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Notes on Neotropical Hymenophyllaceae PDF

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24-37 American Fern Journal 81(1): (1991] on Hymenophyllaceae Notes Neotropical David Lellinger B. DC Department of Botany, Smithsonian Institution, Washington, 20560 Hymenophyllaceae In preparing a treatment of the for the flora of the Guianas, have found several taxonomic problems in the family that lie at least partially I outside the Guianas, and so cannot be included in that Flora, as well as two new, narrowly endemic Trichomanes from need be species of the flora area that to published. Two Hymenophyllum hirsutum and Confused with Species It monograph American Hymenophyllum In his of the species of sect. SphQerocionium (now Morton called sect. Leptocionium), (1947] considered H, hirsutum Swartz be and widely a rather variable distributed species [L.] to He present in virtually the entire neotropics. included 17 basionymns as synonyms under this species, far more than under any other species in the monograph. War Because he prepared treatment during and World this just after he was many unable to see types of of these synonyms. believe that his 11, I inclusive concept of H. hirsutum also arose from his study of Jamaican material (where the type came from), which variable in lamina size and shape. Morton is did indicate [1947, 143, 158) the possibility of recognizing additional species p. from Brazil, especially the robust plants of southeastern Brazil that he placed in H. hirsutum. Hymenophyllum unknown kaiefeurum, an from western essentially species Guyana masquerading as H. hirsutum, and H. vestitum, a species of southeastern Brazil, are distinct species that can readily be separated from H. They hirsutum. differ from H. hirsutum and from each on other characters of lamina indument and division, segment width, and to some extent on frond size and involucre shape, as can be seen in the following key: mm Lamina segments 0.75-1 1. wide, the marginal hairs basally forked with both arms simple; pinnae with only the basal acroscropic segments furcate. Fronds 2-6 cm segments (-10) long; slightly undulate; involucres ovate, much longer than wide, not wider than the segments H. kaieteurum .......2. mm Lamina segments 1-1.5 wide, the marginal more complex 1. hairs (stellate or basally forked with both branches 1 or forked or pinnae with stellate); at segments Marginal lamina hairs basally forked with arm and arm 2. forked sim 1 1 cm !gm long; segments 1-1.1 (-1.25) mm wide; mvolucres much subcircular, a little wider than Ions, not wi um :ments Marginal lamina 2. hairs stellate or bistellate; segments slightly mm cm Fronds folded. (5-)6-18(-30) long; segments 1-1.5 wide NEOTROPICAL HYMENOPHYLLACEAE LELLINGER: 25 involcures subcircular, a little wider than long, as wide as the segments H. vestitum 3. Hymenophyllum ^^^7^ hirsutum 1. (L.) Swartz, Bot [Schrader] 1800[2}:99. 1801. J. Trichomanes hirsutum PL 5"19 L. Sp. 2:1098. 1753.-<rYPE: Based on plate SOB of ( Plumier's "Traite .", which based on a specimen collected by Plumier is , . West in the Indies. — Trichomanes Hymenophylium 3^f ciliatum Swartz, Prodr. 136. 1788. ciliatum HL^ (Swartz] Swartz, 1801.— Sphaerocionium Bot. (Schrader) 1800(2):100. J. ^ciliatum Hymenophyllaceae ^*^/ K. Presl, 34. 1844.-^^ype: Jamaica, Swartz not (S BM seen photo 6185, isotypes B-Hb. Willd. 20222 not seen Tryon photo, not Morton seen photo 6587, US]. ^^^'^ SphaeTocionium Hymenophyllaceae grevilleanum K. Presl, 34. 1844. Xo^ Hymeni \ m superfl. which on specimen on Vincent based collected (Icon. Fil. 35. 1827], a St. is l:t. r by Guilding (K not seen). 1848.-?