Synopsis of Anomobryum and Bryum (Bryaceae, Bryophyta) in Brazil

Bryaceae is a species-rich moss family comprising ten genera and 660 species, with four genera and 54 species occurring in Brazil. Bryum is the largest genus, with 440 cosmopolitan species, while Anomobryum comprises 47 cosmopolitan species. Identifying taxa in Bryaceae is usually a difficult task, mainly due to the lack of the sporophyte in herborized specimens. This study aimed to provide a taxonomic treatment for the species of Anomobryum and Bryum in Brazil. Field expeditions were performed in all Brazilian biomes and States, with specimens deposited in the SP herbarium. We examined type specimens and collections from national and international herbaria, but mostly from the SP herbarium. Two species of Anomobryum and 18 species of Bryum were recorded for Brazil. Twenty species occur in the Atlantic Forest, with Bryum pallescens , Anomobryum conicum and A. julaceum being exclusives. Bryum pallescens is a new record for Brazil, and B. riparioides is a reestablished name also endemic to Brazil. We present the first description of the sexuality of Bryum renauldii . Ten taxa of doubtful occurrence and 20 excluded names for Bryum were reported for Brazil.

Bryum is one of the richest and most complex moss genera in the Neotropics and Brazil (Gradstein et al. 2001;Costa et al. 2011;Costa & Peralta 2015), with 440 cosmopolitan species (Cox & Hedderson 2003;Ochyra et al. 2008;Frey & Stech 2009).The morphology of Bryum is quite variable, with some authors (Frey & Stech 2009) defining this genus as containing green to occasionally red, pink or silver plants, with leaf margins typically limbate, di-, syn-, or autoicous, with usually pendulous capsules, oblong or pyriform to cylindric in shape, and with often complete peristome.In Brazil, only 32 species of Bryum are recorded (Costa et al. 2011).
The primary aim of the present study was to provide a taxonomic treatment for the species of Anomobryum and Bryum in Brazil.Secondary aims were to properly determine which species of Anomobryum and Bryum occur in Brazil, elucidate the distribution and ecology of all recorded species, and provide brief descriptions and illustrations for each species.

Material and Methods
Field expeditions were carried out in every Brazilian biome (vegetation classification system according to IBGE 2012) and State, with specimens deposited at the SP herbarium.We examined herborized specimens from the BM, HAS, HERW, NY, UB and SP herbaria (acronyms according to Thiers 2021).Specimens bearing sporophytes were given preference to ensure proper species identification.Due to the extensive number of examined specimens for most species, only two exsiccatae per state (one per municipality) were selected for the examined specimens lists.A complete list of all examined specimens in this study is provided as an supplementary material.About 40 specimens of Anomobryum and ca.700 of Bryum were analyzed.Glycerin gelatin fixed slides were mounted (Kisser 1935) with the gametophytic and sporophytic morphological structures and analyzed using optical microscopy techniques.Photographs of the specimens were taken using optical and stereoscopic microscopes, and photographic plates were elaborated using Adobe Photoshop CS3 software (Adobe Systems Inc., San Jose, CA).
The classification system adopted here follows Frey & Stech (2009), which accepts both Anomobryum and Bryum.The synonymy for Bryum followed Ochyra et al. (2008) and Frey & Stech (2009), morphological delimitations for genera and species followed Allen (2002), and morphological terminology followed Malcolm & Malcolm (2006) and Gradstein et al. (2001).Only synonyms with type specimens from Brazil were listed.For synonyms from other regions, see the complete checklist in Ochi (1980Ochi ( , 1981Ochi ( , 1982)).Complete morphological descriptions of all species from this study can be found in Allen (2002), Ochi (1980Ochi ( , 1981)), Ochi (1994), and (Bartram 1952).Species distribution classification followed Valente & Porto (2006), considering rarely distributed species occurring in 1-4 States, moderately distributed species occurring in 5-9 States, and widely distributed species occurring in 10 or more States.Costa et al. (2011) and Costa & Peralta (2015) were used for verifying species records in the Brazilian States, and asterisks indicate first records for the States. 2008;Frey & Stech 2009).We observed a central strand in stem cross-section and stereids in leaf cross-section for all Brazilian Anomobryum and Bryum species.
