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Genus Sm. Revision of the Salpichlaena J. (Blechnaceae, Pteridophyta) 1 Gabriela E. Giudice Maria L. Luna*, Cristian Carrion, and ElIas R. dela Sota* , Catedra de Morfologia Vegetal. Facultad de Ciencias Naturales y Museo, Universidad Nacional d La Plata, Paseo del Bosque s/n 1900 La Plata, Argentina , South America. The number of species recognized genus varies ,,m loi this ,|, i authors from one to four. The goal of this work was to provide a systematic revision of the genus Salpichlaena in order to contribute to the knowledge of the fern biodiversity in America. For this purpose morphological, anatomical and palynological characters were analyzed in material from the geographical distribution of Salpichlaena. Herbarium spn imens were Invited according to the LM SEM and The specimens and standard techniques for studies. type the original descriptions Two were consulted to determine the applications of names. species are recognized, S. volubilis (Kaulf.) Sm. and S. hookeriana (Kuntze) Alston. Salpichlaena hookeriana differs from S. }. much volubilis by pronounced foliar dimorphism (the fertile pinnules are reduced), the presence and of foliar buds on sterile basal pinnules, ovate costular scales the presence of glandular hairs on the abaxial surface of the costa. The spores are monolete in both taxa, with rugulate- granulate perispore and superficial spherules. Salpichlaena volubilis widely distributed in is Central and South America, from Guatemala and Caribbean Islands, up to southern Brazil and Bolivia, across a wide altitudinal range of 200 to 1900 Colombia. Venezuelan Guyana, Suriname, British Guia altitudes up to 800 m. The diagnostic characters, illu species are given. The an American endemic and genus Salpichlaena (Blechnaceae) differs is which from other representatives of the family by high climbing leaves, has a its and otLygodium (Tryon Tryon, counterpart only in the leaves (Schizaceae) 1982). According these authors the leaves of Salpichlaena are scandent to at least to m 15 into the top of trees by means of the twining rachis. Other distinguishing genus include: the presence of a long-creeping stem, the leaves features of this monomorphic dimorphic pinnae with narrower segments than the or (the fertile commissure and and on long vascular parallel close to the sterile pinnae), sori a covered by an indusium that envelops the sporangia. Salpichlaena costa that are chromosome most closely related to Blechnum, but differs in its twining habit, is number and with laminae bipinnate (Smith, 1995). fronds The number of taxa included in the genus Salpichlaena varies according to different authors. Until the present three species have been recognized: S. [email protected] Corresponding author, email: de Provincia de Buenos *Comision de L la fi is Tecnicas, Argentir y AMERICAN FERN VOLUME NUMBER JOURNAL: 50 98 2 (2008) hookeriana (Kuntze) Alston, thalassica Grayum & R.C. Moran and S. S. volubilis (Kaulf.) Hook. (Moran, 1995; Jorgensen and Leon, 1999; Prado, 2005). The species thalassica was described by Moran on S. (1990) the basis of a type from Costa Rica, characterized by the presence of longer stalks of the basal when fertile pinnules compared with those of S. volubilis, and by the higher which altitude at S. thalassica grows. Conversely, Smith (1990) proposed the combination S. lomarioidea (Baker) A.R. Smith (= Blechnum voluhile Kaulf. var. lomarioidea Baker) which was distributed from Venezuelan Guayana to synonymized Peru. Later the author lomarioidea hookeriana (Smith to S. S. The two 1995). existence of geotypes of volubilis in Venezuela: euvolubilis S. and maegdefrauiana was established by Vareschi According him, (1969). to monomorphic the former has fronds and