to Elaphoglossum; most have been classified in Bolbitis. Second, to include the clade in Elaphoglossum would drastically alter the characterization of that genus as one with species having mostly simple, entire leaves, free veins, and phyllopodia. This characterization has remained remarkably constant ever since Christ’s (1899) monograph of Elaphoglossum, and it seems unwise to alter it now. On the other hand, one might argue that Mickelia as here defined should be split into more genera. This is, of course, a matter of opinion. At present, we feel it most convenient to recognize a single genus of ten species. Most of the species of Mickelia were thoroughly and admirably treated in a monograph of Bolbitis by Hennipman (1977), who included detailed descriptions, complete synonymy, discussions, and illustrations. The purpose of this paper is not to monograph again these ten species, but to provide a synopsis of them. Below we give a key to the genera of bolbitidoid ferns and a key to the species of Mickelia. For these species we make all the necessary new combinations and present comparative discussions, complete synonymy (taken largely from Hennipman, 1977), and illustrations. Methods and materials Specimens at NY were studied to write the key and comparative discussions. Synonymy and type-specimen information were taken almost entirely from Hennipman (1977). Geographic distributions were obtained primarily from Hennipman (1977), Hennipman and Moran (1995), Mickel and Smith (2004), and Tryon and Stolze (1991), and secondarily by examination of specimens at MO and NY. To obtain spore images, spores were transferred with dissecting needles from herbarium specimens to aluminum SEM stubs coated with an asphalt adhesive. The stubs were then coated with gold-palladium in a sputter-coater for 2.5 min. The spores were imaged digitally using a JEOL jSM-5410LV SEM equipped with JEOL Orion 5410 software interface. The accelerating voltage was 15 kV. The spore images are publicly available at http:// www.plantsystematics.org. Results and discussion Mickelia R. C. Moran, Labiak & Sundue, gen. nov. Type: Acrostichum nicotianifolium Sw. (=Mickelia nicotianifolia (Sw.) R. C. Moran, Labiak & Sundue). Anapausia C. Presl subgen. Euryostichum C. Presl, Abhandlungen der Königlichen Böhmischen Gesellschaft der Wissenschaften, ser. 5, 6: 545. 1851. [= Epimel. Bot. 188. 1851.]. Lectotype, designated by Pichi Sermolli, 1969: Bolbitis nicotianifolia (Sw.) Ching ex C. Chr. (basionym: Acrostichum nicotianifolium Sw.) Hoc genus novum ab Elaphoglosso foliis pinnatis, venis reticulatis, et phyllopodiis carentibus differt. A Bolbitide lamina apicibus conformibus, bulbillis carentibus aut si praesentibus (in speciebus duabus) prope basin pinnarum locatis differt. Plants hemiepiphytic or (3 species) terrestrial; rhizomes short- to long-creeping, dorsiventral, the ventral meristele elongate (as seen in cross-section), the ventral surface rootbearing, the dorsal surface bearing 3 or more ranks of leaves; phyllopodia absent; leaves continuous with rhizome or (in L. ×atrans, L. guianensis, and L. pradoi) articulate; sterile and fertile leaves dimorphic, the petioles of the fertile leaves longer and the fertile pinnae smaller; leaves 1-pinnate or rarely (in M. furcata) with a single free basal segment on the basal pinnae and thus 2-pinnate; laminae usually with a free apical segment similar to (conform) the lateral pinnae; lamina buds absent or (in B. hemiotis and B. oligarchica) present in the axils of the pinnae (buds never associated with the terminal segment as in Bolbitis); pinnae articulate or continuous with (i.e., non-articulate to) the rachis, entire or serrate distally; veins various but never free, either areolate throughout without main lateral veins and without included veinlets, or with main lateral veins connected by slightly arched cross veins, and typically with included veinlets in the areoles; sori acrostichoid; paraphyses 2010] MORAN ET AL.: MICKELIA (DRYOPTERIDACEAE) absent; spores monolete, brown or black (nongreen); perispore broadly folded; x = 41 (Walker, 1966, 1985). Etymology.—The genus name honors John T. Mickel, American pteridologist, for his many contributions to ferns, especially Elaphoglossum, the sister genus of Mickelia. Nearly all species of Mickelia have previously been placed in Bolbitis, with two exceptions. First, M. guianensis has been generally placed in Lomagramma following Ching (1932), although a combination for it is available in Bolbitis (see treatment of the species below). Second, M. scandens has never been provided with a combination in Bolbitis because it has long been considered a synonym of M. guianensis. Mickelia and Bolbitis share certain homoplastic and plesiomorphic characters (such as anastomosing veins, free included veinlets, and sterile-fertile leaf dimorphy) which explain why most Mickelia species have been placed in Bolbitis (Moran et al., 2010). Despite this placement, our molecular phylogenetic analysis recovered Mickelia as sister to Elaphoglossum, a genus that can usually be distinguished by simple entire leaves, free veins, and the presence of phyllopodia (Moran et al., 2010). The morphological similarity between Mickelia and Bolbitis is such that no single character distinguishes the two genera. Nevertheless, the two genera exhibit tendencies to differ. Most species of Mickelia are hemiepiphytic (only three are terrestrial: M. hemiotis, M. oligarchica, and M. pergamentacea), whereas Bolbitis is entirely terrestrial. Except for two of the terrestrial species (i.e., M. hemiotis and M. oligarchica), Mickelia lacks leaf buds; in contrast, Bolbitis typically has leaf buds (in fact, this