Claudio Delgadillo M.

Transcription

1 Crossidium POTTIACEAE 611 GYMNOSTOMIELLA CROSSIDIUM 31. CROSSIDIUM Juratzka, Laubm.-Fl. Oesterr.-Ung., , name conserved [Greek krossos, fringe or tassel, and -idion, diminutive, alluding to tassel-like fringe on adaxial surface of costa] Claudio Delgadillo M. Plants in thin turfs or small cushions. Stem 1 10 mm, central strand present. Leaves evenly distributed or crowded, imbricate to slightly twisted when dry, lingulate, lanceolate to deltoid, mm, 1-stratose, concave in distal half, base rectangular to ovate, margins entire, plane to recurved, apex obtuse, rounded or emarginate; costa excurrent as a whitish hair, in section hydroids present between the stereid and the guide cells, stereid band abaxial, filaments restricted to adaxial surface of costa, filament cells cylindric to subspheric, usually thin-walled, terminal cell cylindric to conic to subspheric, generally thin-walled, often papillose; cells of leaf base rectangular, firm-walled; medial and distal leaf cells quadrate to rectangular, firm-walled, sometimes thick-walled, smooth or with 1 to 8 papillae on either surface. Sexual condition dioicous or monoicous; perigonia budlike, leaves short and broad; perichaetia little or not differentiated. Seta 5 20 mm. Capsule cylindric to ovoid-cylindric, erect; annulus of 1 2 rows of vesiculose cells; operculum conic to rostrate, erect or inclined; peristome short and cribrose to long and twisted. Calyptra cucullate. Spores finely papillose, 9 22 µm. Species 11 (4 in the flora): North America, Mexico, South America, s Europe, n, s Africa, Pacific Islands (New Zealand), Australia.

2 612 POTTIACEAE Crossidium Crossidium rosei R. S. Williams, described from Peru, was reported from North America by T. T. McIntosh (1989) from a specimen collected in British Columbia. This is a small sample containing plants of at least three pottiaceous species, including a fertile plant of Pterygoneurum. The type from Peru shows no relationship to the Canadian plants; i.e., Crossidium rosei is not part of the North American flora. Application of the name C. davidai Catcheside by M. J. Cano et al. (1992, 1993) to the Canadian specimen does not account for the lingulate and piliferous leaves of the American material. SELECTED REFERENCES Delgadillo M., C A new species, nomenclatural changes, and generic limits in Aloina, Aloinella, and Crossidium. Bryologist 76: Delgadillo M., C Taxonomic revision of Aloina, Aloinella and Crossidium (Musci). Bryologist 78: Cano, M. J., J. Guerra and R. M. Ros A revision of the moss genus Crossidium (Pottiaceae) with the description of the new genus Microcrossidium. Pl. Syst. Evol. 188: Terminal cell of filaments subspheric, leaf cells papillose. 2. Cells of distal leaf smooth or with 1( 2) papilla on either exposed surface Crossidium aberrans 2. Cells of distal and mid-leaf pluripapillose Crossidium seriatum 1. Terminal cell of filaments cylindric or conic, leaf cells usually smooth. 3. Leaves lanceolate to lingulate-ovate; filament cells thin-walled; cells of distal leaf with firm walls; dioicous or cladautoicous Crossidium crassinervium 3. Leaves broadly deltoid to oblong-ovate; filament cells thick-walled; cells of distal leaf thick-walled, with lumen nearly obliterated; gonioautoicous Crossidium squamiferum 1. Crossidium aberrans Holzinger & E. B. Bartram, Bryologist 27: 4, plate Crossidium spatulifolium Holzinger & E. B. Bartram Plants mm. Leaves lingulate, lanceolate or ovate, mm, margins recurved from near apex to near base, apex rounded or emarginate, piliferous; costa excurrent, with an abaxial epidermis, filaments of 1 4 cells, cells subspheric, with several papillae per cell, terminal cell subspheric; cells of leaf base µm, medial and distal cells µm, smooth, with 1 2 abaxial papillae or on either side in distal cells. Sexual condition dioicous or cladautoicous. Seta 8 16 mm. Capsule urn ovoidcylindric, mm; operculum mm; peristome strongly twisted, µm. Spores spheric, ovoid or oblong, finely papillose, 9 13 µm. Capsule mature Jan Jun( Aug). Soil and rocks under shrubs, shaded banks or in open sites in dry washes; moderate elevations ( m); Ariz., Calif., Colo., Nev., N.Mex., Tex., Utah; Mexico (Baja California, Sonora); n Africa. Crossidium aberrans is easily distinguished by the smooth or 1-papillose cells of the distal leaf blade and by the low filaments with subglobose-papillose terminal cells. 2. Crossidium seriatum H. A. Crum & Steere, SouthW. Naturalist 3: 117, figs Plants mm. Leaves lingulate to oblong-ovate, mm, margins revolute from near apex to near base, smooth-celled in the middle and base, apex round to slightly emarginate; costa excurrent, with an abaxial epidermis, filaments of 2 6 cells, cells subspheric, with several papillae per cell, terminal cell subspheric; cells of leaf base µm, medial and distal cells µm, with several papillae on either side of distal and median cells. Sexual condition cladautoicous. Seta 9 13 mm. Capsule urn oblong-cylindric, mm; operculum mm; peristome strongly twisted, µm. Spores spheric, finely papillose, µm. Capsules mature Jan Jul. Sandy soil or rocks, along dry washes, in open or shaded places in deserts; moderate elevations ( m); B.C.; Ariz., Calif., Nev., N.Mex.; Mexico (Baja California); Europe (France, Spain, Switzerland). The leaf cells with several papillae on either free surface distinguish Crossidium seriatum from C. aberrans. A poorly preserved specimen from Kansas is perhaps a juvenile form with awned leaves and reduced filaments.

