Marchantiophyta, Lejeuneaceae

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is characterized by asymmetrically ovate-falcate leaf lobes, ventral lobe margin .... diameter, in cross section with 7–11 cortical cells and 5–11 medullary cells, ...
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Nova Hedwigia published online November 2017

PrePub Article

A new and interesting species of Lejeunea Lib. (Marchantiophyta, Lejeuneaceae) from Brazil Cid José Passos Bastos1*, S. Robbert Gradstein2, Silvana B. Vilas Bôas-Bastos1 and Alfons Schäfer-Verwimp3 1 Universidade Federal da Bahia, Instituto de Biologia, Laboratório de Taxonomia de Briófitas – BrioFLORA, Campus Universitário de Ondina, 40170-280 Salvador, Bahia, Brazil

Muséum National d’Histoire Naturelle, Cryptogamie, C.P. 39, 57 rue Cuvier, 75231 Paris cedex 05, France

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Mittlere Letten 11, 88634 Herdwangen-Schönach, Germany

With 2 figures

Abstract: The new species Lejeunea longidentata from Brazil is described and illustrated. This species is characterized by asymmetrically ovate-falcate leaf lobes, ventral lobe margin often upcurved, lobe apex often recurved and by the narrowly elongate, curved, banana-shaped lobule tooth-cell. The specimen was found in Araucaria forest in Minas Gerais growing on bark of living trees. Key words: Leafy hepatics, taxonomy, tropical America.

Introduction Lejeunea Lib. is a large pantropical genus, whose taxonomy is quite complex due to the large number of species, the often scanty or heterogeneous type material of the older names, and the high morphological variability of the gametophytes (Reiner-Drehwald 2010a). Thus, many names, especially the older ones or those only known from type material, need to be confirmed. Söderström et al. (2016) listed 407 species names, of which 96 are good names accepted in taxonomic reviews and monographs; most (246), however, are poorly known names. Costa & Peralta (2015) reported 40 species in Brazil, but the Brazilian species in the genus have not yet been revised. In a study of the Brazilian species of Lejeunea in the framework of the "Flora do Brasil 2020" project, Bastos (in prep.) so far recognized 50 species for Brazil, and Gradstein (in

*Corresponding author: [email protected]; [email protected] © 2017 J. Cramer in Gebr. Borntraeger Verlagsbuchhandlung, Stuttgart, Germany. DOI: 10.1127/nova_hedwigia/2017/0455

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www.borntraeger-cramer.de 0029-5035/2017/0455 $ 1.50

prep.) accepted 59 species in his unpublished manual of the liverworts of Colombia and Ecuador. The most recently published taxonomic treatment of the genus Lejeunea was by Lee (2013) for Malaysia, in which 30 species were recognized. An overview of numbers of Lejeunea species recorded from Asian countries was given in Lee et al. (2014). Our current understanding of the species in tropical America is largely based on the important work of Dr. Elena Reiner-Drehwald, for example treatments of the species of Misiones, Argentina (Reiner-Drehwald 2000) and French Guiana (Reiner-Drehwald 2009a), taxonomic revisions of the species formely placed in Crossotolejeunea (Spruce) Schiffn. (Reiner-Drehwald & Goda 2000), Amphilejeunea R.M.Schust. and Cryptogynolejeunea R.M.Schust. (Reiner-Drehwald 2005a), and Inflatolejeunea S.W.Arnell (Reiner-Drehwald & Schäfer-Verwimp 2008), and the studies on Lejeunea adpressa Nees (Reiner-Drehwald 2009b), L. subg. Nanolejeunea R.M.Schust. (Reiner-Drehwald 2010b), L. rotundifolia Mitt. (Reiner-Drehwald 2005b), L. pulverulenta (Gottsche ex Steph.) M.E.Reiner (Reiner-Drehwald 2005c), and L. deplanata Nees (Reiner-Drehwald 2010a). For Brazil, two new species have been described in recent years: Lejeunea perpapillosa M.E.Reiner & K.C.Pôrto from Pernambuco and Bahia (Reiner-Drehwald & Pôrto 2007) and L. combuensis O.S.Moura, Ilk.-Borg. & M.E.Reiner from Pará (Moura et al. 2012). The genus Lejeunea may be recognized by a set of morphological characters, including: (a) hyaline papilla at the proximal side of the first tooth of the lobule (= apical tooth); (b) ocelli absent; (c) small, granular or homogeneous oil bodies; (d) Lejeunea-type branches; (e) gynoecia with lejeuneoid innovations. Considerable variation is seen in the leaf lobule which can be well-developed (ovate to rectangular) or reduced, the apical tooth which can be unicellular (usually) or pluricellular, the underleaves which can be bifid or entire and highly variable in shape and size, and the perianth which may be keeled (usually 5-keeled) or terete, without keels; when keeled, the perianth keels may be smooth, crenate or variously ornamented with teeth, cilia and/or lacinia. During the study of the Brazilian species of Lejeunea by the first author, an apparently undescribed species from the State of Minas Gerais was found. The new species is here described and illustrated. Description and discussion Lejeunea longidentata C.J.Bastos, Gradst., S.Vilas Bôas-Bastos & Schäf.-Verw., sp. nov. Figs 1–2 Diagnosis: This new species is characterized by: 1) leaf lobe asymmetrical, ovatefalcate, ventral margin often upcurved, apex mostly recurved; 2) leaf lobule welldeveloped, ovate, fully inflated, lateral free margin incurved, 6 cells long, distal free margin cells with oblique walls, lobule tooth cell narrowly elongate, curved, bananashaped; 3) underleaves subimbricate, broadly ovate, sinus V to broadly U-shaped, bases rounded, insertion line curved, a large cell present at either side of the underleaf base adjacent to the stem. Type: Brazil, Minas Gerais: Serra da Mantiqueira, Camanducaia, Monte Verde, Araukarienwald am Caminho do Grande Pinheiro, epiphytisch, 1550 m alt., 10 August 1986, Schäfer-Verwimp &

