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Introduction
The Hedwigiaceae consist of four genera of acrocarpous and mostly epilithic mosses (De Luna, 1995) currently classified in the order Hedwigiales. The monotypic genus Pseudobraunia (Lesq. & James) Broth. is endemic to the western side of North America in the USA and Canada. The other three genera are present in Mexico and Central America (De Luna, 1995). In this geographical area, the genus Braunia Bruch & Schimp. can be set apart by the presence of exserted capsules on a long seta (3-20 mm), whereas the seta is very short (0.7-1.5 mm) in Hedwigidium Bruch & Schimp. and Hedwigia P. Beauv. An immersed sporophyte, surrounded by very long perichaetial leaves without cilia, helps to distinguish Hedwigidium reliably (Allen, 2010; De Luna, 2021a). The whitish leaves and hyaline leaf apices easily identify species of Hedwigia. However, when the sporophyte distinction cannot be used, Hedwigidium imberbe (Sm.) Bruch & Schimp. and the species of Braunia present in Mexico are very difficult to identify, since leaf shape and leaf cell papillae are so similar (Sharp et al., 1978). The genus Braunia is characterized by the ecostate leaves, thick-walled, sinuous leaf cells with multiple papillae over the longitudinal walls, long setae, eperistomate and usually cylindrical capsules, cucullate calyptrae, and an exosporic globular protonema (De Luna, 1990, 1995, 2016). In the area including the southern United States of America, Mexico, Central America and the Caribbean Islands, there are four species of Braunia known: Braunia secunda (Hook.) Bruch & Schimp., B. andrieuxii Lorentz, B. squarrulosa (Hampe) Müll. Hal., and B. plicata (Mitt.) A. Jaeger.
Braunia was established as a genus in the Hedwigiaceae (Bruch et al., 1846) to include the European species, Braunia sciuroides (Bals.-Criv. & De Not.) Bruch & Schimp. (= B. alopecura (Brid.) Limpr.), and two species from Mexico (B. secunda and B. squarrulosa). Later, Müller (1851) described Neckera sphaerocarpa Müll. Hal. and N. liebmanniana Müll. Hal.; eventually both species were added to Braunia. Subsequently, Lorentz (1864) described yet another species from Mexico as B. andrieuxii. In the worldwide catalogue of mosses by Jaeger (1876), only three species were recognized for Mexico: B. secunda, B. andrieuxii, and B. liebmanniana. Early in the 20th century, Brotherus (1909) listed the same three species for Mexico. Shortly after, two additional taxa from Mexico were added to the genus. First, Cardot (1911) described Braunia plicata var. canescens Cardot from two specimens collected by Pringle in central Mexico (Hidalgo and Estado de México). Lastly, Thériot (1926) described B. secunda var. crassiretis Thér. based on a collection by Arsène also in central Mexico (Michoacán).
In modern times, Crum (1951) recognized three species of Braunia in Mexico. He considered B. andrieuxii and B. secunda var. crassiretis as synonyms of B. secunda. Likewise, Crum (1951) treated B. liebmanniana and B. sphaerocarpa as synonyms of B. squarrulosa. Remarkably, he accepted the records of B. plicata as genuine. However, the current understanding of the number of species of Braunia in Mexico has been based on Crum (1994); he re-interpreted the morphological diversity of Braunia in this area as consisting of just two species: B. secunda and B. squarrulosa, suggesting that "Mexican records of B. plicata (Mitt) Jaeger, a South American species, can perhaps also be included in B. secunda." A taxonomic study of the species of Braunia in Mexico had not been undertaken before. A taxonomic treatment and a key to the species of Braunia in Mexico is presented, including illustrations, maps and a summary of the known distribution. This treatment should be useful also for the species of Braunia in the southern United States of America, Central America and the Caribbean Islands.
Material and Methods
In this paper the four Braunia species known in Mexico and neighboring areas in the southern United States of America, Central America and the Caribbean Islands were studied together for the first time. Specimens of over 450 collections from the following herbaria were included: B, BM, DUKE, F, FH, FLA, G, H, JE, LPA, MEXU, MICH, MO, NY, S, US, and XAL (Thiers, 2016). Species distribution maps were prepared with the ZeeMaps software (Zee Source, 2021). Also, examination of original descriptions, protologues, and type specimens in several herbaria worldwide revealed the need of lectotypification for one name. The necessary lectotypification is documented for B. andrieuxii.
Results
Taxonomy
In order to facilitate identification of the four genera of Hedwigiaceae, here I offer the following key.
Key to the genera of Hedwigiaceae
1a. Medial and upper leaf cells with 1-4 papillae per cell, papillae tall, centered over cell lumen; gametophytes without stoloniferous branches ....... 2
1b. Medial and upper leaf cells with 1-6 papillae per cell, papillae low, marginal and arching over cell lumen; gametophytes with stoloniferous branches ....... 3
2a. Seta short, (0.5-)0.8-1(-1.2) mm, capsules immersed; operculum low, plano-convex, rostrum small mammillate; capsule stomata cryptoporic; perichaetial leaf margins ciliate ......... Hedwigia P. Beauv.
2b. Seta long, (4-)6-7(-10) mm, capsules exserted; operculum high-conic, short-rostrate; capsule stomata phaneroporic; perichaetial leaf margins entire ....... Pseudobraunia (Lesq. & James) Broth.
3a. Capsules immersed, urn turbinate-urceolate, subglobose, mouth wide; operculum high-conic, rostrum small mammillate; perichaetial leaf margins undulate; seta short, (0.8-)1-3(-4) mm ......... Hedwigidium Bruch & Schimp.
3b. Capsules exserted, urn cylindric, ellipsoid, ovate, or turbinate, mouth wide or narrow; operculum low-conic, long-rostrate; perichaetial leaf margins not undulate; seta long, (4-)7-12(-17) mm ............. Braunia Bruch & Schimp.
Braunia Bruch & Schimp., Bryol. Eur. 3: 159. 1846. (nom. cons., De Luna, 1999).
TYPE: Braunia sciuroides (Bals.-Criv. & De Not.) Bruch & Schimp. (= Anoectangium sciuroides Bals.-Criv. & De Not. 1836).
Hedwigia sect. Braunia (Bruch & Schimp.) Mitt., J. Linn. Soc., Bot. 12: 404. 1869.
Braunia sect. Macromidium Müll.Hal., Linnaea 42: 380. 1879.
Hedwigia subg. Braunia (Bruch & Schimp.) Kindb., Eur. N. Amer. Bryineae (Mosses) 1: 5. 1897.
Robust plants (3-5 cm tall) 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, cuspidate or acuminate; leaves ecostate; leaf apex concolorous or hyaline, crenulate or serrulate; margins reflexed below or revolute up to the base of acumen; upper leaf cells thick-walled, strongly sinuous, subquadrate or rectangular (28-40 µm long, 8-12 µm wide), with several low rounded, unbranched papillae, marginal, overarching to the lumen; apical cells broad oblong to narrowly elongate, smooth or papillose; leaf basal cells, linear, yellow, cell walls straight, porose, with 3-5 luminal, unbranched papillae or smooth, gradually broad elliptic to subquadrate, cell 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 deep orange, commonly exserted; setae yellow to reddish, short to elongate, straight or twisted near the neck, smooth; neck very short, ampullaceous or long, attenuate; capsule 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 broadly 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.