character is the source of the generic name, which is derived from the Greek bolbition, diminutive of bolbos, bulb, alluding to the foliar buds). The two species of Mickelia with leaf buds have those buds located in a different position than in Bolbitis. In Mickelia the buds occur at the pinna bases on the acroscopic side of the costae where it is joined by the lamina tissue (i.e., the buds are 339 produced just outside of the pinna “axil”). In contrast, the buds in Bolbitis are adaxial and associated with the apical portion or terminal segment of the lamina. Another tendency to differ is that the lamina apex is often conform or subconform in Mickelia (i.e., resembling the lateral pinnae), whereas in Bolbitis it is most commonly pinnatifid and tapered. Although not completely diagnostic, venation helps distinguish Mickelia from Bolbitis. Three general patterns of venation are present in Mickelia, and two of these can occasionally be found in Bolbitis. The first pattern, not found in Bolbitis, is shown by M. guianensis, M. pradoi, and M. scandens. In these species, unlike their congeners, main lateral veins from the costae are absent; instead, the veins are of equal thickness and anastomose completely between the costa and margin to form series of polygonal areoles without free included veinlets. This pattern resembles that of Lomagramma (Holttum, 1937, 1978) and is one reason M. guianensis was formerly placed in that genus (e.g., by Ching, 1932). The second pattern consists of species with main lateral veins that are cross-connected by thinner, slightly arched veins. Within the spaces enclosed by these veins are slightly finer veins that anastomose to form areoles with included free veinlets that are both excurrent and recurrent and sometimes branched. The arched cross veins tend to be slightly thicker than the finer veins between them. This pattern is found in M. furcata, M. hemiotis, M. nicotianifolia, M. oligarchica, and M. pergamentacea. The pattern is rare in Bolbitis but occurs in species such as B. gaboonensis (Hook.) Alston, B. pandurifolia (Hook.) C. Chr., B. sinuata (C. Presl) Hennipman (Hennipman, 1977). The third pattern consists of main lateral veins between which occur more regular, usually polygonal areoles. Free-included veinlets may occur within these areoles, although only occasionally so in M. lindigii. This pattern is found in M. bernoullii and M. lindigii. Similar patterns can be found in species of Bolbitis such as B. aliena (Sw.) Alston, B. serratifolia (Mertens ex Kaulf.) Schott, and B. sculpturata (Fée) Ching. 340 BRITTONIA The perispore morphology of Mickelia does not differ significantly from that of Bolbitis. This can be seen by comparing the spore images in Fig. 1 with that in Tryon and Tryon (1982) and Tryon and Lugardon (1991). Both genera have broadly folded or sharply winged perispores. The finer microstructure found in Mickelia (minute ridges, spines or papillae) can also be found in Bolbitis. [VOL 62 The rhizome ground tissue of most ferns is white (pers. obs.), but in five species of Mickelia observed so far it is greenish: M. lindigii, M. nicotianifolia, M. oligarchica, M. pergamentacea, and M. scandens (see color images of rhizome cross-section of M. nicotianifolia in Fig. 1 of Moran et al. (2010), and of M. lindigii, M. nicotianifolia, and M. pergamentacea at www.plantsystematics. FIG. 1. Spores of nine species of Mickelia. A. M. bernoullii (Mexico, Hernández 2616, NY). B. M. guianensis (Bolivia, Sperling 6518, NY). C. M. hemiotis (Mexico, Hernández 2777, NY). D. M. lindigii (Guyana, Jansen-Jacobs et al. 5903, NY). E. M. oligarchica (Costa Rica, Burger et al. 10733, NY). F. M. pergamentacea (Mexico, Hernández G. 1005, NY). G. M. nicotianifolia (Puerto Rico, Sánchez & Liogier 124, NY). H. M. pradoi (Brazil, Fiaschi et al. 2254 NY). I. M. scandens (Brazil, Gerdes 83, NY). Scale bars = 10 micrometers. 2010] MORAN ET AL.: MICKELIA (DRYOPTERIDACEAE) org). This suggests that green ground tissue might be a synapomorphy for Mickelia, but other species of the genus and more bolbitidoid ferns need to be examined to assess this character. 341 A morphological character matrix representing all bolbitidoid genera and 49 species of bolbitidoid ferns can be found in Moran et al. (2010). The following key will distinguish Mickelia from the other genera of bolbitidoid ferns: Key to the genera of bolbitidoid ferns 1. Plants with differentiated bathyphylls and acrophylls; paraphyses present, simple, or stellate or capitate, long-stalked, the stalk 1 or 2 cells wide. 2. Veins anastomosing; spores smooth to verrucate . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Lomagramma 2. Veins free; spores with broad folds. 3. Leaves 2-pinnate; paraphyses branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Teratophyllum 3. Leaves 1-pinnate; paraphyses simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arthrobotrya 1. Plants without differentiated bathyphylls and acrophylls; paraphyses absent or rarely (in Elaphoglossum) present, lanceolate, short-stalked, the stalk several cells wide. 4. Leaves simple, entire, or rarely (6 species) divided; veins free, rarely (3 species) anastomosing; phyllopodia usually present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Elaphoglossum 4. Leaves 1-pinnate, rarely (2 species) simple; veins anastomosing; phyllopodia absent. 