A Synopsis of Bryoerythrophyllum and Morinia (Pottiaceae) in the New World1

The Bryologist 81(4), 1978, pp. 539-560 Copyright ? 1978 by the American Bryological and Lichenological Society, Inc.

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. recurvum (Griff.) K.Saito, M. crassicuspis (H.Robins.) comb. nov., M. ehrenbergiana (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 redescribed 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, geographic 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 occasionally confused in the herbarium. Gametophytically, the species are similar: reddish 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 morphological 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 recurved 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 margins 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. gen. prior. Plants loosely caespitose to cushion-forming, usually red-brown below. Stems mostly 1.0-2.0 cm long, central strand usually present. Leaves ovate to oblong or lanceolate, ventrally broadly concave or channeled to keeled, margins entire to dentate, plane to recurved, leaf apex rounded- obtuse to acute, leaf base usually sheathing; costa ending a few cells below the apex to percurrent, ventral superficial cells quadrate to short-rectangular, papillose, dorsal superficial cells quadrate and papillose to rectangular and smooth; costa in transverse section elliptical to reniform, with ventral epidermal layer usually differentiated, ventral stereid band usually weak, 2-5 guide cells present, dorsal stereid band strong, dorsal epidermis usually weakly differentiated, often as thick-walled cells with semicircular lumens; upper laminal cells subquadrate to short-rectangular, walls usually evenly thickened, superficially flat, laminal papillae solid to hollow, usually crowded, scattered to occasionally centered over the lumens, bifid to multiplex, flattened, mostly 2-6 per lumen, basal laminal cells usually differentiated, rectangular, occasionally bulging, filling most of the leaf base, smooth to weakly papillose. Dioicous, synoicous or paroicous, occasionally polyoicous. Gametoecia terminal, perigonia gemmate; perichaetial leaves long-oval to long-lanceolate, larger than the cauline leaves, laminal cells usually prosenchymatous below, sheathing the seta in lower 1/3-3/4. Sporophyte seta elongate, twisted clockwise below, often counterclockwise above; urn elliptical to cylindrical, occasionally arcuate; stomata at base of urn, phaneropore; annulus of 1-2 rows of usually vesiculose cells, often revolvable or deciduous in parts; peristome absent, rudimentary, or of 32 straight, linear, usually densely spiculose teeth, basal membrane absent or low. Operculum short-conic to short-rostrate. Calyptra cucullate, smooth. The genus Bryoerythrophyllum is well characterized by Chen (1941), Crum (1957) and Saito (1975). The present study employs (to account for observed clinal variation) 542 THE BRYOLOGIST [Volume 81 a broad species concept that allows recognition of only seven species of Bryoerythro- phyllum in the New World. In comparison, Savicz-Ljubitzkaja and Smirnova (1970) give five species for the U.S.S.R., Chen (1941) lists 12 species in Eastern Asia and Saito (1975) lists six species for Japan alone. The genus Leptodontium is very similar in appearance of gametophyte and sporophyte but is distinguished by the yellow-green coloration, lack of stem central strand, laminal papillae often simple, multiplex or coroniform-capitulate, costa lacking ventral epidermal cells similar to those of the lamina, ventral stereid band exposed and perichaetial leaves usually much elongated and convolute-sheathing. la. Bryoerythrophyllum recurvirostrum (Hedw.) Chen var. recurvirostrum is a relatively common taxon of limey soils distributed across the Northern Hemisphere and also found in Hawaii, northern Africa and Australasia. It is well described by many authors and especially fine illustrations have been done by Flowers (1973). I accept the synonymy of Crum, Steere and Anderson (1973). Steere (1938b) judged from Hed- wig's (1787-1797; 1801) descriptions and plate that the type of Weissia recurvirostra Hedw. was possibly misinterpreted by early authors. Steere suspected, on account of the yellow peristome, absence of an annulus and apparent dioicous condition described in the protologue, that the type did not match the concept of present-day authors. The holotype specimen ("Gottingae," anon., s.n.) from the Hedwig-Schwaeg- richen Herbarium (c!) fits modern notions of B. recurvirostrum var. recurvirostrum. The plants do have yellow, somewhat eroded peristomes but most capsules are deo- perculate and worn. The annulus is differentiated and, though deciduous in parts, is not revolvable; this is not an unusual variation though some authors imply that the annulus is always revolvable. The type is paroicous (only one plant was examined due to the paucity of material), not dioicous. Specimens (e.g. Canada: Newfoundland, St. Johns, Best, 1907-NY) occasionally may be misinterpreted as dioicous as the majority of gametoecia may contain only archegonia; however, gametoecia on separate plants in the same collection that appear to be perigonia, being swollen, with ovate, concave inner leaves, will have at least one archegonium present among the anther- idia. In some alpine and arctic collections of var. recurvirostrum, the plants are flagellate with stems to 3.0 cm long, leaves short, 0.7-1.0 mm long, ovate-triangular, the lamina occasionally bistratose in patches near apex and the costa often broad and ventrally bulging, in transverse section showing up to 11 papillose ventral epidermal cells. Such variation is rather rare. Examples include: U.S.A., COLORADO: Zander 4068 (BUF) and the type, GREENLAND; Heilprin Land, Br0nlund Fjord, Holmen 277d (holotype-c), Independence Fjord, Holmen 7128 (paratype-c) of Barbula recurvi- rostris var. latinervia Holmen, syn. nov. lb. Bryoerythrophyllum recurvirostrum var. aeneum (C.Muell.) Zander (Fig. 1-6) is discussed by Zander (1976). An additional synonym is Didymodon stenopyxis Card., Rev. Bryol. 36: 84, 1909. Type: Mexico, D. F., Cima, Pringle 10518 (Pc-holotype; BM, FH-isotypes). The var. aeneum is at least occasionally polyoicous, the same collection including a mixture of both monoicous plants and apparently separate (but possibly pseudautoicous) male plants with terminal, gemmate perigonia. This is the case in the type of D. stenopyxis, as previously noted by Williams (1913), and in Mexico: Coahuila, Parras, Stanford 1803 (TENN). This variation in sexuality necessitates the determination of the sexual condition of several plants in each collection for ac- curate identification. (I have found that the use of two, rather than the usual single, illuminator for the dissecting microscope greatly facilitates this chore.) Red, multi- 1978] ZANDER: SYNOPSIS OF BRYOERYTHROPHYLLUM & MORINA 543 cellular propagula (Fig. 6), similar in shape to those of B. ferruginascens (Fig. 16) but much larger, may be present on rhizoids from the lower portions of the stems, as in Mexico: D. F., Parres, Orcutt 3817a (FH), Mexico, E of Lerma, Hermann & Crum 20872 (NY) and Guatemala: Quetzaltenago, Volcan Santa Maria, Steyermark 34093 p.p. (NY). Saito (1975) includes some propaguliferous collections within his concept of var. recurvirostrum. The Orcutt 381 7a specimen has only archegonia-bearing perichaetia and is placed with the var. aeneum through the shape of the leaf apex, which appears to be a fairly good character distinguishing the varieties (see key). The dioicous Eurasian species B. rubrum (Geh.) Chen is closely related to B. recurvirostrum by the leaf margins recurved in the lower 3/4 of the leaf but differs by the long-acuminate leaf apices and usual lack of propagula (Saito, 1975, p. 394, reports the rare occurrence of rhizoidial gemmae in B. rubrum var. minus K.Saito). Variation in sexual condition in both varieties of B. recurvirostrum may constitute adaptation toward outbreeding. Steere (1940) described the occurrence of dichogamy (as proterogyny) in the monoicous species Tortula princeps DeNot. Crundwell and Nyholm (1964) found that Bryum tenuisetum Limpr., though often described as polygamous, is usually dioicous in the field. When male and female plants are grown in mixed culture, however, some synoicous gametoecia often develop. Perhaps monoicy in B. recurvirostrum var. aeneum is usually suppressed but is triggered by the environment.