Morphological features most likely to be informative for the distinction of the four species in Mexico are the leaf margins, leaf apices, upper leaf cells, and sporophytes. For example, a detailed morphometric analysis of variation helped to reveal differences between Braunia secunda and B. andrieuxii, in the extent of the revolute leaf margins and upper leaf cell lengths (De Luna and Gómez-Velasco, 2008). The seta is short, up to 5-6 mm long, and the capsule is widely urceolate only in B. squarrulosa. However, sporophytes are of no use for distinguishing B. andrieuxii, B. secunda, and B. plicata, because of overlapping variation in cylindrical to ellipsoid capsule shape and seta lengths (9-14 mm). Without sporophytes, the leaf apex, and the upper leaf cells help to discriminate species. The leaf apex is hyaline and dentate only in B. plicata, while it is concolorous and entire or scabrous in the other three species of Braunia from Mexico. Also, the apical cells are strongly papillose only in B. plicata. The upper leaf cells are quadrate to short-rectangular in B. andrieuxii and B. plicata; whereas the upper leaf cells are elongate in B. secunda and B. squarrulosa.
The morphological features listed above are emphasized in the following key. Since the four species of Braunia present in Mexico are distributed also in the Southern United States of America, Central America and the Caribbean, this key should be useful to identify species of Braunia in those areas.
Key to the species of Braunia in Mexico
1a. Leaf apex dentate, hyaline ; capsule cylindrical …. Braunia plicata (Mitt.) A. Jaeger
1b. Leaf apex entire or scabrous, concolorous ……2
2a. Leaf apex long-acuminate, narrow, flexuose; capsule turbinate, globose; setae (3-)5-6(-7) mm long …. Braunia squarrulosa (Hampe) Müll. Hal.
2b. Leaf apex short-acuminate; capsule cylindrical, narrow; setae (5-)9-14(-20) mm long …. 3
3a. Leaf margin revolute, extending from the base up 3/4 the leaf length; upper leaf cells rectangular, (8.2-)11-16.8(-21.9) (m long, (3.7-)4.9-6.5(-8.2) (m wide, 2-3:1, cell walls sinuose ……….. Braunia secunda (Hook.) Bruch & Schimp.
3b. Leaf margin plane, or revolute, extending from the base up 1/3 the leaf length; upper leaf cells sub-quadrate, (7.2-)9.2-13 (-17.2) (m long, (4.4-)5.3-6.7(-8) (m wide, 1.5-2:1, cell walls smooth …… Braunia andrieuxii Lorentz
Braunia andrieuxii Lorentz, Moosst. 164. 1864. Figs. 1,2.
Figure 1: Lectotype of Braunia andrieuxii Lorentz. A. lectotype specimen at Hooker herbarium (BM), Andrieux 23, Herb. Hooker-BM000960769; B. detail of specimen showing elongate setae and elliptic capsules; C-D. vegetative leaves; E. leaf cells at the apex; F. upper leaf cells (A-F, from Andrieux 23, Herb. Hooker-BM).
Figure 2: Morphological features of Braunia andrieuxii Lorentz in Mexico. A. branching system with a sporophyte; B. detail of a branch showing leaf stance; C. vegetative leaf; D. perichaetial leaf; E. upper leaf cells (A-E, from Cárdenas 348, MEXU, Mexico, Zacatecas).
≡ Braunia secunda (Hook.) Bruch & Schimp. var. andrieuxii (Lor.) Thér., Smith. Misc. Coll. 78(2): 24. 1926. Type citation: “Mexico, in monte St. Felipe prope Oaxacam leg. Andrieux”. TYPE (Lectotype designated here): MEXICO. Oaxaca, in monte San Felipe, prope Oaxacam, IV. 1834, Andrieux 23 (Herb. Hooker-BM000960769!; isolectotypes, BM000960761!, BM000960765!, BM000960766!; H!, Herb. Mitten-NY00913938!, NY00913939!).
Plants medium-sized in dense mats; stems with sympodial branching, plagiotropic, (3-)4-5(-6) cm long, scarcely branched; branches short, erect; flagelliform branches rare; pseudoparaphyllia foliose, with papillose cells; leaves ovate, widest point at or above middle of leaf; acumen abruptly differentiated, to 1/6 length of leaf, short-acuminate, concolorous, 1.7-1.9 × 0.9-1.2 mm, leaf lamina flat or slightly concave, imbricate, erect-patent when dry, spreading when moist, weakly plicate; margins reflexed or recurved from base up 1/2 the leaf length or less, margin plane above (italics show the diagnostic characters), serrulate to erose at acumen; apical leaf cells 13-22 µm long, elliptic (3-4:1), sinuous, with few low marginal papillae; upper and median leaf cells (7.2-)9.2-13(-17.2) µm long, quadrate to rectangular (2-3:1), walls weakly sinuose, papillae low, on side walls, overarching lumina; autoicous; archegonia distal on long vaginula (1.3-1.5 mm); perichaetial leaves oblong lanceolate, 2.3-2.7 mm long, slightly longer than vegetative leaves, acuminate, plicate; perichaetial paraphyses biseriate at base, 2-3 times longer than archegonia, immersed; setae (5-)7-11(-13) mm long; capsule exserted, necks 0.3-0.4 mm long, gradually differentiated, slender, urns 1.2-1.6 mm long, cylindrical, mouth narrow, nearly 1/2 the urn width, capsule walls smooth, or wrinkled when dry, exothecial cells polygonal; stomates superficial; opercula base conical, short obliquely rostrate; calyptrae cucullate, large, 2.8-3.7 mm, hairy; spores unicellular, 25-32 µm in diameter.
Nomenclatural note: There is a single collection, “Andrieux”, cited in the protologue, although without collection number and year. Among several “Andrieux” specimens currently available, there is no certainty as to which particular collection is the type. I examined several “Andrieux” specimens collected in Mexico without collection numbers, or without collection dates (BM000960759, BM000960760, BM000960767, BM000960768); the identity of these specimens is indeed B. andrieuxii. Some of these specimens are from Oaxaca, but there is no additional information to be sure if they are part of the original material. Therefore, here I am designating as lectotype a specimen located at the Hooker herbarium (“Andrieux 23”, BM000960769), clearly labeled as collected in the type locality (Fig. 1). The selection of this numbered specimen as lectotype facilitates the future recognition of additional duplicates in other herbaria. The duplicate of Andrieux 23 in the Mitten herbarium (NY) consists of three small packets, two are good isotypes of B. andrieuxii, but the identity of the third plant (NY00913937) is actually B. secunda. Other specimens of B. andrieuxii collected by Andrieux are very likely not types, since these are not from Oaxaca (Mexico, Santa Fé, Andrieux 18, Herb. Hampe-BM) or there is no precise locality information (Mexique, 1834, Andrieux s.n., Herb. Mus. Paris-BM; Mexico, Andrieux s.n., Herb. Besch.-BM). I have seen an additional non-type specimen collected by Andrieux but the identity is B. secunda (Mexico, Pedregal, Andrieux 20, Herb. Hampe-BM).
