A Synopsis of Bryoerythrophyllum and Morinia (Pottiaceae) in the New World1
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The Bryologist 81(4), 1978, pp. 539-560 Copyright ? 1978 by the American Bryological and Lichenological Society, Inc. A Synopsis of Bryoerythrophyllum and Morinia (Pottiaceae) in the New World1 RICHARD H. ZANDER2 Abstract. Taxa in Bryoerythrophyllum Chen & Morinia Card. (Pottiaceae) in the New World are B. bolivianum (C.Muell.) comb. nov., B. columbianum (Herm. & Lawt.) comb. nov., B. ferruginascens (Stirt.) Giac., B. jamesonii (Tayl.) Crum, B. recurvifolium (Tayl.) Zander, B. recurvirostrum (Hedw.) Chen var. recurvirostrum, B. recurvirostrum var. aeneum (C.Muell.) Zander, B. re- curvum (Griff.) K.Saito, M. crassicuspis (H.Robins.) comb. nov., M. ehrenber- giana (C.Muell.) Ther. var. ehrenbergiana and M. ehrenbergiana var. elongata (Wils. in Mitt.) comb. & stat. nov. The previously poorly understood species B. bolivianum, B. ferruginascens, B. jamesonii and B. recurvum are rede- scribed and illustrated. Sporophytes of B. ferruginascens are describedfor the first time. Bryoerythrophyllumjamesonii is considered a polymorphic complex with many synonyms. Notes on relationships, infraspecific variation, geo- graphic distribution and nomenclature are given for other taxa. Globulina C.Muell., hom. illeg., is a valid name and is a synonym of Bryoerythrophyllum, but Seligeria sect. Globulina C.Muell. is a synonym of Globulinella Steere. Barbula andreaeoides Kindb. is a good species, not a synonym of Bryoeryth- rophyllum ferruginascens. Tortula herzogii is a nom. nov. for T. angustifolium (Herz.) Herz., hom. illeg. Species of four genera of New World Pottiaceae, Bryoerythrophyllum Chen, Erythrophyllopsis Broth. in Herz., Morinia Card. and Rhexophyllum Herz., are oc- casionally confused in the herbarium. Gametophytically, the species are similar: red- dish plants, stems with central strand, leaves usually lanceolate, usually reflexed above a differentiated leaf base, costa usually with papillose ventral epidermal cells (these often similar to the laminal cells), upper laminal cells mostly quadrate with thin to evenly thickened cell walls and laminal papillae apparently scattered over the lumens, often crowded, hollow (o- to c-shaped in optical section) to solid, low and granular, bi- to multifid. Rhexophyllum and Erythrophyllopsis are both monotypic. These are illustrated, the former by Grout (1938) and the latter under synonyms by Sullivant (1860) and Herzog (1916). I have previously discussed synonymy and mor- phological variation in both Rhexophyllum subnigrum (Mitt.) Ther. ex Hilp. (Zander, 1976) and Erythrophyllopsis andina (Sull.) Zander (Zander, 1977a). 'I acknowledge with thanks the helpful correspondence, discussion and specimens com- municated over several years by L. E. Anderson, F. Bowers, M. Crosby, H. Crum, P. Eckel, D. Horton, R. E. Magill, G. Pierce, H. Robinson, W. Schofield, A. J. Sharp, D. K. Smith, W. C. Steere and D. Vitt and the curatorsof the herbaria mentioned for loan of specimens. A portion of this work was supported by a Pre-Doctoral Internship at the Smithsonian Institution, 1967- 1968. 2 Clinton Herbarium, Buffalo Museum of Science, Buffalo, NY 14211. 0007-2745/78/539-560$2.45/0 540 THE BRYOLOGIST [Volume 81 The present publication includes notes on all American species of Bryoerythro- phyllum and Morinia, thoroughly describes four previously little understood taxa and includes a key that should be of service in identification of collections. The species presently in Bryoerythrophyllum and Morinia all have rather similar characters of the areolation, but the genera differ widely in characters of the sporophyte. I agree with Loeske (1910) that areolation is often quite significant but is frequently unde- scribed by taxonomists. Areolation appearance contributes to an often touted "look" or characteristic subtle complexion that is much relied on taxonomically but seldom described in print. Areolation can be overemphasized-Loeske (1910) suggested that species now placed in Bryoerythrophyllum were related to Tortella and Oxystegus. Indeed, the areolation and "look" are quite similar, but other significant characters such as number of amphithecial derivative layers in the capsule (Saito, 1975) and the recurved leaf margin indicate that Bryoerythrophyllum belongs in the Barbuleae. KEY TO NEW WORLD TAXA OF BRYOERYTHROPHYLLUM, ERYTHROPHYLLOPSIS, MORINIA AND RHEXOPHYLLUM 1. Leaf apices acute, entire, leaf base ovate, basal cells quadrate, 1:1 ------ 2 1. Leaf apices acute or obtuse, entire or dentate, leaf base elliptical to long-oblong, basal laminal cells usually rectangular, mostly 2:1 or longer -------_________ 3 2. Leaf apex apiculate by 1-2 clear cells, costa percurrent, narrow, usually sinuose above, in transverse section reniform to