Bryopteris filicina (Sw.) Nees

  • Authority

    Gradstein, S. Robbert. 1994. Lejeuneaceae: Ptychantheae, Brachiolejeuneae. Fl. Neotrop. Monogr. 62: 216. (Published by NYBG Press)

  • Family

    Lejeuneaceae

  • Scientific Name

    Bryopteris filicina (Sw.) Nees

  • Type

    Type. Jamaica, Swartz s.n. (holotype, S).

  • Synonyms

    Jungermannia trinitensis (Lindenb.) Lehm., Bryopteris filicina var. arguta Gottsche, Lindenb. & Nees, Bryopteris tenuicaulis T.Taylor, Bryopteris fruticulosa T.Taylor, Bryopteris liebmanniana Lindenb. & Gottsche, Bryopteris trinitensis (Lehm. & Lindenb.) Lehm. & Lindenb., Bryopteris trinitensis var. intermedia Gottsche, Lindenb. & Nees, Bryopteris galapagana Gottsche, Bryopteris flaccida Lindenb. & Hampe, Lejeunea longispica Spruce, Bryopteris brevis Steph., Bryopteris fruticulosa subsp. monoica Stotler, Jungermannia filicina Sw., Bryopteris filicina var. hookeriana Nees, Lejeunea tenuicaulis (Taylor) Spruce, Bryopteris longispica (Spruce) Steph.

  • Description

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    Species Description - Dioicous or autoicous. Plants up to 25 cm long, usually feather-like, the secondary stems regularly and often densely pinnate to bi- or even tri-pinnate, sometimes branching more sparsely and irregularly pinnate (in very long plants). Branches as in the genus, tiny secondary or tertiary flagelliform branches of the Bryopteris-type sometimes present. Stems as in the genus. Leaves ovate-oblong to narrowly oblong, often falcate, 1.3-2.2 × 0.9-1.3 mm, sharply and irregularly toothed in the upper half to ± entire, ventral margin plane or incurved; median leaf cells 20-35(-40) × 10-15 µm; oil bodies ca. 15-25 per cell, homogeneous. Lobules small, up to l/5× leaf length, ovate, free margin gently curved towards the ventral margin, ± inflexed or plane, with a shallow indentation in the upper half, otherwise entire; hyaline papilla projecting upwards from the inner side of the free margin proximal to the indentation. Underleaves slightly broader than long to distinctly elongated and up to 2× longer than wide, 0.8-1.7 × 0.8-1 mm, 1.5-2× stem width, apex rounded to truncate, sharply toothed, margins plane or narrowly recurved, entire or toothed above the middle, bases narrowly decurrent or subauriculate, insertion line straight or narrowly curved; rhizoids sparse, often lacking. Reproductive structures as in the genus.

  • Discussion

    The species grows in virgin and disturbed very humid rain forests and woodlands on tree trunks, branches and occasionally on moist rocks. It is a good indicator of mist. Most records are from submontane and lower montane elevations but in wet coastal areas the species may approach sea level.

    Bryopteris filicina is one of the most common and conspicuous liverworts of neotropical upland rain forests and because of its large size has more often been collected than any other species of Lejeuneaceae. The material cited above is only a small fraction of the enormous amount of material available in the various herbaria and includes, in particular, recent specimens. Older collections have been cited in extenso in Stotler and Crandall-Stotler (1974). Because of its wide distribution B. filicina shows considerable morphological variation. It has been described in the past under many different names, some of which were accepted by Stotler and Crandall-Stotler (1974) as discrete taxa. I have found the variation in this species to be continuous and have been unable to recognize more than one taxon.

    The species has sometimes been confused with Bryopteris diffusa, which differs in many respects, however. Relationships among these two species and with the African B. gaudichaudii and the fossil B. succinea are briefly discussed under B. diffusa. Old World records of B. filicina usually represent Ptychanthus striatus, which has a similar growth habit. For differences between the two genera see above.

    Branching in Bryopteris filicina is regularly pinnate in rather short or young plants growing away from the substrate in a straight manner, but pendent, more elongated stems and plants growing in deep shade usually have a more laxly and more irregularly pinnate to bipinnate (to tri-pinnate) habit. The secondary and tertiary branches of these plants are rather small and stand at almost right angles with the main branches. Tiny, flagelliform branchlets of the Lejeunea-type are often produced in these plants. Sometimes they are easily detached, e.g. in coll. Gradstein 8651 from Colombia, and function as means of vegetative reproduction. They may also develop on more densely and regularly pinnate forms, however. The laxly branched plants have been described as B. trinitensis and B. tenuicaulis but in my interpretation they are mostly somewhat etiolated shade forms. Leaves in these forms are often rather short ovate and plane and somewhat distant.