Habitat: this species commonly grows on igneous rocks, in exposed habitats and within forests at middle elevations (1300-2500 m).
Distribution: United States of America (Arizona, New Mexico, fide Allred et al., 2016, Texas), Mexico, Guatemala (Allen, 2010), Colombia (as B. secunda in Churchill and Linares, 1995). Fig. 3.
Figure 3: Distribution map of Braunia andrieuxii Lorentz. Each dot is one of the specimens listed in the text.
Illustrations: Bartram (1949, Pl. 106 D-F, as B. secunda); Crum (1994, Fig. 497 a-f, as B. secunda); Churchill and Linares (1995, Fig. 100h-k, as B. secunda); Figs. 1-2.
Selected specimens examined: MEXICO. Aguascalientes, Sierra Fría, Cantrall 54 (MO). Baja California Sur, Sierra La Laguna, S of Pico La Aguja, D. E. Breedlove and D. Axelrod 43286 (G); Sierra de La Laguna, 15 miles E of Todos Los Santos, F. Bowers 5166 (DUKE, MO, XAL); F. Bowers 5201-a (MO); Sierra de la Laguna, C. Delgadillo 3091 (MEXU). Chihuahua, 20 mi N of Las Nieves, F. Bowers 6296-b (DUKE, MO). Municipio Urique between Basigochic and Quirire, R. Bye 6129 (MO); 38 mi S de Creel, Río Urique, Cañón del Cobre, C. Delgadillo 3268 (MEXU); San Pedro Springs, L. N. Godding 2123 (MO); 15 mi WSW of Madera, G. L. Nesom 52 (DUKE, MO); 15 mi WSW of Madera, G. L. Nesom 53 (DUKE, MO); 30 mi W of the La Junta, K. J. Torke 1015 (MO). Ciudad de México (formerly Distrito Federal), Santa Fé, Andrieux 18 (BM); Santa Ana, hacia el Mirador, A. Cárdenas 1438 (DUKE, MEXU); Pedregal de San Angel, C. Delgadillo 1642 (MEXU); C. Delgadillo 2085 (MEXU); near Contreras, A. J. Sharp 1189 (MO, US). Durango, 34.5 mi E of El Salto on Rt. 40, T. Stuessy 409 (DUKE, NY); 45 km W of Durango, F. Bowers 5284 (MO); Otinapa, E. D. Palmer 438 (MO); eastern foothills of the Sierra Madre Occidental, Weber 9856 (MO); Tejamen, E. D. Palmer 475 (MO); Rocky ledge, a few km before Palos Colorados W of Durango, A. J. Sharp 1747 (MO); 34 mi E of El Salto on Rt. 40, Irving 409 (MO); 18 km S. de V. Matamoros, C. Delgadillo 3250 (MEXU). Estado de Mexico, 10.5 km SW villa Nicolás Romero, A. Cárdenas 1855 (DUKE); Sierra de Calpulalpan, A. Cárdenas 2308 (MO). Municipio Tepotzotlan, 4 km de Magu, Sierra de Alcaparrosa, A. Cárdenas 3798 (DUKE, MEXU); Sierra de Alcaparrosa, A. Cárdenas 1820 (DUKE, XAL); Municipio Tepotzotlan, 2 km al W de Magu, Cruz 1698 (XAL), 1700 (XAL). Municipio Texcoco, La Purificación, A. Cárdenas 1886 (DUKE). Guanajuato, 5 km SW de Acámbaro, A. Cárdenas 4461 (DUKE). Municipio Jerécuaro, Sierra de los Agustinos, A. Cárdenas 5351 (MO). Guerrero, municipio Taxco de Alarcón, Caverna "La Joya", J. Castrejon 434 (MEXU, MO). Hidalgo, 2 km NW de San Francisco Sarabia, A. Cárdenas 5636 (MEXU, MO); 5 km NE de Almoloya, C. Delgadillo 5393 (MEXU); 60 km S of San Juan del Río gen Tula, R. Düll 122 (JE); 10 km al W de Alfajayucan, González 2696 (XAL); Cuyamaloya, C. G. Pringle 10627 (B, F, FH, G, S). Jalisco, about 58 km W of Jiquilpan, Mich., on the road to Ciudad Guzman, C. Delgadillo 2761 (DUKE, MEXU); Sierra de Bolaños, García s.n. (MO); mountains above (W of) Hacienda Chinampas, R. McVaugh 17033 (DUKE, MO); Guadalajara, C. G. Pringle 10619 (MO); 8 miles N of los Volcanes, A. Whittemore 2127 (DUKE). Michoacán, Andames, Vicinity of Morelia, G. Arsène 4832 (MO); Dos Tetillas, G. Arsène 7658 (BM, MO); Cerro San Miguel, G. Arsène 4869 (BM, DUKE, MO), 5045 (BM, MO), 5050 (DUKE), 5077 (BM), 5104 (BM, MO); San Miguel prope Morelia in civit Michoacán, G. Arsène 3596 (BM, MO); Punguato, G. Arsène 5047 (DUKE, MO); Uripitio, 11 km NE de Maravatio, C. Delgadillo 4892 (MEXU); 13 km NE de Cotija de la Paz, C. Delgadillo 5053 (MEXU); 2 km N de Acuitzio del Canje, C. Delgadillo 5268 (MEXU); 19 km SSW de Jacona, C. Delgadillo 5339 (MEXU); 20 km N de Cherán, C. Delgadillo 5524 (MEXU); 2 km N de Cuamajo, C. Delgadillo 5475 (MEXU); Los Azufres, 3 km al S del campamento CFE, E. De Luna 1765 (XAL); camino al volcán Paricutín, cerca de San Lorenzo, E. De Luna 1779 (XAL), 1781 (XAL); above Carapan, W. Kiener 18328 (MO); Pátzcuaro, C. G. Pringle 742 (JE, MO), 744 (JE, NY, MO). Morelos, neven Autoban zw. Mexico City u. Cuernavaca, ca 50 km ht. Mex, R. Düll 179 (B). Oaxaca, municipio San Juan, Miahuatlán, E. Hunn OAX-1206 (MEXU); El Punto on Highway 175, A. J. Sharp 2281 (MO); Cerro Jecote, 1.5 km W de Magdalena, Cruz 1890 (XAL); 43 km an der Str. F190 Oaxaca - Huajuapan, Eggers 792022 (MEXU); in Monte San Felipe, Andrieux 23 (lectotype BM, isolectotypes H, NY); 6 km al NW de Mitla, C. Delgadillo 73 (MEXU). Puebla, Cerro Tepoxuchitl, vicinity of Puebla, G. Arsène 4523 (BM, MO), 4851 (BM, MO); loc. cit., Nicolas 5126 (DUKE); along route 119, 8 miles S, F. J. Hermann 26462 (DUKE). Querétaro, El Batán, Argüelles 1459 (MEXU). Sonora, El rancho del Roble, White 4206 (DUKE); NE of El Tigre, White 4211 (MICH); Bavispe región, El Bilito, White 4811 (DUKE, MICH); Cerro Saguarivo, E of San Bernardo, Pennell s.n. (MO). Veracruz, Jalapa, Wagner s.n., 1896 (JE, S); Los Limones, G. Juárez 929 (MO, XAL). Zacatecas, Cerro de la Bufa, A. Cárdenas 348 (MEXU, MO), 349 (MEXU), 355 (MEXU, MO). Municipio Cerro Gordo, 3 km SE de la colonia Hidalgo, A. Cárdenas 741 (MEXU), 745 (MEXU). Municipio Tlaltenango, Cerro del Moro, 29 km W de Jalpa, A. Cárdenas 830 (MEXU, MO). Municipio Tlaltenango, 37 km W de Jalpa, A. Cárdenas 876 (MEXU). Municipio Valparaíso, 14 km N de Valparaíso, A. Cárdenas 796 (MEXU, MO); 7 km S de la Laguna Grande, A. Cárdenas 3034 (DUKE, MEXU). Municipio Fresnillo, 2 km S de San Juan de los Hornillos, A. Cárdenas 751 (MEXU). Municipio Chalchihuites, 2 km E de Gualterio, A. Cárdenas 1080 (MEXU); A. Cárdenas 1075 (MEXU). Municipio Jiménez de Teul, Rancho el Madroño, A. Cárdenas 1101 (MEXU); Road to Huejuquilla de Alto, Jal. 1 mi W of the road-junction, R. McVaugh 17766 (MO).