Two species of Anomobryum were recorded in Brazil (A. conicum and A. julaceum) which corroborates previous studies (Costa et al. 2011;Costa & Peralta 2015).There are seven new records for the Brazilian States.Both species of Anomobryum have moderate distribution (i.e., occurrence in 5 States), inhabit the Atlantic Forest, and occur exclusively in Brazil's Southern and Southeastern regions (also corroborating Costa & Peralta 2015).One species was excluded from Brazil.
We recorded 18 species of Bryum for Brazil.Compared with the literature, Costa & Peralta (2015) reported 19 species, while Costa et al. (2011) mentioned 32 species.This difference is due to the classification system adopted, which may consider Bryum either a broadly or narrowly circumscribed genus.Costa & Peralta (2015) also recorded for Bryaceae the genera Imbribryum, Plagiobryum, Ptychostomum and Rosulabryum, most of them segregated from Bryum (Lindberg 1863;Pedersen 2005;Spence 1996;2005), while Costa et al. (2011) considered just Bryum.Based on the available molecular studies, only Imbribryum and Plagiobryum are currently accepted among the aforementioned genera (Frey & Stech 2009).
Thirty-seven new State distribution records are reported for Bryum in the present study.Regarding the species' geographical distribution, all recognized species occur in the Atlantic Forest.Of these, 13 also occur in the Cerrado, nine in the Caatinga, five in the Pampa, seven in the Amazon Forest, and only one in the Pantanal.Most species occur in Southern and Southeastern Brazil.According to Costa & Peralta (2015), the Atlantic Forest is the most diverse among the Brazilian biomes; and Southern and Southeastern Brazil also represent the richest regions in the country.Bryum pallescens is exclusive to the Atlantic Forest and is reported for the first time in Brazil.Bryum riparioides is a reestablished name and an endemic taxon to Brazil.Costa & Peralta (2015) cited five species of Bryum as endemic to Brazil, while we report only one.Here we present the first description of the sexuality of Bryum renauldii.
Twenty taxa are excluded (i.e., do not occur in Brazil) and ten are considered doubtful occurrences.Four new synonyms with type specimens from Brazil (i.e., B. brasiliense, B. leptocladon, B. oncophorum, and B. pseudomarginatum) are proposed.
Selected specimens examined: BRAZIL.Amazonas: This plant has small-sized (up to 30 mm high), leaves concave, elimbate, with narrowly hexagonal to fusiform upper cells, abruptly larger, and rectangular to quadrate basal cells.Axillary gemmae were observed in a few specimens of B. apiculatum (B.E.Chaves 92 -UB -in the Caatinga; B.K. Canestraro 1598, 1646 -SP -in the Atlantic Forest), as reported by Allen (2002) and Ochi (1994).
Bryum apiculatum is similar to B. orthodontioides in its small-sized (3 mm high), lanceolate leaves with unbordered margins, acuminate apices, short-excurrent costa, and rectangular basal cells.However, B. orthodontioides has even smaller and shorter leaves, entire margins, and upper leaf cells rhomboidal and thin-walled (Fig. 7A-D).Bryum apiculatum is also similar to B. dichotomum due to its smallsized (1-1.7 mm high) and presence of axillary propagules.However, the latter has leaves with narrowly rhomboidal upper cells, sub-quadrate to quadrate basal cells, thick-walled throughout, and capsules with a broad neck (Fig. 6A-6).Bryum subapiculatum differs from B. apiculatum in having narrowly rhomboidal to hexagonal-rhomboidal upper cells, rectangular to sub-quadrate basal cells, and the absence of vegetative propagules (Fig. 10A-D).