5. Bulblets absent, or if present (2 species), borne near the junction at the base of the pinna and the rachis; plants hemiepiphytic or (3 species) terrestrial . . . . . . . . . . . . . . . . . . . . . . . . . . . Mickelia 5. Bulblets present, rarely (2 species) absent, borne at or near the apices of the pinnae or (usually) the terminal segment; plants terrestrial, rarely low hemiepiphytes . . . . . . . . . . . . . . . . . . . . . . Bolbitis Key to the species of Mickelia 1. Areoles without free included veinlets or (in M. lindigii) occasionally present 2. Petioles continuous with (not articulate to) the rhizome; rhizomes ca. 1 cm wide, densely scaly, the scales ca. 10 mm long, not clathrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. M. lindigii 2. Petioles articulate to the rhizome; rhizomes ca. 0.5 cm wide, sparsely scaly, the scales 1.5–5.5 mm long, clathrate. 3. Laminae with (16–)22–25 pairs of pinnae; rhizome scales 4–5.5×0.3–0.5 mm, brown to castaneous, call lumens broad, easily visible; lamina apices often gradually reduced . . . . . . . . . 10. M. scandens 3. Laminae with 9–16 pairs of pinnae; rhizome scales 1.2–2.2×0.3–0.4 mm, blackish, cell lumens narrow, not easily visible; lamina apices abruptly reduced 4. Rhizomes light brown to castaneous; pinnae (1.5–)2–2.7(–3.2) cm wide . . . . . . 3. M. guianensis 4. Rhizomes darkly castaneous to blackish; pinnae 1.4–1.7 cm wide. . . . . . . . . . . . . .9. M. pradoi 1. Areoles with free included veinlets. 5. Pinnae long-stalked, the stalks 1.5–2 cm long; laminae 2-pinnae at base, the basal pinnae often with a stalked pinnule; segments narrowly elliptic, long-cuneate . . . . . . . . . . . . . . . . . . . . . . . 2. M. furcata 5. Pinnae sessile to short-stalked, the stalks up to 0.5 cm long; laminae 1-pinnate at base, the basal pinnae entire; segments elliptic to broadly elliptic, short-cuneate to truncate. 6. Pinna pairs 8–16; pinnae 2.5–3.5 cm wide, lanceolate, equilateral; rhizome scales densely denticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. M. bernoullii 6. Pinna pairs 1–4; pinnae 2.2–9.5 cm wide, broadly lanceolate to elliptic, equilateral or inaequilateral; rhizome scales entire to sparsely denticulate or fimbriate. 7. Buds present at bases of pinnae. 8. Pinna pairs 1, the pinnae opposite or sub-opposite; terminal pinna 15–20 cm wide; rhizome scales brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. M. oligarchica 8. Pinna pairs 2–4, the pinnae alternate; terminal pinnae 10–12.5 cm wide; rhizome scales dark castaneous to blackish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. M. hemiotis 7. Buds absent from bases of pinnae. 9. Rhizome scales firm, opaque, dark brown; lateral pinnae broadly lanceolate to elliptic, widest at the middle, usually symmetrical, with the basiscopic and acroscopic sides of the same width6. M. nicotianifolia 9. Rhizome scales soft, translucent, light brown, sometimes slightly ferrugineous; lateral pinnae ovate, widest at or below the middle, cymbiform, usually inaequilateral, with the basiscopic side wider than the acroscopic side. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. M. pergamentacea 342 BRITTONIA 1. Mickelia bernoullii (Kuhn ex H. Christ) R. C. Moran, Labiak & Sundue, comb. nov. Acrostichum bernoullii Kuhn ex H. Christ, Bull. Soc. Bot. Belg. 35: 224. 1896. Gymnopteris bernoullii (Kuhn ex H. Christ) Diels in Engler & Prantl, Die Natürlichen Pflanzenfamilien 1: 201. 1899. Leptochilus bernoullii (Kuhn ex H. Christ) C. Chr., Botanisk Tidsskrift 26: 296. 1904. Bolbitis bernoullii (Kuhn ex H. Christ) Ching in C. Chr., Index Filic., Suppl. 3: 47. 1934. Type: Guatemala. Dept. Unknown: Costa Grande, between Escamillas and Palahueco (“Palahucco”), K. C. G. Bernoulli & Cario 382 [“282” in error] (lectotype, designated by Hennipman, 1977: B; duplicates: B, fragments BM, NY). (Figs. 1A, 2) Gymnopteris tuerckheimii H. Christ, Bull. Herb. Boissier, sér. 2, 6: 290. 1906. Leptochilus tuerckhemii C. Chr., Index Filic., Suppl. 1: 48. 1913. Type: Guatemala. Alta Verapaz: Cubilquitz, 350 m, Sep 1904, H. Von Türckheim 8831 (holotype: US 2-sheets). Gymnopteris donnell-smithii H. Christ, Bull. Herb. Boissier, sér. 2, 6: 289. 1906. Poecilopteris donnell-smithii (H. Christ) Maxon, Contr. U.S. Nat. Herb. 13: 20. 1909 (as “Poikilopteris”). Leptochilus donnell-smithii C. Chr., Index Filic., Suppl. 1: 48. 1913. Bolbitis donnellsmithii (H. Christ) Ching in C. Chr., Index Filic., Suppl. 3: 48. 1934. Type: Guatemala. Alta Verapaz: Cubilquitz, 350 m, Aug 1904, H. Von Türckheim 8830 (holotype: US 3-sheets; isotype: NY [fragment]). Distribution and habitat.—Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, Venezuela, Colombia, Ecuador; hemiepiphytic in wet forests; 0–1500 m. Mickelia bernoulli is characterized by its venation consisting of main lateral veins between which occur polygonal areoles with included free veinlets (slightly thicker cross veins connecting the lateral ones are lacking). It resembles M. lindigii in habit and pinna shape, but this species differs by the subentire, translucent, and subtortuous rhizome scales, in contrast to the densely denticulate, opaque, and non-tortuous rhizome scales in M. bernoullii. 