Additional specimens examined: COLOMBIA. Antioquia, Río Chico, près de Bogotá, 18.IX.1906, F. Apollinaire s.n. (G); près de Bogotá, 8.IX.1907, F. Apollinaire s.n. (G); Medellín, Andes, 17.VIII.1911, F. Idinöel s.n. (G); Cundinamarca, San Cristóbal, 19.VIII.1909, F. Apollinaire s.n. (G). GUATEMALA. Quetzaltenango, P. C. Standley 83237 (F, FH, NY#02120995). San Marcos, T. B. Croat 41046 (MO). UNITED STATES OF AMERICA. Arizona, ravine on Red Mt., Patagonia Mts., E. B. Bartram 617 (BM, MO); White House Canyon, Santa Rita Mountains, E. B. Bartram 60 (DUKE, MO), E. B. Bartram 4529 (MO), E. B. Bartram 1849 (MO); Patagonia Mountains, E. B. Bartram 13 (DUKE); Sycamore Canyon near Ruby, I. M. Haring 3277 (MO); Santa Cruz Co., near McBeth Campground near Josephine saddle above Madera, D. Norris 76567 (DUKE, MO); trail from Maple Camp to South Fork Campground SW of Portal, D. Norris 75380 (DUKE); Madera Canyon, W. Schofield 105057 (DUKE, MO); Chiricahua Mts., W. Schofield 105088 (DUKE); Chochise County, Chiricahua National Monument, trail from Sugarloaf to Echo Park, C. Weber B-42783 (MO). New Mexico, Fide Allred et al. (2016). Texas, South Rim Trail to Emory Peak, Chisos Mts, Big Bend Nat. Park, L. E. Anderson B-2879 (DUKE, MO); Travis Mountains, Berkman 121 (DUKE); Brewster County, Chisos Mountains, summit of Emory Peak, Big Bend National Park, R. Magill 2008 (MO); Chisos Mountains, shortcut trail between Laguna Meadow and Boot, R. Magill 1855 (MO); Chisos Mountains, R. Magill 1883 (MO); Jeff Davis County, Madera Canyon, R. Magill 184 (MO); 215 (DUKE, MO); NE side of Mount Livermore, SW of SH 118, R. Magill 152 (MO); Rockpile State Park, SH 166, R. Magill and Hatch 531 (MO); Mount Livermore, Madera, Hinckley 1151 (MO); Madera Canyon near Mt. Livermore, Hinckley 1149 (MO); Chisos Mtns., 80 mi S of Marathon, Pedano 837 (MO); Boot Canyon Trail, A. Stoneburner 1791 (DUKE); 1805 (DUKE); Chisos Mtns, lower Boot Spring Canyon, Warnock s.n. (MO); between Boot Springs and South Rim, Warnock s.n. (MO); at Laguna, Chisos Mtns, Warnock 1561 (DUKE); Chisos Mountains, Emory Peak, T. Wieboldt 8050 (DUKE); Chisos Mountains, trail to Boot Springs, E. Whitehouse 24616 (DUKE, MO).
Discussion: in the original description of B. andrieuxii, Lorentz (1864) correctly compared this species with the European B. alopecura: “B. sciuroidi simillima, differ autem folis perichaetialibus angustioribus, longioribus, …” Indeed, leaf shape in B. andrieuxii and B. alopecura is very similar. This morphological similarity and a misunderstanding of the species concept of B. secunda led Frahm (2013) to believe B. alopecura and B. secunda were the same species. The two figures Frahm (2013, Figs. 4-5) presented as B. secunda are in fact B. andrieuxii. Similarities in upper leaf cells and leaf shape are correct, but as Lorentz (1864) correctly pointed out, the perichaetial leaves in B. andrieuxii are elongate, whereas they are short in B. alopecura, almost the same size as vegetative leaves. In Mexico, B. andrieuxii is distinguished from other species in the genus by the leaf margins plane or reflexed to weakly recurved only at the leaf base and an elliptic capsule with polygonal exothecial cells. The distinctions made by Thériot (1926) between B. andrieuxii and B. secunda were correct. Nevertheless, this species was traditionally considered a synonym of B. secunda. Certainly, both species share a cylindrical to ellipsoidal capsule and a somewhat similar leaf shape. However, in B. andrieuxii the upper leaf margins are plane, the leaf cells are shorter and less sinuous, and the short acumen is sharply differentiated from a broader upper leaf lamina. These differences are most evident in mature leaves close to branch tips.
Braunia plicata (Mitt.) A. Jaeger, Ber. St. Gall. Naturw. Ges. 1874-1875: 171. (Adum. 2: 87). 1876.
≡ Hedwigia plicata Mitt., J. Linn. Soc., Bot. 12: 406-407. 1869. Type citation: “Andes Quitensis, Jameson. In Bolivia, Bridges in herb. Hooker”. TYPE (Lectotype designated by De Luna, 2021b): BOLIVIA. 1846, Bridges s.n. (Herb. Hooker-BM000960770!; isolectotypes, Herb. Wilson-BM!, NY!).
= Braunia plicata var. canescens Cardot, Rev. Bryol. 38: 38. 1911. Type citation: “État de Hidalgo: Cuyamaloya, rochers (Pringle, 1908, no. 10627). État de Mexico: Toluca (Pringle, 1908, no. 15171)”. TYPE (Lectotype designated by De Luna, 2021b): MEXICO. Toluca, 4.VIII.1908, Pringle 15171 (Herb. Cardot-PC0701219!); isolectotypes (FH! (pro parte, the specimen at FH contains mostly plants of B. andrieuxi), NY#00792519!, S!,).
