Monographs Details: Frullanoides densifolia Raddi subsp. densifolia
Authority: Gradstein, S. Robbert. 1994. Lejeuneaceae: Ptychantheae, Brachiolejeuneae. Fl. Neotrop. Monogr. 62: 216. (Published by NYBG Press)
Synonyms:Jungermannia bicolor Nees, Phragmicoma rupestris Gottsche, Lejeunea succisa Steph., Brachiolejeunea canaliculata Steph., Marchesinia coniloba Steph., Mastigolejeunea decurrens Steph., Ptychanthus boliviensis Steph., Ptychocoleus boliviensis Steph., Dicranolejeunea boliviensis Steph., Dicranolejeunea nudiflora Steph., Brachiolejeunea appendiculistipa Steph., Brachiolejeunea columbica Steph., Ptychocoleus densifolius (Raddi) Trevis., Brachiolejeunea densifolia (Raddi) A.Evans, Trocholejeunea densifolia (Raddi) R.M.Schust., Phragmicoma bicolor (Nees) Gottsche, Lindenb. & Nees, Brachiolejeunea bicolor (Nees) Mont., Brachiolejeunea rupestris (Gott) Steph., Brachiolejeunea succisa Steph., Brachiolejeunea canaliculata Steph.
Description:Subspecies Description - Dioicous. Plants 3-12 cm long × 2-3 mm wide, green to black when alive, becoming dark brown to black in the dried condition. Branching Frullania-type, occasionally Lejeunea-type. Stems 0.15-0.3 mm in diam., in cross section composed of 20-35 epidermal cells surrounding ca. 50-100 smaller medullary cells, dorsal epidermal cells distinctly larger than the ventral epidermal cells, dorsal and lateral epidermis rather thinwalled, ventral epidermis and medulla strongly thickwalled; ventral merophyte 8-10 (-14) cell rows wide. Leaves imbricate, dorsal lobe ovate-falcate, 1-1.8 × 0.8-1.4 mm, apex acute-apiculate or narrowly rounded, usually recurved, dorsal margin auriculate at base, ventral margin curved upwards, when spread out forming an angle of 160-180° with the keel; median cells 25-40 × 18-32 µm, base cells larger, margin cells subquadrate, 14-22 µm high; oil bodies homogeneous, small, ellipsoid to fusiform, ca. 2 × 3-6 µm, 15-25 per cell. Lobules ovate-triangular, 0.5-0.8 × 0.3-0,7 mm, 2/5-l/2× lobe length, apex oblique, keel almost straight, free margin plane, with 6-10 teeth, the teeth 1-3 cells long, longest and more widely spatiated towards the apex of the lobule, erect with the upper cell(s) inflexed. Underleaves imbricate, broadly ovate-subquadrate, 0.5-1.1 × 0.8-1.3 mm, 4-5× stem width, apex rounded to truncate, recurved, margins recurved or almost plane, bases distinctly auriculate, the auricles ca. 0.1-0.3 mm long, appressed to the stem, insertion line deeply arched. Androecia on short or long branches, bracts in 3-20 series, lobule swollen, epistatic, with reduced teeth; antheridia two per bract. Gynoecial bracts bifid to 1/2 or more, keel with a large wing, lobes ovate-obovate, ca. 2 mm long, squarrose, apex acute-apiculate or narrowly rounded, lobules narrow rectangular, l/3-l/2× lobe length, apex acute; bracteoles erect, rectangular, 1-1.4 mm long, apex truncate-emarginate to short bifid, plane or slightly recurved, margins plane. Perianths immersed, occasionally exserted to 1/4, obovoid-cylindrical, 1.2-1.7 mm long, 10-keeled over much of its length, beak 8-10 cells long. Spores ca. 45-55 µm long; elaters 68 per capsule, 435-550 × 18 µm, with a single brown, ca. 4 µm wide spiral.

Discussion:Chemistry: the sesquiterpenes bicyclogermacrene, 13-caryophyllene and various pinguisanines, and the free flavonol aglycone kaempferol-3-methylether are the main secondary metabolites detected in this species (Kruijt et al., 1986; Asakawa & Inoue, 1987).

Frullanoides densifolia subsp. densifolia varies considerably in size and plants from high altitudes tend to be more robust. Small plants from low altitudes often have a narrowly rounded leaf apex and rather small underleaf auricles and may therefore be confused with F. liebmanniana. The latter may be recognized, however, by the fewer lobule teeth and the thin-walled ventral epidermal cells.

Discovery and Ecology: Mexico, Costa Rica, Panama, Andes (Venezuela to Argentina), SE Brazil; a common montane species, occurring at altitudes of 500-3500 m in the Andes and Central America, in SE Brazil approaching sea level. Frullanoides densifolia grows in rather open, exposed habitats on bark, rock or soil in natural and degraded submontane and montane forests, scrubby vegetations, meadows, along roads and in rather dry cerrado vegetation of southeast Brazil. In dense forests the species is restricted to canopy branches

Distribution:Mexico North America| Veracruz Mexico North America| Costa Rica South America| Cartago Costa Rica Central America| Puntarenas Costa Rica Central America| San José Costa Rica Central America| Panama Central America| Chiriquí Panamá Central America| Colombia South America| Antioquia Colombia South America| Boyacá Colombia South America| Casanare Colombia South America| Cundinamarca Colombia South America| Huila Colombia South America| Magdalena Colombia South America| Risaralda Colombia South America| Santander Colombia South America| Tolima Colombia South America| Venezuela South America| Lara Venezuela South America| Mérida Venezuela South America| Táchira Venezuela South America| Ecuador South America| Carchi Ecuador South America| Chimborazo Ecuador South America| Pichincha Ecuador South America| Peru South America| Amazonas Peru South America| Cajamarca Peru South America| Cusco Peru South America| Huánuco Peru South America| San Martín Peru South America| Brazil South America| Bahia Brazil South America| Espirito Santo Brazil South America| Minas Gerais Brazil South America| Rio de Janeiro Brazil South America| Paraná Brazil South America| Rio Grande do Sul Brazil South America| Santa Catarina Brazil South America| São Paulo Brazil South America| Bolivia South America| Beni Bolivia South America| Cochabamba Bolivia South America| La Paz Bolivia South America| Santa Cruz Bolivia South America| Argentina South America| Salta Argentina South America| Tucuman Argentina South America|