circular, usually up to 4 epidermal cells across ventral surface of costa, upper laminal cells entirely papillose; Nepal, India, U.S.A., Mexico ---- - Bryoerythrophyllum recurvum (Griff.) K.Saito 2. Leaf apex ending in a sharp, stout mucro, costa short-excurrent, broad, straight throughout, in transverse section elliptical, usually up to 8 epidermal cells across ventral surface of costa, upper marginal laminal cells epapillose in 2-6 rows; northwestern U.S.A. Bryoerythrophyllum columbianum (Herm. & Lawt.) Zander 3. Laminal cells bistratose in large or small patches or along margin --- 4 3. Laminal cells unistratose -------------- 8 4. Laminal cells bistratose in a narrow marginal band, leaf margins narrowly re- curved at midleaf ------------- 5 4. Laminal cells bistratose in medial patches, leaf margins plane ---- 7 5. Leaves 1.8-2.2 mm long, apex usually expanded as an elliptical propagulum, caducous early, smooth or with large hemispherical protuberances; Mexico, Guatemala ---- ----------------------------- Morinia crassicuspis (H.Robins.) Zander 5. Leaves 2.0-5.0 mm long, apical propagulum, if present, oblong or spatulate, merely fragile or at least caducous late, margins and dorsal surface of propagulum distantly blunt-dentate -------------- 6 6. Leaves short-lanceolate, mostly 2.0-3.0 mm long, apex blunt to narrowly acute, upper margins strongly serrate, apical propagulum not differentiated; Mexico, Guatemala, Venezuela, Colombia, Ecuador Morinia ehrenbergiana (C.Muell.) Thir. var. ehrenbergiana 6. Leaves long-lanceolate, mostly 2.5-5.0 mm long, apex narrowly acute, upper mar- gins weakly serrate, apical propagulum often differentiated; Ecuador - ------------- Morinia ehrenbergiana var. elongata (Mitt.) Zander 7. Leaf margins entire, peristome present; Andes--- ----------------------- Erythrophyllopsis andina (Sull.) Zander 7. Leaf margins dentate to erose-dentate, peristome absent; Latin American cordillera, southwestern U.S.A. ---- Rhexophyllum subnigrum (Mitt.) Ther. ex Hilp. 8. Leaves oval to ovate-triangular, seldom longer than 1.0 mm, apex not apiculate, broadly rounded to narrowly obtuse, somewhat cucullate; Peru, Bolivia -- Bryoerythrophyllum bolivianum (C.Muell.) Zander 8. Leaves ovate-triangular to lanceolate, usually longer than 1.9 mm, apex often apiculate by 1-4 clear cells, narrowly obtuse to acute, not cucullate --- 9 9. Leaf margins strongly recurved above midleaf, often to near apex; usually monoicous, sometimes dioicous ---------------- 10 1978] ZANDER: SYNOPSIS OF BRYOERYTHROPHYLLUM & MORINA 541 9. Leaf margins plane above midleaf or seldom weakly recurved to near apex; dioicous 11 10. Capsules mostly to 2.0 mm long; synoicous or paroicous; leaf apex acute to broadly acute; without propagula; widespread in Northern Hemisphere, Africa, Austra- lasia - Bryoerythrophyllumrecurvirostrum (Hedw.) Chen var. recurvirostrum 10. Capsules mostly 2.0-3.0 mm long; dioicous or polyoicous; leaf apex broadly acute to rounded; often with red, multicellular propagula on lower portion of stem; Mexico, Guatemala Bryoerythrophyllum recurvirostrum var. aeneum (C.Muell.) Zander 11. Plants green, leaf base short, squarish, leaves with pale border of 1-2(4) rows of yellowish, epapillose, short-rectangular, thick-walled cells, stem central strand absent; western Canada, western Europe ----- Bryoerthrophyllum recurvifolium (Tayl.) Zander 11. Plants reddish, at least below, leaf base long, elliptical to oblong, leaves not bordered or occasionally marginal cells weakly papillose in 1-2 rows, yellowish, stem central strand present ----------------12 12. Leaves oblong to long-lanceolate, 1.0-3.5(-5.0) mm long, apex usually broadly acute to broadly obtuse, upper margins often dentate, often weakly bordered by thick-walled cells, propagula absent; western Canada, Scotland, Latin American cordillera, West Indies, China, India, Tasmania, New Zealand --- --------Bryoerythrophyllum jamesonii (Tayl.) Crum 12. Leaves ovate-triangular to short-lanceolate, 1.0-1.8 mm long, apex narrowly acute to narrowly obtuse, margins entire, not bordered, propagula often present on basal rhizoids; Arctic circumpolar regions; Canada; Greenland; U.S.A.: Alaska, Rocky and Appalachian Mts.; Mexico; Europe; northern Asia; possibly Japan - --Bryoerythrophyllum ferruginascens (Stirt.) Giac. Bryoerythrophyllum Chen, Hedwigia 80: 4. 1941. Lectotype species: B. recurvirostrum (Hedw.) Chen Barbula sect. Erythrophyllum Lindb. in Braithw., Brit. Moss Fl. 1: 260. 1887. Globulina C.Muell., Nuov. Giorn. Bot. Ital., N. S. 4: 39. 1897, hom. illeg. non Lind in Nees, Hor. Phys. Berol. 4. 1820, nec Spegaz. in Sacc., Syll. Fung. 9: 993. 1891. Erythrophyllum (Lindb.) Loeske, Hedwigia 47: 175. 1908, hom. illeg. non J. Agardh, Bidr. Florid. Syst. 10: 237. 1872. Erythrobarbula Steere, THE BRYOLOGIST54: 191. 1951, hom. illeg. incl.