3 Crossidium POTTIACEAE Crossidium crassinervium (De Notaris) Juratzka, Laubm.-Fl. Oesterr.-Ung., Tortula crassinervia De Notaris, Mem. Reale Accad. Sci. Torino 40: Varieties 2 (1 in the flora): North America, Europe, n Africa, Asia (China, India), Atlantic Islands (Canary Islands). 3a. Crossidium crassinervium (De Notaris) Juratzka) var. crassinervium Crossidium desertorum Holzinger & E. B. Bartram; C. erosum Holzinger & E. B. Bartram Plants 1 5 mm. Leaves lanceolate, lingulate to lingulate-ovate, mm, margins recurved to revolute from near apex to near base, apex obtuse, rounded or emarginate, piliferous; costa excurrent, with an abaxial epidermis, filaments 2 12 cells, cells cylindric, thin-walled, with 2 4 hollow papillae on apical cell, terminal cell cylindric or conic; cells of leaf base 7 58 µm, medial and distal cells 7 22 µm, smooth. Sexual condition dioicous or cladautoicous. Seta 6 13 mm. Capsule urn ovoid-cylindric, mm; operculum mm; peristome strongly twisted, µm. Spores spheric, nearly smooth to conspicuously papillose, 9 15 µm. Capsule mature Dec Jul. Soil and rocks, on banks and dry washes, under shrubs in desert areas; moderate to high elevations ( m); Ariz., N.Mex., Tex., Utah; Mexico; Europe; Asia (China, India); n Africa; Atlantic Islands (Canary Islands). Crossidium crassinervium is characterized by the smooth-walled leaf cells and long, thin-walled filaments with cylindric to conic terminal cells. Papillae are present on the apex of the terminal cell and, occasionally, on the distal leaf cells. 4. Crossidium squamiferum (Viviani) Juratzka, Laubm.- Fl. Oesterr.-Ung., (as squamigerum) Barbula squamifera Viviani, Ann. Bot. (Genoa) 1(2): ; Tortula squamifera (Viviani) De Notaris Plants 6 10 mm. Leaves deltoidovate, oblong-ovate or lanceolate, mm, margins recurved to erect distally, usually undifferentiated, apex obtuse or rounded, serrulate, with abaxial papillae, piliferous; costa excurrent, with an inconspicuous abaxial epidermis, filaments of 4 10 cells, cells cylindric to subspheric, [thick-walled, terminal cell rectangular, subspheric or conic with 2 5 solid papillae; cells of leaf base µm, medial and distal cells 9 33 µm, smooth, thickwalled. Sexual condition gonioautoicous. Seta 5 20 mm. Capsule urn short-cylindric to ovoid-cylindric, mm; operculum mm; peristome short, nearly straight, cribose at base to long and strongly twisted, µm. Spores spheric, 9 22 µm. Varieties 2 (2 in the flora): North America, nw Mexico, Europe, Asia (India), n Africa, Atlantic Islands (Canary Islands). Crossidium squamiferum is a distinctive species characterized by a whitish marginal area on the distal half of the leaves, thick-walled distal leaf cells, a long hyaline hair point, thick-walled filaments ending in a papillose terminal cell, and the gonioautoicous sexual condition. North American specimens without sporophytes from Arizona, California, Colorado, and Utah have been studied, but these cannot be satisfactorily referred to an infraspecific category. The type specimen of C. succulentum is also sterile. 1. Capsule ovoid-cylindric with a long and twisted peristome; operculum long-rostrate a. Crossidium squamiferum var. squamiferum 1. Capsule cylindric with a short, cribrose peristome; operculum conic b. Crossidium squamiferum var. pottioideum 4a. Crossidium squamiferum (Viviani) Juratzka var. squamiferum Crossidium succulentum Holzinger & E. B. Bartram Leaves with margins recurved to erect distally, undifferentiated in North American plants. Seta 7 20 mm, urn ovoid-cylindric, mm; operculum mm; peristome long, strongly twisted, sometimes cribrose at base, µm. Capsules mature (Aug )Dec Mar. Soil, sandy soil or rocks, dry washes; moderate elevations ( m); Ariz., Calif.; Europe; Asia (India); n Africa; Atlantic Islands (Canary Islands). North American specimens of var. squamiferum lack the differentiated border at the leaf base that is distinctive of the taxon elsewhere. Some specimens may show an indistinct proximal leaf border or a peristome somewhat cribrose at base, indicating intergradation with var. pottioideum. Despite this, var. squamiferum is recognized in North America by long setae, ovoid-cylindric capsules, and a long and twisted peristome.

4 614 POTTIACEAE Crossidium Aloina CROSSIDIUM ALOINA 4b. Crossidium squamiferum var. pottioideum (Notaris) Mönkemeyer, Laubm. Eur., Tortula squamifera var. pottioidea De Notaris, Musci Ital., ; Crossidium griseum (Juratzka) Juratzka Leaves with margins recurved to erect and whitish distally, undifferentiated proximally. Seta 5 8 mm. Capsule urn shortcylindric, mm; operculum mm; peristome short, nearly straight, cribrose, µm. Capsule mature May Aug( Dec). Banks, sandy soil or rocks along dry washes; m; Ariz., Calif., Colo., Nev., Utah; Mexico (Baja California); Europe; Asia; n Africa; Atlantic Islands (Canary Islands). M. J. Cano et al. (1993) did not recognize var. pottioideum as a distinct taxon. However, the character states given in the key offer good reasons to retain it, despite some intergradation with var. squamiferum. 32. ALOINA (Müller Hal.) Kindberg, Bih. Kongl. Svenska Venensk.-Akad. Handl. 6(19): , name conserved [Genus Aloë and Latin -ina, resembling, alluding to fleshy leaves] Barbula sect. Aloina Müller Hal., Syn. 1: Claudio Delgadillo M. Plants in thin turfs or scattered. Stem mm, central strand present. Leaves forming a rosette, lingulate, suborbicular or deltoid, deeply concave, mm, infolded, partly 2- stratose, base auriculate, sheathing, margins entire to irregularly crenulate distally, apex cucullate,

