taxonID	type	description	language	source
03EDC1190E2D620FFF4F9010466D3C2D.taxon	materials_examined	Type: — BRAZIL. Pará: Serra do Cachimbo an der BR 163 nahe Cachimbo, epiphytisch in feuchtem Sekundär-Regenwald; ca. 500 m, 7 July 1989, A. Schäfer-Verwimp & Verwimp 11356 (holotype ALCB; isotype SP, hb. Schäfer-Verwimp). Main axis including leaves 340 − 430 µm wide, vegetative branches Lejeunea - type with conspicuous basal collar. Stem 50 µm in diameter, in cross section with 7 epidermal cells and 3 − 4 medullary cells; ventral merophyte two cells wide. Leaves imbricate, suberect; lobe suborbicular, 170 − 215 µm long × 150 − 195 µm wide, dorsal margin slightly arched, entire, ventral free margin straight, usually continuous with the keel of the lobule, entire, apex obtuse to acute to rounded, plane; cells hexagonal to oblong to isodiametric, 18 − 30 µm long × 13 − 18 µm wide, strongly mammillose, trigones minute, intermediate thickenings absent; oil bodies not observed; ocelli absent; lobule inflated, ovate, 95 − 120 µm long × 70 − 85 µm wide, ca. 1 / 3 − 1 / 2 of the leaf length, free margin involute, second and first teeth evident, contiguous or parallel, generally the first tooth is prominent, hyaline papilla situated in the notch between the two teeth or not conspicuous, keel arched with cells strongly mammillose. Underleaves distant, small, 80 − 105 µm long × 75 − 110 µm wide, slightly longer than wide, sometimes slightly wider than long, ca. 1.0 − 2.5 × stem width, sinus V-shaped to U-shaped, lobes triangular-acute, bases cuneate, insertion line slightly curved. Autoicous. Androecia on main branch or on lateral branches or intercalary and proliferative, 2 − 4 pairs of bracts, 1 − 2 bracteoles at the base of the branch. Gynoecia terminal on main branch or on short lateral branch, bract lobes obovate, 430 µm long × 370 µm wide, margins entire, apex rounded, mostly curved, bracteole narrowly ovate, bifid, 400 µm long × 280 µm wide. Perianths pyriform, 530 µm long × 550 µm wide, 5 - keeled, beak short, 25 µm long, 1 − 2 cells long. Vegetative multiplication by cladia, sometimes arising from the stem of androecial branch. Additional specimens examined: — BRAZIL. Amazonas: between Manaus and São Gabriel, São Luiz, 00 ° 10 ’ S, 63 ° 00 ’ W, in secondary forest, 28 July 1979, R. M. Schuster 79 - 4 - 103, 79 - 4 - 117 (F); NW of São Gabriel, near mouth of Rio Uaupés, 00 ° 08 ’ S, 67 ° 10 ’ W, riverside forest, 21 July 1979, R. M. Schuster 79 - 20 - 1025 (F). Manaus, Rio Negro, Praia da Lua, 3 ° 02 ’ 04.9 ” S 60 ° 09 ’ 32.5 ” W, floresta sazonalmente inundada (Igapó), epífita, 25 November 2016, A. M. Sierra 4592 (INPA; ALCB).	en	Bastos, Cid José Passos, Sierra, Adriel M., Zartman, Charles E., Schäfer-Verwimp, Alfons (2025): Novelties in Cheilolejeunea (Spruce) Steph. sect. Trachylejeunea (Lejeuneaceae: Jungermanniopsida) from the Brazilian Amazon. Phytotaxa 715 (3): 279-286, DOI: 10.11646/phytotaxa.715.3.7, URL: https://doi.org/10.11646/phytotaxa.715.3.7
03EDC1190E2D620FFF4F9010466D3C2D.taxon	etymology	Etymology: — In reference to the production of cladia in the androecial branch.	en	Bastos, Cid José Passos, Sierra, Adriel M., Zartman, Charles E., Schäfer-Verwimp, Alfons (2025): Novelties in Cheilolejeunea (Spruce) Steph. sect. Trachylejeunea (Lejeuneaceae: Jungermanniopsida) from the Brazilian Amazon. Phytotaxa 715 (3): 279-286, DOI: 10.11646/phytotaxa.715.3.7, URL: https://doi.org/10.11646/phytotaxa.715.3.7