2. Mickelia furcata R. C. Moran, Labiak & Sundue, sp. nov. Type: Ecuador. Pichincha: [VOL 62 Reserva de ENDESA, NE of Vicente Maldonado, 0°6′N, 79°02′W, 600 m, 29 Apr–1 May 1985, S. Laegaard 45213 (holotype: QCA; isotypes: AAU, QCNE). (Fig. 3) A speciebus congeneribus petiolulis pinnarum 1–2 cm longis, pinnis 2–2.5 cm latis, ascendentibus, in dimidio inferiore anguste cuneatis, pinnis basalibus cum segmento singulari usque ad 11 cm longo distinguitur. Roots inserted ventrally; rhizomes dorsiventral, with an elongate ventral meristele, long-creeping, about 1 cm wide, dark brown, densely scaly throughout, the scales dark, lanceolate; leaves presumably dimorphic; sterile leaves 40×20 cm, petiole 12 cm long, 2.5 mm wide, stramineous, articulate to the rhizome, glabrous or with a few scales like those of the rhizome, grooved adaxially; lamina 2-pinnate at basal pinnae, 1-pinnate distally, with 4 pinna pairs, the base truncate, the apex conform; rachis narrowly alate distally, adaxially grooved, the groove with a central carina, neither the carina or groove continuing onto the pinnae, abaxially rounded, with scattered scales or glabrescent; medial pinnae 15–20 × 1.5–2 cm, linear-elliptic, stalked, the stalks 1.5–2 cm long, the pinna bases narrowly cuneate, the pinna apices longattenuate, the margins slightly serrate distally; costae rounded abaxially and adaxially; lamina surfaces glabrous both abaxially and adaxially; veins anastomosing throughout, with main lateral veins connected by slightly arched cross veins, with 3 areoles between the costa and the margins, with free included veinlets both recurrent and excurrent, the costal areoles measuring about two to three times longer than wide; fertile leaves unknown. Distribution and habitat.—NW Ecuador, known only from the type; rheophytic, on rocks in river; 600 m. Although the type lacks fertile leaves, Mickelia furcata can be confidently placed as a bolbitidoid fern because of its elongate ventral meristele, and furthermore placed in Mickelia because of its lack of bulblets, conform terminal pinna, venation consisting of main lateral veins connected by slightly arched cross veins, and areoles with included veinlets. Among its congeners it is unique by forked basal pinnae (thus the specific epithet) with a 2010] MORAN ET AL.: MICKELIA (DRYOPTERIDACEAE) 343 FIG. 2. Mickelia bernoullii. A. Rhizome and sterile leaf. B. Rhizome scale. C. Adaxial surface of rachis-costa juncture. D. Venation of sterile pinna. (Mexico, Hernández 850, NY.) 344 BRITTONIA [VOL 62 FIG. 3. Mickelia furcata. A. Rhizome and sterile leaf; note furcate basal pinna on left side of leaf, and the apparently aborted lobe on the basal pinna on the right side. B. Venation of sterile pinna. (Ecuador, Laegaard 45213, AAU.) 2010] MORAN ET AL.: MICKELIA (DRYOPTERIDACEAE) single basal basiscopic segment up to 11 cm long. It is also distinctive by pinna stalks 1–2 cm long, and lateral pinnae 2–2.5 cm wide, ascending, and narrowly cuneate for about half their length. 3. Mickelia guianensis (Aubl.) R. C. Moran, Labiak & Sundue, comb. nov. Polypodium guianense Aubl., Hist. Pl. Guiane 2: 962. 1775. Acrostichum guianense (Aubl.) Baker, Fl. Bras. (Martius) 1(2): 591. 1870. Gymnopteris guianensis (Aubl.) H. Christ, Bull. Herb. Boissier, sér. 2, 1: 70. 1901. Leptochilus guianensis (Aubl.) C. Chr., Bot. Tidsskr. 26: 286, fig. 1. 1904. Lomagramma guianensis (Aubl.) Ching, Amer. Fern J. 22: 17. 1932. Bolbitis guianensis (Aubl.) Vareschi, Flora Venezuela 1: 376. 1969 (Vareschi cited Kramer as the combining author, Acta Bot. Neerl. 3: 486. 1954, but Kramer’s combination was invalid; Art. 33.2). Type: French Guiana. s.d., F. Aublet s.n. (holotype: BM-n.v., photos GH, US-n.v.). (Figs. 1B, 4A–E) Lomagramma guianensis var. antillensis C. Chr., Kungl. Svenska Vetenskapsakademiens Acad. Handlingar, ser. 3, 16(2): 43. 1936. Type: Dominican Republic. Samaná: Península de Samaná: Playa de Rincón, on trail to Laguna, near La Yautía, 23 May 1930, E. L. Ekman H15083 (holotype: S; isotype: NY). Distribution and habitat.—Greater Antilles (except Jamaica), French Guiana, Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil (Amazonia); hemiepiphytic, wet forests; 0–800 m. The venation of Mickelia guianensis differs from most of its congeners by lacking main lateral veins from the costae. It resembles that of M. pradoi and M. scandens, which are further similar by veins completely areolate, the areoles more or less polygonal and without included veinlets. The other species of Mickelia with areoles lacking (or nearly so) included veinlets is M. lindigii, but that species has prominent main lateral veins from the costae. For a long time, Mickelia scandens was considered a synonym of this species (e.g., Christensen, 1904), but it is distinguished here on the basis of lamina apex shape, pinna number, and number of areoles between the costa and margin (see additional comments 345 under M. scandens). See comments under M. pradoi for comparisons with that species. Plants of Mickelia guianensis from the Greater Antilles differ slightly from plants of the continent. Most conspicuously, the Greater Antillean plants have laminae more tapered towards the apex, and in this they resemble M. scandens, from southeastern Brazil. The rows of areoles between the costae and margins are 1 or 2 as in M. scandens, as opposed to 3 or 4 in M. guianensis. In describing the Greater Antillean plants as Lomagramma guianensis var. antillensis, Christensen (1936) noted that the veins became pale reddish, a character present in almost all the specimens we have seen from the Greater Antilles; however, this can also be found occasionally in typical M. guianensis (e.g., Bolivia, Jiménez 1941, NY). The Greater Antillean plants agree with typical M. guianensis as to size of the leaves, size and number of pinna pairs, serration of the pinna margins, and rhizome scale color. For that reason they are treated here as M. guianensis as opposed to M. scandens. The growth architecture and rhizome morphogenesis of this species were studied by Gay (1993) and Hebant-Mauri and Gay (1993). They found differences in the anatomy, morphology, and growth behavior between the terrestrial versus climbing rhizomes. Some of these differences might be synapomorphies for Mickelia, but extensive fieldwork is needed to confirm this not only for other species of Mickelia, but also for other genera of bolbitidoid ferns. 4. Mickelia hemiotis (Maxon) R. C. Moran, Labiak & Sundue, comb. nov. Leptochilus hemiotis Maxon, Amer. Fern J. 14: 101. 