Plants in dense mats; main stems sympodial, plagiotropic, (3-)4-5(-7) cm long, closely branched; branches elongate, ending in obtuse, rosulate tips; flagelliform branches common; pseudoparaphyllia foliose, with papillose cells; leaves ovate or broadly ovate, 1.9-2.2 mm long, 0.8-1.1 mm wide, concave, imbricate, spreading when moist, strongly plicate; margins recurved to acumen; high base, widest point at or above middle of leaf; apex cuspidate, hyaline, margins sharply dentate; apical leaf cells homogeneously elongate, narrow elliptic (5-7:1), 18-28 µm long, not sinuose, with many marginal papillae; upper and median leaf cells 14-18 µm long, sub-quadrate to short-elliptic (2:1), walls sinuose, papillae 2-3, low, on side walls, overarching lumen; autoicous; archegonia medial on short vaginula (1.4-1.6 mm); perichaetial leaves narrowly lanceolate, 2.4-2.9 mm long, less than twice the length of vegetative leaves, covering 1 mm of seta, acuminate, plicate, perichaetial paraphyses biseriate at base, immersed; setae (7-)9-10(-12) mm long; capsule exserted, necks 0.3-0.4 mm, broad, gradually attenuated, urns 1.7-2.5 mm long, narrowly elliptic, mouth narrow, nearly 1/2 as wide as urn, capsule walls smooth, exothecial cells rectangular; stomates superficial; opercula base conical, short oblique rostrate; calyptrae cucullate, large; spores 29-31 µm in diameter.
Habitat: this species commonly grows on trees, but also colonizes andesitic rocks in exposed habitats within forests at middle to high elevations (1500-2700 m).
Distribution: Mexico (Aguascalientes, Estado de México, Hidalgo, Nuevo León, Oaxaca, Querétaro, Tlaxcala), Peru, Bolivia (De Luna, 2021b); Fig. 4.
Figure 4: Distribution map of Braunia plicata (Mitt.) A. Jaeger. Each dot is one of the specimens listed in the text.
Illustrations: De Luna (2021b, figs. 1-4); Fig. 5.
Figure 5: Morphological features of Braunia plicata (Mitt.) A. Jaeger in Mexico. A. branching system with a stoloniform branch and a sporophyte; B. detail of a branch showing leaf stance and leaf plications (A-B, from De Luna 2720, XAL, Mexico, Tlaxcala); C-F. vegetative leaves (C, from Pringle 10627, JE, Mexico, Hidalgo; D, from Pringle 10627, NY, Mexico, Hidalgo; E, from Sharp 433, MICH, Mexico, Tlaxcala; F, from Pringle 15171, NY, Mexico, Toluca); G-H. perichaetial leaves (G, from Pringle 15171, NY, Mexico, Toluca; H, from Pringle 10627, NY, Mexico, Hidalgo); I. upper leaf cells; J. leaf apex cells (I-J, from De Luna 2720, XAL, Mexico, Tlaxcala).
Selected specimens examined: MEXICO. Aguascalientes, San José de Gracia, La Congoja, Villalobos 8 (MEXU, NY). Estado de México, San Martín de las Pirámides, Cerro Gordo, B. Allen 12943 (MO); Cerro Gordo, A. Cárdenas 2200 (DUKE). Toluca, C. G. Pringle 15171 (FH pro parte, NY, S). Hidalgo, Cuyamaloya, C. G. Pringle 10627 (BM, H, JE, MO, NY, S, US); ridge above Real del Monte, near Pachuca, A. J. Sharp 797 (MICH, US); S of San Juan del Rio, Rauh and R. Düll 122 (H). Nuevo León, Cerro del Viejo, 15 mi W Dulces Nombres, Meyer and Rogers 149 (G, NY); 3 mi NW of Picacho Onofre, G. L. Nesom and Wells 34 (DUKE). Oaxaca, procedente de Trellan, C. Conzatti 4739b (F). Querétaro, cerro El Pinguical, cerca de Pinal de Amoles, G. Ocampo 620 (XAL). Tlaxcala, cerca de Villareal, al NE de Huamantla, E. De Luna 2646 (XAL); 2718 (XAL); 2720 (XAL); Mt. Tlacuapanga, A. J. Sharp 433 (MICH).
Additional specimens examined: BOLIVIA. 1846, Bridges s.n. (BM, NY); alrededores de Sorata, E. De Luna 2094 (DUKE, LPB); in market at Cochabamba, F. J. Hermann 25280 (NY); Santa Cruz, Comarapa und Abra de Catalina, T. Herzog 3703 (JE, S); in der Cordillere von Santa Cruz, T. Herzog 3482 (JE, S); Carasani and Niño Karine, NNW of Chuma, M. Lewis 79-947 (F). PERU. Cusco, Calca, Pisac, Marin 622 (F); Laguna Urpicancha, near Huacarpay, H. Inoue 828 (JE).
Discussion: This is the same species concept from that of Mitten (1869). Morphological characters distinguishing B. plicata in Mexico are the leaves imbricate, strongly plicate, and concave (Fig. 5B). The leaf apex is cuspidate, hyaline, and dentate (Fig. 5J). Shoots are often dark green to brownish, especially darker green along the leaf plications, which contrasts sharply with the hyaline leaf acumens (Fig. 5A-B). Braunia plicata is mostly montane Andean. In Mexico B. plicata is not as frequent as B. secunda or B. andrieuxii, but it is common in Peru and Bolivia.
Braunia secunda (Hook.) Bruch & Schimp., Bryol. Eur. 3 (fasc. 29-30): 161. 1846.
≡ Hedwigia secunda Hook., Musci Exot. 1: 46. 1818 (Anoectangium secundum Brid., Bryol. Univ. 2: 165. 18. 1827, nom. inval. Harrisonia secunda Spreng., Syst. Veg. IV, I: 145. 1827, nom inval.); ≡ Neckera secunda (Hook.) Müll. Hal., Syn. Musc. Frond. 2: 103. 1851. Type citation: "In regni Mexicani montosis apricis, juxta Toluccam ad radices montis perpetua nive obtecti, regione frigida, altitudine 1640 hexapodarum". TYPE (Lectotype designated in De Luna, 2016): MEXICO. In Nevado de Toluca, leg. Humboldt & Bonpland s.n. (Herb. Hooker no. H.1784-BM000960762!; isolectotypes, herb. Hampe-BM!, NY!, P!).
= Braunia secunda var. crassiretis Thér., Smiths. Misc. Coll. 78(2): 24. 1926. Type citation: “(Mexico), Morelia, Campanario (Arsène) (7448)”. TYPE (Lectotype designated in De Luna, 2016): MEXICO. Vicinity of Morelia, Campanario, 2200 m, XI.1911, leg. Arsène 7448 (BM001226211!; isolectotypes, DUKE!, F!, US!).
Not: Anoectangium secundum Mont., Musc. Neelgher. Ann. Sci. Nat. Bot. Ser. 2, t17: 251. 1842. 32. Patria. India Orientalis, in montibus Neelgheriensibus, ad rupes basalticas prope Kaitie lectum = Braunia macropelma (Müll. Hal.) A. Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Ges. Gen. Sp. Musc. 2: 87. 1876.