2. Bryoerythrophyllum ferruginascens (Stirt.) Giac., Atti Ist. Bot. Univ. Lab. Critt. Pavia, Ser. 5, 4: 210. 1947. (FIG. 7-20) Basionym: Barbula ferruginascens Stirt., Ann. Scott. Nat. Hist. 9: 176. 1900. Barbula rubella var. ruberrima Ferg. in Braithw., Brit. Moss Fl. 1: 261. 1887. Barbula botelligera Moenk. in Murr.,Allg. Bot. Zeitschr. 20: 24. 1914. Didymodon botelliger (Moenk.) Hag., K. Norsk. Vid. Selsk, Skrift. 1928: 70. 1929. Leptodontium arsenii Ther., Smiths. Misc. Coll. 85(4): 10. 1931. Type: Mexico, Michoacan, Morelia, Arsene 5073 (us-isotype). Erythrophyllumrubellum var. ruberrimum (Ferg. in Braithw.)Sav., Act. Inst. Bot. Acad. Sci. U.S.S.R., Ser. 2, 3: 522. 1936. Plants in cushions, red-brown to yellow-brown above, red-brown below; stems branching by subgametoecial innovations, stems 0.5-2.5(3.0) cm long, rounded-pentagonalin transverse section with a central strand, cortex of 1-2 layers of substereid cells to 3-4 layers of stereid cells, epidermis not differentiated or of a single layer of superficially thin-walled cells; axillary hairs of 4-6 uniseriate cells, all clear or the basal cell yellow-brown. Leaves increasing in size towards the stem apex, appressed-incurvedto weakly spreading and weakly twisted when dry, spreading to 45-60? from top of sheathing base when wet, ovate-triangularto short-lanceolate,mostly 1.0- 1.8 mm long, ventral surface broadly concave to nearly flat, margins recurved in lower Y2,entire, apex not apiculate to apiculate by 1-3 clear, weakly papillose cells, apiculus occasionally weakly serrulate, leaf base differentiated, distinctly sheathing, oblong, decurrent below costa; costa percurrent,ventral and dorsal superficial cells quadrateto short-rectangularand papillose above, in transverse section hemispherical to reniform, laminal insertion lateral to 60?, ventral stereid band weak, dorsal distinct, mostly 4 epidermal cells ventrally and 2-3(4) guide cells in one layer; upper laminal cells quadrate, with evenly thickened walls, 10-13 ,umwide, weakly patterned, papillae c-shaped to low and flat, about 4 per lumen; basal cells differentiated juxtacostally or across the leaf, hyaline to yellowish, usually papillose, to 12 ,umwide, mostly rectangular,2:1, walls evenly thickened; asexual propagula red to red-brown, obovoid to irregularly bulging, mostly 45-115 tim long, usually of several multiseriate cells, borne on basal rhizoids. Dioicous. Perichaetia terminal, inner leaves oblong-lanceolate, to 2.0 mm long, lower ?2 sheathing the seta. Perigonia terminal, weakly gemmate, inner leaves ovate-lanceolate, paraphyses uniseriate. Seta 1 per perichaetium, 0.7-0.8 cm long, brown, twisted clockwise; urn 2.2-2.5 mm long, elliptical, brown, smooth when dry, microstomous; exothecial cells 20-25 ,tm wide, 2-3:1, walls evenly thickened; stomataat base of urn, phaneropore; annulus of 1-2 rows of highly vesiculose cells, revolvable and deciduous in parts; peristome absent to rudimentary,teeth subulate, to 25 um 544 THE BRYOLOGIST [Volume 81

FIGURES1-21. Bryoerythrophyllumrecurvirostrum var. aeneum (C.Muell.) Zander and B. ferruginascens (Stirt.) Giac. - 1-6. B. recurvirostrum var. aeneum. - 1-3. Leaves, x30. - 4. Leaf medial transverse section, x300. - 5. Leaf apex, x300. - 6. Propagula, x77. - 7-21. B. ferruginascens. - 7-14. Leaves, x30. - 15. Leaf apex, x300. - 16-17. Propagula, x77. - 18- 19. Perichaetial leaves, x30. - 20-21. Capsules, x 17. Figures 1-6 from Mexico: Orcutt 3817a; 7-8 from Greenland: Holmen 16766; 9-12, 15-21 from Mexico: Sharp 1014; 13-14, 16 from Guatemala: Steyermark36732. long, pale yellow, not twisted, without basal membrane, spiculose; spores 12-15 Atmin diameter, weakly papillose, light brown; operculum 0.5-0.9 mm long, cells not twisted, short- to long- conic; calyptra not seen. Habitat: limestone, dolomite, thin soil over rock, cliffs, scree and fell field, bluffs, in seep, 100-3960 m elevation. Range: Arctic circumpolar regions, Canada, Greenland, Rocky and Appalachian Mountains of U.S.A., Mexico, Europe, northernAsia, possibly Japan. Bryoerythrophyllum ferruginascens is redescribed above from New World material to include characters of the sporophyte, which previously was unknown in this species. It is restricted in range in the New World to arctic and montane regions south into Middle America where it was reported from Mexico (Zander, 1972) from the type 1978] ZANDER: SYNOPSIS OF BRYOERYTHROPHYLLUM & MORINA 545 of the synonym Leptodontium arsenii Th6r. I have not seen collections from the Rocky Mountains in the U.S.A. but Weber (1973) reports B. ferruginascens from Col- orado mentioning presence of propagula. The geographic range has been poorly understood due to confusion in the herbarium with B. recurvirostrum. Small forms of B. recurvirostrum have much the same habit as B. ferruginascens but the leaves of the former are flaccid when wet and contorted when dry, the margins are recurved to near the apex and lateral teeth near the apex can usually be found in some leaves. The rhizoidial propagula used by various authors as a primary diagnostic character in the identification of B. ferruginascens are not always found, as noted by Dixon and Jameson (1924), but when present provide a good character (see key). Typical propagula were found in Mexico: Sharp 1014 (Fig. 16-17) and Virginia: Zander 4673. The illustration and description of the Japanese species B. gymnostomum (Broth.) Chen by Saito (1975) fit the present concept of B. ferruginascens, though propagula are not noted. An isotype collection of B. obtusissimus (Broth.) Chen (Japan: lishiba, 1907-NY), sterile but propaguliferous, is well within the limits of variation of New World B. ferruginascens. It is probable that fruiting plants of B. ferruginascens occur in Japan as well as in Mexico and have been previously described by Saito (1975) under another name or names. Barbula andreaeoides Kindb. (type: Canada, British Columbia, Macoun, 1904, CANM-isotype) and its synonym B. subandreaeoides Kindb. (type: Canada, British Columbia, Macoun, 1904, CANM-isotype) have been referred to the synonymy of Bryoerythrophyllum ferruginascens (Lawton, 1971). Barbula andreaeoides is, however, a good species easily distinguished by its large, simple, hemispherical to granular laminal papillae, 1(-3) per lumen and is actually closely related to Didymodon nigrescens (Mitt.) K. Saito (described and illustrated by Saito, 1975), being distinguished from the latter by the absence of sporophytes, compact growth form and presence of fragile, filiform branchlets with tiny, cochleariform leaves. Representative specimens examined. Greenland, Head of S0ndre Str0mfjord,67?N, Holmen 4196 (CANM);Upernivik Island, Inukavail Fjord, Niaquoranquaq, Holmen 16766 (us). U.S.A.: Alaska: Wiehl Mt., 17?39'N, 149?40'W, Murray 76-691A (ALA); Philip Smith Mountains, "Slope Mt.," 68?40'N, 149?03'W, Murray 76-410 (ALA).Virginia: Giles Co., Little Stoney Cascades Park, near Cascades, Zander 4673 (DUKE,NY). Mexico. Distrito Federal: near Morelos line, Sharp 1014 (NY). Mexico: W slope of Popocat6petl, Hermann 20841 (NY). Guatemala. San Marcos: Volcan Tajumulco, Steyermark35732 (FH).