Bryum apiculatum and Anomobryum conicum share their small-sized (15 mm high), imbricate and lanceolate leaves, short-excurrent costa, and unbordered margins.Nevertheless, A. conicum has linear and thick-walled upper cells that gradually become narrowly hexagonal to rectangular and sub-quadrate towards the base (vs. the abrupt transition from narrowly hexagonal to fusiform upper cells to enlarged, rectangular to quadrate basal cells in B. apiculatum) (Fig. 1A-E).Brym apiculatum is similar to Pohlia elongata Hedw.due to the aspect of their gametophytes, leaves lanceolate, erect and plane, margins not bordered and serrulate, and percurrent to short-excurrent costa.However, Pohlia elongata has paroicous inflorescences, upper cells longer and narrower (4:1 proportion), thick-celled walls, and a more elongate capsule (Allen 2002).Bryum apiculatum is similar to B. nanoapiculatum Ochi & Kürschner (Ochi & Kürschner 1988).However, the last occurs in Yemen, is smaller in size, has short ovate and mucronate leaves, and short-excurrent costa.Hodgetts et al. (2020) proposed a new combination of B. apiculatum to Anomobryum apiculatum (Schwägr.)D. Bell & Holyoak.However, the authors based their analysis on few samples from Europe, not taking into account the global distribution of the species.Therefore, we do not follow their classification.Plants small, up to 10 mm high, silvery-green.Leaves imbricate, erect, evenly spaced, concave, ovate, oblong to elliptic; apices hyaline, aristate; margins not bordered, entire, plane; costa strong, long-excurrent; upper cells rhomboidal-hexagonal to narrow rhomboidal, basal cells rectangular, sub-quadrate to quadrate, non-porose, firm-and thin-walled.Dioicous.Capsules reddish-brown, pendulous, oblong-cylindric, neck short, corrugate, broader than the urn; opercula conic-apiculate; peristome complete; exostome It is a species with imbricate leaves with hyaline apices, not bordered margins, long-excurrent costa, and capsules with a broad neck.This species is very similar to B. argenteum.However, it has weak, evanescent and sub-percurrent costa, and capsules with a slender neck (Fig. 3A-E).The shape of the capsules and width of the neck is comparable to B. coronatum, but it has longer leaves with green apices (Fig. 5A-D).It differs from B. lanatum (P.Beauv.)Brid.due to the plants' silvery color, ovate leaves, and excurrent and reflexed costa (the excurrent part ⅓ of the length of the lamina) (Frahm 2002).Plants small, 3-10 mm high, silvery-green, leaves imbricate, erect, evenly spaced.Leaves concave, oblong-ovate to elliptic; apices hyaline, apiculate; margins not bordered, entire, plane; costa weak, evanescent, sub-percurrent; upper cells rhomboidal to rhomboidal-hexagonal, basal cells rectangular to sub-quadrate, non-porose, firm-and thin-walled.Dioicous.Capsules green to orange-brown, pendulous, cylindric to conic, neck slender; opercula conicapiculate; peristome complete; exostome well-developed, endostome with segments well-developed, cilia present.

Bryum argenteum
Selected  Bryum argenteum is a weedy species (Allen 2002) that thrives in disturbed environments (Pisa et al. 2014).It is characterized by silvery-green gametophytes, leaves elimbate with hyaline apices, costa weak and evanescent, and capsules with a slender neck.Bryum arachnoideum is similar to B. argenteum due to the color and aspect of the gametophyte, areolation and shape of the leaves.However, that species usually has larger gametophytes and capsules oblong-cylindric with a broad and corrugate neck (Fig. 2F-I).Bryum argenteum is similar to B. lanatum, but this is a plant hoary white, with stems evenly foliate and inconspicuously julaceous, long-excurrent costa, awns slender and recurved when dry (Spence 2015).Bryum argenteum may be similar to Anomobryum conicum and A. julaceum due to the julaceous stems, imbricate and ovate leaves, and small-sized (10 mm high) (Spence & Ramsay 2002).In contrast, A. julaceum has obtuse apices, entire or serrulate margins, and vermicular and thick-walled upper leaf cells (Fig. 1F-I), while A. conicum has entire to denticulate margins, strong, percurrent to short-excurrent costa, and linear to narrowly rhomboidal and thick-walled upper cells (Fig. 1A-E).These morphological similarities among Anomobryum and Bryum argenteum result from convergent evolution since both taxa are molecularly independent (Holyoak & Pedersen 2007;Pedersen et al. 2007).Plants small-to medium-sized, 3-4 mm high, light green, dark to yellowish-green, leaves imbricate, spirally-twisted when dry, evenly spaced or in inconspicuously rosulate tufts.