distributed in the Coastal is Cordillera (900-1500 m) whereas the has dimorphic fronds and occurs latter Amazonas in the state of (100 m). Other studies focused on cytological and palynological aspects of Salpi- chlaena. Walker determined chromosome number X=40 (1973) a for this genus, the highest recorded Blechnaceae. The in structure of the spore wall of was described by Tryon and Lugardon volubilis S. (1991) as papillate-rugulate, with scattered spherules on the surface and in section with a plain exospore and a laminate perispore. They considered the spore wall be characteristics to useful in distinguishing Salpichlaena from other Blechnaceae. A comprehensive work including the analyses of morphological, anatomical and palynological aspects of Salpichlaena across range Thus lacking. the its is aim of study provide this is to a systematic revision of Salpichlaena in order to knowledge contribute to the of the fern biodiversity in the Americas. Materials and Methods Nearly 500 specimens from GH, LP, NY, and SP were analyzed F, P, SI during this study. Type specimens and the were original descriptions consulted to determine application of names. Vegetative well reproduc- as as were tive characters analyzed and included: rhizomes, fronds, rachis, petiolules, pinnules, indument, indusia and spores. For microscopy light observations (LM) the herbarium material was rehydrated with soft cold water-detergent then dehydrated through an ethanol and embedded series in Paraplast. Sections 8-12 urn thick were stained with was safranin-fast green. Material also cut employing a cryotome 20 urn at and thickness with stained safranin. For scanning electron microscope (SEM) was analysis of the spores, material 3% with sodium treated hot carbonate two minutes for preserve the to perispore, then cleaned with water and distilled transferred ethanol to (Morbelli, 1980). Specimens were placed on acetate plates and sputter coated SEM with made gold-palladium. Observations were using JEOL, JSM-35 CF, a of "Servicio de Microscopia Electronica de Barrido", Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata. GIUDICE ET REVISION OF THE GENUS SALPICHLAENA AL.: hookeriana S. A m 1800 A A A thalassica S. (cfr. 1600 A Moran) " " * A *X 1400 A ? ±A a S. volubilis * 1200 | 1000 ^i A g 1 800 A * i g 600 A *i* t A H A 400 AtM *A 200 l^t A i 4* : 0.5 1.5 2 2.5 3 3 5 1 Petiolules length of basal fertile pin nules (cm) and the bibliographic Results / Moran Analyzing specimens from Costa Rica and Panama, (1990) described new on Salpichlaena thalassica as a species the basis of the correlation and which between pinnule the stalk length of the basal fertile the elevation at we the plant grew. In the present work, expanded the scope of the study and found a complete overlap of these features with those of S. volubilis (Fig. 1). we For reason do not consider the ratio of altitude to stalk length a this from same diagnostic character to distinguish S. thalassica S. volubilis. In the work, Moran mentioned thalassica has blue-green colored (1990) that S. However, pinnules whereas possesses dark-green ones. Murillo volubilis S. (2001) stated that the color of the fronds varied in Salpichlaena based on the As which our area in grows, thus rejecting this characteristic as diagnostic. it we make study was based on herbarium material were not able to definitive color observations. We buds and recognize the presence of foliar the reduction of the fertile pinnules in Salpichlaena hookeriana as diagnostic characters to differentiate which buds and has not this species from volubilis, foliar their fertile S. pinnules wider than the pinnules. These characters were also are sterile we observed by Alston (1932) and Smith (1995). Based on our observations, are who considered in disagreement