3. Bryoerythrophyllum bolivianum (C.Muell.) Zander, comb. nov. (FIG. 22-39) Type: Bolivia, Cochabamba, Choquecamata, Germain, 1889 [det. c. Mueller sub. n. 1071] (H-BR-lectotype). Globulina boliviana C.Muell., Nuov. Giorn. Bot. Ital., N.S. 4: 39. 1897. Barbula boliviana (C.Muell.) Hilp., Beih. Bot. Centralbl. 50(2): 649. 1933. Plants loosely caespitose or gregarious to cushion-forming, red-brown to yellow-brown above, dark brown below; stems branching by subperichaetial innovations, to 1.0(-1.5) cm long, transverse section oval to pentagonal, central strand strong to weak, cortex weakly differentiated to 2-3 layers of stereid cells, epidermis not differentiated; axillary hairs of 4-8 uniseriate, clear cells, the basal 1-2 clear to light yellow. Leaves larger and more crowded near stem apex, appressed, terete, not contorted or twisted and apices often incurved when dry, weakly spreading to 30-45?, reflexed at top of short leaf base when wet, oval to short-triangular, ventral surface broadly concave across leaf, margins recurved from leaf base to near apex, occasionally weakly recurved to plane, entire, apex broadly rounded to narrowly obtuse, often somewhat cucullate or narrowly channeled, not apiculate; leaf base sheathing. Costa broad and flat, percurrent or ending 1-4 cells below apex, upper ventral and dorsal superficial cells quadrateto short-rectan- 546 THE BRYOLOGIST [Volume 81

2 -2A3 24 .25 26

22 _ _

FIGURES 22-46. Bryoerythrophyllum bolivianum (C.Muell.) Zander, B. recurvum (Griff.) - K.Saito and Morinia ehrenbergiana var. elongata (Wils. in Mitt.) Zander. 22-39. B. bolivianum. - 22. Stem apex, x 17. - 23-33. Leaves, x30. - 34. Leaf medial transverse section, x300. - 35. Leaf x300. - 36-37. Perichaetial leaves, x30. - 38. Capsule, x 17. - 39. Portion apex, - of and capsule mouth, x77. - 40-44. B. recurvum. - 40-44. Leaves, x30. 45-46. peristome - M. ehrenbergiana var. elongata. - 45. Leaf from sporophyte-bearing plant, x30. 46. Propag- uloid apex of leaf from sterile plant, x30. Figures 22, 27-30, 34, 36, 38-39 from Bolivia: Herzog 4853; 23-24 from Bolivia: Herzog 2985; 25-26, 35, 37 from Bolivia: Herzog 2576; 31-33 from Bolivia: Herzog 4126a; 40-41 from Mexico: Dull 2/62; 42 from U.S.A.: Steere 10100; 43-44 from U.S.A.: Zander 1089; 45 from Ecuador: Jameson s.n.; 46 from Ecuador: Jameson 30.

band gular, papillose, in transverse section elliptical, laminal insertion lateral, ventral stereid absent or weak, dorsal stereid band distinct, ventral epidermal cells 3-7 in one layer, 2-3 guide cells in one layer; upper laminal cells subquadrate, walls thin to evenly but weakly thickened, sub- superficially flat, 9-11 ,/m wide, occasionally longitudinally elongated 2:1 marginally or to marginally especially at apex when leaf apex is broadly rounded, papillae hollow, irregular multiplex, 2-6 per lumen, scattered; basal laminal cells differentiated medially or throughout lower Y2 of leaf, clear, smooth, 12-14 ,um wide, quadrate to short-rectangular, mostly 1-2:1, 1978] ZANDER: SYNOPSIS OF BRYOERYTHROPHYLLUM & MORINA 547 occasionally transverselyelongate, walls thin to evenly thickened; asexual reproductionprobably through fragile branches. Dioicous. Perichaetia terminal, inner leaves long-oval to oblong-triangular,apex often broadly obtuse, to 1.8 mm long, leaf base strongly sheathing the seta, laminal cells strongly differentiated, hexagonal-rhomboidial,smooth, throughout or in lower 3/ of leaf. Male plants about same size as female, perigonia terminal, weakly gemmate, perigonial leaves little differentiated. Seta (3.5-)0.5-0.6 cm long, red to dark brown, twisted clockwise below, counterclockwise above; urn 1.0-1.2 mm long, elliptical, microstomous,red-brown, smooth when dry; exothecial cells (15-)20-30 ,um wide, about 2:1, walls evenly thickened; stomata at base of urn, phaneropore;annulus of 1-2 rows of vesiculose cells, quickly deciduous in large portions; peristome apparentlyabsent to rudimentary,of 32 linear teeth, to 90-100 Amlong, white to light yellow, straight,with low basal membrane to 25 ,um high, spiculose-papillose; spores 12-16 Am in diameter, lightly papillose, yellow-brown; operculum 0.2-0.4(-0.6) mm long, quickly deciduous, cells not twisted, short-rostrateto short-conic; calyptra not seen. Habitat: "Red earth," high mountain pastures, 2500-4200 m elevation. Range: Peru and Bolivia. The genus Globulina was published validly by C. Mueller (1897) in a combined genus and species description with G. boliviana (Art. 42, I.C.B.N., 1969). (It is, however, the interpretation of the authors of Index Muscorum, Wijk et al., 1959-1969, that Brotherus (1902) first described the genus.) Although Mueller noted that Seligeria globifera Hampe (= Globulinella globifera (Hampe) Steere) was closely related and apparently in the same genus ("Globulina mihi"), he did not make the appropriate combination, leaving the genus monotypic with G. boliviana. No type was designated for the species but a specimen at H collected in Bolivia by Germain and determined as G. boliviana by Mueller provides a lectotype. Four years later, Mueller (1901) created Seligeria subg. Globulina C.Muell. with S. globifera the only species and therefore the type of that, different, taxon. Steere (Steere & Chapman, 1946) proposed Globulinella as a new genus, citing Globulina C.Muell., 1897 non Spegaz., 1889, in synonymy. Later in the publication he excluded the type of Globulina, G. boliviana. Apparently he did not intend Glo- bulinella as a nomen novum for Globulina as was suggested by Magill (1977). Ac- cepting Hilpert's (1933) transfer of G. boliviana to Barbula, Steere conceived of his new genus as a segregate including only Globulinella globifera and G. peruviana (Williams) Steere. Thus, Globulina C.Muell., hom. illeg., is not a synonym of Glo- bulinella Steere (Art. 7, I.C.B.N., 1969) but is instead a synonym of Bryoerythro- phyllum Chen. Seligeria subg. Globulina C.Muell. is a synonym of the valid genus Globulinella Steere, as they have the same type species, G. globifera (lectotype fide Wijk et al., 1959-1969). Magill (1977), in a treatment of Globulinella in the New World, did not mention study of Bryoerythrophyllum bolivianum. Globulinella species, however, are easily distinguished by the smooth or only medially papillose leaf lamina with usually in- crassate cell walls. Bryoerythrophyllum bolivianum is, additionally, illustrated by Brotherus (1902). It is apparently an Andean endemic but is closely related to B. ferruginascens. Fruit- ing plants usually are gregarious, have short stems, large leaves with plane, broadly obtuse apices; sterile plants are rather flagellate but are usually in dense turf, have relatively long stems, small leaves with weakly cucullate, narrowly channeled, ob- tusely acute apices. Bryoerythrophyllum bolivianum looks much like and is probably best considered a vicariad of the Asian B. rotundatum (Lindb. & Arn.) Chen, which differs mainly in being synoicous and eperistomate. Specimens examined. PERU. PASCO-JUNIN: Huamatanga and Cerro de Pasco, Mathews 2408 (NY). BOLIVIA. CHUQUISACA:Huaillas, Herzog, 1908 (JE). COCHABAMBA:Chocayatal, Herzog 3573 (NY); Llavetal, Herzog 4853 (FH, JE); Pojo, Herzog 4162a (JE); Valle de Llave, Herzog, 1908 548 THE BRYOLOGIST [Volume 81

(JE). LA PAZ: Araca, Herzog 2985 (FH, JE); Cacaltaya, Buchtien 5 (jE), 7 (E). Dept. (?): Challa, Herzog 2576 (JE).