Bryum atenense
Acta Botanica Brasilica, 2022, 36: e2021abb0283 Bryum atenense is characterized by imbricate leaves that become spirally-twisted when dry, long-excurrent costa, leaf basal cells distinctly quadrate, and axillary bulbils sometimes present.This species resembles B. capillare due to the leaves that become spirally-twisted when dry, the presence of bulbils, and excurrent costa.However, the latter has leaves with rectangular basal cells and bordered margins with 1-4 rows of narrow, rectangular cells (Fig. 4E-I Plants medium-to large-sized, 15-25 (50) mm high, dark green to yellowish-green, leaves imbricate, spirally-twisted and crisped when dry, evenly spaced with leaves congested at apex or in dense rosulate tufts, 1-2 (3) congested rosettes.Leaves obovate to spatulate; apices green, cuspidate; margins distinctly bordered by 1-3 rows of narrow, rectangular cells, serrate to denticulate in the upper half, teeth occasionally paired, recurved at base; costa strong, short-excurrent; upper cells rhomboidal-hexagonal, basal cells rectangular, porose, Acta Botanica Brasilica, 2022, 36: e2021abb0283 This is a medium-to large-sized plant (15-25 (50) mm high), with leaves evenly spaced and congested at the apex or in dense 1-2 (3) rosulate tufts, obovate, distinctly bordered margins serrate above, and short-excurrent costa.Sterile plants usually have evenly spaced leaves, which are smaller and scattered below and larger and congested above, not distinctly rosulate (Ochi 1994).The species is extremely variable in stature, leaf size and arrangement, border width, costa excurrence, and presence of axillary filaments (Ochi 1994;Allen 2002).Axillary filaments were not observed in the studied Brazilian samples.The presence of rhizoidal tubers is rare among Brazilian specimens (i.e., Peralta 4056 -SP385870).
The type material of Rhodobryum stenothecium and Bryum liebmannii Schimp.(synonym of B. billarderii acc.Allen, 2002 -BM000873712 image!, BM000873738 image!) were analyzed.The gametophytes show variable morphology, with leaves evenly spaced and congested at the apex or in up to 3 dense leaf whorls.
Selected specimens examined: BRAZIL.Goiás: Alto Paraíso de Goiás, 12/II/2011, D. Bryum capillare has leaves spirally-twisted when dry, bordered margins, costa short-to medium-excurrent, lax upper cell walls, and axillary gemmae and bulbils are occasionally present.This species occurs in the mountain regions of Southern and Southeastern Brazil, especially in the Serra da Mantiqueira and Serra do Mar ranges.This species is very polymorphic (Casas et al. 2006), with the cell row number in the margins varying from 1-2 (Ochi 1967) to 3-7 rows of thick-walled cells (Allen 2002), depending on its geographic location.
The species resembles B. atenense due to the longexcurrent costa and spirally-twisted leaves, but it has leaves with quadrate basal cells and bordered margins by 1-2 narrow rectangular cells (Fig. 3F-J).Bryum capillare differs from B. pallescens because the latter is synoicous, and the leaf has thick cells at the base, and recurved margins (Fig. 8E-H).Bryum capillare is similar to B. pseudocapillare Besch.However, this species has filiform axillary gemmae, percurrent to short-excurrent costa and leaves not conspicuously spirally-twisted (Ochi 1980;Allen 2002).The features leaf areolation (hexagonal upper cells and rectangular basal cells) and gametophyte aspect (leaves lax, spirally-twisted, and in inconspicuously rosulate tufts) of Bryum capillare could be confused with Brachymenium consimile and B. radiculosum.However, both species of Brachymenium have spatulate leaves, erect capsules, and rudimentary endostome (Canestraro & Peralta unpubl.res.).

Bryum coronatum
Selected Bryum coronatum is characterized by the ovate to ovate-lanceolate leaves with unbordered margins and long-excurrent costa and oblong-cylindric capsules with a corrugate and broad neck.Ochi (1980) mentions that this species presents axillary gemmae.However, this feature was not observed in the Brazilian specimens.The gametophyte aspect, leaf shape, excurrence of the costa and shape of the basal cells are plastic features (Ochi 1980), probably due to the wide distribution of the species.