5 Aloina POTTIACEAE 615 mucronate or piliferous; costa broad, percurrent to long-excurrent, in section hydroids absent, stereid band abaxial, occasionally wanting, filaments present on costa and adjacent 2-stratose area of leaf blade, filament cells cylindrical to subspheric, thin-walled, terminal cell subspheric to nearly conic, with an apical thickening; cells of leaf base thin-walled, rectangular, sometimes differentiated marginally, medial and distal leaf cells thick-walled, mostly transversely elongated, smooth. Sexual condition monoicous or dioicous, perigonia distinct, with noncucullate unfolded leaves, perichaetia sometimes partly differentiated, the inner leaves with unfolded blades. Seta mm. Capsule cylindric to ovoid-cylindric, erect or slightly inclined, annulus of 1 2 rows of vesiculose cells, deciduous or persistent, operculum conic to long-rostrate, erect or inclined, peristome generally long and twisted, papillose, with a basal membrane. Calyptra cucullate. Spores spheric, finely papillose, 9 25 µm. Species 9 (5 in the flora): North America, Mexico, Central America, South America, Mediterranean Basin, s Africa, Pacific Islands (New Zealand), Australia. Aloina is sparsely distributed in the New World. It is generally a very distinct genus characterized by differentiated leaf bases, infolded partly 2-stratose laminae, broad costae, and filaments ending in a terminal cell with an apical thickening; the filaments cover the costa and part of the lamina. Plants of Aloina sometimes resemble Indusiella (Grimmiaceae) in habit and general leaf structure although no filaments are found on the leaves of the latter. SELECTED REFERENCES Delgadillo M., C A new species, nomenclatural changes, and generic limits in Aloina, Aloinella, and Crossidium. Bryologist 76: Delgadillo M., C Taxonomic revision of Aloina, Aloinella and Crossidium (Musci). Bryologist 78: Leaf apex piliferous; costa undifferentiated in the distal two thirds, in section with no stereid band, but with a few stereid or substereid cells in mid-costal region Aloina bifrons 1. Leaf apex muticous; costa percurrent to subpercurrent, in section with 1 7 stereid layers. 2. Leaf base without a differentiated margin of thin-walled hyaline cells. 3. Costa with strong solid papillae on the abaxial side at and near the apex; capsule ovoid-cylindric, peristome short (to 900 µm), nearly straight Aloina hamulus 3. Costa smooth abaxially; capsule cylindric, peristome long ( µm) and twisted Aloina aloides 2. Leaf base with a marginal area of narrow thin-walled hyaline cells. 4. Leaves suborbicular to short-lingulate; synoicous (or dioicous); capsule cylindric, operculum conic to short-rostrate Aloina brevirostris 4. Leaves ligulate; dioicous; capsule ovoid-cylindrical, operculum long-rostrate Aloina rigida 1. Aloina bifrons (De Notaris) Delgadillo, Bryologist 76: Tortula bifrons De Notaris, Mem. Reale Accad. Sci. Torino 40: ; Aloina pilifera (De Notaris) H. A. Crum & Steere; A. rigida var. pilifera (De Notaris) Limpricht Plants to 3 mm. Leaves lingulate to ovate-lingulate, mm, margins entire to serrulate, undifferentiated at base, apex piliferous, open, broadly obtuse or rounded; costa undifferentiated, with few or no stereid cells, filaments of 4 8 cells, cells subspheric to cylindric; cells of leaf base µm, medial and distal cells 9 22 µm, papillae none. Sexual condition dioicous. Seta 6 14 mm. Capsule urn cylindric to ovoid cylindric, mm; operculum conical, rostrate, erect or slightly inclined, mm; peristome µm, strongly twisted; spores 9 24 µm. Capsules mature Jan Jul( Aug). Sunny sandy soil or soil over limestone in dry areas; moderate elevations ( m); Ariz., Calif., Idaho; Mexico (Baja California, San Luis Potosí, Nuevo León, Puebla and Zacatecas); s South America; s Europe; sw Asia (Jordan); Africa (Egypt, South Africa); Pacific Islands (New Zealand); Australia. Aloina bifrons is readily identified by the hyaline hair of the leaf apex and the poorly developed costa that is only evident toward the leaf base, but inconspicuous in the distal half of the leaf. No other North American species of the genus exhibits a piliferous leaf apex.

6 616 POTTIACEAE Aloina 2. Aloina hamulus (Müller Hal.) Brotherus in H. G. A. Engler and K. Prantl, Nat. Pflanzenfam. 214[I,3]: Barbula hamulus Müller Hal., Bull. Herb. Boissier 5: Plants to 6 mm. Leaves lingulate, mm, margins entire to irregularly crenulate, undifferentiated at base, apex cucullate; costa subpercurrent to percurrent, filaments of 3 7 cells, cells cylindric to subspheric; cells of leaf base µm, medial and distal cells 9 26 µm, large solid papillae on abaxial surface distally. Sexual condition dioicous. Seta 7 16 mm. Capsule urn ovoid-cylindric, mm; operculum conical to long rostrate, erect or inclined, mm; peristome to 900 µm, twisted to nearly straight. Spores 9 13 µm. Sporophytes unknown in flora area. Sandstone or calcareous rocks, sandy soil in dry sunny sites; moderate to high elevations ( m); Colo., Ill., Kans., La., Nebr., Okla., Tex.; Mexico; Central America (El Salvador, Guatemala). The undifferentiated marginal cells of the leaf base and the large abaxial solid papillae near the leaf tip distinguish Aloina hamulus from A. rigida, which, in addition, has a strongly twisted and longer peristome. The ovoid-cylindric capsule and abaxially papillose leaves distinguish this species from A. aloides var. ambigua. Young leaves should be checked when papillae are not evident in mature leaves. 3. Aloina aloides (Schultz) Kindberg, Bih. Kongl. Svensk. Vetensk.-Akad. Handl. 7: Trichostomum aloides Schultz, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 11: ; Barbula aloides (Schultz) Bruch; Tortula aloides (Schultz) De Notaris Varieties 2 (1 in the flora): sw United States, Mexico, Europe, n Africa, Asia, Atlantic Islands (Canary Islands), Australia. 3a. Aloina aloides var. ambigua (Bruch & Schimper) E. J. Craig in A. J. Grout, Moss Fl. N. Amer. 1: Barbula ambigua Bruch & Schimper, Bryol. Europ. 2: 76, plate Plants to 2.5 mm. Leaves ligulate to lingulate, mm, margins entire, undifferentiated at base, apex cucullate to nearly open; costa subpercurrent to percurrent, filaments of 3 6 cells, cells subspheric; cells of leaf base µm, medial and distal cells µm, occasionally with large solid papillae at leaf tip. Sexual condition dioicous or rhizoautoicous. Seta mm. Capsule urn cylindric, mm; operculum campanulate, mm; peristome µm, strongly twisted. Spores µm. Capsules mature Jan May. Low, desert areas, banks and dry washes, soil and limestone; moderate elevations ( m); Ariz., Calif.; Mexico (Baja California); Europe; Asia (India, Israel, Jordan, Lebanon, Turkey); n Africa; Australia. In var. ambigua, the cylindric capsules with a long and twisted peristome are distinctive. Forms with shortlingulate leaves may be confused with Aloina hamulus when sterile. Outside North America, var. ambigua may occur as high as 2100 m. 4. Aloina brevirostris (Hooker & Greville) Kindberg, Bih. Kongl. Svenska Vetensk.-Akad. Handl. 7(9): Tortula brevirostris Hooker & Greville, Edinburgh J. Sci. 1: 289, plate Plants mm. Leaves lingulate to suborbicular, mm, margins entire, undulate-denticulate distally, differentiated at base, apex cucullate to nearly open; costa subpercurrent, filaments of 2 5 cells, cells subspheric to cylindric; cells of leaf base µm, medial and distal cells µm, papillae none. Sexual condition synoicous or dioicous. Seta 6 17 mm. Capsule urn cylindric, 1 2 mm; operculum conic, short-rostrate, erect or inclined, mm; peristome µm, twisted. Spores µm. Capsules mature (Mar )May Aug. Bare or disturbed soil or silt, roadside banks, calcareous boulders or gravel; low to moderate elevations ( m); Greenland; Alta., B.C., Man., N.W.T., Nunavut, Ont., Yukon; Alaska, Mont.; Europe; Asia (Siberia).