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Fig. 1. Lejeunea longidentata. A. Part of plant with gynoecium, ventral view. B. Underleaf. C. Leaf lobe, dorsal view. D. Upper part of plant with gynoecium and two innovations, ventral view. E. Female bract. F. Perianth, ventral view. G–I. Cross sections of stem. J. Central leaf cells. All from the holotype.

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Fig. 2. Leaf lobule apices of Lejeunea longidentata. A–E. Leaf lobule apices, showing the long apical tooth. In A hyaline papilla indicated. F. Inner leaf lobule surface. All from the holotype.

Verwimp 7532 (holotype: ALCB; isotype: JE). Paratype: Brazil, Minas Gerais, Serra da Mantiqueira, Piranguçu bei Campos do Jordão, Bachtal bei Centro Vale Verde, epiphytisch in lichtem, feuchtem Wald, 1450 m, 19 April 1986, Schäfer-Verwimp & Verwimp 6893/A (ALCB; GOET; JE).

planTs 850–975 µm wide, vegetative branches Lejeunea-type. Stem 100–110 µm in diameter, in cross section with 7–11 cortical cells and 5–11 medullary cells, thin-walled; cortical cells 23–30 µm long × 13–15 µm wide, medullary cells 15–20 µm long × 10– 15 µm wide; ventral merophyte two cells wide. leaves with a long J-shaped insertion, widely spreading; leaf lobe asymmetrically ovate-falcate, 400–510 µm long × 280– 400 µm wide, dorsal margin strongly arched, ventral margin straight, often upcurved, apex obtuse to subacute, plane or recurved, margin crenate; leaf lobe cells with small, simple-triangular trigones, intermediate thickenings scarce, cuticle smooth; ocelli lacking; lobules well-developed, ovate, 130–160 µm long × 80–90 µm wide, 1/4–1/3 × leaf length, fully inflated, lateral free margin incurved, 6 cells long, distal free margin cells with oblique walls, lobule apex with a long (ca. 40 µm) slightly falcate tooth, the tooth one-celled, ca. 2.5 × longer than wide, banana-shaped, tip rounded, hyaline 4 eschweizerbart_xxx

papilla proximal-marginal, in a notch at proximal side of the tooth, apical free margin concave, terminating at the tip of the keel, not continuing along the ventral margin of the lobe, margin cell distal to the tooth not larger than inner cells, orifice narrowly rounded; keel curved, crenulate (often conspicuously so), forming a narrow or wide angle with the leaf lobe. UnDerleaves subimbricate, broadly ovate, wider than long, 200–300 µm long × 270–400 µm wide, 3–4 × stem width, bifid to 1/3, lobes upright, broadly triangular, apices rounded to subacute, sinus V-shaped to broadly U-shaped, margins crenulate, outer margins rounded, without tooth, bases rounded, insertion line curved, a large cell present at either side of the underleaf base adjacent to the stem; underleaf cells as in the lobe. anDroecia not seen. gynoecia with 1–2 lejeuneoid innovations, bracts and bracteole upright, somewhat undulate, margins crenulate, bract lobe ovate-triangular, 600 µm long × 360 µm wide, apex acute, bract lobule oblong, extended beyond the keel, 250 µm long × 90 µm wide, 1/2–2/3 × lobe length, apex rounded; bracteole widely ovate, bifid, slightly shorter than bract, 430 µm long × 400 µm wide, lobe apex mostly recurved. Perianth obovate, 680 µm long × 470 µm wide, slightly exserted, with 5 smooth keels and a ca. 20–50 µm long beak. DisTribUTion anD habiTaT: Lejeunea longidentata is so far known only from two collections from elevations between 1450 and 1550 m. The type specimen was collected in Araucaria forest, growing on a living tree trunk, mixed with Radula fendleri Gottsche ex Steph. and Lejeunea cancellata Nees & Mont. As pointed out by Fiaschi & Pirani (2009) the Araucaria forest is a subtropical forest included in the Atlantic Forest Domain, which extends far inland in some areas of southeastern Brazil, eastern Paraguay, and Misiones Province of Argentina. The paratype collection is from Atlantic Forest not very far from the type locality, growing on bark of living tree and mixed with Lejeunea oligoclada Spruce, another Southeast Brazilian endemic. DiscUssion: The asymmetrically ovate-falcate leaf lobes with the recurved apex and the narrowly elongate, curved, banana-shaped lobule tooth-cell are the most distinctive features of the new species. The wider than long underleaves (3–4× stem width) with a large cell at the base, at either end of the insertion, resemble those of L. puiggariana Steph. (Reiner-Drehwald 2000), however, the leaves in the latter species have plane margins and the lobule tooth is much shorter. Acknowledgements We thank Dr. M.E.Reiner-Drehwald, Germany, for carefully reviewing the manuscript and valuable suggestions. The first author is grateful to Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the grant of the Research Productivity Fellowship.