This species is similar to B. subapiculatum in aspect to leaf shape and areolation.However, the latter has cylindric to conic capsules and a slender neck (Fig. 10A-D).Bryum coronatum and B. arachnoideum share the oblong-cylindric capsules with a corrugated and broad neck, but this species is silvery-green with leaves imbricated with hyaline apices (Fig. 2F-I).Bryum dichotomum also has capsules with a broad neck.However, it has thick cell walls, pyriform to ovoid capsules, and sometimes vegetative propagules (Fig. 6A-E).Bryum densifolium is a robust plant (20-70 mm high), with leaves evenly spaced along the stem, which have a distinct sharply serrate border in the upper half and are often reflexed in the lower half, upper cells narrow rhomboidalhexagonal, and basal cells rectangular and porose.Bryum densifolium is variable in plant size and leaf shape, size, and leaf arrangement (Ochi 1967).The size (15-25 mm high) and leaf arrangement are similar to Bryum billarderii.However, the latter has leaves varying from evenly spaced to rosulate (up to 3 rosettes), with obovate to spatulate leaves (Fig. 4A-D).Bryum procerum Schimp.ex Besch.can be distinguished from B. densifolium in its broader, longdecurrent and conspicuously limbate leaves, inconspicuous or absent dorsal stereids and generally plurisetous condition (Allen 2002).
According to Ochi (1980), B. pseudomarginatum Geh.& Hampe can be distinguished by the leaves conspicuously crisped when dry, denticulate at the apex, and teeth irregular in size, shape and arrangement.Alternatively, B. densifolium has leaves wrinkled or flexuose and spreading when dry, serrate at the apex, and teeth regular in shape and size.However, the type of B. pseudomarginatum (Puiggari 1829 -BM, G, PC) has flexuose and spreading leaves, which is a character of B. densifolium.Furthermore, B. densifolium and B. pseudomarginatum show variable leaf upper cell shape, and the rows of elongate cells at the margin can vary from 1-2 up to 3-5.Thus, we believe the characters used by Ochi (1980) to separate the species are not supported.Since we have analyzed images of both type specimens and a vast collection from Brazil (including a specimen seen and identified by Ochi (1980)  Acta Botanica Brasilica, 2022, 36: e2021abb0283 = Bryum angusticymba Müll.Hal., Gen. Musc. Frond. 208. 1901, nom. nudum. Original Material: (Brazil. Santa Catharina) E. Ule (944) (PC, H?), syn.acc.Ochi (1980, in the examined material).
Selected This is a weedy species (Allen 2002) distinguished by the thick leaf cell walls, pyriform to ovoid capsules with a broad neck, and by the occasional presence of axillary gemmae and bulbils.Bryum dichotomum could be confused with B. subapiculatum due to the leaf shape, width of cell walls, and scarcely bordered leaf margins.However, B. subapiculatum has hexagonal to narrowly romboidal upper leaf cells, rectangular to sub-quadrate basal cells, thin cell walls, conic to cylindric capsules with a slender neck, and lacks vegetative propagules (Fig. 10A-D).Bryum dichotomum is similar to B. coronatum due to the width of the neck.However, the latter has thin cell walls, oblong-cylindric capsules, corrugate neck, and does not produce gemmae (Fig. 5A-D).Bryum dichotomum and B. orthodontioides share the small-sized (1-2 mm high), imbricate, concave and elimbate leaves, and short-excurrent costa.However, this has thin leaf cell walls, capsules with a slender neck, and lacks vegetative propagules (Fig. 7A-D).Bryum dichotomum and B. apiculatum have axillary gemmae, but the latter presents narrowly hexagonal to fusiform leaf upper cells and abruptly larger rectangular to quadrate basal cells (Fig. 2A-E).Plants large-sized to robust, 10-40 (60) mm high, light green to yellowish-green, leaves crisped and spirallytwisted when dry, in dense rosulate tufts, 2-4 distinctly storied rosettes.Leaves obovate to elliptic; apices green, mucronate; margins distinctly bordered by 3-5 rows of narrow, long-rectangular cells, conspicuously serrate at apex, plane or recurved at base; costa strong, short-excurrent; upper cells rhomboidal to rhomboidal-hexagonal, basal cells rectangular, porose, firm-and thin-walled.Dioicous.Capsules green to orange-brown, pendulous to horizontal, cylindric to conic, neck slender; opercula conic-apiculate; peristome complete; exostome well-developed, endostome with segments well-developed, cilia present.