7 Aloina POTTIACEAE 617 ALOINA SYNTRICHIA The leaves of Aloina brevirostris are usually short, and cucullate with a reduced lamina; forms with larger laminae may be confused with A. rigida, but the synoicous condition and the cylindric capsules with a conical operculum are diagnostic. The latter species is dioicous, and has ovoid-cylindrical capsules with a long-rostrate operculum. 5. Aloina rigida (Hedwig) Limpricht, Laubm. Deutschl. 1: Barbula rigida Hedwig, Sp. Musc. Frond., Varieties 3 (1 in the flora): North America, Mexico, South America, Europe, Asia, Africa. 5a. Aloina rigida (Hedwig) Limpricht var. rigida Plants to 2.5 mm. Leaves shortlingulate to ligulate, mm, margins entire to irregularly denticulate, differentiated at base, apex cucullate; costa subpercurrent to percurrent, filaments of 3 9 cells, cells cylindric to spheric; cells of leaf base µm, medial and distal cells 9 40 µm, papillae none. Sexual condition dioicous. Seta mm. Capsule urn ovoid-cylindric, mm; operculum conical to subulate, long-rostrate, mm; peristome µm, strongly twisted. Spores µm. Capsules mature Jun Aug( Oct). Rocks, banks, clay, sandy or gravelly soil in deserts, plains or conifer forests; moderate to high elevations ( m); Alta., B.C., N.B., N.S., Nunavut, Ont., Yukon; Colo., Ill., Iowa, Kans., Mont., Nebr., N.Mex., Tex.; Mexico (Nuevo León, Querétaro, Zacatecas); South America; Europe; Asia; Africa. The ovoid-cylindrical capsule with a long peristome and long-rostrate operculum as well as the differentiated basal leaf margins are the distinguishing features of var. rigida. Despite recent reports from California, this species has not been confirmed from that state.

8 618 POTTIACEAE Syntrichia 33. SYNTRICHIA Bridel, J. Bot. (Schrader) 1801(1): [Greek syn, plus, and trichos, hair, alluding to twisted peristome united by a basal membrane] Brent D. Mishler Plants small to large tufts, green to yellow- or red-brown. Stems simple or forked, epidermal cells differentiated in 1 3 layers, central strand usually present. Leaves appressed and variously twisted when dry, erect-spreading to squarrose when moist, obovate, spatulate, or lingulate, rarely broadly oblong-lanceolate; margins plane or revolute, usually entire; apices acute to truncate or emarginate, and often apiculate; costa generally strong, percurrent to long-excurrent as a hyaline or colored awn, usually rounded abaxailly, in section showing 1 median layer of large guide cells, 1 3 layers of smaller adaxial cells, a weak or strong abaxial stereid band, and sometimes a group of hydroids, but no abaxial epidermis of larger cells; basal cells abruptly differentiated, large and rectangular, smooth, hyaline, thin-walled, often lax; distal cells quadrate or sometimes slightly longer than wide, small and obscure to large and lax, usually pluripapillose on both surfaces, rarely 1-papillose only on the abaxial surface; marginal cells sometimes differentiated, thicker-walled and colored or pale. Specialized asexual reproduction often present, propagula borne on the lamina or costa, in leaf axils, or on rhizoids, in shape leaflike, spherical, or cylindric. Sexual condition dioicous, autoicous, or synoicous; perigonial bracts often short, ovate or oblong-lingulate, blunt, with costa often ending before the apex, perichaetia terminal, the bracts not differentiated. Seta red or brown, elongate, smooth. Capsule erect, straight or slightly curved, red or brown, cylindric, sometimes with a distinct neck, usually striolate when dry; operculum conic, straight or slightly oblique, shorter than the urn; peristome consisting of 32 filiform, reddish, papillose divisions slightly to strongly twisted together above a pale, tessellated basal membrane 1 /6 2 /3 the total length of the peristome. Calyptra long-cucullate, acute, yellow or brown. Spores finely to strongly papillose. KOH laminal color reaction brickred. Species ca. 80 (17 in the flora): North America, Mexico, Central America, South America, Europe, Asia, Africa, Atlantic Islands, Pacific Islands (Hawaii, New Zealand), Australia, Antarctica. Syntrichia appears to be a monophyletic group, segregated from Tortula by synapomorphic leaf characters, in particular the large basal cells abruptly differentiated from the small distal cells, usually forming an inverted U-shaped group on each side of the costa. Also, the costal cross section of Syntrichia lacks an abaxial epidermis, unlike the case in Tortula. The high basal membrane of the peristome, while responsible for the name of the genus, is not diagnostic for Syntrichia since an equally high basal membrane has evolved in Tortula as well. SELECTED REFERENCES Mishler, B. D Tortula. In: A. J. Sharp et al., eds. The moss flora of Mexico. Mem. New York Bot. Gard. 69: Steere, W. C. 1939c. Tortula. In: A. J. Grout Moss Flora of North America, North of Mexico. 3 vols. in 12 parts. Newfane, Vt. and New York. Vol. 1, pp Costa percurrent or short-excurrent into a mucro or apiculus. 2. Laminal cells collenchymatous (sometimes weakly so; often best observed in the region where the basal and distal cells meet). 3. Leaves strongly bordered by thicker-walled, often colored cells; propagula present, cylindric Syntrichia amphidiacea 3. Leaves not bordered; propagula absent Syntrichia cainii 2. Laminal cells with evenly thickened walls, not collenchymatous. 4. Propagula present, leaflike, papillose, borne on stalks in leaf axils 5. Leaves 2-stratose Syntrichia chisosa 5. Leaves 1-stratose Syntrichia ammonsiana 4. Propagula absent or, if present, cylindric, smooth, borne on leaf surface.