03EDC1190E2D620FFF4F9010466D3C2D.taxon	distribution	Distribution and habitat: — The species so far known to be restricted to the states of Amazonas and Pará, since no other specimens were found in collections examined. Notes: — Due to morphology of the gynoecial bracts and bracteoles, appearance of the cells of the dorsal leaf lobe, and number of stem medullary cells, the new species is similar to Cheilolejeunea papulosa. However, C. androproliferans presents more strongly mammillose keel cells, suberect leaves with a lobe ca. 2 × the length of the lobule, underleaves with broadly acute to lunate sinus (U-shaped), smaller trigones and asexual reproduction by cladia. A striking feature of the new species in relation to asexual reproduction is the production of cladia on the androecial branch. Although the holotype presented a monoicous sexual condition, the material from the State of Amazonas had only androecial branches and no production of cladia. Due to the strongly mammillose keel cells, small underleaves and small size of the gametophyte, C. androproliferans may resemble some phenotypes of Cheilolejeunea savannae Macedo, Ilkiu-Borges & Bastos (2020: 79) (Macedo et al. 2020). However, the presence of two adjacent or overlapping teeth in the leaf lobule and the absence of innovations indicates that they are different species.	en	Bastos, Cid José Passos, Sierra, Adriel M., Zartman, Charles E., Schäfer-Verwimp, Alfons (2025): Novelties in Cheilolejeunea (Spruce) Steph. sect. Trachylejeunea (Lejeuneaceae: Jungermanniopsida) from the Brazilian Amazon. Phytotaxa 715 (3): 279-286, DOI: 10.11646/phytotaxa.715.3.7, URL: https://doi.org/10.11646/phytotaxa.715.3.7
03EDC1190E2F6208FF4F92FA404735BF.taxon	materials_examined	Type: — BRAZIL. Amazonas: Manaus, Rio Negro, Praia da Lua, 3 ° 02 ’ 04.9 ” S 60 ° 09 ’ 32.5 ” W, floresta sazonalmente inundada (Igapó), epífita, 25 November 2016, A. M. Sierra 4590 (holotype INPA; isotype ALCB). Main axis with leaves 0.9 − 1.2 mm wide, branches Lejeunea - type. Stems 80 µm in diameter, in cross section with 7 epidermal cells and 10 − 11 medullary cells, ventral merophytes 2 − 3 cells wide. Leaves imbricate, spreading to suberect; leaf lobes ovate-oblong, convex, 410 − 560 µm long × 320 − 490 µm wide, dorsal margins slightly arched, crenulate to entire, ventral free margins straight, entire to crenulate, apices rounded, curved; lobe margins formed by two rows of quadrate cells; leaf cells oblong to hexagonal, basal cells 20 − 30 µm long × 15 − 23 µm wide, median cells 15 − 20 µm long × 13 µm wide, strongly mammillose, trigones small to median, intermediate thickenings 0 − 1 per cell; oil bodies not seen; ocelli absent; lobules oblong to oblong-subrectangular, large, 160 − 230 µm long × 95 − 115 µm wide, ca. 1 / 2 − 2 / 3 of the leaf lobe length, weakly inflated, free antical margin slightly involute, 8 − 10 cells long, lobule apices with two small, tightly associated teeth, apical free margin curved to oblique, keel slightly arched to straight, crenate due to mammillose cells; reduced lobules not observed. Underleaves contiguous to distant, 200 − 230 µm long × 190 − 280 µm wide, ca. 2.0 − 3.0 × stem width, ovate to suborbicular, bifid to 1 / 2, lobes triangular, sinus V-shaped, bases cuneate, insertion line slightly curved. Autoicous. Androecia terminal in the main stem or intercalary, or on short to long lateral branches, 3 − 4 pairs of bracts, bracteoles present throughout the spike. Gynoecia on short or long lateral branches, or on the main stems, without innovations, occasionally with one subgynoecial branch Lejeunea - type (pseudo-innovation), bract lobes obovate, margins entire, apices