1925. Bolbitis hemiotis (Maxon) Ching in C. Chr., Index Filic., Suppl. 3: 48. 1934. Type: Trinidad. 1877–1880, A. Fendler 101 (holotype: US; isotypes: B, BM, G, K, MICH, MO, NY 4-sheets, S, UC, US). (Figs. 1C, 5) Distribution and habitat.—Mexico, Belize, Costa Rica, Panama, Trinidad, Venezuela, Colombia, Ecuador; terrestrial or on rocks, wet forests; 100–1000 m. Mickelia hemiotis is characterized by buds located near the bases of the distal pinnae, the main lateral veins connected by slightly 346 BRITTONIA [VOL 62 FIG. 4. Comparison of sterile leaves from three similar species of Mickelia. A–E. M. guianensis. A. Sterile leaf of climbing rhizome (Guyana, Jenman s.n., NY). B. Sterile leaf of climbing rhizome (Colombia, Killip & Smith 14868, NY). C. Sterile leaf of terrestrial rhizome (from Bolivia, Boom 4743, NY). D. Sterile leaf of terrestrial rhizome (Bolivia, Kessler et al. 8811, NY). E. Detail of the venation (Guyana, Jenman s.n., NY). F–J. M. scandens. F. Sterile leaf of climbing rhizome (Brazil, Prado & Hirai 2025, SP). G. Sterile leaf of climbing rhizome (Brazil, Boldrin 174, SP). H. Sterile leaf of terrestrial rhizome (Brazil, Luederwaldt 21567, SP). I. Sterile leaf of terrestrial rhizome (Brazil, Athayde 226, SP). J. Detail of the venation (Brazil, Prado & Hirai 2025, NY). K–O. M. pradoi. K. Sterile leaf of climbing rhizome (Brazil, Fiaschi et al. 2045, CEPEC). L. Sterile leaf of climbing rhizome (Brazil, Sant’Anna et al. 788, CEPEC). M, N. Sterile leaf of terrestrial rhizome (Brazil, Matos 1841, CEPEC). O. Pinna base showing slightly swollen articulation (Brazil, Fiaschi et al. 2045, NY). P. M. scandens, pinna base showing slightly swollen articulation (Brazil, Prado & Hirai 2025, NY.) arched cross veins, and the areoles with included free veinlets that are both excurrent and recurrent. It greatly resembles M. pergamentacea but differs primarily by the presence of leaf buds. In Trinidad, Walker (1985) found that M. hemiotis had diploid and tetraploid cytotypes, the latter he suspected of being autoploid. In general appearance, the two cytotypes did not differ, but he noted differences in the size of 2010] MORAN ET AL.: MICKELIA (DRYOPTERIDACEAE) 347 especially by the number, size, and shape of the pinnae. Mickelia lindigii is distinguished from M. bernoullii by the rarity or lack of included veinlets in the areoles and rhizome scales that tend to be subentire, translucent, and subtortous (versus densely denticulate, opaque, and straight). FIG. 5. Mickelia hemiotis. A. Bud at base of pinna. B. Fertile pinnae. C. Pinnae with buds at base. (Trinidad, Hart 6924, NY; reproduced from Mickel & Smith, 2004.) their stomata and spores, and slight differences in the color and length of the rhizome scales. 5. Mickelia lindigii (Mett.) R. C. Moran, Labiak & Sundue, comb. nov., Chrysodium lindigii Mett. in Triana & Planchon, Ann. Sci. Nat. Bot., sér. 5, 2: 205. 1864. Acrostichum lindigii (Mett.) Baker in Hooker & Baker, Syn. Fil. 423. 1868, nom. illeg. non Karsten (1858). Leptochilus lindigii (Mett.) C. Chr., Bot. Tidsskr. 26: 290. 1904. Bolbitis lindigii (Mett.) Ching in C. Chr., Index Filic., Suppl. 3: 48. 1934. Type: Colombia. Boyacá: Muzo, 700–900 m, A. A. Lindig 258 (lectotype, designated by Hennipman, 1977: B 2sheets; duplicates: BM, K [fragment], P [fragment]). (Figs. 1D, 6) Chrysodium pellucens Mett. in Triana & Planchon, Ann. Sci. Nat. Bot., sér. 5, 2: 205. 1864. Bolbitis pellucens (Mett.) Ching in C. Chr., Index Filic., Suppl. 3: 49. 1934. Type: Colombia. Magdalena: Santa Anna, 1851, Lewy s.n. (lectotype, designated by Hennipman, 1977: B; duplicate: P). Gymnopteris costaricensis H. Christ, Bull. Herb. Boissier, sér. 2, 4: 964. 1904. Leptochilus lindigii (Mett.) C. Chr. var. costaricensis C. Chr., Bot. Tidsskr. 26: 291. 1904. Type: Costa Rica. Without locality, 1903, C. Wercklé & Brune s.n. (lectotype, designated by Hennipman, 1977: P; duplicate: BM). Distribution and habitat.—Costa Rica, Panama, Colombia, Ecuador, Peru, Bolivia, Brazil (Acre); hemiepiphytic, wet forests; 50–1000 m. Mickelia lindigii is included in the genus based on its similarity to M. bernoullii, 6. Mickelia nicotianifolia (Sw.) R. C. Moran, Labiak & Sundue, comb. nov. Acrostichum nicotianifolium Sw., Syn. Fil. 13, 199. 1806 (as “nicotianaefolium”). Gymnopteris nicotianifolium (Sw.) C. Presl, Tent. Pterid. 244. 1836. Anapausia nicotianifolia (Sw.) C. Presl, Epimel. Bot. 189. 1851. Chrysodium nicotianifolium Mett., Fil. Lips., 22. 1856. Leptochilus nicotianifolius (Sw.) C. Chr., Bot. Tidsskr. 26: 285. 1904. Bolbitis nicotianifolia (Sw.) Alston, Bull. Misc. Inform. Kew 310. 1932. Type: St. Thomas: s.d., E. P. Ventenat s.n. (holotype: S). (Figs. 1G, 7) Acrostichum acuminatum Willd., Spec. Pl. 5: 116. 1856, non Jussieu ex Poiret in Lam. 1810. Gymnopteris acuminata (Willd.) C. Presl, Tent. Pterid. 244, 1836. Anapausia acuminata (Willd.) C. Presl, Epimel. Bot. 188. 1851. Chrysodium acuminatum (Willd.) Mett., Fil. Lips. 22, 1856. Photinopteris acuminata (Willd.) C. V. Morton, Contr. U.S. Nat. Herb. 38: 31. 1967. Type: Plumier, Tract. Fil. Amer. 100, pl. 115, 1705, Lingua cervina, scandens, Citrei foliis, major, drawn from C. Plumier s.n., Lesser Antilles, eastern Martinique, St. Maria (P-JU). Gymnopteris acuminata C. Presl var. heterophylla Fée, Hist. Acrost. 86. 1845. Type: Guadeloupe. s.d., Anon. (holotype: P, 3-sheets). Nephrodium sodiroi Baker, J. Bot. 15: 16. 1877. Type: Ecuador. Pichincha: “Andes of Ecuador,” s.d., A. L. Sodiro s.n. (lectotype, designated by Lellinger, 1993: K, photo US). Acrostichum nicotianifolium Sw. var. saxicolum Jenman, Bull. Bot. Dept. Jamaica (Kingston) n.s. 5, 153. 