9 Syntrichia POTTIACEAE Leaves not fragile; propagula present Syntrichia latifolia 6. Leaves fragile, often broken and appearing eroded; propagula absent Syntrichia fragilis 1. Costa excurrent as an awn. 7. Laminal cells collenchymatous, with single papilla on abaxial surfaces Syntrichia papillosa 7. Laminal cells with evenly thickened walls, not collenchymatous, with multiple papillae on both surfaces. 8. Leaf margins plane or slightly recurved at mid leaf. 9. Plants sometimes bearing green, fusiform, axillary propagula; costa smooth abaxially; awn smooth or with a few short teeth Syntrichia laevipila 9. Plants without propagula; costa strongly papillose abaxially; awn serrulate Syntrichia bartramii 8. Leaf margins strongly revolute at least in proximal part of leaf. 10. Leaf 2-stratose or thicker Syntrichia caninervis 10. Leaf 1-stratose. 11. Costa lacking hydroids but having a row of large-lumined cells abaxial to guide cells; stem lacking a central strand; basal leaf cells µm wide; leaves tapered to the apex from their widest point about one-third the way up from the base, without a constriction near mid leaf. 12. Distal cells with tall, bulging mammillae, µm high, as high as the thickness of the lamina, sometimes higher, also bearing 1 2 papillae per cell Syntrichia papillosissima 12. Distal cells with mammillae much lower than thickness of the lamina. 13. Leaf margins plane in the distal 1 /4; distal cells µm Syntrichia norvegica 13. Leaf margins recurved nearly to the apex; distal cells 8 12 µm Syntrichia ruralis 11. Costa with hydroids; stem with a central strand; basal leaf cells µm wide; leaves constricted near mid leaf. 14. Awns short, to 0.5 mm, smooth or with a few short teeth Syntrichia sinensis 14. Awns long, mm, serrate. 15. Distal laminal cells 8 12 µm Syntrichia montana 15. Distal laminal cells µm. 16. Leaf apex truncate, retuse, or emarginate; costa yellowish to brownish, not serrulate near the apex because of projecting cell ends; distal cells µm; autoicous Syntrichia obtusissima 16. Leaves acute (or sometimes truncate); costa reddish and serrulate near the apex because of projecting cell ends; distal cells µm; synoicous Syntrichia princeps 1. Syntrichia amphidiacea (Müller Hal.) R. H. Zander, Bull. Buffalo Soc. Nat. Sci. 32: Barbula amphidiacea Müller Hal., Linnaea 38: ; Tortula amphidiacea (Müller Hal.) Brotherus; T. caroliniana L. E. Andrews Stems 2 15( 25) mm. Leaves infolded, twisted, and only slightly contorted when dry, erectspreading when moist, variable in shape and size, narrowly or broadly spatulate, (1.5 )2 3( 4) mm; margins revolute in the proximal 1 /2 3 /4, entire; apices acute to acuminate; costa ending a few cells before the apex or percurrent, yellow or brown, smooth; basal cells abruptly differentiated, narrower toward the margins; distal cells rounded, polygonal, or quadrate, µm (or slightly elongate and µm), with 3 6 low or high papillae per cell, thick-walled and collenchymatous; marginal cells in several rows smaller and smoother with thicker, often colored walls. Specialized asexual reproduction as gemmae on the adaxial or occasionally the abaxial leaf surface of leaves, cylindric, rounded at the ends, up to 50 (rarely 200) µm, brown when mature, multicellular. Sexual condition dioicous. [Seta red, 6 8( 10) mm. Capsule red, 2 3 ( 3.5) mm, straight or slightly curved, with a distinct neck; operculum mm; peristome mm, basal membrane 1 /6 1 /5 the total length, teeth irregularly twisted and contorted (crumpled) when dry, widely spreading or

