Efrain De Luna,

Biodiversidad y Sistemática,

INECOL. Xalapa. México.



Contents of this website


Phylogenetic classification

Hedwigiaceae Taxonomic treatment

Keys to genera




Pseudobraunia (link to Flora of North America)



BRAUNIA Bruch & Schimper. Bryol. Eur. 3: 159 (fasc. 29-30). 1846.

Type: Braunia sciuroides (Bals.-Criv. & De Notaris) Bruch & Schimper, (= Anoectangium sciuroides Bals.-Criv. & De Notaris. 1836).

Nom. cons. (De Luna 1999).


Description. Robust plants in loose or dense, dull, rigid tufts or mats, dark green or red-brown. Stems sympodially branched, plagiotropic, tips ascending; most branches terete and blunt, some stoloniform. Pseudoparaphyllia foliose, often deeply dentate, cells papillose.

Leaves imbricate, spreading or squarrose, concave, plicate, ovate to wide oblong, apiculate, cuspidate or long acuminate; ecostate; apex concolorous or hyaline, crenulate or serrulate; margins reflexed below or revolute up to the base of acumen; upper cells thick-walled, strongly sinuous, subquadrate or rectangular (28-40 mm long,  8-12 mm wide), with several low rounded, unbranched papillae marginal, overarching to the lumen; apical cells broad oblong to narrowly elongate, smooth or papillose; basal cells, linear, yellow, cell walls straight, porose, with 3-5 luminal, unbranched papillae or smooth, gradually broad elliptic to subquadrate, lumina subcruciform at basal margins.

Autoicous or rarely synoicous; perichaetia terminal on sympodial stems, becoming lateral because of innovations, perichaetial leaves erect, elongate, plicate, long acuminate to cuspidate; paraphyses uniseriate, long, yellowish to strong orange, commonly exserted.

Setae yellow to reddish, short to elongate, straight or twisted near the neck, smooth; neck very short, ampullaceous or long, attenuated; capsule emergent to exserted, erect, symmetric, rarely inclined, urceolate, subglobose, broadly ellipsoid to cylindric, smooth or longitudinally ribbed, slightly constricted below the rim, commonly microstomous; operculum base conic, rostrum slender, long, oblique; exothecial cells broadl subquadrate, rectangular or polygonal, thick walled, gradually subquadrate or oblate and darker in several suboral rows; annulus and peristome absent; stomata superficial, restricted to neck.

Spores yellow-brown, vermiculate papillose or rugulose, unicellular, weakly tetrahedral, or pluricellular.

Calyptrae large, cucullate, smooth or rarely hairy, beak narrow, long, base broad open or strongly concave, naked, 4-7 cell layers thick.

Species: 23.

Nomenclatural notes: The type specimen of the earliest described species now included in Braunia was collected in central México (Humboldt and Bonpland s.n.) and it was named in 1818 as Hedwigia secunda Hook.

Bruch and Schimper (Bruch et al, 1846) established Braunia as a new genus in the Hedwigiaceae. They classified three species: B. secunda from Mexico, B. schimperi Bruch. & Schimper from Africa, and B. sciuroides (Balsamo & De Notaris) Bruch. & Schimper from Europe.

In his insightful treatment of the genus, Jaeger (1876) accepted 11 species: B. secunda, B. sciuroides, B. andrieuxii Lor., B. schimperi, B. squarrulosa (Hampe) Müll. Hal, (as B. liebmanniana (Müll. Hal.) Besch.), B. cirrhifolia (Mitt.) Wils. ex Jaeger, B. plicata (Mitt.) Jaeger, B. attenuata (Mitt.) Jaeger, B. diaphana, B. macropelma, and B. rupestris. At this point, the taxonomic concepts of species within Braunia were clearly delineated.

Brotherus (1909) recognized 24 species in Braunia.

A preliminary cladistic analysis of the worldwide morphological variation in Braunia (De Luna, 1992) provided the basis for the current list of 23 species.