rounded, 550 µm long × 380 µm wide, bract lobules oblong, apices acute, 480 µm long × 130 µm wide, bracteoles oblong, bifid, margins entire; perianths obovoid, 610 µm long, 480 µm in diameter, 4 - keeled, keels smooth to crenulate, beak long, 110 µm long × 50 µm wide, 3 − 4 elongated cells high.	en	Bastos, Cid José Passos, Sierra, Adriel M., Zartman, Charles E., Schäfer-Verwimp, Alfons (2025): Novelties in Cheilolejeunea (Spruce) Steph. sect. Trachylejeunea (Lejeuneaceae: Jungermanniopsida) from the Brazilian Amazon. Phytotaxa 715 (3): 279-286, DOI: 10.11646/phytotaxa.715.3.7, URL: https://doi.org/10.11646/phytotaxa.715.3.7
03EDC1190E2F6208FF4F92FA404735BF.taxon	etymology	Etymology: — In reference to the type locality (Praia da Lua): a white sands beach along the Rio Negro near Manaus.	en	Bastos, Cid José Passos, Sierra, Adriel M., Zartman, Charles E., Schäfer-Verwimp, Alfons (2025): Novelties in Cheilolejeunea (Spruce) Steph. sect. Trachylejeunea (Lejeuneaceae: Jungermanniopsida) from the Brazilian Amazon. Phytotaxa 715 (3): 279-286, DOI: 10.11646/phytotaxa.715.3.7, URL: https://doi.org/10.11646/phytotaxa.715.3.7
03EDC1190E2F6208FF4F92FA404735BF.taxon	distribution	Distribution and habitat: — Restricted to the State of Amazonas, Northern Brazil. The holotype was collected on the base of trees in a seasonally inundated forest on a fluvial island of the Rio Negro near the Praia da Lua ~ 20 km from Manaus. However, other specimens were collected near São Gabriel in terra firme forest and in Serra do Aracá. Notes: — The new species resembles the neotropical Cheilolejeunea aneogyna, but differs from the latter by the following characters: (1) leaf lobe margins formed by two rows of quadrate cells; (2) androecial branches with bracteoles present throughout the spike (bracteoles only at base of the spike in C. aneogyna); (3) large, subrectangular leaf lobules (160 − 230 µm long × 95 − 115 µm wide), ca. 1 / 2 − 2 / 3 leaf lobe length (leaf lobule shorter, triangular-ovate to subrectangular, 100 − 140 µm long × 80 − 92 µm wide, ca. 1 / 3 of the lobe length in C. aneogyna); (4) leaf lobe cells strongly mammillose (cells weakly mammillose in C. aneogyna) (5) ventral merophyte 2 − 3 cells wide (only two cells wide in C. aneogyna). Asexual reproduction by caducous leaves, sometimes with rhizoids on the margin, was reported for C. aneogyna by Gradstein & Costa (2003) and Bastos (2008, 2012). This characteristic has not yet been observed in the specimens of C. luensis studied. An interesting characteristic of C. aneogyna, also shared with C. luensis, is the greater development of the first tooth of the leaf lobule in relation to the reduced second tooth. This is unusual in Cheilolejeunea. Mizutani (1961), Bastos (2010) and Renner (2012) stated that the genus Cheilolejeunea is diagnosed not by the position of the hyaline papilla (this is always positioned at the proximal base of the first tooth and, consequently, distal to the second tooth), but rather by the greater development of the second tooth, in relation to the first tooth, which is reduced. Specimens of section Trachylejeunea with a more developed first tooth could be confused with species belonging to the genus Pycnolejeunea (Spruce 1884: 246) Schiffner (1893: 124), especially due to the greater development of the first tooth and the strongly mammillose cells of the leaf lobe, since the absence of ocelli would not be unusual in Pycnolejeunea (some specimens of C. aneogyna were found with ocelli, as reported by Bastos 2012).	en	Bastos, Cid José Passos, Sierra, Adriel M., Zartman, Charles E., Schäfer-Verwimp, Alfons (2025): Novelties in Cheilolejeunea (Spruce) Steph. sect. Trachylejeunea (Lejeuneaceae: Jungermanniopsida) from the Brazilian Amazon. Phytotaxa 715 (3): 279-286, DOI: 10.11646/phytotaxa.715.3.7, URL: https://doi.org/10.11646/phytotaxa.715.3.7