10 620 POTTIACEAE Syntrichia reflexed when moist, red. Spores µm, strongly papillose.] Bark of trees, logs, acidic and basic rocks, occasionally soil; low to high elevations (to 3000 m); N.C., Tenn., Va.; Mexico; Central America; n South America. The peristome of Syntrichia amphidiacea is unusual, being irregularly twisted and crumpled when dry but widely spreading or reflexed when moist. The species has a distribution disjunct between the southern Appalachians and Mexico, similar to that of a number of other mosses (including S. fragilis) and flowering plants that appear to be relicts of an ancient tropical flora. Sporophytes are known only from southern Mexico and southwards. Syntrichia amphidiacea is a rare species in the southeastern United States but common in Mexico. 2. Syntrichia papillosa (Wilson) Juratska, Laubm.-Fl. Oestrr.-Ung., Tortula papillosa Wilson, London J. Bot. 4: Stems 1 4( 8) mm. Leaves incurved and slightly twisted when dry, erect to wide-spreading when moist, spatulate, (1.5 ) mm; margins incurved when dry, plane to erect when moist, entire or occasionally serrulate near the apex; apices acute; costa excurrent into a short, smooth or serrulate, yellowish or hyaline awn 1 /8 1 /5 the leaf length, yellow or red, rounded and sharply papillose-serrate abaxially, smooth on the adaxial surface; basal cells gradually differentiated; distal cells isodiametric, rounded-hexagonal, µm, papillae abaxial, single, simple, rarely forked near the costa, cells rather thick-walled, collenchymatous; cells elongate near leaf apex. Specialized asexual reproduction by gemmae borne on the adaxial surface of the costa, spherical or ovoid, 4 10-celled, brown when mature, smooth. Sexual condition reportedly dioicous. Sporophytes not known from the flora region. Bark of trees or in rock crevices; low to high elevations; N.S., Ont.; Ariz., Calif., Colo., Conn., Ga., Ill., Maine, Mass., Mich., Mo., Nev., N.J., N.Mex., N.Y., N.C., Ohio, Pa., Tenn., Tex., Utah, Wash.; Mexico; South America (Brazil, Colombia, Ecuador); Europe; s Africa; Atlantic Islands (Falkland Islands); Pacific Islands (New Zealand); Australia. The leaves of Syntrichia papillosa have unipapillose cells, with the papillae only on the abaxial surface, strongly papillose-serrate costae, and small, smooth, brown propagula borne on the adaxial surface of the costa. The plants superficially resemble S. laevipila in the field, but the position and nature of the propagula as well as the incurved leaf margins and roughened back of the costa distinguish S. papillosa, even with a hand lens. Sporophytes are known only from Australia, Tasmania, and New Zealand. 3. Syntrichia cainii (H. A. Crum & L. E. Anderson) R. H. Zander, Bull. Buffalo Soc. Nat. Sci. 32: Tortula cainii H. A. Crum & L. E. Anderson, J. Elisha Mitchell Sci. Soc. 74: 35, figs Stems mm. Leaves infolded, twisted, and slightly contorted when dry, erectspreading with recurved tips when moist, variable in shape and size, usually oblong-ovate, mm; margins revolute in the proximal 1 /2 2 /3, entire; apices obtuse but abruptly narrowed to a blunt tip; costa ending well before the apex, brown, smooth; basal cells abruptly differentiated, narrower toward the margins; distal cells rounded, polygonal, or quadrate, µm, with 3 5 papillae per cell, thick-walled and collenchymatous. Specialized asexual reproduction absent. Sexual condition dioicous (perigonia and sporophytes unknown). Limestone; moderate elevations; Ont. Plants identified as Syntrichia cainii from elsewhere in North America (e.g., Kansas, Colorado, and California) are probably incorrectly identified. The difficulty comes from several other species (especially S. norvegica) having scattered neotenic races in which the mature leaves lack awns and thus appear superficially similar to S. cainii. More detailed research, including culture studies, is needed to better understand and distinguish these forms. 4. Syntrichia latifolia (Bruch ex Hartman) Huebener, Muscol. Germ., Tortula latifolia Bruch ex Hartman, Handb. Skand. Fl. ed. 2, Stems 4 12 mm. Leaves infolded, twisted, and slightly contorted when dry, erect-spreading when moist, narrowly or broadly spatulate, mm; margins revolute in the proximal 1 /2, entire; apices obtuse; costa ending a few cells before the apex or percurrent, brown, smooth; basal cells abruptly differentiated, narrower toward the margins; distal cells rounded, polygonal, or quadrate, µm, with 4 6 papillae per cell. Specialized asexual reproduction by gemmae on the adaxial or occasionally the abaxial leaf surface of leaves,

11 Syntrichia POTTIACEAE 621 ovoid and rounded, µm in diameter, green to brown, multicellular. Sexual condition dioicous. Seta red, 6 mm. Capsule red, 3 mm, straight, with a distinct neck; operculum 1.5 2mm, red; peristome 1 mm, basal membrane 1 /2 2 /3 the total length, red. Spores µm, papillose. Bark of trees, rarely on rocks; low to high elevations; B.C.; Ariz., Calif., Oreg., Wash.; Europe. 5. Syntrichia laevipila Bridel, Muscol. Recent., suppl. 4: Syntrichia pagorum (Milde) J. J. Amann; Tortula laevipila var. meridionalis (Schimper) Wijk & Margadant; T. pagorum (Milde) De Notaris Stems 1 3( 5) mm. Leaves infolded and somewhat twisted when dry, erect-spreading when moist, spatulate, mm; margins plane (or somewhat erect), crenulate; apices obtuse, blunt, or emarginate; costa excurrent into a conspicuously tapered, smooth, yellowish or hyaline awn 1 /5 1 /2 the length of the leaf, yellow or red, rounded and smooth on abaxial surface; basal cells rather abruptly differentiated, often with thickened cross-walls, usually extending slightly farther up the margin than near the costa; distal cells quadrate to hexagonal, µm, bulging, with about 4 papillae per cell, the outermost row of cells less papillose. Specialized asexual reproduction as propagula borne at the tip of the stem and in axils of distal leaves, fusiform, green, ecostate, papillose, with an elongate terminal cell, occasionally as several cells. Sexual condition dioicous or autoicous. Seta red, 5 15 mm. Capsule red, 3 5 mm, straight, with a distinct neck; operculum 1 2 mm, red; peristome mm, basal membrane 1 /2 the total length, red. Spores µm, papillose. Occasional on bark of trees, rarely on rock; low to high elevations; B.C.; Ala., Ariz., Ark., Calif., Colo., Conn., Ga., Kans., Ky., La., Md., Mo., Nebr., Nev., N.Mex., N.C., Ohio, Okla., Oreg., S.C., S.Dak., Tenn., Tex., Va., Wash., W.Va.; Mexico; South America (Chile); s Europe; e Asia (Japan); s Africa; Pacific Islands (New Zealand); Australia. This treatment follows M. T. Gallego et al. (2006) in including Syntrichia pagorum with forms of S. laevipila reported in the literature from North America, but this is tentative pending a careful reexamination of this complex of small, corticolous plants. These plants might be confused superfically with S. papillosa, but the leaf margins are plane, the abaxial surface of the costa is smooth and shiny, the propagula are leaf-like, and the leaf cells are pluripapillose in S. laevipila. The other taxa with leaf-like propagula, S. chisosa and S. ammonsiana, are never corticolous. 6. Syntrichia bartramii (Steere) R. H. Zander, Bull. Buffalo Soc. Nat. Sci. 32: Tortula bartramii Steere in A. J. Grout, Moss Fl. N. Amer. 1: 241, plate 115, fig. B Stems 2 10 mm. Leaves infolded and spirally twisted around the stem when dry, wide-spreading when moist, lingulate to spatulate, mm; margins plane, entire, except for papillose crenulations, not bordered; apices acute to truncate or occasionally emarginate; costa excurrent into a conspicuously tapered, serrate awn mm, red or yellow, sparsely to densely spinulose abaxially; basal cells abruptly differentiated, with somewhat thickened cross walls, those at the margins narrower; distal cells irregularly polygonal, isodiametric, 9 13 µm, obscure, bulging and densely papillose, with 4 6 papillae per cell, moderately thick-walled and not collenchymatous. Specialized asexual reproduction absent. Sexual condition dioicous (perigonia and sporophytes unknown). Dry soil and rocks; moderate to high elevations; Ariz., Calif., Colo., Idaho, N.Mex., Tex.; Mexico (Baja California). Diagnostic features of the rare Syntrichia bartramii include plane leaf margins and the costa excurrent into a tapered, serrate awn. In contrast with those of S. fragilis, the leaves of S. bartramii are not fragile and have awns, somewhat smaller, more obscure distal cells, less welldefined groups of basal cells, and plane margins. Without propagula, plants of S. laevipila can be separated from S. bartramii by the abaxially smooth costae and smooth awns, and also by larger and less obscure leaf cells. It should be noted that leaves of S. bartramii sometimes show 2-stratose patches (one or a few cells in width). In contrast, S. chisosa has leaves that are almost completely 2-stratose distally, less twisted when dry, and merely mucronate at the apex.