^ Kunze, Linnaea 21:240. ifoJium Surinam, Weigeli Hymenophyllum €%'} nom. Based on Trichomanes ciliatum ultimately superfl.-t^TYPE: based on name. so the type of that [M Hymenophyllum '^H^l elegantissimum Fee, Hist. Foug. Antill. RB 1866.-^ype: Guadeloupe, L'Herminier in 1861 (P or not seen]. 29, 2. t. f. z.0^ [Mem. Hymenophyllum & Foug. Fee L'Herminier in Fee, Hist. Antill. atrovirens nom. Foug. 1866, non Colenso, 1844, nee Christ, 1904, 11]:120, 30, 4. t. f. RB Guadeloupe, Germain in 1864 (P or not seen]. illeg.-t^TvPE: Pitou Caraibe, 1904. 'i^^/ Hymenophyllum dimorphum Herb. Boissier 4:941. Christ, Bull. II, Costa Mano Puntarenas, Type: Alto de Tigre, basin of the Rio Diquis, Pcia. US]. 700 m, 12109 not seen, isotype Rica, (P? Pittier and Venezuela Colombia Common Mexico Panama, to Bolivia, Distribution: the Antilles, to in Minas Amazonas, Gerais). Para, to French Guiana. Rare in northern and central Brazil (Acre, %Q^l\ 1898.^ W. Guianas Hymenophyllum Ferns Ind. 15. kaieteurum Jenm. Brit. /enman (K).— Type: Guyana, Potaro River, Figs. 1, 2. mm Fronds the fronds distant. Rhizome diam., 0.25 in wide-creeping, ca. mm apex from the cm diam., black, in 0.2 determinate; 0.8-2.2 long, ca. stipes margms, on the sparsely pilose to the middle alate, the ala abruptly contracted, laminae ovate-lanceate, simple; base or the the hairs pluricellular, furcate at (3-pmnate the proximally at cm cm pinnate wide. 2 [2-]3-6 long, (1.5-]2-3.5 pmnae uniformly alate; rachises pinnate acroscopic base of the pinnae] distally; , segments slightly revolute, the and pinnae (2-]4-8 pairs, pinnules the alternate, mm margins pilose, the hairs furcate or emarginate; 0.75-1.25 wide, not f-1.5i — VOLUME NUMBER AMERICAN FERN JOURNAL: 81 1 (1991] 26 t L * ^ V ^ -^ ® ->^ ® ;irHAJ' i % f ^*^^^ Df^HBMJt BfllTllt* Ci^«|»A M*« Figs. 1-2. Holotype of Hymenophyllum kaieteurum Jenm., /enman (K). Fig. i. Holotype. Fig. 2 two Detail of fronds. simple, acicular, unicellular; indusial valves ovate or broadly ovate, obtuse or truncate the base, pilose the apex, the hairs 1- or 2-celled, the receptacles at at not exserted. m Distribution: Rare endemic at ca. 400 elevation in the uplands of western Guyana. GUYANA, Specimens ex.\mined: Potaro River, Sheenabowa [Chenapowu], ]enman 12^% (P). & 12^7 Mazaruni under 250[0]-3G00 McConneJl QueJch 596 Mount Raywa, [K); River, (K); ft, /enman (NY). Hymenophyllum vestitum Bosch, Ned. Kruidk. Arch. 3. [K. Presl) v. d. 5(3):193. — IPS%^ 1863- Sphaerocionium vestitum K. Presl, Hymenophyllaceae 58. 1844. i-^Lectotype: Rio de Janeiro, Est. Rio de Janeiro, Brazil, Beyrich [PRC? not seen), chosen by Morton and inferentially (Contr. U.S- Natl. Herb. 29:155. 1947} confirmed here. Hymenophyllum gardnerianum Sturm %ti\^ in Martins, Fl. Bras. 1[2):297. 1859.-*^Type: Rio de Rio de Gardner 213 Janeiro, Est. Janeiro, Brazil, BR {holotype not seen, isotypes K, P not seen Morton photo 4563). Hymen Hum ^n caulopteron Fee, Crypt. Vase, Bresil 1:197, 70, 3< t. f. 1869 Rio 1713 Morton NEOTROPICAL HYMENOPHYLLACEAE LELLINGER: 27 J Morton Hymenophyllum microcarpon %39f Fee, Crypt. Vase. Bresil 1:245, 69, 1869, 3. t. f. non micTocarpum nom. H. Desv., illeg.-^SYNTYPEs: Rio de Rio Janeiro, Est. de Glaziou 2268 not seen Morton photo and 3356 Janeiro, Brazil, (P 4558J not (P seen). Hymenophyllum & 1899.-^ lo51\ ulei Christ Giesenh. Flora 86:85, f ype: 6, 7. I Teresopolis, Serra dos Orgaos, Est. Rio de Janeiro, Brazil, 1000 m. Ule 4510 (presumably P not seen, isotypes L not seen Morton photo 2528, US]. Hymenophy] Jum Schwacke, Nov. Mineir. 1900.-*^ -L*s"^'Z- eJatius Christ in 2:13. Pi. Type: Sao Antonio, Ule 206 Est. Sta. Catarina, Brazil, (P). 'Xcb'I^HymenophylJum tuberosum Hedwigia ciliatum Rosenst. 46:74. 