1898. Type: Jamaica. “1874–79,” G. S. Jenman s. n. (holotype: K). Leptochilus nicotianifolius (Sw.) C. Chr. var. simplex Rosenst., Repert. Spec. Nov. Regni Veg. 22: 23. 1926. Type: Costa Rica. Alajuela: Llanuras de San Carlos, 200 m, Apr 1910, A. C. Brade & C. Brade 552 (holotype: S). Leptochilus killipii Maxon, Amer. Fern J. 21: 138. 1931. Bolbitis killipii (Maxon) Lellinger, Fern Gaz. 11: 107. 1975. Type: Panama. Panamá: 4 miles above Juan Díaz, in wet forest ravine, 75 m, 13 348 BRITTONIA [VOL 62 FIG. 6. Mickelia lindigii. A. Sterile leaf. B. Fertile leaf. C. Detail of venation from sterile lamina. D. Scales from the petiole base. E. Rhizome cross-section. (Costa Rica, Rojas et al. 4568, NY.) 2010] MORAN ET AL.: MICKELIA (DRYOPTERIDACEAE) 349 FIG. 7. Mickelia nicotianifolia. A. Venation of sterile pinna (Puerto Rico, Sánchez & Liogier 124, NY). B. Rhizome, sterile and fertile leaves. C. Rhizome and sterile leaf. D. Rhizome scales. (B–D, Ecuador, Øllgaard 99724, AAU.) 350 BRITTONIA [VOL 62 Jan 1918, E. P. Killip 2778 (holotype: US; isotypes: B, P, S) tagua, and Jivaria de Pintuc, 1000 m, s.d., M. A. Stübel 906 (holotype: B 2-sheets). Bolbitis riparia R. C. Moran, Nord. J. Bot. 15: 178. 1995. Ecuador. Sucumbíos: trail at Río Ushaué N of Cooperativa Flor del Valle, 2.5 kmW of bridge over Río Aguarico at Lumbaquí, 00°05′ N, 77°20′W, 500 m, 29 Jan 1992, B. Øllgaard 99724 (holotype: AAU; isotypes: QCA, QCNE). Leptochilus bradeorum Rosenst., Repert. Spec. Nov. Regni Veg. 9: 70. 1910. Bolbitis bradeorum (Rosenst.) Ching in C. Chr., Index Filic., Suppl. 3: 47. 1934. Type: Costa Rica. San José: La Palma, 1400 m, 18 Aug 1909, A. C. Brade & C. Brade 367 p.p. (lectotype, designated by Hennipman, 1977: UC; duplicate NY). Distribution and habitat.—Jamaica, Dominican Republic, Puerto Rico, Virgin Islands, Guadeloupe, Martinique, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, French Guiana, Suriname, Trinidad, Guyana, Venezuela, Colombia, Ecuador, Peru, Brazil; hemiepiphytic, wet forests; 0–600 m. Distribution and habitat.—Nicaragua, Costa Rica, Panama, Guyana, Colombia, Ecuador, Peru, Bolivia, Brazil (Acre); terrestrial, wet forests; 200–1200 m. Mickelia nicotianifolia resembles M. pergamentacea, and the two are frequently confused in herbaria. Mickelia pergamentacea differs by asymmetrical pinnae (slightly wider on the basiscopic side) and laminae that tend to dry a light yellow-green. In M. nicotianifolia the pinnae are symmetrical and the laminae tend to dry a darker green. Mickelia nicotianifolia is variable in its habitat preference; it may occur terrestrially, on logs, or as a low epiphyte climbing up to 2 m on tree trunks. When terrestrial, the rhizomes are narrower (usually less than 1 cm) and longer creeping than the climbing ones, and their leaves are either simple or, if pinnate, with only one or two pairs of pinnae (pers. obs.). Bolbitis riparia, here placed in synonomy, represents such a terrestrial form. 7. Mickelia oligarchica (Baker) R. C. Moran, Labiak & Sundue, comb. nov. Acrostichum oligarchicum Baker, Syn. Fil. 418. 1868. Leptochilus oligarchicus (Baker) C. Chr., Bot. Tidsskr. 26: 285. 1904. Bolbitis oligarchica (Baker) Hennipman, Amer. Fern J. 65: 30. 1975 [as “oligarchia”]. Type: Peru. San Martín: Mt. Guayrapurima, near Tarapoto, Sep 1856, R. Spruce 4737 (lectotype, designated by Maxon, 1931: K, photo US). (Figs. 1E, 8) Hypoderris stuebelii Hieron., Hedwigia 46: 323. 1907. Leptochilus stuebelii (Hieron.) Maxon, Proc. Biol. Soc. Wash. 46: 142. 1933. Bolbitis stuebelii (Hieron.) C. Chr., Index Filic., Suppl. 3: 50. 1934. Type: Ecuador. Tungurahua: between Abi- Mickelia oligarchica is easily recognizable by its trifoliolate leaves with relatively large terminal segments, and buds at the bases of the two lateral pinnae. The buds are present on both sterile and fertile leaves. Some specimens from South America differ from the Central American ones by having more densely scaly rhizomes and petioles, but no other differences appear to correlate. The single specimen known from Guyana (Henkel 155, NY) is unusual because it lacks buds but is otherwise typical. In the wild, the sterile laminae are often oriented more or less parallel to the ground. Their upper surfaces traps fallen organic matter in the shallow depression at the juncture of the two lateral pinnae and terminal segment. The organic matter decays to form a humus in which the buds proliferate. As the leaf senesces and inclines toward the ground, the buds are lowered to the soil where they take root. In the field it is common to find young plantlets on senesced, partially decayed leaves still attached to the parent plant (Moran, pers. obs.). The only other species of Mickelia with buds is M. hemiotis (also terrestrial), which differs from M. oligarchica by having 2 or 3 pairs of lateral pinnae. 8. Mickelia pergamentacea (Maxon) R. C. Moran, Labiak & Sundue, comb. nov. Leptochilus pergamentaceus Maxon, J. Wash. Acad. Sci. 14: 144. 1924. Bolbitis pergamentacea (Maxon) Ching, Index Fil., Suppl. 3: 49: 144. 1934. Type: Jamaica. Green River Valley, 750 m, 11–12 Feb 1903, L. M. Underwood 1426 (holotype: US 2-sheets; isotype: NY). (Figs. 1F, 9) Leptochilus turrialbae Rosenst., Repert. Spec. Nov. Regni Veg. 22(206–608): 22. 1925. Bolbitis turrialbae 2010] MORAN ET AL.: MICKELIA (DRYOPTERIDACEAE) 351 FIG. 8. Mickelia oligarchica. A. Rhizome, sterile and fertile leaves. B. Adaxial surface of rachis. C. Abaxial surface of rachis. D. Bud on acroscopic side of pinna base. (Ecuador, Moran 6033, NY.) 352 BRITTONIA [VOL 62 FIG. 9. Mickelia pergamentacea. A. Fertile leaf (Jamaica, Maxon & Killip 550, NY). B. Sterile leaf. C. Detail of the venation. (B–C, Jamaica, Underwood 1426, NY.) 2010] MORAN ET AL.: MICKELIA (DRYOPTERIDACEAE) (Rosenst.) Ching, Index Fil., Suppl. 