12 622 POTTIACEAE Syntrichia SYNTRICHIA 7. Syntrichia chisosa (Magill, Delgadillo & L. R. Stark) R. H. Zander, Bull. Buffalo Soc. Nat. Sci. 32: Tortula chisosa Magill, Delgadillo & L. R. Stark, Ann. Missouri Bot. Gard. 70: 200, figs Stems 2 6 mm. Leaves infolded and spirally twisted around the stem when dry, wide-spreading to squarrose-recurved when moist, spatulate, ( 1) mm, distally 2-stratose, occasionally with 1-stratose patches; margins plane, entire; apices acute to truncate; costa excurrent into a short, mm mucro, red, rounded and sharply serrate with hyaline teeth abaxially; basal cells abruptly differentiated, those near the margin smaller, short-rectangular; distal cells hexagonal to irregularly polygonal, or isodiametric, 8 12( 13) µm, densely papillose with 3 5 papillae per cell, bulging, moderately thick-walled, not collenchymatous, obscure. Specialized asexual reproduction by propagula borne on stalks in axils of distal leaves, leaf-like, squarrose-recurved, mm, green, papillose, costate, with an apiculus of 1-several elongate, smooth cells. Sexual condition dioicous (perigonia and sporophytes unknown). Locally common in desert scrub, thin soil in crevices of rocks, soil around bases of shrubs and boulders; moderate to high elevations (about m); N.Mex., Okla., Tex.; Mexico; s Africa. Distinguishing features of Syntrichia chisosa include the 2-stratose laminae, strongly serrate abaxial surface of the costa, plane leaf margins, mucronate apex, and costate, foliose propagula. In the northern part of its range, the species frequently occurs with S. bartramii (see comments thereunder). Syntrichia chisosa also occurs with S. fragilis, and the two may be confused because of a superficially similar habit, color, and dry leaf stance, but S. chisosa can be distinguished with a handlens by non-fragile leaves, a fuzzy appearance of the back of the costa, and usually abundant propagula, and, under a compound microscope, by smaller, 2-stratose, and relatively obscure laminal cells.

13 Syntrichia POTTIACEAE Syntrichia ammonsiana (H. A. Crum & L. E. Anderson) Ochyra, Fragm. Florist. Geobot. 37: Tortula ammonsiana H. A. Crum & L. E. Anderson, Bryologist 82: 469. figs Stems 5 10 mm. Leaves infolded and twisted when dry, recurved when moist, broadly spatulate, mm; margins plane, entire before but with a few teeth near the apex; apices acute; costa percurrent, yellow or brown, smooth; basal cells abruptly differentiated, narrower toward the margins; distal cells quadrate to hexagonal, µm, bulging, with about 4 papillae per cell. Specialized asexual reproduction propagula borne on stalks in axils of distal leaves, leaf-like, mm, green, papillose, costate. Sexual condition dioicous (perigonia and sporophytes unknown). Sandstone in deep shade; moderate elevations; N.C., Tenn., W.Va.; South America (Peru); Africa (South Africa). Syntrichia ammonsiana can easily be distinguished from S. chisosa by its 1-stratose laminae. 9. Syntrichia fragilis (Taylor) Ochyra, Fragm. Florist. Geobot. 37: Tortula fragilis Taylor, London J. Bot. 6: Stems (5 )10 25 mm. Leaves crowded, longitudinally folded and spirally twisted around the stem (but little crisped) when dry, wide-spreading when moist, oblong-lingulate to spatulate, mm, fragile, with sheets of cells breaking off, sometimes along lines of weakness; margins revolute before to as much as 1 /2 the leaf length, to plane, entire or crenulate; apices truncate to acute; costa percurrent or slightly excurrent as a mucro 1 3( 5) cells long, yellow or red, finely to strongly papillose abaxially, abruptly tapered; basal cells abruptly differentiated, becoming rather abruptly shortrectangular near the margins; distal cells quadratehexagonal, 9 15 µm, bulging, with 3 8 papillae per cell, with moderately thick walls, not particularly collenchymatous; distal marginal cells not or weakly to strongly differentiated as a border of brownish, thickerwalled cells in about 2 3 rows. Specialized asexual reproduction absent. Sexual condition dioicous. Seta red, mm. Capsule red, (1.5 )2 4 mm, slightly curved, rather gradually tapered to the seta or with a distinct neck; operculum mm, red; peristome mm, teeth twisted about 1 /2 turn, the distal divisions yellow, the basal membrane pale, about 1 /4 the total length. Spores µm, densely papillose. Bark of trees, rock (usually calcareous), occasionally soil; low to high elevations; Ariz., N.Mex., N.C., Okla., Tenn., Tex., Va., W.Va.; Mexico; Central America; South America; Europe; Asia (China, Iran); Pacific Islands (Hawaii). Syntrichia fragilis shows considerable variability in size and leaf shape (apparently correlated with habitat conditions). In some small, corticolous plants, extreme fragmentation is evident, with an abnormal development of the lamina beyond the usual outline of the leaf. In such plants, several regions of each leaf seem to remain meristematic and produce scalloped or ruffled extensions that eventually break off. Fragility of leaves should not be considered an infallibly diagnostic character, however, since some plants of S. fragilis have quite firm leaves, and the leaves of some other species, particularly when old, may become broken. 10. Syntrichia sinensis (Müller Hal.) Ochyra, Fragm. Florist. Geobot. 37: Barbula sinensis Müller Hal., Nuovo Giorn. Bot. Ital., n. s. 3: ; Tortula sinensis (Müller Hal.) Brotherus Stems 4 15 mm. Leaves longitudinally folded and spirally twisted around the stem but little crisped when dry, wide-spreading when moist, oblong-lingulate to spatulate, mm; margins revolute in the proximal 1 /2, entire; apices acute; costa excurrent into a smooth to slightly toothed, hyaline awn, brown or reddish, smooth; basal cells abruptly differentiated, narrower toward the margins; distal cells polygonal, or quadrate, µm, with 8 10 papillae per cell; marginal cells not differentiated. Specialized asexual reproduction absent. Sexual condition autoicous. Seta red, mm. Capsule red, mm, straight or slightly curved, with a distinct neck; operculum mm, red; peristome mm, red, the basal membrane pale, about 1 /4 the total length. Spores µm, papillose. Vertical limestone faces; high elevations; Colo., N.Mex.; Europe; Asia; n Africa. Only recently discovered in the Rocky Mountains, Syntrichia sinensis may have been overlooked elsewhere in the flora area. For example, it is to be expected in the northern part of the Sierra Nevada of California where limestone is exposed.