Distribution: The 23 species of Braunia known worldwide (De Luna 1992) are mainly distributed in the mountains of subtropical and tropical regions around the world, in five important geographic regions:

Region 1: There are four species known in the area including the southern United States of America, Mexico, Central America and the Caribbean Islands: Braunia secunda, B. andrieuxii, B. squarrulosa and B. plicata.

Region 2: In the Andes of Colombia, Ecuador, Peru, Bolivia, Chile and Argentina, in South America, there are 12 species known: B. nephelogenes, B. cirrhifolia, B. secunda, B. subplicata, B. plicata, B. canescens, B. incana, B. subincana, B. exserta, B. reflexifolia, B. tucumanensis, and B. argentinica. In this area, there is a revision of the three species of Braunia in Tucuman (Biasuso, 1992).

Region 3: In eastern and western Africa there are six species listed: B. rupestris, B, diaphana, B. arbuscula, B. entodonticarpa, B. camptoclada, and B. schimperi.

Region 4: There is only one species in southern Europe: B. alopecura.

Region 5: In Asia there are three species, B. attenuata and B. alopecura are known from the Himalayan mountains of India, Nepal, and western China (He & De Luna, 2004; Dalton et al., 2013), wheras B. macropelma is abundant in India (listed as B. secunda in Dalton et al., 2013).


Biasuso, A. B. 1992. El género Braunia (Hedwigiaceae, Musci) en Tucumán. Lilloa, 38: 5-21.

Brotherus, V. F. 1909. Hedwigiaceae. In: A. Engler & K. Prantl, eds. Die natürlichen Pflanzenfamilien 1(3): 712–722. W. Engelman, Leipzig.

Bruch, P., Schimper, W. P. & Gümbel, W. v. T. 1846. Bryologia Europaea, seu Genera Muscorum Europaeorum monographice illustrata. Vol. III. E. Schweizerbart, Stuttgart.

Dalton, N.J., Kungu, E. M. & Long, D. G. 2013. A taxonomic revision of Hedwigiaceae Schimp. from the Sino-Himalaya. Journal of Bryology, 35: 96–111.

De Luna E. & W.R. Buck. 1991. An undescribed species of Braunia (Hedwigiaceae) from the Andean cloud forest. The Bryologist 94: 401-403.

De Luna E. 1992. Developmental and systematic studies in the Hedwigiaceae (Musci). PhD thesis. Duke University, Durham, NC.

De Luna, E. 1999. Proposal to conserve the name Braunia Bruch & Schimp. (Musci) against Braunia Hornsch. (Musci). Taxon 48: 385-386.

De Luna E & G. Gómez-Velasco. 2008. Morphometrics and the identification of Braunia andrieuxii and B. secunda (Hedwigiaceae: Bryopsida). Systematic Botany 33(2): 219-228.

De Luna, E. 2009. Typification, taxonomy and distribution of Braunia squarrulosa (Hedwigiaceae) in Mexico and Central America. The Bryologist 112(1): 202–207.

De Luna E. 2016. Typification, taxonomy, and worldwide distribution of Braunia secunda (Hook.) Bruch & Schimp. (Hedwigiaceae). Journal of Bryology 38: 286-294.

He, S. & E. De Luna. 2004. The Genus Braunia (Bryopsida, Hedwigiaceae) in China. The Bryologist 107: 373–376.

Jaeger A. 1876. Genera et species muscorum systematice disposita seu Adumbratio florae muscorum totius orbis terrarum. VI, p 85-188.

Sharp, A. J., Gardner, R. & Giesy, R. M. 1978. SEM separation of sterile specimens of Hedwigidium integrifolium, Braunia secunda and B. squarrulosa. Phytomorphology, 28: 329-331.



Extensive mats of Braunia growing on rocks. Camino a Valle Colorado, Depto, Valle Grande, Jujuy, Argentina (photo courtesy of Beatriz Biasuso, Tucumán)

Braunia secunda, on volcanic rock. México


Braunia andrieuxii

Braunia secunda


Upper leaf cells, Braunia squarrulosa.


Braunia exserta

Braunia plicata

Braunia plicata


Braunia specimens at MEXU

Braunia specimens at FRULLANIA collaborative research network (US herbaria)

Braunia specimens at the MNHN