    Leaves in Bryopteris filicina may be toothed all along the upper half of the dorsal and ventral leaf margins but more often there are only few teeth and some populations have mostly entire leaves. The latter were described as B. flaccida. The teeth are always sharp but vary considerably in size and on a single stem leaves with many or with few teeth may be found. Occasionally, small primary shoot-type leaves and underleaves with entire margins are produced on elongated branches, resembling stolons! Spreading of the leaves may be oblique or wide and in the latter case they may become strongly falcate. The ventral leaf margin is usually distinctly upcurved but in weaker populations with small, spatiated leaves described as B. tenuicaulis margins are plane.

    Underleaves of Bryopteris filicina vary considerably in length and in some populations they are very short and broader than long whereas in others (e.g. Colombia, Bischler 6110) they are elongated rectangular and twice as long as wide. Size of ther male barcteoles also varies considerably and in some populations they are very tiny and become reduced towards the apex of the spike. The specialized male spikes of Bryopteris are unusual for members of the subfamily Ptychanthoideae and are a feature reminiscent of the Lejeuneoideae. Female shoots, finally, are normally short-specialized as usual in the genus. I have seen a collection from Brazil (Burchell 2515), however, in which the female branches were distinctly elongated and had several series of vegetative leaves preceding the involucre.

    Chemistry: Terpenoids: bazzanene, bicyclogermacrene and norpinguisone-methylether have been recored as main constituents; the chemical constitution of the species seems to be considerably variable (Gradstein et al., 1981,1985).

  • Objects

    Specimen - 613523, J.-P. Frahm s.n., Bryopteris filicina (Sw.) Nees, Lejeuneaceae (45.0), Marchantiophyta; North America, Mexico, Chiapas

    Specimen - 710741, W. W. Thomas 3769, Bryopteris filicina (Sw.) Nees, Lejeuneaceae (45.0), Marchantiophyta; North America, Mexico, Guerrero, General Heliodoro Castillo Mun.

    Specimen - 613532, C. Conzatti 3822, Bryopteris filicina (Sw.) Nees, Lejeuneaceae (45.0), Marchantiophyta; North America, Mexico, Oaxaca

    Specimen - 613518, W. W. Thomas 3546, Bryopteris filicina (Sw.) Nees, Lejeuneaceae (45.0), Marchantiophyta; North America, Mexico, Veracruz, Hidalgotitlán Mun.

    Specimen - 574655, Fr. León 10251, Bryopteris filicina (Sw.) Nees, Lejeuneaceae (45.0), Marchantiophyta; West Indies, Cuba

    Specimen - 574656, L. M. Underwood 667, Bryopteris filicina (Sw.) Nees, Lejeuneaceae (45.0), Marchantiophyta; West Indies, Cuba

    Specimen - 574635, A. W. Evans 666, Bryopteris filicina (Sw.) Nees, Lejeuneaceae (45.0), Marchantiophyta; West Indies, Jamaica

    Specimen - 574638, L. M. Underwood 2883, Bryopteris filicina (Sw.) Nees, Lejeuneaceae (45.0), Marchantiophyta; West Indies, Jamaica

    Specimen - 574637, W. R. Maxon 2205, Bryopteris filicina (Sw.) Nees, Lejeuneaceae (45.0), Marchantiophyta; West Indies, Jamaica

    Specimen - 574633, E. G. Britton 214a, Bryopteris filicina (Sw.) Nees, Lejeuneaceae (45.0), Marchantiophyta; West Indies, Jamaica

  • Distribution

    Very common throughout tropical America, from sea level up to 2500 m. In addition, there are a number of doubtful records from the palaeotropics (Mauritius, Nepal, Java, Solomons Is., and Tahiti; see Stotler & Crandall-Stotler, 1974). All of these are based on old, 18th or 19th century specimens and may have been mislabelled or misidentified. The specimen from Nepal, described as Bryopteris nepalensis Steph., belongs to Ptychanthus striatus according to Mizutani (1976).

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