Bryum huillense
Selected This is a very distinct species among the Brazilian taxa of Bryum.It is large to robust in size (10-40 (60) mm high), leaves in dense rosulate tufts with 2-4 storied rosettes, obovate to elliptic leaves with distinctly bordered margins that are serrate at the apex, short-excurrent costa, and porose basal cells.Allen (2002) mentions filamentous axillary propagula in B. huillense.However, we did not observe this character among the Brazilian specimens.African and Asian individuals of this species are up to 3 cm high (vs.up to 6 cm in Neotropical samples) and show less morphological variation than the Neotropical individuals (Ochi 1972;1974).This species can be confused with B. billarderii in having large-sized (15-25 (50) mm high), rosulate leaves, which are obovate, limbate and serrate at the apex.However, B. billarderii is smaller (medium-to large-sized), with leaves evenly spaced and crowded at the apex (when sterile) or in dense rosulate tufts, 1-2 (3) storied rosettes, margins bordered by 1-3 narrow cells, and single seta (Fig. 4A-D).Polysety was observed in a single specimen of B. huillense.
Selected Among the Brazilian Bryum species, B. orthodontioides exhibits the smallest gametophyte (up to 3 mm high) and leaves.This species is also distinguished by the usually imbricate and concave leaves, leaf margins entire and unbordered, short-excurrent costa, lax upper cell walls, and horizontal to suberect capsules.Some sterile individuals of B. orthodontioides present more elongate stems and lax leaf arrangement.After careful analysis of the type material of B. orthodontioides, we noticed that this species is conspecific to B. leptocladon.According to the principle of priority of the International Code of Nomenclature for algae, fungi and plants (Turland et al. 2018), we reduce B. leptocladon to a synonym of B. orthodontioides.
Bryum orthodontioides seems to B. apiculatum due to the lanceolate leaves with cuspidate apex, unbordered margins, short-excurrent costa, and rectangular to sub-quadrate basal cells.However, B. apiculatum has more elongate leaves with serrulate apices, red base, narrowly hexagonal to fusiform upper cells, and abruptly larger, rectangular to quadrate basal cells (Fig. 2A-E).Bryum orthodontioides is similar to B. subapiculatum, but the latter has leaves with serrulate margins, firm cell walls, and pendulous and conic to cylindric capsules (Fig. 10A-D).The species is also close to B. dichotomum because both have small-sized (1-2 mm high), imbricate leaves, and short-excurrent costa.However, B. dichotomum has thick cell walls, broad capsules neck, and vegetative propagules (Fig. 6A-E).
Acta Botanica Brasilica, 2022, 36: e2021abb0283 Acta Botanica Brasilica, 2022, 36: e2021abb0283 confused with B. subapiculatum due to the gametophyte aspect and shape of the leaves.However, this has dioicous inflorescences, and rectangular to sub-quadrate basal leaf cells (Fig. 10A-D).Ochi (1980) pointed out that B. pabstianum and B. subapiculatum have rhizoidal gemmae and the shape and size of the gemmae are a diagnostic feature.This character was not observed in both examined material, therefore we consider the best diagnostic feature is sexuality.Bryum pallescens is also synoicous.However, it has recurved and distinct bordered leaf margins (Fig. 8E-H).The gametophyte aspect and leaf shape of B. pabstianum also resemble B. coronatum.However, the last has ovatelanceolate leaves with long-excurrent costa, oblong-cylindric capsules, and a broad neck (Fig. 5A-D).Plants medium to large-sized, up to 30 mm high, light green to dark green to reddish-green, leaves imbricate, crisped when dry, evenly spaced or in inconspicuously rosulate tufts.Leaves ovate, ovate-lanceolate to elliptic; apices green, acuminate or cuspidate; margins distinctly bordered by 2-4 rows of narrow, long-rectangular cells, entire to serrulate at apex, usually recurved at base; costa strong, short-excurrent; upper cells rhomboidal-hexagonal, thin-walled, basal cells rectangular, non-porose, firm-and thick-walled.Synoicous.Capsules green to orange-brown, pendulous, cylindric to conic, neck slender; opercula conicapiculate; peristome complete; exostome well-developed, endostome with segments well-developed, cilia present.