Plants in dense mats; main stems sympodial, plagiotropic, (3-)4-5(-6) cm long, scarcely branched; branches short; flagelliform branches rare; pseudoparaphyllia foliose, with papillose cells; leaves ovate or ovate-lanceolate, 1.9-2.2 mm long, 0.8-1.1 mm wide, flat or slightly concave, imbricate, erecto-patent, spreading erect when moist, weakly plicate; margins recurved extending from the base up 3/4 the leaf length, sometimes reflexed; low base, widest point below middle of leaf; apex gradually differentiated, short acuminate to aristate, concolorous, margins erose-dentate; apical leaf cells mixed short (3-4:1) and narrow elliptic (5-7:1), 18-28 μm long, sinuose, with few low marginal papillae; upper and median leaf cells 14-20 μm long, elliptic (3-5:1), walls nodulose, sinuose, papillae low, on side walls, overarching lumen; paroicous; archegonia distal on long vaginula (1.4-1.6 mm); perichaetial leaves oblong lanceolate, 2.4-2.9 mm long, longer than vegetative leaves, covering 2 mm of seta, acuminate, plicate, perichaetial paraphyses biseriate at base, exserted; setae (8-)13-15(-20) mm long; capsules exserted, necks 0.3-0.4 mm long, gradually differentiated, broad, urns 1.7-2.5 mm long, narrowly elliptic to broadly ellipsoid, mouth narrow, nearly 1/2 as wide as urn, capsule walls smooth, or wrinkled when dry, exothecial cells polygonal; stomata superficial; opercula base conical, long oblique rostrate; calyptrae cucullate, large; spores unicellular, 29-31 μm in diameter.
Habitat: this species commonly grows on igneous rocks, in exposed habitats and within forests at middle elevations (1300-2500 m).
Distribution: United States of America (Arizona, Texas), Mexico (Chiapas, Chihuahua, Ciudad de México, Durango, Estado de México, Guanajuato, Hidalgo, Jalisco, Michoacán, Morelos, Nuevo León, Oaxaca, Puebla, Tlaxcala, Veracruz), Guatemala, Dominican Republic, Haiti, Venezuela, Bolivia; Fig. 6.
Figure 6: Distribution map of Braunia secunda (Hook.) Bruch & Schimp. Each dot is one of the specimens listed in the text.
Figure 7: Morphological features of Braunia secunda (Hook.) Bruch & Schimp. in Mexico. A. detail of a branch showing leaf stance; B. upper half of vegetative leaf; C. upper leaf cells; D. leaf cells at the apex. (A-D, from De Luna 2643, XAL, Mexico, Tlaxcala).
Illustrations: De Luna (2016, Figs. 1-2); Fig. 7.
Selected specimens examined: MEXICO. Chiapas, municipio of Motozintla, D. E. Breedlove 25803 (MO); Barrio de San Felipe de Ecatepec, G. Gómez-Velasco 277 (XAL); 2 km N of San Cristóbal de Las Casas, den Held et al. HM34 (MO). Chihuahua, Ejido de Bocyna, W of Creel, R. Bye 7306 (MO); 30 mi N de San Juanito, C. Delgadillo 3302b (MEXU). Ciudad de México (formerly Distrito Federal), Santa Rosa, Amable 1510 (BM); Pedregal, Andrieux 20 (Herb. Hampe-BM); Desierta Vieja, 1860, Bourgeau s.n. (BM, NY); Pedregal, Valle de México, Bourgeau 687 (BM); Ajusco, 1868, Hahn s.n. (BM); Cima, C. G. Pringle 10549 (BM, MO); Contreras, 4˚ Dinamo, M. Vivas 307 (DUKE). Durango, 9 miles W of La Ciudad, F. Bowers 5067 (MO, XAL); Hwy. 40 about 9 mi W of La Ciudad. F. Bowers 5075-b (MO); near Puerto Buenos Aires, D. E. Breedlove 69153 (DUKE); 69172 (DUKE); 13 km SE de La Ciudad, C. Delgadillo 3160 (MEXU); C. Delgadillo 3163 (MEXU); Metate, N of Cueva, Pennell 18400a (MO); near Salto, A. J. Sharp 1817 (DUKE, MO); 34.5 km E of El Salto on rt 40, T. Stuessy 407 (NY, MO). Estado de México, municipio San Martin de Las Piramides, Cima del Cerro Gordo, valle de Teotihuacán, B. Allen 12938 (DUKE, MO); Valle de México, Amable 1291 (DUKE). Municipio Ixtapaluca, 1 km W de la Colonia Agrícola Manuel Ávila Camacho, A. Cárdenas 4360 (MO). Municipio Temascalapa, Cerro Gordo, A. Cárdenas 2199 (MO); Nevado de Toluca, Humboldt and Bonpland s.n. (type, BM, NY); Sierra de Las Cruces, C. G. Pringle 25a (BM, JE, MO, S). Guanajuato, 13 km NE de la ciudad de Guanajuato, A. Cárdenas 5312 (MEXU, MO); 9 km NE de Guanajuato, C. Delgadillo 5698 (MEXU); Mountains east of Comanja and 20 mi by road N of Leon, R. McVaugh 10426 (DUKE). Hidalgo, Casas Quemadas, Real del Monte, Arreguín 853 (CAS, MO). Municipio Epazoyucan, Peñas Largas, A. Cárdenas 3277b (MEXU, MO); El Cerezo al NW de la ciudad de Pachuca, A. Cárdenas 3331 (DUKE); 6 km N de Pachuca, Parque Nacional El Chico, García 101 (MO); Agua Blanca, Noroeste de Tulancingo, Gimate s.n. (XAL); Real del Monte, I. Mexía 2770 (BM, MO); Cuyamaloya, C. G. Pringle 10627 (MICH, NY); Tepeapulco, M. Vivas 81 (XAL); Parque Nacional "El Chico", M. Vivas 34 (XAL); 151 (XAL); 9 (XAL). Jalisco, slopes of La Ferrería, above Manantlán, H. A. Crum 776 (MICH); 1167 (S). Michoacán, Campanario, vicinity of Morelia, G. Arsène 7442 (BM, MO); 7448 (BM, MO, US); 7571 (MO); 9 km W de Mil Cumbres, hacia Puerto Garnica, C. Delgadillo 496 (MEXU); Paracho, about 6 km Northward along highway to Cheran, T. C. Frye and E. W. Frye 3073 (DUKE, MO). Morelos, municipio Tepoztlán, Sierra de Chichinautzin, A. Cárdenas 1059 (MEXU, MO); Tres cumbres, Cuernavaca, R. Düll 186 (B, MO); Mirador near kilometer marker 67, Cuernavaca, R. Magill 2508 (MO). Nuevo León, Picacho Onofre, Peña Nevada, G. L. Nesom and Wells 6 (DUKE). Oaxaca, distrito de Coixtlahuacán, Cerro de Tamazulapan, 3 km al E de San Francisco Teopa, Cisneros 2241 (XAL); 16 km al E. de Mitla, hacia San Lorenzo, C. Delgadillo 68 (MEXU, XAL); 37-38 km NE de Oaxaca, hacia Ixtlan, C. Delgadillo 129 (MEXU); 698 (MEXU, XAL); 2 km E de Ixtlan, C. Delgadillo 738b (MEXU); 20 km NNE de Tlacolula, C. Delgadillo 106 (MEXU); roadside at km 218 of Hwy. 175 at La Cumbre, Jim and Rosemary 3107 (MO). Puebla, on cerro Pinar, 1945, Alexander #A (DUKE); Tepoxuchitl frie Puebla, Nicolas s.n. (JE); Las cuevas on Ixtaccihuatl above Huejotzingo, A. J. Sharp 4321 (MO); above Río Frío, A. J. Sharp 229 (MO). Tlaxcala, cerca de Villareal al N de Huamantla, E. De Luna 2643 (XAL), 2644 (XAL). Veracruz, Cofre de Perote, C. Delgadillo 3930 (MEXU, XAL); Rincón de Atotonilco, laderas del Pico de Orizaba, E. De Luna 1655 (XAL); Las Vigas, entre Perote y Jalapa, E. De Luna 1803 (XAL); Cima del Cofre de Perote, E. De Luna and R. Kéller 2245 (XAL); 25 km W of Orizaba im pedregal, R. Düll 43 (JE, MO); Los Pescados, Cofre de Perote, G. Gómez-Velasco 3 (XAL); La Joya, G. Gómez-Velasco 1 (XAL); Camino al Cofre de Perote, D. Hernández 129 (MO, XAL); "El Volcancillo" entre la "Joya" y Rafael Ramírez, G. Juárez 554 (MO, XAL); Santa Marta del Monte, G. Juárez 830 (MO, XAL); Pedregal de las Vigas, above La Joya, Rickett 23 (MEXU); 33 (DUKE, MEXU); 17 km NE of Jalapa, R. Pursell 5005 (MO).