1906, as var. A "tuberosa."-^TYPE: renaming of H. ulei Christ & Giesenh., and so based on name. the type of that ^«6^y Hymenophyllum Hedwigia cihatum abbreviatum Rosenst. 56:360. 1915, var. Brade 5169 as "abbreviata."—Type: Ribiera, Est. S. Paulo, Brazil, A. C. (S not seen). Distribution: Occasional endemic southeastern Brazil (Rio de Janeiro, Sao Paulo, Parand). in name bv Morton cited Kru commutatum on Sphaerocionium K. Arch. based Presl., 4:413. 1859, boryanum and Hymenophyllaceae This based on H. sensu Raddi, 1844. 34. is on "Plantarum Brasiliensium ." This figure so plate 79, 4 of Raddi's is fig. . . somewhat Morton thought too lackine in detail and aouears to be stylized. it and lamina division seems be like H. gJaziovii Baker in Hook. Icon. PL 17, 1612. 1886. If this proves to t. name with H. name must be replaced the earlier the H. gJaziovii true, commutatum. Selected Specimens Examined (All from Brazil): Rio de Janeiro: Near Rio de Janeiro, Wilkes woods, Burkart 17453, 17465 (both Exped. Sao Paulo: Alto da Serra, Biological Station n. (US). s. do 800 m, A. Brade US); Serra da Bocaina, Segadas-Vianna 2746, 2832 (both US); Serra Itatiaia, C. 8826 [US), 1000 m, Dus^n 713 (US). Parana: Mun. Cerro Azul, Morro Grande. Hatschbach 7109 Morro Kummrow Mun. Quatro Mun. 1703, 1931 (both US); Barras, [VSy, Morretes. Pilao de Pedra, Kummrow Anhangava, 2476 (US). Hymenophyllum nigrescens The Identity of HymenophyiJum Liebm. usually has been thought to be a fairly nigrescens common with wide range (Mexico to Venezuela independent species a rather and Mickel and Recently, Beitel (1989, and perhaps Colombia Bolivia Brazil). to synonym myriocarpum Hook., a species H. considered be a of 216) to p. it and Tryon and known from Mexico Venezuela and Colombia to Bolivia, to On hand. myriocarpum. the other H. placed a variety of Stolze (1989. 63) as p. it FERN JOURNAL 28 synonym Smith thought be a of H. axiJIare Swartz, a species [1985, 127) to p. it known and only from Cuba, Hispaniola, Jamaica. myriocarpum, H. the believe that H. nigrescens a juvenile state of for is I and plants are always small, the pinnae congested, the fronds rarely fertile. In addition, the ranges of these two species overlap and juveniles of the H. common from Mexico Venezuela. nigrescens type are to fairly Although H. axillare very close to H. myriocarpum, prefer to maintain the is I more two with they as separate species until evidence indicates certainty that indeed HymenonphyJium myriocarpum does not overlap with H. are one. know and only one H. nigrescens-type juvenile specimen from the axiJiare, of I & West (Maxon The have found Indies Kiliip 1107, US). only differences are I and involucres mostly longer than wide, rachis alae relatively narrow, fronds narrowly rhombic and subdeterminate in H. axillare, versus involucres mostly wider than long, rachis alae wider (about 2 times as wide as the rachis), and fronds mostly lanceolate and determinate in H. myriocarpum. The LeCTOTYPE HyMENOPHYLLUM XRICHOMANOmES of As new typical of van den Bosch's H. trichomanoides Bosch (Ned. is taxa, v. d. Kruidk. Arch. was from 5(3):158. 1863) described several syntypes, including Cuming specimens of from Ecuador, Moritz from "Colombia" Venezuela], [i.e., Schomburgk from Venezuela and Guyana, and Spruce from Only Peru. the Spruce collection precisely localized and widely and is distributed, so best is it Monte Pampana Aue totype: near Taraooto. Deoto. Martin. Peru. S. 1856, Spruce 4696 (K not seen; isolectotypes GH, NY, L none seen, P not seen Morton photo Although 4620, US). have not seen the lectotype, presume I I it Kew, which present houses main is at the set of Richard Snruce's South American Qacq.) Swartz