BFNA Title: Brachymenium
Author: J. R. Spence 
Date: August 25, 2011
Edit Level: R 
Version: 2

Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden
BFNA Web site: http://www.mobot.org/plantscience/BFNA/bfnamenu.htm

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XX. BRACHYMENIUM Schwägrichen, Sp. Musc. Frond. Suppl. 2(1): 131. 1824 * [Latin brachy, short, and menium, a membrane, alluding to poor development of endostome]

John R. Spence

 

Plants small, forming dense turfs or cushions, green to red-green. Stems short, 0.3--1.5 cm, sometimes rosulate, sparsely to strongly branching by subfloral innovations, rhizoids sparse to abundant, variously colored, smooth to papillose, micronemata often present on stems. Leaves irregularly contorted to spirally twisted around stem when dry, erect-spreading when wet, broadly ovate, 1--2.5 mm, smaller proximally, leaf base not curved at insertion, not or only weakly decurrent; margins plane or revolute proximally, 1-stratose, limbidium absent or present, distal margins serrulate to serrate, apex acute to broadly rounded, differentiated apiculus absent, costa shortly to moderately long-excurrent in a stout, denticulate, colored awn, transverse section with stereid band single and well developed, guide cells present; adaxial supracostal cells irregularly to regularly quadrate or short-rectangular at leaf base, laminal cells heterogeneous, distal and medial cells generally similar, short-rectangular to irregularly hexagonal, 1.5--3(--4):1, not in rows oblique to the costa, thin- to somewhat thick-walled, proximal cells usually quadrate, distinctly different in shape, walls not pitted, alar cells usually similar to juxtacostal cells. Specialized asexual reproduction by spherical rhizoidal tubers and bulbils in leaf axils. Sexual condition dioicous; perigonia and perichaetia terminal, perichaetial leaves same size as vegetative leaves or typically larger, sometimes forming a rosette, inner leaves highly differentiated, often narrower with a weaker costa. Seta single, variously colored, elongate, straight. Capsule erect to suberect, ovate to cylindric, wrinkled when mature, hypophysis slender to well-differentiated, often rugose, operculum short to tall-conic; peristome double, exostome red-brown proximally, pale distally, papillose, teeth linear-lanceolate, separated at base, endostome of basal membrane hyaline to pale yellow, low, segments absent or short and blunt, cilia rudimentary or absent. Spores shed singly, 12--18 \um, smooth or papillose, pale tan to brown.

 

Species ca. 30 (2 in the flora): worldwide in tropical and subtropical regions, primarily Paleotropical.

 

The genus Brachymenium was originally based on the erect capsule with a variously reduced peristome, but recent research indicates the genus to be highly polyphyletic.  Apparently this sporophytic syndrome has evolved independently several times in groups with distinctive gametophytes.  I have used gameophyte morphology as the basis for re-assigning sections and species to other genera. Typical Brachymenium consists of subtropical-tropical epiphytes with gametophytes similar to those of Rosulabryum. Brachymenium andersonii has been re-assigned to Rosulabryum, while the small species B. exile Dozy & Molkenboer is now placed in Gemmabryum near the G. bicolor complex.  Brachymenium niveum and B. systylium have been transferred to the newly erected genus Leptostomopsis, of which the latter is the type. The only remaining North American species, B. macrocarpum and B. mexicanum, are probably not closely related to the type section of Brachymenium.  H. Ochi (1980) placed B. macrocaprum in sect. Peromnium, along with B. regenellii Hampe and B. jamesonii Taylor. This group is similar to some species of Gemmabryum, especially the Paleotropical G. coarctatum (Bosch & Sande Lacoste) J. R. Spence & H. A. Ramsay, but differ in the presence of bulbils and strongly rosulate and twisted leaves.  Because of uncertainty in the phylogenetic relationships of the section, I have tentatively retained B. macrocarpum in Brachymenium pending further study. The species B. mexicanum seems to be related to B. macrocarpum, although also showing similarities to species of the type section. The genus description above is largely based on Brachymenium sect. Peromnium (Mitten) Brotherus.