with Murillo (2001) these features as variations in the phenotypic expression due to changes in the environmental According our observations the "geotypes" proposed by conditions. to AMERICAN FERN JOURNAL: VOLUME NUMBER 98 2 (2008) n u-/c\ - attenuated apex and dentate-glandular margin. C. Portion of scale base that shows the foot 1 D-H: (arrow). fertile pinnule. D. Outline of transverse section showing extremely reduced i Margin inae. E. in detail with sporangia and indusium. F. Transverse section with a marked G a. and H. Details of the abaxial surfaces that show glandular, 2-3 cellular hairs. I-K: SEM GIUDICE ET REVISION OF THE GENUS SALPICHLAENA AL.: 53 dimorphism Vareschi concerning correspond (1969) the foliar to the species Salpichlaena volubilis {"euvolubilis") and hookeriana (" maegdefrauiana"). S. Salphichlaena hookeriana also differs from S. volubis by the presence of which glandular hairs on the pinnules, are absent in S. volubilis. Additionally, morphology between two the of the costular scales differs the species. In S. hookeriana the scales are ovate, with attenuate apices and dentate-glandular whereas with margins, in volubilis they are lanceolate, long attenuated S. and apices ciliate-glandular margin. All analyzed material showed two linear coenosori on the comisural veins by open The parallel to the costa, protected elongate indusia to the costa. The number indusia were 1-4-cellular layers thick. of layers decreased toward the distal portion of the indusia. The cells of the outermost layer had unevenly and thickened walls take part in the indusial fragmentation the maturity. at (Fig. E, F; Fig. E, G). 2, 3, We among consider spore characteristics insufficient to differentiate species within this genus. However, they are useful in distinguishing Salpichlaena from other Blechnaceae genera because species of Salpichlaena have a poorly and developed perispore (apparently one-layered in section) a granulate- whereas Blechnaceae have complex papillate-rugulated surface, other a perispore that psilate and generally folded. The spherical bodies observed is composed Tryon on the spore surface are "spherules" of perispore material [cf. and Lugardon, 1991, p. 9). two According our observations taxa of Salpichlaena are distinguishable to morphology based on dimorphism, the of the costular scales, the foliar presence/absence of glandular hairs on the pinnules and the presence/absence of foliar buds on the sterile basal pinnules. Systematic treatment Sm. Hook. Gen. T.93.1841 Salpichlaena in Fil. J. Salpichlaena volubilis (Kaulf.) Sm., in Hook., Gen. Fil., t.93.1841 J. Enum. 1824 Blechnum 159. volubile Kaulf., Fil. Rhizomes long-creeping and scaly, scales dark-brown to blackish, basipel- m, pinnae dimorphic, Fronds 15 the margins bi-pinnate, to ca. tate, entire. m Rachis climbing pinnules narrower than the sterile ones. to 15 in fertile below abraded perispore K. Surface details with granulate- jxospore (E) (P). G ;culpture and spherules (arrow). Scale bars: A: 3 cm. B, E and F: 200 urn. C, a f=wr — M: J Salpichlaena LM Fig. 3. volubilis. A. Rep resentative herbarium sample. B-D. micrographs of largely attenuated apex and ciliate-glandular margin. C. Scale base and margins in detail. D. Sea e margins that show a glandular hair. E-G: fertile pinnulae in transverse section. pinnu E. Detail of ae at the costa. F. Outline showing laminae expansion. G. Detail of multilayered indusium, SEM the o itermost layer shows cells with thicker walls. H-K. GIUDICE ET REVISION OF THE GENUS AL.: SALPIi micrographs of spores. H. Monolete spore in equatorial view. Spore in proximal view. and K. I. J show and Spore surface details that a granulate- papillae- rugulate sculpture spherules (arrow). K. A smooth exospore (E) below abraded perispore (P) is evident. Scale bars: A: 3 cm. B and C: 100 urn. D, E and G: 0.5 cm. H: 10 urn, I: 20 urn. and K: 5 fim. J AMERICAN FERN VOLUME NUMBER JOURNAL: 56 98 2 (2008) trees, twining with occasional scales similar to those of the lamina. Pinnules linear-elliptic to linear-lanceolate, bases asymmetric, cordate to roundish, apex Lamina acuminate to attenuate. Veins simple or proximally once forked. scales on brown present the costa abaxially, light to yellowish, deciduous, basipeltate; glandular, 2-3-celled hairs, present along the costa abaxially. buds Foliar absent or present the bases of basal pinnules of tropophylls. at Cenosori linear, positioned on the comisural veins parallel to the costa, covered by elongate indusia open to the costa; paraphyses absent. Spores view and monolete, plane-concave in equatorial elliptic in polar view; and equatorial diameter 67.5 97.2 |im polar diameter 48.6 (78.9) (55.43) 25-43 2- 70.2 urn; laesura urn long, tenuimarginate, scarcely evident; exospore um 3.5 |im thick, to 5.5 along the proximal face, smooth, apparently two- layered, inner: outer exospore ratio, perispore 1-1.4 jim thick, apparently 1:2; one- layered in section, easily detached from the exospore, granulate-papillae- and with rugulate sculpture superficial spherules isolated or grouped. Key to the of Salpichlaena species n wide; buds foliar in axils of sterile basal costal scales ovate, with attenuate apices ; volubilis hookeriana S. (Fig. 2) Blechnum volubile \ Salpichlaena lomi 1990 250. Rhizomes long- creeping, scales dark-brown margins to blackish, basipeltate, m Fronds entire. bi-pinnate, Fronds ca.15 dimorphic, pinnules fertile cm extremely reduced, 14-40 cm long, 0.2-1.5 wide. buds on Foliar sterile basal pinnules; hairs along abaxial surface of the glandular 2-3 costa, celled; scales along costa brown abaxially, apex light yellowish, to basipeltate, ovate, margin attenuate, dentate-glandular. Coenosori paired on comisural veins the parallel to the costa, covered by elongated indusia open Spores to the costa. monolete, granulate-papillae-rugulate sculpture. REVISION OF THE GENUS SALPICHLAENA : (Fig. 4). Note: Salpichlaena hookeriana is a combination proposed by Alston in 1932, taking into account the species Spicanta hookeriana basionym. as a Spicanta hookeriana was cited for the first time in 1891 by Kuntze as sinonymus Lomaria The specimen of volubilis. type of Spicanta hookeriana we was Thus never established. consider that the type of Salpichlaena hookeriana correspond to the type specimen of Lomaria volubilis Isotype: : Brasil, Barra by Igarape dos Manaos, Spruce 1263 (P 347482), Feb. 1857. volubilis S. (Fig. 3) Enum Blechnum Salpichlaena voluhile Kaulf., 159. 1824. volubilis Filic. TYPE.— Hook., Gen. 1841. BRAZIL, Chamisso (Kaulf.) J.Sm. in 93. Fil. s.n. t. LZ— (HT, destroyed). Blechnum scandens Bory, Dup. Voy. Bot. 272 36. 1828. Salpichlaena 1. t. DU TYPE.— CATHERINE scandens (Bory) C. Presl, Epim. bot.122. 1849. ST. BRASIL, 1827 P Durville s/n, (Coquille n° 92, 347471!). Grayum & Salpichlaena thalassica R.C. Moran, Ann. Missouri Bot. Gard. COSTA TYPE.— 77:591. 1990. RICA: Heredia: forest between Rio Peje and Sardinalito, Atlantic slope of Volcan Barba, 10° 17' N, 84° 4.5'W. 800- & 1000 m, Grayun Chazdon 6833 (MO). Rhizomes dark-brown margins long-creeping, scales to blackish, basipeltate, 11- entire. Fronds bi-pinnate, ca. 15 m. Fronds dimorphic, fertile pinnules cm cm buds and 35 long, 0.5-3 wide. Foliar glandular hairs absent, scales margin along costa abaxially, lanceolate, largely attenuated apex, ciliate- glandular. Coenosori paired on the comisural veins parallel to the costa, open Spores monolete with covered by elongated indusia to the costa. sculpture. granulate-papillae-rugulate Salpichlaena volubilis widely distributed in Central and South America, is and from Guatemala and the Caribbean Islands, to southern Brazil Bolivia, m m wide 200 1900 across a elevation range, to (Fig. 4). Salpichla m Examined.