have been thought artz) Farwell protrusum (Hook.) Farw. believe the former two more species I are closely related to each other than either one H. polyanthos, segments is to for of the somewhat latter species are often folded or undulate and have involucres that mm Segments of the former two on species, the other that 1 mm known lyanthos from throughout is America, tropical and H. trichomanoides have more restricted ranges, known rens principally from Choco is the Colombia of icent northern with few Brazil, a from collections the Molina US) and all the mountains range for the Colombia LELLINGER: NEOTROPICAL HYMENOPHYLLACEAE 29 Rica Maxon Molina 529, Trichomanes and ekmanii kapplerianum T. Tryon and Stolze (1989, p. 88} considered T. ekmanii Wessels Boer be to a synonym kapplerianum Sturm many of examining T. in Mart. In specimens for the Flora of the Guianas found comments Project, their well taken with respect I the south American which seem to material, does be kapplerianum. all of to T. However, I do not believe the two species are synonyms, for one character does them distinguish America and in Central the Antilles: T. ekmanii has a usually continuous submarginal false vein that is as thick as the true veins that meet it, whereas kapplerianum somewhat T. has a usually discontinuous submarginal vein None false that is thinner than the true veins that meet of the distinctions it. mentioned by Wessels Boer nor by (1962, p, 317) Lellinger (1989, p. 208) serve two to distinguish the species consistently. Although Wessels Boer (1962, p. new 320) ascribed his species ekmanii Cuba, Hispaniola, Honduras, T. to British Honduras, Panama, Bolivia, Peru, Colombia, Venezuela, and northeastern Bra- zil, think likely to be absent from continental South America and Trinidad. I it The Identity of Trichomanes guianense Trichomanes guianense Sturm in Mart, a rare species described from is Guyana. most members Like Lacostea Bosch) of sect. (v. d. Christ, it is hemiepiphytic with the fronds adherent to tree trunks by means of '*rhizoids" and narrow on rhizomes and (actually hairs scales) the abaxial surface of the lamina axes. The involucres are long-stalked and oblique to the plane of the laminae, doubtless an adaptation in this section to facilitate spore dispersal from adherent laminae. It is certain that T. guianense is juvenile material of T. ankersii Parker ex & Hook. continuum from narrow, divided fronds Grev., for a exists the less of more The juvenile specimens to the wider, divided fronds of adults. smallest and most Appun, K) juvenile specimens of this species {Sagot 751, K, P; common, apparently are for they are represented in collections at least as frequently as adult specimens They have barely repand segments (Fig- are. 3). The laminae of T. guianense, on the other hand, are narrowly triangular to oblong and rather deeply pinnatifid with the segments slightly lobed, as in the specimens type of this species (Schomburgk 1215 B; isotype K); these are p. p., Guyana, Jenman from Potaro River subjuvenile (NY), the of (Fig. n. 4). s. and Adult ankersii Parker ex includes both juvenile subjuvenile material. T. Hook. & segment margins Rarely, Grev. has crenate or crenate-serrate (Fig. 5). adult specimens more deeply lobed are (Fig. 6). For specimens phases instance, Identification of in certain is difficult. pedicellatum elaborated adults of T. ankersii are similar to subjuveniles of T. more The have narrower fronds with attenuate (Fig. 6 vs. Fig. former longer, 8). segments with more lobes and shallower lobes, apices, often longer segments, and more veins distant. AMERICAN FERN JOURNAL: VOLUME NUMBER 30 81 1 (1991) 0"i^"«^ °* «"fce'-«"^ Fig. ° ^"f 3. Juvenile