 

SELECTED REFERENCES: Ochi, H. 1980. A revision of the Neotropical Bryoideae, Part 1. J. Faculty Educ. Tottori Univ., Nat. Sci. 29: 49--154.  Spence, J. R. 2006. New combinations in the Bryaceae (Bryophyta) for North America. II. Phytologia 89: 110--114.

 

 

1. Leaves strongly spirally twisted around stem when dry, limbidium present; large red spherical tubers present in sterile material ......................................................................1. Brachymenium macrocarpum

1. Leaves imbricate to irregularly contorted when dry, limbidium absent; tubers lacking ....................................................................................................................2. Brachymenium mexicanum

 

 

1. Brachymenium macrocarpum Cardot, Rev. Bryol. 38: 6. 1911

 

Plants small, forming dense dark green to olive green cushions, not shiny. Stems 1--2 cm, strongly rosulate, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller.  Leaves ovate, 0.6--1.5 mm,  spirally twisted around stem when dry, forming rosettes when wet, somewhat concave, margins weakly revolute proximally, plane distally, distinctly serrate, limbidium usually present, of 1--2 rows of somewhat elongate incrassate cells, apex broadly rounded-acute; costa short-excurrent into a short stout somewhat yellowish-hyaline denticulate hairpoint; distal and median laminal cells hexagonal to short-rectangular, 20--50 x 15--20 \um, 2--3:1, proximal cells quadrate,  Specialized asexual reproduction of  rhizoidal tubers, spherical, 100--300 \um, red to orange-red, commonly on rhizoids at base of stem. Sexual condition dioicous. Seta 1--2 cm, light brown to red-brown. Capsule 1--3 mm, elongate-ovate to subcylindric, distinctly wrinkled when dry.

 

Capsules mature spring--summer (Apr--Aug). Uncommon on tree trunks, wood and occasionally rock, rarely on mineral soil; 0--1000 m; Ariz., Fla., La., N.Mex., N.C., Tex; Mexico.

 

Brachymenium macrocarpum is a distinctive species that, within its range, should be easy to identify. It forms dense turfs or cushions on trees and rocks.  The very similar B. klotzschii (Schwägrichen) Paris is found to the south in Central and South America.  Many workers lump the two, but there are differences in capsule morphology, leaf shape, lamina areolation, and asexual reproduction, so I have retained the name B. macrocarpum here, based on the Mexican type.  In Mexico, the species is known to produce bulbils in leaf axils, but I have not seen these in material from the flora area. Rosulabryum andicola is similar and its range overlaps with B. macrocarpum, but it has strongly serrate upper margins with very strong limbidium, filiform gemmae in leaf axils, and nodding capsules with a well developed peristome.

 

 

2. Brachymenium mexicanum Montagne, Ann. des Sci. Natur., Bot., sér. 2, 9: 54. 1838

 

Plants in open to dense turfs on soil or rock, green, yellow-green, to golden, often shiny-lustrous. Stems 0.5--1(--2) cm, gemmiform to somewhat rosulate, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller. Leaves 1--2(--3) mm, strongly imbricate, slightly twisted when dry, concave, not decurrent, apex acute, margins plane to weakly recurved proximally, distal margins smooth to serrulate, limbidium absent, costa strong, short to long-excurrent in an awn, distal and mid-laminal cells rhomboidal to elongate-hexagonal, 40--60 x 10--16 \um, mostly 3--4:1, moderately incrassate, proximal laminal cells abruptly quadrate to short-rectangular, 1--2:1, wider and shorter than cells above. Specialized asexual reproduction lacking.  Sexuality dioicous. [Capsule 2--3(--4) mm, erect, pyriform to cylindric with somewhat thickened and rugose apophysis; peristome double, reduced, endsotome low, segments and cilia absent or rudimentary.]

 

Capsules unknown in flora region. Rare on dry rock outcrops or soil covered crevices, 1000 m;  AZ; Mexico, Central America.

 

Brachymenium mexicanum is distinguished from B. macrocarpum by the irregularly contorted leaves, lack of a limbidium, and absence of rhizoidal tubers. The sporophyte description is based on Mexican material.