4. Bryoerythrophyllum columbianum (Herm. & Lawt.) comb. nov. Basionym: Didymodon columbianus Herm. & Lawt., Bull. Torr. Bot. Club 95: 388. 1968. Type: U.S.A.: Oregon, Wasco Co., 10 km NE of The Dalles, S bank of Columbia River, along U.S. Rt. 30, ledge of basalt bluffs, dense sod on soil, Hermann 18751 (us-holotype); Washington, Walla Walla Co., Wallula, bluffs of Columbia R. gorge, sandy ridge, Foster, 1906 (Holz., Musc. Bot.-Amer. 233) (us-paratype). Barbula columbiana (Herm. & Lawt.) Herm. & Lawt., Bull. Torr. Bot. Club 99: 310. 1972. This species is described and illustrated by Hermann and Lawton (1968) and Lawton (1971). The weakly developed peristome, reddish coloration of the leaves, papillose ventral costal epidermis and large, mostly bifid laminal papillae usually 2- 3 per cell lumen are character states indicating proper placement in Bryoerythro- phyllum. It is intermediate in morphology between B. bolivianum and B. recurvum. The short leaves and broad, flattened costa are similar to those of the former while the firm leaves with acute apices and basal cells mostly quadrate are reminiscent of the latter. The acute leaf apex ending in a sharp mucro and the epapillose upper leaf margin are salient character states that, in combination, distinguish B. columbianum from other species of the genus. This species is known only from the type and paratype collections and is unusual in not being closely related to any species in the Old World. Other New World Bryoerythrophyllum species are either also found in the Old World or are presented there by closely related species.

5. Bryoerythrophyllum recurvum (Griff.) K.Saito, Univ. Mus. Univ. Tokyo Bull. 8: 254. 1975. (FIG. 40-44) Basionym: Gymnostomum recurvum Griff., Calcutta Jour. Nat. Hist. 2: 482. 1842. Desmatodon recurvus (Griff.) Mitt., Jour. Linn. Soc. Bot. Suppl. Jour. Linn. Soc. Bot. Suppl. 1: 37. 1859. Didymodon recurvus (Griff.)Broth., Nat. Pfl. 1(3): 405. 1902. Plants in turf or cushions, vivid-green to red-brown above, brown to red-brownbelow. Stems seldom branching, to 2.5 cm long, in transverse section rounded-triangularto rounded-pentagonal, central strand present, cortex of 1-2 rows of substereid to stereid cells, epidermis undifferentiated; axillary hairs of 3-4 cells, the basal one yellowish; stems weakly radiculose. Leaves distant to crowded, often distinctly larger near stem apex, when dry appressed to spreading to 30? from the insertion and apices incurved, when wet spreading to 45?, ovate-triangularto lanceolate, 0.9-1.5(-1.9) mm long, ventrally broadly channeled, margins plane to recurved in lower 3/4, entire, apex narrowly acute, occasionally acuminate, often weakly sinuose, apiculate by a single conical clear cell; leaf base ovate, usually not sheathing the stem, not decurrent; costa percurrent,usually sinuose above midleaf, ventral superficial cells quadrateto short-rectangular, 2:1, papillose, dorsal superficial cells short-rectangular,papillose, mostly 2-3:1, costa in transverse section elliptical, lamina inserted laterally, ventral epidermal cells 4-8 in one layer, ventral stereid band indistinct to strong, guide cells 2-4 in one layer, dorsal stereid band weak to strong, dorsal epidermis usually differentiated; upper laminal cells subquadrate, 10-12(-15) ,umwide, 1:1, walls thin to evenly thickened, longitudinal cell rows distinct, transverse rows indistinct, laminal papillae hollow, o- to c-shape in optical section, simple to bifid, mostly 4-6 per lumen, scattered to weakly centered over lumens; basal laminal cells similar to upper laminal cells, only differentiatedin a few rows near insertion orjuxtacostallyshort-rectangular and weakly papillose. Dioicous. Perichaetia terminal, apparently immature, leaves ovate-triangular,little larger than 1978] ZANDER: SYNOPSIS OF BRYOERYTHROPHYLLUM & MORINA 549 cauline leaves, cells prosenchymatousin lower /2, base sheathing. Perigonia terminal, gemmate. Sporophyte not seen. Habitat: rock, spray of falls, rocky slopes in montane oak forest, 600-2800 m elevation. Range: Nepal, India, U.S.A., Mexico. Bryoerythrophyllum recurvum is described and illustrated by Gangulee (1972) as Didymodon recurvus. It is here newly reported for the New World. The Appalachian populations differ from Old World specimens by a less deep-red coloration, the rather wider acumination and leaves more distant on the stem, with mostly plane margins. They have, however, the essential characters of leaves spreading from the insertion, not the top of the leaf base as in the similar B. ferruginascens, apices incurved when dry, leaf base ovate to weakly cordate, basal leaf cells not or little differentiated and costa usually sinuose above midleaf. The collections cited from Mexico approach Chen's (1941) illustration and description of B. tenerrimum (Broth.) Chen, which also has little-differentiated basal leaf cells, but these are here considered short-leaved forms of B. recurvum. Bryoerythrophyllum recurvum is similar in size and most morphological details to B. ferruginascens but differs most markedly in characters of the leaf base, which affect leaf stance. In dry plants the protruding, cup-like leaf bases and incurved leaf apices are distinctive. Variation occurs in leaf shape, amount of marginal flexion and in degree of acumination of the apex-well developed Asian specimens having highly recurved margins and long, whip-like acuminations.

Specimens examined. U.S.A. NORTH CAROLINA, MACON CO.: Highlands, Horse Cove, Steere 10100 (MICH); RUTHERFORD CO.: Broad R. gorge, Rainbow Falls, Zander 1089 (DUKE), 1095 (DUKE). MEXICO. MEXICO: El Rosal, between San Juan del Rio and Jilotepec, Dull 2/62 (TENN); 10 km W of Ixtapalucca, Dull 2/95 (TENN). NEPAL. Between Dor and Chauke, Iwatsuki 437 (NY). INDIA. ASSAM:Between Gauhati and Loharghat, Srivastava 2223i (BUF).

6. Bryoerythrophyllum jamesonii (Tayl.) Crum, Svensk. Bot. Tidskr. 51: 200. 1957. TYPE: Ecuador, Pichincha, Quito, Jameson 88 (NY-isotype). (FIG. 47-69) Basionym: Barbula jamesonii Tayl., Lond. Jour. Bot. 5: 48. 1846. Syrrhopodonjamesonii Tayl., Lond. Jour. Bot. 6: 331. 1847. Type: Ecuador, Pichincha,Jame- son, s.n. (NY-isotype). Trichostomumcampylocarpum C.Muell., Bot. Zeitung 9: 261. Apr., 1851. Type: Costa Rica, s. loc., Oersted, s.n. (NY-syntype). Syrrhopodoncrispatus Hampe, Ann. Sci. Nat. Bot. 5(5): 335. 1866. Type: Colombia, Distrito Especial, Bogota, San Juan, anon., s.n. (NY-isotype). Tortula arcuata Mitt., Jour. Linn. Soc. Bot. 12: 163. 1869. Type: Ecuador, Pichincha, Quito, Jameson 98 (NY-lectotype fide Crum, 1957; CAN-M-isotype). Trichostomumamblystegium C.Muell., Nuov. Gior. Bot. Ital., N.S. 4: 118. 1897. Trichostomum arcuatum (Mitt.) C.Muell., Nuov. Giorn. Bot. Ital., N.S. 4: 119. 1897, hom. illeg. non Cand. in Lam. & Cand., Fl. Franc. ed. 3, 6: 218. 1815. Trichostomum campylopyxis C.Muell., Nuov. Gior. Ital., N.S. 4: 118. 1897. Type: Bolivia, Cochabamba, Choquecamata, Germain, 1889 [det. C. Mueller sub. n. 1195] (NY-lectotype). Didymodon campylocarpus (C.Muell.) Broth., Nat. Pfl. 1(3): 405. 1902. Leptodontium albovaginatum Herz., Beih. Bot. Centralbl. 26(2): 61. 1909. Type: Bolivia, Cochabamba,Incacorral, Herzog, 1908 (M-holotype). Didymodon calymperidictyon Broth. in Skottsb., Nat. Hist. Juan Fernandez 2: 419. 1924. Didymodon linearis Broth. in Skottsb., Nat. Hist. Juan Fernandez 2: 419. 1924, hom. illeg, non Sw., Adnot. Bot. 100. 1829. Leptodontium subplanifolium Ther., Rev. Bryol. Lichenol. 9: 17. 1936. Type: Ecuador, Pi- chincha, Rumipamba, Benoist 2896 (BM, NY, S-PA-syntypes). Bryoerythrophyllumarcuatum (Mitt.) Crum, Svensk. Bot. Tidsk. 51: 199. 1957. Bryoerythrophyllumcampylocarpum (C.Muell.) Crum, Svensk. Bot. Tidsk. 51: 200. 1957. 550 THE BRYOLOGIST [Volume 81

