20. BRYOERYTHROPHYLLUM Plates 24– 25.

Bryoerythrophyllum Chen, Hedwigia 80: 4, 1941. Type: Bryoerythrophyllum recurvirostrum (Hedw.) Chen.

Globulina C. Müll., Nuov. Giorn. Bot. Ital., N.S. 4: 39, 1897, hom. illeg. non Link in Nees 1820, nec Turpin 1827, nec Velen. 1834, nec Spegaz. in Sacc. 1891. Type: Globulina boliviana C. Müll.

Erythrophyllum (Lindb. in Braithw.) Loeske, Hedwigia 47: 175, 1908, hom. illeg. non J. Ag., 1872.

Erythrobarbula Steere, Bryologist 54: 199, 1951, nom illeg. incl. gen. prior.

Didymodon subg. Erythrophyllum Limpr., Laubm. Deutschl. 1: 405, 1887. Type: Didymodon rubellus BSG.

Barbula subg. Erythrophyllum (Lindb. in Braithw.) C. Jens., Skand. Bladmfl. 243, 1939.

Barbula subg. Erythrobarbula Szafr., Fl. Polska Mchy 1: 213, 1957 [1958], nom. inval. descr. polon.

Barbula sect. Erythrophyllum Lindb. in Braithw., Brit. Moss Fl. 1: 260, 1887. Type: Barbula rubella Mitt. in Lindb.

Didymodon sect. Orthocarpae Broth., Nat. Pfl. 1(3): 405, 1902.

Didymodon sect. Amblystegioideae Broth., Nat. Pfl. 1(3): 405, 1902.

Pl ants turf-forming or loosely caespitose, usually green above and red-brown below. Stems occasionally branching, short or to several cm in length, rounded-pentagonal in transverse section, usually with central strand, sclerodermis usually present, hyalodermis usually absent; axillary hairs several cells in length, sometimes with 1–3 brownish basal cells. Leaves appressed when dry, spreading when wet, ovate, elliptical or lanceolate, 1–5 mm in length, often grooved along the costa, marins usually recurved below or to near apex, seldom plane, entire or more often dentate above, often with a border of 3–4 rows of thick-walled cells; apex rounded-obtuse to acute; base usually ovate to oblongand sheathing; costa ending a few cells below the apex to short-excurrent, with quadrate, papillose cells ventrally, cells elongate dorsally, in transverse section showing a differentiated epidermis ventrally or on both sides, ca. 2–6 rows of cells across costa ventrally at midleaf, two stereid bands present, guide cells 2–4 in 1(–2) layers, one or more hydroid strands occasionally present; upper laminal cells subquadrate to short-rectangular, ca. 10–15 µm in width, 1(–2):1, walls evenly thickened, weakly sinuose, superficially nearly flat, occasionally bulging; papillae solid, or occasionally hollow, bifid, or occasionally flattened or compound, mostly 4–6 per lumen, obscuring the lumens; basal cells usually hyaline and rectangular, filling most of the base medially, little wider than upper cells, occasionally bulging, 2–5:1. Asexual reproduction occasional, by unicellular propagula in masses in the leaf axils or obovoid brood bodies borne on rhizoids in the soil. Dioicous or occasionally monoicous. Perichaetia and perigonia terminal. Perichaetial leaves usually sheathing in the lower half, larger than the stem leaves, long-oval to long-lanceolate, lower cells long rhomboidal in lower half. Perigoniate plants occasionally small, bud-like. Seta elongate, twisted clockwise below, often counterclockwise above, usually 1 per perichaetium; theca ellipsoidal to cylindric, usually red-brown, 0.7–2.5(–4.0) mm in length, sometimes curved; stomates phaneropore, at base of theca, annulus of 1–2 rows of vesiculose cells, often revoluble or deciduous in pieces; operculum short-conic to short-rostrate, cells in straight or oblique rows; peristome none, rudimentary, or consisting of 16 or 32 linear rami, erect to twisted to 2–4 times counterclockwise, usually densely spiculose, basal membrane low or absent. Operculum short-conic to short-rostrate, 0.2–1.2 in length. Calyptra cucullate, smooth, ca. 2 mm in length. Spores ca. 8–15 µm in diameter, light brown, lightly papillose. Laminal KOH color reaction red to orange-red. Reported chromosome number n = 13, 13+m, 14, 26+2m.

An essentially cosmopolitan genus. More extensive treatments of this taxon for the New World have been done by Zander (1978g, 1980a, 1981a).