—VENEZUELA: Depto Amazonas, Alto Rio Orinoco, 125 Specimens & Wurdack 41477 Cerro Paraque Sipapo, Maguire m., Maguire, Keith (F); 8> s. km 28776 Depto Atures, 9 por arriba del Raudal Remo. 04 34' N, 67 Politi (F); & GUIANA: BRITISH 18' W. 120 m. Foldats Velazco 9542 (NY). Region Potaro- Pakaraima Mutchnick, Henkel &- Williams 240 (NY), Tumatu- Siparuni, Mts., COLOMBIA: mari, Potaro River, 5° 20' N. A.S. Hitchcock 17374 (NY). Depto AMERICAN FERN VOLUME NUMBER 58 JOURNAL: 98 2 (2008) Amazonas, & Vaupes, Rio Apaporis, 800 feet m, 0° 5' S, 70° 30' W, Schultes Cabrera 15410 Vaupes, W, & (NY), Rio Piraparana. 0°15'S, 70 30' Schultes Cabrera 17449 (GH); Depto Vichada, alrededores de "Gaviotas", Cano Urimaca. 130-160 m, Murillo 1582 (P). PERU: Depto Loreto, Rio Nanay, Shiriara, NW Plowman km 2550 (GH, BOLIVIA: Depto Beni, Guayaramerin, F). 5 of Anderson 11824 (F). BRAZIL: Edo Amazonas, Rio Xie. 0° 55' N, 67° 15' W, & Stevenson, Daly Guede 815 Municipio km (NY); de Humaita, Velho Porto 60, W, Mc & 8 S, 63 Teixeira, Farland, Mota, dos Santos, Gomes Nelson 208 Fife, Manaus, (NY); Reserva Forestal Ducke, 02 53' 59 58' W, Assungao 385 26. S, A San (NY), Antonio, de 2458 la Sota (LIL); Reserva Forestal Ducke, Manaus- An Itacoatiara, 02 53'S, 59 58'W, Ribeiro 1025 Alto et (SP), al. Schwacke Herb. 4110 (P 347479), Igarape, Drake 1263 (P 347482). Salpichlaena volubilis HONDURAS: Specimens Examined.— Depto Guan Yoro, Rio Guan. 300-380 m. & 15 30' 00" N, 87° 27' 20" W, Hawkins Merello 820 NICARAGUA: (NY). Depto Zelaya, Bluefields, Nichols 893 (GH); Cano Costa Riquita. 11 43' N, 84 W. 18' 150-180 m, & COSTA km Stevens 4959 Krukoff RICA: (LP). Pcia Alajuela, 20 NW of San Ramon, 10 13' N, 84 32' W, 850 msm, Smith, Grayum, Beliz, & Dickie Carvajal 2239 (NY); San Ramon, 1000 m, Mickel 2927 Idem, de (LP); Sota 5155 la (LP); Pcia Cartago, Canton de Paraiso, Valle del Reventazon, 09 3 44' 40" N, 83° 50' 00" W, 1700-2000 m. Rojas, Jimenez Aguilar 2462 Br (NY); m Mountains 5 miles S of Cartago, 1800 m. Maxon 512 (NY); 9 42' N, 83 47' s. & W, 1500 m, Burger Liesner 6807 (NY); Refugio Nacional de Fauna Silvestre & Tapanti, 1500-1620 m, Almeda, Anderson Zamora 5736 (NY); Valle of Rio Grande & del Orosi, Tryon Tryon 7028 (GH); Pcia Heredia: Finca La Selva, Grayum 1822 E de (NY); Puerto Viejo. R.K. Godfrey 67300 (GH); Pcia Limon, Lomas de 22 Sierpe, 10 83° 31'W, Robles 2198 'N, (NY); Pcia Puntarenas, San m 1500 de 5217 Vito. Sota San la Pcia (LP); Jose, Vicinity of El General. 915 m. A.F. Skutch 2166 (GH); Los Angeles de Siquirres. 1000 m. Gomez, Liesner & PANAMA: Judziewicz 20541 (LP). Pcia Chiriqui, 8° 43' N, 82 14' W. 1300 m., Hampshire Er Whitefoord 370 (NY); Pcia Code, 600 m. 8 45'N, 80 30' W, m Hamilton SW Davidse 2806 &. (LP); Pcia Colon, 10 10- of Puertobelo. 200 m. Liesner 1106 Panama, Pcia Campana, Cerro 14755 (P); Croat (NY); ( Idem, 900 m, Madison 768 (GH); Pcia Veraguas, de Distrito Montijo, Coiba, Isla Playa Hermosa, Arduz GUADALUPE: 455 (NY). et al. Base-Terre, foret de Choisy, Barrier 2356 Moscou (NY); district, south of La Citerne, 650-660 m, Proctor 20130 (GH); Hugues trace Victor au depart de Montebello, 2885 Barrier (P); Foret de Sofaia, Rodriguez 4257 DOMINICAN REPUBLIC: Concorde (P). Valley, moist forests bordering Pegoua River Deux in vicinity of Branches, Hodge 3482 (GH); Rainforest bordering Imperial Road, Sylvania Mahaut to Hodge River, 459 m., 98 MARTINICA: (GH). Joseph, 450-660 m, Duss 1901 St. SANTA LUCIA (NY). Barre de 800-1400 Ridge, L' Isle Webster, feet, Ellis fr : Miller 9285 (GH); Forest between Quilesse and head Muray of Road, Hill