3: 50. 1934. Type: Costa Rica. Cartago: Turrialba, 650 m, s.d., A. C. Brade & C. Brade 370 (holotype: S). Distribution and habitat.—Greater Antilles, S. Mexico, Belize, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru; terrestrial, wet forests; 200–700(–1500) m. Mickelia pergamentacea resembles M. hemiotis and M. nicotianifolia. It differs from the former by lack of buds at the base of the pinnae; otherwise, the two species are nearly identical in pinna shape and venation. From the latter species it differs by relatively broader terminal segments and pinnae, and the laminae tending to dry a lighter color. Also, the lateral pinnae tend to be broadest at or below the middle and slightly wider on the basiscopic side. In Puerto Rico, Mickelia pergamentacea often forms extensive colonies by means of creeping rhizomes that frequently branch. The rhizomes are generally 2–3 cm under the soil surface (Sundue, pers. obs.). In Jamaica, Walker (1966) found Mickelia pergamentacea to be diploid (2n=42). 9. Mickelia pradoi R. C. Moran, Labiak & Sundue, sp. nov. Type: Brazil: Bahia: Uruçuca, Estrada de Itacaré para Serra Grande, pouco após km 43, ramal à direita após acesso para a cachoeira do Tijuípe, 14°23′12″S, 39°04′45″W, 4 Apr 2004, P. P. Fiaschi, A. M. Amorim, A. Matini & L. L. Paixão 2254 (holotype: CEPEC; isotype: NY). (Figs. 1H, 4K–O) Ab Mickelia guianensi rhizomatibus atrantibus, pinnis 1.4–1.7 cm latis, areolis biseriatis vel rariter triseriatis et apicibus laminarum subconformatis differt. Plants hemiepiphytic, climbing up to ca. 2 m; roots inserted ventrally; rhizomes dorisventral, with an elongate ventral meristele, the cortex whitish or sometimes drying reddish, long-creeping, 5–7 mm wide, darkly castaneous to blackish, lustrous, densely scaly at apex and petiole bases, the scales 1.2–1.6×0.2–0.3 mm, lanceolate, subpeltate, blackish, clathrate, the lumens translucent but very narrow and not easily seen, the base cordate, the apex acuminate, the margins 353 castaneous, fimbriate; leaves dimorphic; sterile leaves 25–50×12–20 cm, petioles 7–10 cm long, 2–2.5 mm wide, stramineous, articulate to the rhizome, glabrous or with a few scales like those of the rhizome, grooved adaxially; lamina 1-pinnate throughout, with 14–16 pinna pairs, the base truncate, the apex abruptly reduced and ending in a terminal segment that is subconform to the lateral pinnae and which is not articulate to the rachis; rachis narrowly alate distally, adaxially grooved, the groove with a central carina, neither the carina or groove continuing onto the pinnae, abaxially rounded, with scattered scales or glabrescent, the scales 1–1.2×0.1–0.2 mm, linear-lanceolate, dark brown, the margins slightly fimbriate; pinnae 7–13×1.4–1.7 cm, linear-lanceolate, stalked, the stalks articulate to the rachis, the pinna bases cuneate, the pinna apices attenuate, the margins serrate distally; costae rounded abaxially and adaxially; lamina surfaces glabrous both abaxially and adaxially; veins proximally anastomosing, distally free, with 2(3) areoles between the costae and the margins, lacking free included veinlets, the costal areoles measuring about twice as long as wide, the veins terminating submarginally in elongate hydathodes; fertile leaves 20–25×4–8 cm, with ca. 10 pairs of pinnae, the lamina reduced to only the sporangium-bearing tissue; sori acrostichoid; sporangia glabrous; paraphyses absent; spores light brown, monolete, elliptic, with a crested perispore. Distribution and habitat.—Brazil (Bahia); hemiepiphytic, Atlantic rain forest; 0–200 m. Additional specimens examined. BRAZIL. Bahia: Mun. de Una, Reserva Biológica do Mico-Leão, picada paralela ao Rio Maruim, 15°09′S, 39°05′W, 8–12 Mar 1993, Amorim et al. 1147 (CEPEC, NY); Mun. de Uruçuca, estrada de Serra Grande para Uruçuca, 14°29′ 59″S, 39°06′54″W, 380 m, 18 Mar 2004, Fiaschi et al. 2045 (CEPEC); . Porto Seguro, Parque Nacional do PauBrasil, 16°30′S, 39°15′W, 90 m, 22 Sep 2009, Matos et al. 1841 (CEPEC); Mun. de Una, Reserva Biológica do Mico-Leão, 15°09′S, 39°05′W, 30 Apr 2000, Sant’Ana et al. 788 (CEPEC, NY); Mun. de Ilhéus, 3 kmN of Rodoviária, Mata da Esperança, forest N of dam and reservoir, 14°46′56″S, 39°04′09″W, 50 m, 19 Sep 1994, Thomas et al. 10546 (CEPEC, NY). Mickelia pradoi is included in Mickelia based on its similarity to M. guianensis and M. scandens. It can be recognized by dark rhizomes, linear-lanceolate pinnae, and lami- 354 BRITTONIA nae abruptly reduced to a subconform terminal segment that resembles the lateral pinnae. It most resembles M. guianensis, which differs by reddish brown rhizomes, 9–13 pinna pairs, wider pinnae ([1.5–]2–2.7[–3.2] cm), more areoles (3, rarely 4) between the costa and the margin, and proximal costal areoles about as long as wide. Whereas M. pradoi is restricted to southeastern Brazil (Bahia), M. guianensis occurs in Amazonian and premontane Andean forests. The two species do not overlap in distribution. Mickelia pradoi hybridizes with M. scandens to form M. ×atrans, described above. Mickelia scandens differs from M. pradoi by reddish brown rhizomes, rhizome scales 4–5.5 mm long, brown, and clathrate with broad cell lumens, leaves on the climbing rhizomes with 18–25 pairs of pinnae, and laminae gradually reduced. The specific epithet honors Jefferson Prado, Brazilian pteridologist, for his many contributions to the study of tropical American ferns. 10. Mickelia scandens (Raddi) R. C. Moran, Labiak & Sundue, comb. nov. Acrostichum scandens Raddi, Syn. Fil. Bras. 6, tab. 34. 1819 [Opusc. Sci. 3: 284. 1819.]; Pl. Bras. Nov. Gen. 1: 6, tab. 18. 