Additional specimens examined: BOLIVIA. Auf Felsblocken uri Chocayatal, T. Herzog 2585 (JE). DOMINICAN REPUBLIC. San Juan, Pico Duarte, W. R. Buck 8436 (FLAS), along trail up Pico Duarte, W. R. Buck 14174 (NY). GUATEMALA. Quetzaltenango, S of llano de Pinal, W slopes of Volcán Santa María, T. S. Quedensley 4566 (NY); mtns. above San Juan Ostuncalco, P. C. Standley 85253 (NY). HAITI. La Vega, Constanza, von Türckheim s.n. 1910 (H). UNITED STATES OF AMERICA. Arizona, Madera Canyon, Santa Rita Mts., Santa Cruz Co., W. Schofield 103112 (MO). Texas, Brewster Co., Upper Boot Canyon, Big Bend National Park area, Muller C4a (BM); 21270 (MO); Davis Co., Davis Mountains, E. D. Palmer 34297 (MO); Mt. LiverMore, Davis Mts., E. D. Palmer 34278 (DUKE, MO); Davis Mountains, Madera Canyon, Worthington 25422 (MO). VENEZUELA. Mérida, Quebrada Gavidia, S de Mucuchíes, D. Griffin and J. Duarte 1143 (F, FLAS, H, MICH, NY).
Discussion: paroicous stems are a character that clearly circumscribes this species. Other characters that distinguish B. secunda are the nearly flat, narrowly lanceolate leaves, with margins revolute up to acumen. The apex is acuminate, scabrous dentate; apical cells are a mix of oval and elliptic cells (Fig. 7). Also, the perichaetial paraphyses are frequently exserted and the capsules are ellipsoid, inflated at the base and slightly constricted below the mouth. The capsule wall is irregularly wrinkled when dry.
Braunia secunda, as recognized by De Luna (2016), is exclusively Neotropical and commonly grows on rocks in forests at middle elevations (1500-2000 m). This is a much narrower circumscription than previously accepted (Dixon, 1920; Brotherus, 1925). It does not include populations distributed in Africa and India. In Mexico it is indeed difficult to separate sterile material of Braunia squarrulosa, Hedwigidium imberbe and B. secunda as discussed by Sharp et al. (1978). Leaf margins are revolute up to the acumen in B. secunda and H. imberbe, but the margin is revolute only at the base in B. squarrulosa. In Mexico, it is also difficult to distinguish gametophytes of B. secunda from those of B. andrieuxii and B. plicata. Leaves are broader, obovate and the upper leaf cells are shorter in the latter two species. Additionally, B. andrieuxii has the leaf margins erect or reflexed but only at the base. On the other hand, B. plicata has strongly plicate leaves with a broad, hyaline, strongly papillose, and dentate acumen.
Braunia squarrulosa (Hampe) Müll. Hal., Linnaea 42: 380. 1879.
≡ Harrisonia squarrulosa Hampe, Linnaea 12: 349-350. 1838 ≡ Hedwigidium squarrulosum (Hampe) Bruch & Schimp., Bryol. Eur. 3: 161. 1846 ≡ Hedwigia squarrulosa (Hampe) Mitt., J. Linn. Soc. Bot. 12: 407. 1869. Type citation: "Mexico, Inter muscos Mexicanos, terrestris?". TYPE (Lectotype designated in De Luna, 2009): MEXICO. Schlechtendahl s.n. (Herb. Hampe-BM!; isolectotypes, BM!, NY!).
= Neckera sphaerocarpa Müll. Hal., Syn. Musc. Frond. 2: 105. 1851 ≡ Braunia sphaerocarpa (Müll. Hal.) Müll. Hal., Bull. Herb. Boiss. 5: 201. 1897. Type citation: "Mexico: C. Ehrenberg. E Guatemala copiose communicavit Kegel." TYPE (Lectotype designated in De Luna, 2009): MEXICO. Mineral del Monte, in arbor. C. Ehrenberg s.n. (BM!).
= Neckera liebmanniana Müll. Hal., Syn. Musc. Frond. 2: 668. 1851 ≡ Hedwigidium liebmannianum (Müll. Hal.) Schimp., Syn. Musc. Eur. Ed. 1, p. 239, 1860 ≡ Hedwigia liebmanniana (Müll. Hal.) Mitt., J. Linn. Soc., Bot. 12: 407. 1869 ≡ Braunia liebmanniana (Müll. Hal.) Besch., Mém. Soc. Sci. Nat. Cherbourg 16: 185. 1872 ≡ Braunia liebmanii Kindb., Enum. Bryin. Exot. 8, 1888, nom. illeg. Type citation: "Mexico. In monte Orizabensi: Liebmann". TYPE (Lectotype designated in De Luna, 2009): MEXICO. Orizaba, Liebmann s.n. (BM!; isolectotypes, BM!, NY!).
Plants robust in dense mats; stems with sympodial branching, plagiotropic, (3-)5-7(-10) cm long, scarcely branched; branches arcuate, flexuose-flagelliform branches frequent; pseudoparaphyllia foliose, with papillose cells; leaves ovate or oblong-lanceolate, widest point below middle of leaf; large 2-2.5 × 0.8-1.3 mm, concave, erect-spreading to squarrose, strongly plicate; leaf acumen gradually differentiated, to 1/4 length of leaf, acuminate to cuspidate, concolorous, sometimes ending in a flexuose hyaline point; margins reflexed, sometimes recurved in lower 1/2, entire or papillose-crenulate at acumen; apical leaf cells elliptic, 22-31 µm long, narrow (4-7:1), sinuous, with few low marginal papillae; upper and medial cells elliptic, 12-20 µm long, narrow (3-5:1), walls nodulose, sinuous, papillae low, on side walls, overarching lumina; autoicous; archegonia distal on short vaginula (1.4-1.6 mm); perichaetial leaves oblong lanceolate, 2.4-2.6 mm long, slightly longer than vegetative leaves, acuminate, plicate; paraphyses biseriate at base, 2-3× longer than archegonia, immersed; setae (3-)4 - 5(-8) mm long; capsule exserted; necks 0.3-0.4 mm long, sharply differentiated, broad; urns 1.2-1.6 mm long, urceolate to globose, mouth broad, nearly as wide as urn; capsule walls smooth, or wrinkled when dry; exothecial cells polygonal; stomata superficial; operculum base conical, short oblique-rostrate; calyptrae cucullate, large, hairy; spores multicellular, 35-42 µm in diameter.