14 624 POTTIACEAE Syntrichia 11. Syntrichia montana Nees, Flora 2: Syntrichia intermedia Bridel; Tortula intermedia (Bridel) De Notaris; T. ruralis var. crinita De Notaris Stems 2 10 mm. Leaves infolded and twisted when dry, erectspreading when moist, lingulate or spatulate, mm; margins revolute in the proximal 1 /2, entire; apices broadly acute or obtuse, sometimes emarginate; costa excurrent into a short, serrate, hyaline awn, brown or red, often papillose abaxially and serrulate near the apex because of projecting cell ends; basal cells abruptly differentiated, narrower toward the margins; distal cells polygonal or quadrate, 8 12 µm, with 4 6 papillae per cell. Specialized asexual reproduction absent. Sexual condition dioicous. Seta red, 8 14 mm. Capsule red, mm, straight or slightly curved; operculum mm, red; peristome mm, the basal membrane to 1 /2 the total length, red. Spores µm, papillose. Widespread on soil and rock, occasionally on tree bark; moderate to high elevations; Alta., B.C.; Ariz., Calif., Idaho, Maine, Mont., Nev., N.Mex., N.Y., Oreg., Wash.; Europe; Asia. This treatment follows M. T. Gallego et al. (2006) in employing Syntrichia montana to encompass what has been called S. intermedia or S. ruralis var. crinita in North America. This is, however, a tentative decision as there remains an incompletely understood complex of small, dioicous plants with hydroids in the costa, and some currently undescribed taxa may yet need to be recognized. One recently described species that falls in this complex is S. rupicola B. H. Allen. Further field sampling, molecular characterization, and culture studies are necessary before these taxa are fully understood. 12. Syntrichia obtusissima (Müller Hal.) R. H. Zander, Bull. Buffalo Soc. Nat. Sci. 32: Barbula obtusissima Müller Hal., Syn. Musc. Frond. 1: ; Tortula obtusissima (Müller Hal.) Mitten Stems (5 )10 20 mm. Leaves clasping at base, infolded, and twisted around the stem when dry, wide-spreading to squarrose when moist, lingulate to spatulate, mm, keeled; margins revolute in the proximal 3 /4 7 /8, entire, often laxly undulate; apices emarginate to retuse, occasionally truncate or acute; costa excurrent into a toothed, hyaline awn (often brown at base) usually longer than the leaf, yellow or brown, often minutely papillose abaxially (but not serrate because of projecting cell ends); basal cells abruptly differentiated, rectangular, 70 90( 100) µm, often with yellow walls, narrowly rectangular at the margins; distal cells quadrate to polygonal, µm, thin-walled, pellucid, bulging and bearing 4 7 papillae per cell. Specialized asexual reproduction absent. Sexual condition autoicous, although perigonia often lacking, apparently sometimes dioicous. Seta red-brown, mm. Capsule yellowish or reddish brown, mm, straight or slightly curved, with an abrupt neck; operculum mm, reddish brown; peristome ca. 1.5 mm, the distal divisions twisted ca. 2 turns, yellow or red, the basal membrane white, 1 /3 the total length. Spores 9 16 µm, papillose. Soil, rock, rarely the bases of trees; moderate to high elevations; Ariz., N.Mex., Tex.; Mexico; South America (Bolivia, Peru). Syntrichia obtusissima, with a distinctive Southwest and Mexican-Andean disjunction, has been much confused with S. ruralis, leading to many misidentified specimens in herbaria, but the large, pellucid leaf cells (about µm) contrast with those of S. ruralis (about 8 12 µm), and the back of the costa lacks serrations from projecting cell ends. Furthermore, it has a clear central strand in the stem, and hydroids in the costa, neither of which is found in S. ruralis. 13. Syntrichia princeps (De Notaris) Mitten, J. Proc. Linn. Soc., Bot., suppl. 1: Tortula princeps De Notaris, Mem. Reale Accad. Sci. Torino 40: Stems 5 20 mm. Leaves usually in distinct whorls, infolded, somewhat contorted, and weakly to strongly twisted around the stem when dry, wide-spreading to slightly recurved when moist, concave, spatulate, mm; margins revolute in the proximal 1 /2 3 /4, entire; apices acute or sometimes truncate; costa excurrent into a long, serrate, hyaline awn (reddish at base), often strongly papillose abaxially and serrulate near the apex because of projecting cell ends, red; basal cells abruptly differentiated, long-rectangular, µm, short-rectangular to quadrate at the margins; distal cells quadrate to hexagonal, µm, slightly bulging, bearing 4 6 papillae per cell. Specialized asexual reproduction absent. Sexual condition synoicous