626 · Syntrichia

Sexual condition dioicous. Seta 6–15 mm, brown. Dry soil, rock; moderate to high elevations; B.C.; Ariz., Capsule red, 2.5–3.2 mm, straight or slightly curved, with Calif., Colo., Idaho, Mont., Nev., N.Mex., Oreg., Utah, a distinct neck; operculum 1.5–2mm, red; peristome 0.7– Wash.; Mexico (Coahuila, Nuevo León); s Europe; c Asia. 1 mm, the basal membrane about 1/2 the total length. Syntrichia papillosissima is primarily a species of the Spores 7–8 µm, lightly papillose. Great Basin Desert north into the shrub-steppe ecosystems Soil, deserts and steppe, often forming extensive of the Columbia Basin, where it often occurs as a co- carpets; moderate to high elevations; Alta., B.C.; Ariz., dominant with S. ruralis and S. caninervis. It is similar Calif., Colo., Idaho, Mont., Nev., N.Mex., Oreg., Utah, to a robust S. ruralis, differing most conspicuously in the Wash., Wyo.; s, sw Asia (Afghanistan, Iran, Lebanon, extremely tall mammillae on the distal leaf cells, each Russia, Turkey); n Africa. crowned by only one or two papillae, unlike the shorter Syntrichia caninervis is most common in the colder bulging cell surface bearing four or five papillae deserts and steppes of the flora area, particularly in the characterizing other species in the S. ruralis complex. Mojave and Great Basin deserts and the Columbia Basin. Syntrichia papillosissima also has larger distal laminal It can be confused in the field with S. ruralis, but good cells, which are more pellucid than those of S. ruralis. field distinctions for S. caninervis include the blackish or olive green color, the imbricate, weakly twisted leaf stance 16. Syntrichia norvegica F. Weber, Arch. Syst. when dry, and the back of the costa showing no trace of Naturgesch. 1(1): 130, plate 5, fig. 1. 1804 red and often having a frosty appearance because of the norvegica (F. Weber) stellate papillae. Microscopically, S. caninervis is unique 5 Lindberg with its combination of 2-stratose laminae, non-bulging 5 cell surfaces, and costal cross-sections with sub-stereid Stems 8–25 mm. Leaves clasping at base, infolded and twisted 5 cells. 5

5 5 around the stem when dry, 5 5 5 5

5 5 squarrose-recurved when moist, 15. Syntrichia papillosissima (Coppey) Loeske, 5 5 5 Hedwigia 49: 44. 1910 lingulate-ovate, 2.5–3.5 × 1–1.2 mm, canaliculate to keeled; Barbula papillosissima Coppey, Bull. margins tightly revolute in the Séances Soc. Sci. Nancy, sér. 3, 8: proximal 3/4; apices acute to acuminate; costa excurrent 314, plate 2, figs. f, g. 1908; B. into a serrulate or sometimes serrate, hyaline awn that is ruralis var. hirsuta Venturi; Tortula often red at base or throughout but sometimes broadly 5 papillosissima (Coppey) Brotherus; 5 hyaline at base, papillose abaxially and sometimes 5 5 5 T. ruralis var. hirsuta (Venturi) Paris 5 5 serrulate near the apex because of projecting cell ends, 5 5