Habitat: this species is mainly epiphytic on the base of trees, on oak trunks, on bark of Juniperus L, on trunk of Cupressus L, on fallen tree branches. Also forming carpets on very decayed logs, and on volcanic rocks, rarely on soil. It is common in coniferous and broadleaf forests at middle elevations (1500-2900 m).
Distribution: Mexico (Chiapas, Chihuahua, Ciudad de México, Durango, Estado de México, Guerrero, Hidalgo, Jalisco, Michoacán, Nayarit, Oaxaca, San Luis Potosí, Veracruz, Zacatecas), Guatemala, Honduras, Costa Rica, Panama (De Luna, 2009). Fig. 8.
Figure 8: Distribution map of Braunia squarrulosa (Hampe) Müll. Hal. Each dot is one of the specimens listed in the text.
Figure 9: Morphological features of Braunia squarrulosa (Hampe) Müll. Hal. in Mexico. A. branch with sporophytes (A, from Dieterle 3192, DUKE, Mexico, Guerrero); B-D. vegetative leaves; E. apical leaf cells (B-E, from Schlechtendahl s.n., Herb. Hampe-BM, 1846, lectotype).
Illustrations: Hampe (1844, Icon. Musc. 19, as Harrisonia squarrulosa); Bartram (1949, p. 235, Fig. 106, A-C); Crum (1994, p. 666, Fig. g-i); De Luna (2009, Figs. 1-17); Fig. 9.
Selected specimens examined: MEXICO. Chiapas, 1 km above Aguacatenango, D. E. Breedlove 68944 (DUKE); 68973 (DUKE); 9 km südlich San Cristobal, R. Düll 8 (B, MO); NE of Cerro Boquerón, on road from El Rosario to Niquivil, D. E. Breedlove 68923 (DUKE); 10 km NW of Comitán on side road to Zaragoza, D. E. Breedlove 70423 (DUKE); Laguna Chamula, between Amatenango and Comitán, D. E. Breedlove 70073 (DUKE); between Teopisca and Comitán, J. P. Frahm 79-2277 (B); along route 190, F. J. Hermann 26429 (BR, DUKE, NY); Lake Jusnajay, NW of Comitán, A. J. Sharp 3538 (DUKE). Chihuahua, 29 mi along road from San Francisco del Oro to Guadalupe y Calvo, Kimnach and Brandt 1040 (MEXU). Ciudad de México (formerly Distrito Federal), mountain Tlaxpehualco, 26.VIII.1926, M. S. Antipovich s.n. (MO); Cima, C. G. Pringle 10550 (BM, F, NY). Durango, near Puerto Buenos Aires, D. E. Breedlove 69154 (DUKE). Estado de México, Temascaltepec, Dziekanowski et al. 2005 (BM, MO); Valle de Bravo, R. Düll 55a (JE). Guerrero, about 40 km W of Chilpancingo, Dieterle 3192 (DUKE, MO); about 30 km W of Chilpancingo, A. J. Sharp 1106 (MO, TENN). Hidalgo, Mineral del Monte, Ehrenberg s.n. (lectotype, BM); Jacala, T. C. Frye and E. W. Frye 2713 (DUKE, G). Jalisco, El Cuartón, above Manantlán, H. A. Crum 1175 (DUKE); Sierra Madre W of Bolaños, Rose #e (NY). Michoacán, vicinity of Morelia, Cerro Azul, G. Arsène 4536 (BM, F, MO); 4539 (BM, F), G. Arsène 4776 (DUKE); W of Ciudad Hidalgo, C. Delgadillo 457 (MEXU, XAL); Mil Cumbres, cerca de Puerto Garnica, 20 km al E de Morelia, E. De Luna 1772 (DUKE, XAL); 1777 (DUKE, XAL); Uruapan, C. G. Pringle 15085 (F, NY); San Miguel del Monte Morelia, 1908, Solorzano s.n. (JE). Nayarit, La Cienega, on ridge about 10 miles NW of Mesa del Nayar, D. Norris and D. J. Taranto 14452 (FLAS). Oaxaca, El Punto, NE of Oaxaca, J. Conrad and R. Conrad 3160 (DUKE); 23 km NE of Oaxaca, C. Delgadillo 628 (MEXU, XAL); al Este de Ixtlán, C. Delgadillo 709 (MEXU, XAL); Sierra de Juárez, R. Düll 66 (JE); La Cumbre, between Oaxaca and Ixtlán de Juárez, A. J. Sharp et al. 2609 (F); towards cerro San Felipe from Oaxaca, A. J. Sharp 4015 (DUKE, F). San Luis Potosí, Alvarez, C. Orcutt 6530 (F). Sinaloa, about 6 mi W of Las Palmitas, D. Norris et al. 20444 (BR). Veracruz, Tlacotepec near Huayacocotla, G. Juárez 1014 (XAL); Orizaba, Liebmann s.n. (lectotype BM, isolectotypes BM, NY). Zacatecas, municipio Chalchihuites, Gualterio, A. Cárdenas 1068 (MEXU, MICH, XAL).
Additional specimens examined: COSTA RICA. San José, Copey De Dota, Fournier and Fournier 1030b (USJ); Valle de Copey, ca. 30 km S de Cartago, D. Griffin and I. Morales B 97 (NY); Copey De Dota, I. Morales 704 (USJ). GUATEMALA. No locality, Kegel s.n. (BM); 3/48, Kegel #10004 (syntypes BM, JE, NY); Huehuetenango, near Paquix, above Huehuetenango, A. J. Sharp 4766 (DUKE); N of Chiantla, P. C. Standley 82548 (NY); Quetzaltenango, mountains N of Olintepeque, P. C. Standley 85220 (NY); cerro Pixpix, above San Ildefonso Ixtahuacán, J. A. Steyermark 50604 NY). HONDURAS. Lempira, Montaña de Celaque, Filo Seco, 13 km SW of Gracias, B. Allen 12095 (DUKE, MO). PANAMA. Chiriquí, E slope of volcán Chiriquıí (Barú), WNW of Boquete, G. Davidse and D’Arcy 10190A (G, MO).
Discussion: a short seta (5-6 mm), globose, smooth-walled capsules, leaves wide spreading to squarrose, and multicellular spores help in distinguishing this from all other species in the genus. Without the sporophyte, the variability in leaf shape and leaf cells makes it difficult to identify it. In general, the leaves are more strongly plicate, the leaf base is broader, the acumen is slender, long cuspidate, and the apical cells are narrower in B. squarrulosa than in B. secunda and Hedwigidium imberbe. These differences are more evident when comparing juvenile leaves in the three species.