frond (Appun. Guyana, F r 4^, K ? °'t^'""'' K). ^°*"° ^^^-^- F ^*^^^' ^'YJ- Typical adult lldiAcZ'^S^m^n^h^^^^ 5. frond '- (A. C. 2824a, Guyana. NY). Elaborated aduh pinnae Uenman. Pomeroon Guyana River. LELLINGER: NEOTROPICAL HYMENOPHYLLACEAE 31 Lamina Variation Trichomanes in pedicellatum A study South American of the Trichomanes species pedicellatum and Desv. members allied species, also of sect. Lacostea, has revealed great diversity in frond form. Juvenile specimens merely [Fig. are pinnatifid with nearly 7) entire lobes. Subjuvenile specimens have pinnae (Fig. 8) v^ith shorter lobes and correspondingly wider, uncut median Such portions. specimens have been Trichomanes called commutatum suhsessile Sturm Splitg. or T. in Mart. specimens are divided]. Intergradation from most the juvenile adult forms unbroken, and to is may somewhat unt ^ „^^^ ^ ,,, , French Guiana. These strikingly condensed plants have been [Fig. 10] called T. furcatum v. d. Bosch. In addition, atypical adult plants with more divided, uncommon elaborated laminae (Fig. 11) are not in the Guianas and The Brazil. pinnae have and some the basal lobes suprabasal lobes pinnately divided, in addition to having typical simple and furcate lobes. The plants are robust and appear very These have been name full. called T. voiubile Vellozo, a sometimes who attributed to Antonio de Arrabida, edited Vellozo's "Flora Fluminensis." Trichomanes radicans and collariatum T. Trichomanes radicans Swartz based on a type from Jamaica, but widely is is distributed throughout tropical America. Trichomanes coJJariatum Bosch v. d. is an allied species with a narrower range, Mexico to Venezuela and Peru. Specimens from outside this range identified as this species are likely to be T. radicans. Trichomanes collariatum notable widely spreading for is its involucral which seem form wide apex labia, to a collar at the of the involucre. In Mesoamerica, T. radicans said to differ in having involucral labia is rudimentary However, and South America, or lacking. in the Antilles in the labia of T. radicans are well developed. The two species can be distinguished by the following key: cm 3-4 10-20 Involucres times longer than wide; mature laminae wide, 1. usually widest near the base, the pinnae acute to acuminate at the apex; stipes cm 3-12 long radicans ......T. Involucres 2-3 times longer than wide; mature laminae mostly no more than 1. cm round 10 wide, usually widest near the middle, the piimae mostly nearly cm 1-6 or nearly obtuse (acute pinnae also seen]; stipes long... collariatum T. , , may Mesoamerican Because the material of the difference in involucral labia, known mexicanum ^ be Trichomanes Bosch) radicans Lellingert'^-cjS^f as var. (v. d. 3^7 which based on mexicanum Bosch, Ned. Kruidk. Arch. 5(2):164. 1861, ^ T. v, d. L based on twcKsyntypes from Mexico, Schiede 806 (B not seen fragm not is K seen); and Scha;^ner 7 (P or RB not seen fragm L; probable isosyntype not seen Morton photo 19052). FERN JOURNAL 32 ® omanes t-21B8, Guyana, K). Fig. Subjuvenile 8, "T. subsessile" pinnae [Sagot 742, French Guiana Typical pmnae adult Nov 9. (Leprieur 229 in 1837, French Guiana, P). Fig. 10 Condensed e Pi"^^« (i^P"eur 229 Dec ,}T" in 1830. French Guiana, P). Fig. 11. Elaborated a volubile pinnae (Granville al et 6180, French Guiana, NY). New lU \H jenmaii nov.— Lellinger, sp. i Fig. 12. Rhizoma mm diametro, basibus stipitum mm LELLINGER: NEOTROPICAL HYMENOPHYLLACEAE 33 T * ' " ' t I ? '^ t I I Inches I tfl *•.. f ,N -f ^ £*.^i /i *.t-t i% ;^n /i* ', ^ . 1-^4, / -;r-7- . ^.' . . . <' '.-4 «. / * ^ . # tf Fig. 12. Holotype of Itichomanes jenmanij, Jenmaii (NY).

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