65 K C(J{/ 62 63\( 671l/ 68 1 FIGURES47-68. Bryoerythrophyllumjamesonii (Tayl.) Crum. - 47-55. Leaves, x30. - 56. Upper marginal leaf cells, x733. - 57-59. Medial costal transverse sections, x300. - 60-61. Perichaetial leaves, x30. - 62-68. Capsules, x 17. Figures 47-49, 64 from Colombia: Killip & Smith 18069; 50 from Colombia: Cleef4229a; 51, 60-61, 62-63 from Ecuador: Pichincha, Spruce s.n.; 52 from Ecuador: Cargnairazo, Spruce s.n.; 53 from Jamaica: Crosby 3527; 54, 57 from Ecuador: Jameson s.n.; 55, 59 from Colombia: Cleef 4834; 56 from Holm-Nielson & Jeppeson 1086; 58 from New Zealand: Sainsbury, 1929; 65 from Bolivia: Herzog 2929; 66 from Ecuador: Jameson 98; 67-68 from Jameson 88. 1978] ZANDER: SYNOPSIS OF BRYOERYTHROPHYLLUM & MORINA 551

Plants loosely caespitose to cushion-forming, red-brown to yellow-brown; stems branching by subperichaetial innovations, mostly 0.2-2.0(-3.5) cm long, in transverse section pentagonal to occasionally rounded-triangular,central strand present, cortex of 1 layer substereid cells to 2- 3 layers stereid cells, epidermis not differentiated or occasionally with superficial cell walls thin and collapsed; axillary hairs of 4-14 uniseriate clear cells; stems weakly radiculose, rhizoids smooth to aculeate. Leaves of nearly equal size along stems, often ratherdistant, erect-spreading, occasionally incurved, twisted when dry, spreading to 30-45? when wet, long-oblong to oblong- lanceolate, 1.0-3.5(-5.0) mm long, lower leaves occasionally bearing rhizoids at apices, ventral surface broadly channeled to keeled, margins recurved at top of sheathing leaf base, otherwise plane, entire to distantly strongly dentate by clear, smooth cells, 1-2 rows of marginalcells often less papillose and with somewhat thicker walls, forming a yellowish border; apex narrowlyacute to obtuse; leaf base sheathing the stem, ovate-oblong, occasionally narrowly decurrent at the margins; costa short-excurrentin a mucro, percurrent or ending 1-3 cells below apex and apex apiculate, ventral superficial cells quadrate, rhomboidal, short- or long-rectangular,papillose to smooth and dorsal superficial cells long-rectangular,papillose to smooth, in transverse section reniform, laminal insertion 45-90?, ventral stereid band weak to strong, dorsal stereid band strong, ventral epidermal cells absent to 4 in one layer, guide cells 4-5 in one layer; upper laminal cells quadrate to rounded-quadrate,occasionally short-rectangular,2:1, elongated longitudinally or transversely,walls evenly thickened, mostly 7-9(-12) ,tm wide, patterned in rather even longitudinal rows and more irregulartransverse rows; papillae crowded, low, flat, multiplex, 1-2 per lumen to irregular,low, scablike, small, simple to bifid, 6-12 per lumen, seldom hollow, centered to scattered across the lumens, mostly 4-8 salients per lumen; basal cells differentiated medially to occasionally across the leaf base, often as hyaline fenestrations, clear to yellow, smooth to occasionally weakly papillose, little wider than upper laminal cells to bulging, to 22- 24 ,tm wide, mostly 2-4:1, thin- to occasionally thick-walled, basal marginal cells not differentiated or more commonly short- to long-rectangular,not bulging, smooth. Dioicous. Perichaetia terminal, inner leaves long oblong-lanceolate, 2.0-2.8 mm long, lower ?2 sheathing seta. Male plants somewhat smaller than female, perigonia gemmate, inner leaves with sheathing bases, ovate-triangularto ovate-lanceolate, paraphyses uniseriate, numerous. Seta (0.9-)1.2-1.5(-2.1) cm long, 1(-2) per perichaetium, red- to yellow-brown, twisted clockwise below, often counterclockwise above; urn (0.7-)1.0-2.0(-4.0) mm long, macrostomous, straight to strongly curved, cylindrical or occasionally ovoid, red-brown, smooth when dry; exothecial cells mostly 18-28 ,tm wide, 2:1, walls evenly thickened or longitudinal walls often greatly thickened, 4-5 rows of cells below capsule mouth often quadrate and with very thick walls, forming a sturdy ring; stomata at base of urn, phaneropore;annulus of 1-2 rows of weakly vesiculose cells, not deciduous or occasionally deciduous in pieces; peristome teeth 32, linear, often short, fragile, often broken off above base, wider and perforatebelow, with up to 10-12 articulations,(30-)100-275(- 700) ,tm long, red-brownto orange, occasionally yellow-brown, straight,basal membraneabsent, densely spiculate or spiculate grading to closely spirally ridged; spores (12-)14-16(-19) ,um in diameter, weakly papillose, yellow-brown; operculum 0.4-1.2 mm long, cells not twisted, short- conic to conic; calyptra 2.0-2.1 mm long, smooth. Habitat: Rock, soil, clay, black volcanic rock and dust, dry mountain slope, ridge top, rain forest cloud forest; near sea level to 4100 m but commonly 2400-3000 m elevation. Range: Canada(British Columbia), Mexico, Guatemala,Costa Rica, Jamaica,Haiti, Colombia, Venezuela, Ecuador, Peru, Bolivia, Chile (Juan Fernandez), China, India, Australia(Tasmania), New Zealand, Scotland. Bartram (1949), Brotherus (1924), Crum (1957), Herzog (1909) and Sainsbury (1955) discuss and illustrate this species under various synonyms. Bryoerythrophyllum jamesonii is a highly variable species. I include in its synonymy, as did Bartram (1949) tentatively, two names recently recognized by Crum (1957), B. campylocarpum and B. arcuatum. A large-small "stature gradient" (Zander, 1977b) of phenotypic variation is evident. Very robust plants (including the type of B. jamesonii) are distinctive in the extreme by the following combination of character states, not all of which are always present: plants often 3-5 cm tall, with green branches, stems rounded-pentagonal in transverse section, leaves oblong to long-lanceolate, mostly 3.0-3.5(-5.0) mm long, reflexed above an oblong leaf base, margins distantly dentate above, revolute along leaf base, laminal cells weakly papillose, costa dorsally smooth, with quadrate, papillose ventral epidermal cells, sporophyte urn straight, without a circumstomal 552 THE BRYOLOGIST [Volume 81 ring, to 4.0 mm long, operculum to 1.1 mm long, peristome to 700 ,/m long, annulus strong, deciduous in pieces. Small plants (including the type of B. arcuatum) approach this combination of character states: plants mostly to 1.5 cm tall, entirely red, stems rounded-triangular in transverse section, leaves long-oblong, occasionally wasp- waisted, mostly 1.5-2.0 mm long, not or weakly reflexed above an elliptical leaf base, margins entire except for an apiculus of smooth, conical cells, plane to recurved along leaf base, laminal cells strongly papillose, costa dorsally papillose, with elongate, smooth ventral epidermal cells, sporophyte urn commonly curved, with a rigid circumstomal ring of 4-5 rows of thick-walled exothecial cells, urn almost as short as the operculum is long, peristome rudimentary, annulus weak, not deciduous. Variation in the sporophyte is also described by Crum (1957). Intermediates, with various com- binations or expressions of the above character states, are too common to allow recognition of infraspecific categories. The details of variation are rather unlike those described (Zander, 1977b) for stature gradient character variation in the Pleuroweisieae. It is too early yet to say whether related species of mosses that exhibit stature gradients vary in similar ways, as a parallel to Vavilov's (1951) "Law of Homologous Variation" described for cultivated plants. Besides, the great probability of extensive environmental modification has been a bugaboo in bryophyte taxonomy. Lodge (1960) demonstrated through culture study that parallel series of variation between two species of Drepanocladus were environmental modifications. Corley (1976), in culture studies of Campylopus species, showed that the limits of variation were not the same for every species, and variants tended to retain only size differences, losing other varietal characters. Other recent workers, e.g. Briggs (1965), Longton (1974) and Wigh (1975), have demonstrated that, in some unrelated moss taxa, many characters in the past considered taxonomically significant are environmental modifications. However, some of these authors and others, e.g. Crundwell (1956), Crundwell and Nyholm (1964) and Whitehouse (1966), demonstrate that certain taxonomic forms, varieties and closely related species are at least partially stable in culture and therefore presumably genetically based. The culture work of Hatcher (1967) gives evidence that both environmental modification and widespread differentiation of genetic races contribute to the morphological and de- velopmental variability of the hepatic Lophocolea heterophylla-the environmental factors affecting the same taxonomic characters and producing about the same range of variation as do the genetic factors. In any case, taxa with gradients in morphological variation are apparently rather common in mosses and should be expected and allowed for in taxonomic treatments. For example, During (1977) found that some species of Garovaglia show a range of variation (in a series of subspecies) parallel to that of other species but patterns of variation in the largest sections were different. Steere (1938a) suggested that Barbula brachyphylla Sull. was the ". . . most reduced form in the extremely variable B. vinealis-B. cylindrica complex ...." Pursell (1976) noted that the Fissidens bryoides complex is ". . . an intricate morphologically intergrading series .... Beginning with F. exiguus, morphologically the smallest and least differentiated of the variants . . . ," supporting the opinion of Crum (1973). The stature gradient in B. jamesonii is far more complex than that of B. recurvirostrum, which has no wide variation in morphology between large and small forms. This may be due to a possible higher level of out-breeding in the former, dioicous, species than in the latter, monoicous, species, as discussed by Gemmell (1950) for species of the British moss flora, or to the patchy "insular" (Carlquist, 1974) nature of the mostly montane range of the former. In some collections (e.g. Jamacia: Robbins 53 and Australia: Tasmania, Weymouth 1978] ZANDER: SYNOPSIS OF BRYOERYTHROPHYLLUM & MORINA 553