References COSTA, D.P. & D.F. PERALTA 2015: Bryophytes diversity in Brazil. – Rodriguésia 66: 1063–1071. FIASCHI, P. & J.R. PIRANI 2009: Review of plant biogeographic studies in Brazil. – J. Syst. Evol. 47: 477–496. LEE, G.E. 2013: A systematic revision of the genus Lejeunea Lib. (Marchantiophyta: Lejeuneaceae) in Malaysia. – Cryptog. Bryol. 34: 381–484.

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LEE, G.E., T. PÓCS, S. CHANTANAORRAPINT, A. DAMANHURI & A. LATIFF 2014: An account of the genus Lejeunea (Marchantiophyta: Lejeuneaceae) in Thailand, including seven newly recorded species. – Cryptog. Bryol. 35: 247–259. MOURA, O.S., A.L. ILKIU-BORGES, & M.E. REINER-DREHWALD 2012: A new species of Lejeunea Lib. (Lejeuneaceae) from Low Várzea forest in lower Amazon (Pará, Brazil). – Nova Hedwigia 95: 197–202. REINER-DREHWALD, M.E. 2000: Las Lejeuneaceae (Hepaticae) de Misiones, Argentina. VI. Lejeunea y Taxilejeunea. Trop. Bryol. 19: 81–131. REINER-DREHWALD, M.E. 2005a: On Amphilejeunea and Cryptogynolejeunea, two small genera of Lejeuneaceae (Jungermanniopsida), and two common neotropical Lejeunea species. – Nova Hedwigia 81: 395–411. REINER-DREHWALD, M.E. 2005b: On Lejeunea rotundifolia and Dicladolejeunea (Lejeuneaceae, Jungermanniopsida). – Systematic Botany 30: 687–692. REINER-DREHWALD, M.E. 2005c: Taxilejeunea pulverulenta (Lejeuneaceae, Jungermanniopsida), a poorly known species from Neotropics, is transferred to Lejeunea. – Cryptog., Bryol. 26: 59–65. REINER-DREHWALD, M.E. 2009a: Lejeunea. – In: GRADSTEIN, S.R. & A.L. ILKIU-BORGES: Guide to the Plants of Central French Guiana IV. – Mem. N.Y. Bot. Garden 76: 95–101. REINER-DREHWALD, M.E. 2009b: Lejeunea adpressa Nees (Lejeuneaceae), a widely distributed species of tropical America. – Cryptog. Bryol. 30: 329–336. REINER-DREHWALD, M.E. 2010a: A taxonomic revision of Lejeunea deplanata (Lejeuneaceae, Marchantiophyta) from tropical America. – Nova Hedwigia 91: 519–532. REINER-DREHWALD, M.E. 2010b: On Lejeunea subgenus Nanolejeunea (Lejeuneaceae, Jungermanniopsida). – Nova Hedwigia 138: 117–128. REINER-DREHWALD, M.E. & A. GODA 2000: Revision of the genus Crossotolejeunea (Lejeuneaceae, Hepaticae). – J. Hattori Bot. Lab. 89: 1–54. REINER-DREHWALD, M.E. & K.C. PÔRTO 2007: Lejeunea perpapillosa (Lejeuneaceae), a new species from north-eastern Brazil. – Nova Hedwigia 85: 541–546. REINER-DREHWALD, M.E. & A. SCHÄFER-VERWIMP 2008: On Inflatolejeunea, Lejeunea species with eplicate perianths and Lejeunea talamancensis sp. nov. from Costa Rica (Lejeuneaceae). – Nova Hedwigia 87: 387–420. SÖDERSTRÖM, L., A. HAGBORG, M. VON KONRAT, S. BARTHOLOMEW-BEGAN, D. BELL et al. 2016: World checklist of hornworts and liverworts. – PhytoKeys 59: 1–828.

Manuscript submitted July 28, 2017; accepted October 13, 2017.

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