03EDC1190E2F6208FF4F92FA404735BF.taxon	materials_examined	Additional specimens examined: — BRAZIL. Amazonas: Between Manaus and São Gabriel, along Rio Curicuriari near mouth of the Rio Cariua; terra firme; 00 ° 20 ’ S, 66 ° 55 ’ W, 14 July 1979, R. M. Schuster 79 - 17 - 880 (F); Barcelos, Serra do Aracá, Rio Aracá, Igarapé Madixi, 0 ° 14 ’ 29 ” S, 63 ° 06 ’ 03 ” W, 18 August 2014, C. E. Zartman 9613 (INPA / ALCB). Cheilolejeunea remotistipula (C. J. Bastos & C. E. Zartman) A. M. Sierra, C. J. Bastos & C. E. Zartman, comb. nov. Basionym: Pycnolejeunea remotistipula C. J. Bastos & C. E. Zartman, Neodiversity 10: 2. 2017. Type: ― BRAZIL. Amazonas: Rio Negro, between Manaus and São Gabriel, along BR 307, from São Gabriel, just N of igarapé-Mirim, near Jerusalém; primary forest, 00 ° 20 ’ N, 66 ° 35 ’ W, 17 July 1979, R. M. Schuster 79 - 18 - 835 (holotype, F!). Description and illustration: Bastos & Zartman (2017). Notes: ― The Cheilolejeunea remotistipula was previously described as Pycnolejeunea remotistipula Bastos & Zartman (2017: 2) by Bastos & Zartman (2017) because the first tooth of the leaf lobule is well developed, and the second tooth is reduced. However, this possibility (the first tooth being elaborated in relation to the second tooth) was observed in some Cheilolejeunea species belonging to sect. Trachylejeunea, as observed in Cheilolejeunea aneogyna, for example. More detailed studies of the holotype of P. remotistipula, mainly in relation to the morphology of the leaf lobule, as well as the absence of innovation showed that P. remotistipula is a member of Cheilolejeunea Sect. Trachylejeunea. Due to the long, subrectangular leaf lobule and lack of innovations, C. remotistipula also resembles the new species described here, C. luensis. However, the new species can be separated from C. remotistipula based on the following characteristics: (a) perianth with long beak (shorter to very short in C. remotistipula); (b) androecial branches with bracteoles present throughout the spike (bracteoles only at base of the spike in C. remotistipula); (c) free margins of the leaf lobule 8 − 10 cells long (15 − 17 cells long in C. remotistipula); and (d) underleaves with acute lobe apices (rounded to obtuse in C. remotistipula). In Cheilolejeunea luensis sp. nov. often the first tooth is more developed (as is also common in C. aneogyna), and this tooth may be straight or vertically oriented, as is observed in C. remotistipula. In this condition the two species can be confused, and only careful examination of the four characters listed above can ensure correct identification of the specimens. Additional specimens examined: — BRAZIL. Amazonas: São Gabriel da Cachoeira, Rio Negro, Ilha Cariuari, 20 km descendo o Rio desde a cidade de São Gabriel da Cachoeira, 0 ° 10 ’ 47.5 ” S 67 ° 01 ’ 25.0 ” W, floresta sazonalmente inundada (Igapó), epífita, 16 December 2016, A. M. Sierra & C. E. Zartman 4702 (INPA).	en	Bastos, Cid José Passos, Sierra, Adriel M., Zartman, Charles E., Schäfer-Verwimp, Alfons (2025): Novelties in Cheilolejeunea (Spruce) Steph. sect. Trachylejeunea (Lejeuneaceae: Jungermanniopsida) from the Brazilian Amazon. Phytotaxa 715 (3): 279-286, DOI: 10.11646/phytotaxa.715.3.7, URL: https://doi.org/10.11646/phytotaxa.715.3.7