1220) the ventral surface of the costa is only partially covered by parenchyma leaving the ventral stereid band superficially exposed (Fig. 58) in a narrow medial strip. Small forms with elongate ventral superficial cells on the costa may have the ventral stereid band completely exposed, a character that is typical of and constant in the easily confused genus Leptodontium (Zander, 1972). Bryoerythrophyllum jamesonii may be distinguished from Leptodontium species with similar appearance by the red coloration, the central strand of the stem present and often large, the oblong leaves when dry reflexed below and incurbed above, and the laminal papillae often bifid, usually 4-6 crowding the surface over each cell lumen, apparently scattered. The peristome teeth of B. jamesonii are red and grade from closely spiculose to spirally grooved, often in the same collection, e.g. Mexico: Purpus 4287, Amable 1248, while the capsule may be straight or variously curved, again often in the same collection. Such arcuate, amblystegioid capsules are considered minor variations in Bryoerythrophyllum by Bartram (1949), in Hymenostylium by Zander (1977b) and in Fissidens by Pursell (1976). A sterile collection from the western coast of Canada (British Columbia: Schofield 15257) originally identified as B. alpigenum (Vent.) Chen but annotated by H. Crum as B. recurvirostrum, differs from Andean B. jamesonii only in the rather long leaves, to 5 mm, about 1 mm longer than those of most of the larger Latin American specimens but closely matched by Colombia: Cleef & Cleef 4834. This is the only New World specimen of B. jamesonii from north of the Mexico-U.S.A. border. This northern disjunction is unusual but not incongruous in view of the known stations in China and India, evidence that B. jamesonii may be widely distributed in mesic, montane cir- cum-Pacific areas. The collection from Scotland: Binsted & Dixon, 1898, is discussed by Dixon and Jameson (1924) as a reddish form of Leptodontium flexifolium (Dicks. ex With.) Hampe in Lindb. but has a strong central strand and otherwise matches robust Andean B. jamesonii. The known geographic distribution (Fig. 69) of Bryoerythrophyllum jamesonii is similar to that of Molendoa sendtneriana (Zander, 1976). The latter species is apparently able to colonize more harsh environments but has the same general restriction to mountainous areas, especially along axes of Tertiary folding. A similar disjunction of range between montane Latin America and the Himalayas is noted (Zander, 1972) for the two most widespread species of Leptodontium. I suspect that an Andean- Himalayan distribution pattern is not unusual in the Pottiaceae; however, little is known of the floristic relationships of the Andean moss flora (Delgadillo, 1971). Ful- ford (1951), in a discussion of the distribution patterns of South American leafy hepaticae, pointed out that disjunctions at least at the generic level between South America and the Sikkim-Nepal region of Asia are well known and possibly ultimately related to continental drift. The east-west disjunction of B. jamesonii between northwestern North America and Great Britain in western Europe is the same as that of B. recurvifolium and essentially that of several other species of a floristic element characteristic of northern hyperoceanic climates, as discussed by Schofield (1969) and Schofield and Crum (1972). Perhaps some of these other northern hyperoceanic species may also occur in the Andes. The presence of B. jamesonii in Great Britain is remarkable but not anomalous. Both Orthodontium gracile (Wils.) Schwaegr. (Bry- aceae) and Leptodontium flexifolium were first described from British material but were later found to be widespread throughout the Latin American cordillera. As An- drews (1950) pointed out, such should be considered essentially Andean species that are also present in Europe, not vice versa as conceived by earlier authors. The geographic distribution of B. jamesonii, basically a Latin American cordillera-Himalayan 554 THE BRYOLOGIST [Volume 81

FIGURE69. Known geographic distribution of Bryoerythrophyllumjamesonii (Tayl.) Crum.

disjunction with outliers in hyperoceanic areas including northwestern North Amer- ica, northwestern Europe and New Zealand, may well prove to be similar to that of other Andean species. Possible historical factors that may have determined the world-wide disjunctive range of Bryoerythrophyllum jamesonii, i.e. past fragmentation by continental drift and climate change, must be viewed only as additional influences to long distance dispersal because of the presence of this species on the oceanic island of Juan Fer- nandez. I thank H. H. Iltis (pers. comm.) and G. Pierce (pers. comm.) for pointing out that species with oceanic island stations cannot be used to demonstrate effects of continental drift (= Iltis' "Long Distance Dispersal Exclusion Principle"). Although Van Zanten's (1976) experiments do not indicate that long distance dispersal is easily effected in mosses, the many species of continental distribution that are also found in the bryoflora of Hawaii (Hoe, 1974) are evidence that long distance dispersal is of significance in determining at least portions of some disjunctive distribution patterns. Historical factors are often over-emphasized by bryologists. For instance, I have previously indicated (Zander, 1972) that the presence in both New and Old Worlds of Leptodontium species that are not closely related is evidence that the genus is old and most patterns of dispersal were established long ago. This assertation is valid only if historical factors are most important in determining distribution. However, of the seven American species of Leptodontium that are disjunct in range to the Old World, four are also found on oceanic islands. Overlapping disjunctive distribution patterns are also found for Bryoerythrophyllum species and vicariads and the same caveat as to their bryogeographical significance is necessary. Relative robustness in B. jamesonii, especially in length of stems and of leaves, is correlated with montane habitat, especially paramos, or otherwise with very mesic climate. Likewise, judging from specimens at NY and elsewhere, Andean populations of Barbula replicata Tayl. are strikingly more robust than Mexican populations (of the synonym B. spiralis Schimp. ex C.Muell.), although there are many exceptions. The single known Andean collection of Rhabdoweisia crenulata (Mitt.) Jameson reported and discussed by Zander (1977a) is far larger than specimens examined from North America or Europe. Montane gigantism is also associated with populations in the southern Appalachian Mountains of the eastern United States of Oxystegus tenuirostris (Hook. & Tayl.) A.J.E. Sm., discussed here under Bryoerythrophyllum recurvifolium and by Crum and Anderson (1958), and in Diphysciumfoliosum (Hedw.) Mohr. In the latter, montane collections have leaves 1-2 mm longer than usual, as is 1978] ZANDER: SYNOPSIS OF BRYOERYTHROPHYLLUM & MORINA 555