Bryoerythrophyllum has much the appearance of Barbula or Didymodon species, but is easily distinguished by a combination of features, most of which are usually present: red coloration (at least in older parts of stems—Bryoerythrophyllum fuscinervium, for instance, generally having immature leaves at the stem apex with a yellow KOH reaction), the red color usually evident even without KOH; bifid, crowded laminal papillae obscuring the upper laminal cell lumens; usually flat superficial laminal cell walls (easily seen in section, Pl. 24, f. 7; 25, f. 6); and well-differentiated leaf basal cells (Pl. 24, f. 6; 25, f. 5). Like Barbula and Didymodon, Bryoerythrophyllum exhibits a full range of peristome development, some species eperistomate (e.g. B. boliviana, Pl. 24, f. 11), some with rudimentary peristomes (e.g. B. rotundifolia), others with straight, well-developed teeth (e.g. B. binsii, Pl. 25, f. 8, and B. recurvirostrum), and finally those with long, twisted peristomes (e.g. B. inaequalifolium, Pl. 24, f. 10). Only recently has it been recognized (Saito 1975a; Zander 1978g, 1980a) that this genus is rather large and includes many species previously placed in Barbula and Didymodon. Bryoerythrophyllum fuscinervium and the similar B. columbianum have a rather distinctive broad and bulging ventral costal epidermis of a single layer of cells and may be confused with Pseudocrossidium species (such as P. chilense) but the red KOH reaction and distinct ventral stereid band are diagnostic. Bryoerythrophyllum differs from Syntrichia, which is also KOH red, in having two stereid bands, a usually narrower leaf shape and dorsal costal epidermis commonly differentiated. The upper areolation of rather large cells with evenly thickened walls is similar to that of Trichostomum subg. Oxystegus but the red color in KOH, presence of a stem central strand and costal hydroid strand(s), and the basal cells extending farthest up the leaf medially distinguish Bryoerythrophyllum (e.g. cf. B. chimborazense). The genus Mironia is very similar, but differs in the keeled leaves with bistratose upper margins (the upper lamina of Bryoerythrophyllum, in one species, B. sharpii, may be bistratose throughout) and, in two species of Mironia, the presence of propaguloid leaf apices.

The genus Globulina was published validly by C. Müller (1897a) in a combined genus and species description. Although Müller indicated that Seligeria globifera Hampe (║ Globulinella globifera (Hampe) Steere) was related to and was probably also a member of the genus (“Globulina mihi”), he left the genus monotypic with G. boliviana. Later, Müller (1901) established Seligeria subg. Globulina C. Müll., with S. globifera the only species, being thus the type of that, different, taxon. Steere (Steere & Chapman 1946) proposed Globulinella as a new genus, with Globulina C. Müll. non Link in Nees as a synonym, but excluded the type of Globulina, G. boliviana, later in the publication. Steere apparently did not intend Globulinella as a nom. nov. for Globulina as was suggested by Magill (1977a). This confusion is reflected in the Index Nominum Genericorum (Farr et al. 1979). Actually, Globulina C. Müll., hom. illeg., is not a synonym of Globulinella Steere, but is instead a synonym of Bryoerythrophyllum Chen. Seligeria subg. Globulina C. Müll. is a synonym of the valid genus Globulinella Steere, both taxa having the same type species, G. globifera (lectotype fide van der Wijk et al., 1959–69).

The type of Husnotiella glossophylla Herz. is not at L, JE or W and may have been destroyed during the Second World War (pers. comm. F. K. Meyer at JE). Specimens collected by Hosseus in Argentina (FH, W) identified by Bartram as this or as H. baueri are eperistomate collections of the rather variable Didymodon tophaceus. The description and illustrations of H. glossophylla itself indicate that it is probably the same as Bryoerythrophyllum calcareum.

Additional literature: Chen (1941), Crum (1957a), Long (1982a, 1982b), Ratkowsky (1980).

Number of accepted species: 27.

Species examined: B. andersonianum (BUF, TENN), B. binnsii (BUF), B. bolivianum (BUF, JE, TENN), B. byrdii (Bartr.) Zand. (NY), B. calcareum (BUF, FH, MICH, TENN, US), B. campylocarpum, B. chimborazense (NY), B. columbianum (UBC, US), B. ferruginascens, B. fuscinervium (BUF, NY), B. hostile (JE), B. inaequalifolium (BUF, DUKE, NY, TENN), B. jamesonii, B. ligulare (NY), B. recurvirostrum, B. rotundatum (H), B. rubrum (BUF, NY), B. sharpii (BUF), B. wallichii (NY, US).

New heterotypic synonymy: Bryoerythrophyllum columbianum var. atacamense Zand. & Lewis in Lewis = Bryoerythrophyllum fuscinervium (Mitt.) Zand. Didymodon integrifolius Broth. in Mildbr. var. paucidentatus Thér. = Bryoerythrophyllum campylocarpum (C. Müll.) Crum. Didymodon luzonensis Bartr. = Bryoerythrophyllum ferruginascens (Stirt.) Giac. (the type of the former at FH is of fruiting material and includes the characteristic propagula). Didymodon merceyoides Broth. in Herz. = Bryoerythrophyllum campylocarpum (C. Müll.) Crum. Didymodon pelichucensis Williams = Bryoerythrophyllum campylocarpum (C. Müll.) Crum. Hyophila calymperoides Thér. & Nav. = Bryoerythrophyllum campylocarpum (C. Müll.) Crum.

New combinations: Bryoerythrophyllum byrdii (Bartr.) Zand., comb. nov. (Barbula byrdii Bartr., Ann. Missouri Bot. Gard. 25: 720, 1938). Bryoerythrophyllum chimborazense (Mitt.) Zand., comb. nov. (Tortula chimborazensis Mitt., J. Linn. Soc. Bot. 12: 163, 1869; Didymodon chimborazensis (Mitt.) Broth. in Par.). Bryoerythrophyllum fuscinervium (Mitt.) Zand., comb. nov. (Tortula fuscinervia Mitt., J. Linn. Soc. Bot. 12: 154, 1869; Barbula fuscinervia (Mitt.) Jaeg. Bryoerythrophyllum ligulare (Mitt.) Zand., comb. nov. (Tortula ligularis Mitt., J. Linn. Soc. Bot. 12: 156, 1869; Barbula ligularis (Mitt.) Jaeg.).