Bryum pallescens
Selected Bryum pallescens is a synoicous and medium-to largesized (up to 30 mm high) plant with leaves imbricate, crisped, ovate-lanceolate to elliptic, distinctly limbate and recurved margins, short-excurrent costa, and thick basal cell walls.This species is reported for high altitudes (Ochi 1982).Bryum pallescens is very variable in gametophyte size and can also be autoicous, but the most easily distinguishable character is its sexuality (Ochi 1959;Allen 2002).Peralta et al. (2008) and Yano & Peralta (2011a) made the two first records of B. pallescens in Brazil (Pirani 1575 -SP230701!; Yano 21790 -SP274708!, respectively).However, the correct identifications are B. coronatum and B. subapiculatum, respectively.Therefore, we actually present the first record of B. pallescens for the country.
This species is similar to B. capillare by the imbricate and crisped leaves.However, B. pallescens is dioicous, the leaf margins are usually plane, and the upper cell walls are lax (Fig. 4E-I).Bryum pallescens and B. limbatum have distinctly limbate leaves.However, the latter has thin-walled basal cells, dioicous inflorescences and horizontal to sub-erect and pyriform capsules (Fig. 7E-H).Bryum pallescens and B. pabstianum are synoicous.However, the latter has plane and unbordered leaf margins (Fig. 8A-D).Bryum pallescens may be confused with B. pseudotriquetrum (Hedw.)Gaertn.However, the latter has decurrent leaves, percurrent to short-excurrent costa, and dioicous inflorescences (Allen 2002).This species was excluded from Brazil in the present study (see excluded species section).Bryum renauldii is characterized by lax and evenly spaced leaves, which are ovate to elliptic, with obtuse apices, unbordered margins, weak, and evanescent and subpercurrent costa.During the herbaria and fieldwork, we could not find sporophytes, but we did observe sex organs.We analyzed eight specimens for Brazil, of which six were dioicous, one synoicous, and three unknown.Spence (2015) mentioned it is "apparently dioicous".Therefore, this is the first description of the sexuality of B. renauldii (Ochi 1980;1994;Allen 2002).Besides this observation by Spence (2015), previous studies mentioned that the sporophyte is unknown (Ochi 1980;1994;Allen 2002), and we could not find any information about the sporophytic generation of this species.
Selected Bryum riparioides is a large plant (10-20 mm high) and has imbricate, elongate and slightly concave leaves, serrulate, recurved and indistinctly bordered margins, weak, evanescent and sub-percurrent to percurrent costa, and thick-walled basal cells.Bryum riparioides and B. limbatum share the sub-percurrent to percurrent costa, and recurved leaf margins.However, the last has wider leaves (ovate to broadly elliptic), distinctly bordered margins and pyriform capsules (Fig. 7E-H).The species is also similar to Imbribryum alpinum (Huds.ex With.)N. Pedersen, but the latter has conspicuously imbricate, smaller and narrower leaves, with acute apex, and narrow and vermicular upper cells (Spence 2015).Bryum riparioides is commonly confused with Imbribryum muehlenbeckii (Bruch & Schimp.)N. Pedersen.However, the latter has smaller, ovate and conspicuously concave leaves with a decurrent base and weak costa (Ochi 1980;Spence 2015).This species was excluded from Brazil (see excluded species).
This species is endemic to Brazil (Matteri 2003;Larraín 2016).Bryum riparioides was described by Bartram (1952) and recorded for Brazil by Yano (1981;2011) and Bordin & Yano (2010).The name was rarely mentioned in the literature, and after careful analysis, we decided to reestablish the name.
The name was cited for Brazil by Yano (1981) based on the protologue of Müll.Hal. (1900).However, the protologue lacks both a voucher and a diagnosis.The name is insufficiently known (Crosby et al. 1999) and is considered a nom.nud.
It was cited by Yano (1981) and Costa et al. (2005).The last study does not mention the voucher.It was not possible to find the type specimen of B. multiflorum, and information on this taxon is scarce.The species is listed by Costa et al. (2011) in the excluded taxa section.
Fig. 2A-E Geographic distribution: United States, Mexico, Central America, the Caribbean, western, northern, and southern South America; Europe; Asia; Africa; Australia and the Pacific Islands (Allen 2002; Holyoak 2009); and Brazil (wide
, which was not observed in the Brazilian specimens.