well documented in collections at DUKE. In some species, larger, montane specimens are distinguishable at the varietal level by additional characters as in Morinia ehrenbergiana, q.v. Blunt-leaved forms of Anoectangium aestivum (Hedw.) Mitt., when sterile, may be mistaken for B. jamesonii but lack the red coloration, have only one stereid band in the costa and have weakly differentiated basal leaf cells. The species Bryoerythro- phyllum angustulum (Herz.) H.Robins., regarded by Robinson (1967) as close to B. campylocarpum (= B. jamesonii) recently has been referred (Robinson, 1975) to Tri- chostomopsis australasiae (Hook. & Grev.) H.Robins.

Representative collections examined. CANADA. BRITISH COLUMBIA:Queen Charlotte Is., Moresby I., Hibben I., Schofield 15257 (CAN-M). MEXICO. DISTRITO FEDERAL: Vallee de Mexico, Desierto, Amable 1248 (us). JALISCO: La Ferreria, above Manantlan, Crum 1161 (TENN). MtX- ICO: Popocatepetl, Barkley, Rowell & Webster 2348 (CAN-M). OAXACA: Sierra Juarez, Sharp, 1962 (us). PUEBLA: Boca del Monte, Purpus 4287 (us). VERACRUZ: El Puerto, above Acultzingo, Sharp 594 (CAN-M).GUATEMALA. QUETZALTENANGO: Volcan Santa Maria, Steyermark 34092 (FH). COS- TA RICA. SAN JOSE:Cordillera de Talamanca, Cerro de la Muerte, Cleef & Fournier 10220 (BUF). PIEDRA BLANCA, Valerio 90 (us). HAITI. Above Bodaric, Orcutt 8315 (CAN-M). JAMAICA. Blue Mt. Peak trail, Robbins 53 (CAN-M);Portland, Orcutt 5154a (CAN-M); Crosby 3527 (DUKE). VENE- ZUELA. TRUJILLO: Vajo-La Morita, Medina & Aristequieta 1328 (CAN-M). COLOMBIA. BOYACA: Paramo del Pisva, Socha-La Punta highway, km 72, small valley, Cleef 4229a (BUF); Chiquin- quira, Lindig, 1863 (NY). CUNDINAMARCA:Valle Rio Teusaci, 5 km SW of La Calera, Cleef & Cleef 4834 (BUF); Zipaquiri, Schultes & Bell 11447 (CAN-M). SANTANDER: La Baja, Killip & Smith 18069 (NY). ECUADOR.Cargnairazo, Spruce s.n. (NY). AZUAY: Cuenca, Harling 2190 (CAN- M). COTOPAXI: Pilalo, Holm-Nielsen &Jeppesen 1083 (us), 1086 (us). PICHINCHA: Pichincha Mt., Spruce 163 (NY, US), s.n. (NY). PERU. CUZCO: Machu Picchu, above Urubamba R., Worthley, 1962 (us). BOLIVIA. COCHABAMBA: Tablas, Herzog 2929 (cu). LA PAZ: Unduavi, Rusby 3126 (NY); Sorata, Williams 1722 (NY). CHILE. JUAN FERNANDEZ ISLANDS: Masatierra, Hatcher & Engel 186 (us). GREAT BRITAIN. SCOTLAND: Argyll Co., Ben Douran, Binstead & Dixon, 1898 (BM). CHINA. YUNNAN: Akalii, near Lidjiang, Handel-Mazzetti 6837 (us). INDIA. SIKKIM:Jongri, Hooker 167 p.p. (NY-segregated from syntype of Bryoerythrophyllum wallichii). AUSTRALIA.TASMANIA: Hobart Rivulet, Weymouth 1220 (NY). NEW ZEALAND. North Island, Wairoa, Sainsbury, 1948 (NY), 1929 (NY).

7. Bryoerythrophyllum recurvifolium (Tayl.) Zander, THE BRYOLOGIST 75: 277. 1972.

Basionym: Bryum? recurvifolium Tayl., Ann. Mag. Nat. Hist. 11: 208. 1843. Didymodon? recurvifolius (Tayl.) Wils., Bryol. Brit. 110. 1855. Leptodontium? recurvifolium (Tayl.) Lindb., Ofvers. Forh. Kongl. Svenska Vet.-Akad. 21: 227. 1864.

This species, described and illustrated by Dixon and Jameson (1924), is known only from sterile material from western Europe and northwestern North America, growing on wet rock, soil, cliff, talus slopes and along streams, often among grasses. It is closely related to B. wallichii (Mitt.) Chen of Asia (illustrated by Saito, 1975), which differs in the reddish coloration and presence of a central strand. Both B. recurvifolium and B. wallichii differ from B. jamesonii by the highly differentiated leaf border and rather narrow geographical ranges. Certain robust, montane forms of Oxystegus tenuirostris that match Dixon's (Dixon & Jameson, 1924) description ofvar. holtii (Brithw.) A.J.E. Sm. (e.g. U.S.A.: North Carolina, Macon Co., Dry Falls, Zander 4499; Glen Falls, 4554; Highlands Falls, 4543; Transylvania Co., Looking Glass Falls, Zander & Mansberg 4594, all in BUF & DUKE) with dark green, broad, oblong leaves becoming blackish on lower portions of stems and having weakly papillose, weakly dentate upper leaf margins approach closely the appearance of 556 THE BRYOLOGIST [Volume 81

B. recurvifolium. However, I agree with Crum and Anderson (1958) that forms such as these intergrade completely with the more usual expression of 0. tenuirostris with long-lanceolate leaves, undifferentiated or distantly notched leaf margins and comparatively massive laminal papillae. Specimens examined. CANADA. BRITISH COLUMBIA: Queen Charlotte Is., Moresby I., Scho- field 24950 (CAN-M),24980 (CAN-M),25146 (CAN-M),34457 (CAN-M,TENN), 37377 (CAN-M,us), Schofield & Sharp 25878 (CAN-M,TENN); Graham I., Schofield & Krajina 39737 (CAN-M,TENN). U.S.A. ALASKA:Adak I., Finger Bay, Smith 3693a (TENN). GREAT BRITAIN.SCOTLAND: s. loc., Jack, 1910 (FH); Inverness: West Inverness, Nicholson, 1909 (FH); Ross and Cromarty, Ben Dearg, Baleall 3479 (CAN-M);Perth, Ben Vorlich, anon., 1865 (BUF). WALES:Merioneth, Rhaiadr Du, Tyn-y-Graes, Evans et al., 1954 (us). IRELAND.KERRY: Brandon Mt., Binstead 896 (us).