1825. Cyrtogonium scandens (Raddi) Baker, J. Bot. (Hooker) 4: 154. 1841. Heteroneuron raddianum Fée, Mém. Foug. [Hist. Acrost.] 2: 94. 1845 (an illegitimate renaming of Acrostichum scandens Raddi). Poecilopteris scandens (Raddi) C. Presl, Epimel. Bot. 175. 1849. Lomariopsis raddiana (Fée) Mett., Fil. Hort. Bot. Lips. 22. 1856. Neurocallis scandens (Raddi) T. Moore, Index Fil. XIX. 1857. Acrostichum raddianum (Fée) Kunze ex Hook., Sp. Fil. 5. 264. 1864. Gymnopteris scandens (Raddi) H. Christ, Farnkr. 51. 1897. Type: Brazil. Rio de Janeiro: Mt. Estrella, s.d., G. Raddi s.n. (holotype: PI-n.v.; isotype: FI, photo US). (Figs. 1I, 4F–J) Distribution and habitat.—Coastal region of SE Brazil; hemiepiphytic, wet forests; 0–800 m. This species has long been treated as a synonym of Mickelia guianensis (e.g., Christensen 1904; Pichi Sermolli & Bizzarri, 2005). [VOL 62 It differs from the similar M. pradoi by laminae often gradually reduced toward the apex (vs. abruptly reduced) and (16–)22–25 (vs. 9–13) pairs of pinnae on the leaves of the climbing rhizomes. The lamina differences are also evident in the terrestrial leaves (Fig. 4). The leaves of the climbing rhizomes have 2(–3) series of areoles between the costa and margin, whereas M. guianensis has 3 or 4 (Fig. 4). 11. Mickelia ×atrans R. C. Moran, Labiak & Sundue, hybr. nov. Type: Brazil. Bahia: Parque Nacional de Monte Pascoal, on the NW side of Monte Pascoal, 39°25′W, 16° 53′S, 100–200 m, 11 Jan 1977, R. Harley, S. J. Mayo, R. M. Storr, T. S. Santos & R. S. Pinheiro 17837 (holotype: CEPEC; isotypes: K-n.v., NY). Hybrida inculta ex Mickelia pradoi et M. scandenti genita, intermedia inter parentes; scandentes et terrestres; rhizoma atrans, longe repens; sporae abortivae, irregulariter formatae. Plants hemiepiphytic, climbing up to ca. 4 m; roots inserted ventrally; rhizomes dorsiventral, with an elongate ventral meristele, the cortex drying whitish, long-creeping, 6–8 mm wide, dark brown to blackish, lustrous, densely scaly at apex and petiole bases, the scales 1.5–2.2×0.2–0.3 mm, lanceolate, subpeltate, brown to dark brown, clathrate, the lumens translucent but very narrow and not easily seen, the base cordate, the apex acuminate, the margins castaneous, fimbriate; leaves dimorphic; sterile leaves 40×11–13 cm, petiole 6–7 cm long, 3 mm wide, stramineous, articulate to rhizome, glabrous or with a few scales like those of the rhizome, grooved adaxially; lamina 1-pinnate throughout, with 21–23 pinna pairs, the base truncate, the apex gradually reduced and ending in a terminal segment that is pinnatifid or sub-conform to the lateral pinnae and which is not articulate to the rachis; rachis narrowly alate distally, adaxially grooved, the groove with a central carina, neither the carina or groove continuing onto the pinnae, abaxially rounded, with scattered scales or glabrescent, the scales 1–1.2×0.1– 0.2 mm, linear-lanceolate, dark brown, the margins slightly fimbriate; pinnae 6–7×1.4– 2010] MORAN ET AL.: MICKELIA (DRYOPTERIDACEAE) 1.7 cm, linear-lanceolate, stalked, the stalks articulate to the rachis, the pinna bases cuneate, the pinna apices attenuate, the margins serrate distally; costae rounded abaxially and adaxially; lamina surfaces glabrous both abaxially and adaxially; veins proximally anastomosing, distally free, with 2(3) areoles between the pinna costa and the margins, lacking free included veinlets, the costal areoles measuring about at long as wide, the veins terminating submarginally in elongate hydathodes; fertile leaves essentially the same size as the sterile leaves, and with the same number of pinna pairs, the lamina reduced to only the sporangium bearing tissue; sori acrostichoid; sporangia glabrous; paraphyses absent; spores light brown, aborted. Distribution and habitat.—Brazil (Bahia), known only from the type; hemiepiphytic, Atlantic rain forests; 100–200 m. Mickelia ×atrans appears to be a hybrid between M. pradoi and M. scandens. Its spores are aborted, being of irregular sizes and shapes. In number of pinna pairs (21– 23) it resembles M. scandens ([16-]22–25) instead of M. pradoi (14–16), but by its dark rhizome it resembles M. pradoi. The hybrid’s rhizome scales are short (1.5– 2.2 mm long) like those of M. pradoi (1.2–1.6 mm long) instead of the longer ones (4–5.5 mm) found in M. scandens, but in color and visibility of the cell lumens they appear intermediate between the two putative parents. The specific epithet atrans, refers to the dark (at least when dried) rhizomes. Acknowledgments This research was funded by a grant to the first author from the United States National Science Foundation (DEB 0717056). Labiak’s research at the New York Botanical Garden was partially funded by the Brazilian government (CNPq/PDE 201782/2008-1). We thank Alan R. Smith, Jefferson Prado, and Fernando Matos for helpful information about specimens, and Judith Garrison Hanks for help in taking the SEM photomicrographs of the spores. We are grateful to David Barrington and Thomas Janssen, who provided many helpful comments on 355 the manuscript. We thank John Mickel for allowing use of the line illustration for Fig. 5, drawn by Bobbi Angell, from The Pteridophytes of Mexico. The line illustrations in Figs. 2, 6, 8, and 9 were prepared by Haruto Fukuda. Literature Cited Ching, R. C. 1932. The genus Lomagramma in America. American Fern Journal 22: 15–18. Christ, H. 1899. Monographie des genus Elaphoglossum. Neue Denkschriften Allgemeinen Schweizerishcen Gessellschaft für die Gesammten Naturwissenschaften 36: 1–159, + tab. 1–4. Christensen, C. 1904. On the American species of Leptochilus sect. Bolbitis. 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