Morinia Card., Rev. Bryol. 37: 124. 1910. Type species: M. trichostomoides (Besch.) Card. (= M. ehrenbergiana (C.Muell.) Ther.). Plants usually 0.5-4.0 cm tall, yellow- to red-brown, seldom branching, stem central strand usually strong. Leaves of about equal size to near base, ovate-lanceolate to oblong-lanceolate, keeled, with a narrow groove ventrally along the costa, margins narrowly recurved in lower 4, narrowly bistratose in upper Y3, entire or dentate, leaf apex occasionally thickened as a multistratose propagulum,leaf base usually oblong and sheathing; costa percurrent,ventral superficial cells quadrate to rectangular, papillose, dorsal superficial cells rectangular, smooth, costa in transverse section semicircular to reniform, ventral surface usually concave, lamina inserted ventrally at 60-90? angle, ventral stereid band usually small but dorsal strong, with 2-5 guide cells, dorsal epidermis thick-walled, with small, semicircular lumens; upper laminal cells subquadrate to short-rectangular,to 2:1, with walls evenly thickened, superficially flat, mostly homogeneous in shape and size; laminal papillae usually solid, flattened, 2-6 per lumen, granular to multiplex, crowded; basal laminal cells differentiated across leaf base, rectangular, weakly papillose to smooth. Asexual reproductionwhen present probably through deciduous, thickened leaf apices. Dioicous. Perichaetia and perigonia terminal,perichaetial leaves elongate and sheathing. Sporophyte seta elongate, urn long-cylindrical, peristome of 32 red, densely spiculose teeth, twisted 2.5 times above a short, tesselated basal membrane, annulus of 1-2 rows of vesiculose cells, operculum long-conic, calyptra not seen. This distinctive genus has been discussed by Bartram (1949), Brotherus (1924) and Cardot (1910). It has most of the characters of Bryoerythrophyllum except that the peristome teeth are long and twisted, the leaves are commonly keeled, with margins narrowly bistratose and the apex is often differentiated as a multistratose propagulum. Laminal papillae are usually solid in mature leaves.

la. Morinia ehrenbergiana (C.Muell.) Ther. var. ehrenbergiana, Smiths. Misc. Coll. 85: 22. 1931.

Basionym: Barbula ehrenbergiana C.Muell., Synop. Musc. 1: 636. 1849. Type: Mexico, s. loc., Ehrenberg s.n. (BM-isotype). Barbula trichostomoides Besch., Mem. Soc. Natl. Sci. Nat. Cherbourg 16: 182, 1872. Type: Mexico, D.F., Desierta Vieja, Bourgeau 1335 (PC-holotype; NY-isotype). Morinia trichostomoides (Besch.) Card., Rev. Bryol. 37: 124. 1910. Plants in cushions or mats; stems mostly 2-4(-8) mm long, in transverse section rounded- pentagonal, cortex of 1-2 layers of substereid or stereid cells, epidermis not differentiated; axillary hairs to 8 cells in length, hyaline, basal cell usually somewhat thicker-walled and weakly yellowish; stem occasionally matted with rhizoids. Leaves crowded near stem apex, when dry erect-twisted, when wet erect-spreadingto 45? from top of sheathing leaf base, ovate- to oblong- lanceolate, 2.5-3.2(-3.5) mm long, margins narrowly recurved in lower 3/4, occasionally to near apex, sharply dentate in upper 1/5 to Y2, occasionally obscurely dentate or entire; leaf apex broadly to narrowlyacute, not differentiated as a propagulum;leaf base oblong, sheathing, mar- 1978] ZANDER: SYNOPSIS OF BRYOERYTHROPHYLLUM & MORINA 557 gins occasionally narrowly decurrent; costa excurrent as a short, sharp, stout mucro, ventral superficial cells short-rectangular and papillose to long-rectangular and smooth; upper laminal cells mostly 8-10 gm wide, 1-2:1, weakly patterned; laminal papillae solid, low, spiculose-bifid to flattened multiplex, crowded, mostly 2-4 per lumen, apparently scattered; basal laminal cells light yellow-brown, smooth to weakly papillose with scattered bifid papillae, 14-16 tm wide, not bulging, 5-8:1, walls thin to evenly thickened. Dioicous. Male plants same size as female. Perichaetial leaves oblong-lanceolate, prosenchymatous in lower 3/4,5-6 mm long, strongly sheathing. Perigonia gemmate, inner leaves ovate-lanceolate, paraphyses abundant, cells uniseriate. Sporophyte seta about 2.0 cm long, red-brown, twisted clockwise below, counterclockwise above; urn 5-6 mm long, red-brown, smooth when dry, neck short, exothecial cells rectangular, 20-30 ,um wide, 4-6:1, walls often longitudinally thickened; stomata phaneropore, at base of urn; annulus of 1-2 rows of vesiculose cells; peristome often broken off at base, twisted counterclockwise, basal membrane about 90 /m high. Operculum about 1.5 mm long, cells strongly twisted about 2.5 times counterclockwise. Calyptra not seen. Spores 9-11 tm in diameter, weakly papillose, light brown.

Morinia ehrenbergiana var. ehrenbergiana is illustrated by Bartram (1949). I have seen collections from Mexico, Guatemala, Venezuela (Griffin, 1975) and Colombia; Robinson et al. (1977) report it from Ecuador. The synonymy presented above agrees with that of Theriot (1931). The habitat includes paramos, cloud forests, soil, rock, bark, walls, banks, from 2700-4200 m elevation.

lb. Morinia ehrenbergiana var. elongata (Wils. in Mitt.) Zander, comb. & stat. nov. (FIG. 45-46) Basionym: Barbula elongata Wils. in Mitt., Kew Jour. Bot. 3: 51. 1851. Type: Ecuador, Quito, Jameson 7 (BM-holotype). Tortula elongata (Wils. in Mitt.) Mitt., Jour. Linn. Soc. Bot. 12: 164. 1869, hom. illeg. non Funck, Flora 15: 482. 1832. Morinia ecuadorensis Bartr.,Bull. Brit. Mus. Nat. Hist. 2(2): 55. 1955. Type: Ecuador, San Juan, Bell 727 (FH-isotype). The var. elongata has overlapping geographic range and some intergradation of characters with the typical variety, but the characters noted in the key will distinguish most collections. The type of M. ecuadorensis has propaguliferous and non-propaguliferous leaves on the same stems. Specimens of the var. elongata at NY lack propagula on stems that bear sporophytes. The sporophytes are little different from those of var. ehrenbergiana except for a somewhat longer urn, to 6 mm. Small, non-propaguliferous specimens of the var. elongata are probably indistinguishable from the typical variety. Tortula herzogii, nom. nov. for T. angustifolia (Herz.) Herz. (Basi- onym: Calyptopogon angustifolius Herz., Beih. Bot. Centralbl. 26(2): 63, 1909), of Bolivia and Peru (Peru: Ancash, Laguna Llanganuco, Hegewald & Hegewald 7522- BUF) has similar leaf shape and channeled, fragile, apical propagulum but has only one stereid band in the costa, leaf marginal cells differentiated but mostly unistratose, epapillose, yellow and short-rectangular and upper leaf cells very thin-walled, with 5-7 large, hollow, simple papillae per lumen. Additional collections examined. ECUADOR. PICHINCHA: Quito, Jameson 7b, 89, s.n., 1874 (all at NY), Jameson 30 (BM).

2. Morinia crassicuspis (H.Robins.) Zander, comb. nov. Basionym: Barbula crassicuspis H.Robins., THE BRYOLOGIST 67: 446. 1964. Type: Mexico, Oaxaca, Sierra Juarez, Llano de las Flores, Sharp, 1962 (us- holotype; TENN-isotype). 558 THE BRYOLOGIST [Volume 81

This species is described and illustrated by Robinson (1964). It has been found in Mexico and Guatemala in habitats including: parklike area at edge of grassland, bark, oak bark, 30004200 m elevation. Both species of Morinia in the New World have in common narrowly bistratose leaf margins and propagula consisting of ventrally narrowly channeled fleshy leaf apices. Morinia crassicuspis is known only from sterile material, and the bistratose leaf margins are of only 1-2 cell rows, difficult to distinguish except in section. The propagulum is quite different from that of M. ehrenbergiana var. elongata as noted in the key. Plants of M. crassicuspis may have some leaves non-propaguliferous, ending in a sharply acute, entire apex, and others with propagula (sometimes only partially differentiated) on the same stems (e.g. Guatemala: Steyermark 34093). Propagula never remain on mature leaves and probably are caducous early. In M. ehrenbergiana var. elongata, propagula usually remain on mature leaves and are apparently not caducous, merely fragile. Additional specimens seen. MEXICO.JALISCO: La Ferreria, above Manantlan, Crum 1097a (MICH);Puebla, Esperanza, Purpus 42906 (MICH). GUATEMALA.QUETZALTENANGO: Volcan Santa Maria, Steyermark 34093 (FH, NY).

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