5 5 Stems 10–25 mm. Leaves clasping red-brown; basal cells abruptly differentiated, rectangular, at base, infolded and twisted 45–100 × 16–23 µm, quadrate to narrowly rectangular around the stem when dry, at the margins; distal cells quadrate, polygonal, or squarrose-recurved when moist, lingulate-ovate, 2.5–4 rectangular, 13–18 µm, with 3–6 papillae per cell, bulging, × 1–1.6 mm, canaliculate to keeled; margins tightly somewhat collenchymatous. Specialized asexual revolute in the proximal 3/4–7/8, entire; apices obtuse to reproduction absent. Sexual condition dioicous. Seta acute; costa excurrent into a serrate, hyaline awn that is brown, 15–20 mm. Capsule red-brown, 3–4 mm, slightly often brown, sometimes broadly hyaline at base, strongly curved, with an abrupt neck; operculum ca. 1.8 mm, papillose abaxially and serrate because of projecting cell brown; peristome ca. 1.8 mm, the upper divisions twisted ends, yellow-brown; basal cells abruptly differentiated, ca. 2 turns, red, the basal membrane white, 1/3–1/2 the rectangular, 45–90 × 15–23 µm, quadrate to narrowly total length. Spores 11–15 µm, papillose. rectangular at the margins; distal cells quadrate to Soil, rocks; high elevations; Greenland; Alta., B.C., polygonal, 11–18 µm, with tall, bulging mammillae, Ont.; Alaska, Ariz., Calif., Colo., Idaho, Mich., Mont., bearing 1–2 papillae per cell, thick-walled and sometimes Nev., N.Mex., Oreg., Wash.; Mexico; n, c Europe; Asia; collenchymatous. Specialized asexual reproduction Africa (South Africa). absent. Sexual condition dioicous. Seta brown, 12–18 Syntrichia norvegica can be distinguished from S. mm. Capsule brown, 3–5 mm, curved, with an abrupt ruralis by its larger laminal cells, leaf margins less recurved neck; operculum ca. 2 mm; peristome ca. 1.8 mm, the distally, and apices consistently acute to acuminate. The upper divisions twisted ca. 2 turns, yellow-brown, the awn is often partially to completely red, but the amount basal membrane white, ca. 1/2 the total length. Spores of color seems to vary with shade and is not considered 10–14 µm, papillose. Syntrichia· · POTTIACEAE 627 definitive. Another potential recognition feature in the condition dioicous. Seta red, 5–10 mm. Capsule red- field is that the stereid band in the costa often disappears brown, 2–3.5 mm, straight, with an abrupt neck; near the apex, making the normally reddish costa appear operculum 1.25–1.75 mm, brown; peristome ca. 1.25 green just before the awn. Note that there are frequent mm, the upper divisions twisted ca. 2 turns, red, the basal neotenic forms of this species that lack an awn. membrane white, about 1/3 the total length. Spores 11– 15 µm, papillose. 17. Syntrichia ruralis (Hedwig) F. Weber & D. Mohr, Dry to moist soil and rock; low to high elevations; Index Mus. Pl. Crypt., [2]. 1803 Greenland; Alta., B.C., Nfld. and Labr. (Nfld.), N.S., Ont.; Barbula ruralis Hedwig, Sp. Musc. Alaska, Ariz., Calif., Colo., Idaho, Maine, Mass., Mich., Mont., Mo., Nev., N.Y., Okla., Oreg., S.Dak., Tex., Utah, 5 Frond., 121. 1801; Syntrichia ruraliformis (Bescherelle) Dixon; Wash., Wyo.; Mexico; s South America; Eurasia; s Africa. Tortula ruraliformis (Bescherelle) Syntrichia ruralis generally has conspicuously

5 5 squarrose-recurved leaves when wet, with margins 5 W. Ingham; T. ruralis (Hedwig) P. 5 5 5 5 5 5 recurved nearly to the apex, distal portions of the costa 5 5 5 5 5 Gaertner, B. Meyer & Scherbius 5 5 5 5 5 5 toothed abaxially because of projecting cell ends, and 5 5 5 5 Stems 5–15 mm. Leaves clasping relatively small laminal cells. The decurrent, hyaline base at base, infolded and twisted of the awn sometimes used to distinguish S. ruraliformis around the stem when dry, wide- is not reliable and can, on occasion, be found in S. spreading (in smaller forms) to squarrose-recurved when princeps, S. papillosissima, and S. norvegica. Specific moist, lingulate-ovate, 1.5–3.5 × 0.75–1.25 mm, distinctions in the S. ruralis complex are subtle, for the canaliculate to keeled; margins tightly revolute in the most part, requiring cross sections of leaves and stems, 7 proximal /8 or more, entire; apices emarginate to acute; and careful measurements. The leaves of S. princeps and costa excurrent into a serrate (or occasionally only faintly S. obtusissima are narrowed near the middle, whereas serrulate), hyaline awn that is often red or sometimes those of S. papillosissima, S. norvegica, and S. ruralis are broadly hyaline at base, weakly to strongly papillose on widest about one-third the way up from the base and the abaxial surface and often serrate near the apex then taper to the apex. The stem of S. princeps and S. because of projecting cell ends, red-brown; basal cells obtusissima has a strong central strand of thinner-walled abruptly differentiated, narrowly rectangular, 35–70 cells, the costa has a group of hydroids just abaxial to (–90) × 11–18 µm, quadrate to narrowly rectangular at the guide cells, and the basal cells in the leaf are relatively the margins; distal cells quadrate to polygonal, 8–12 µm, wide. In S. papillosissima, S. norvegica, and S. ruralis, with 3–6 papillae per cell, bulging, somewhat obscure. the stem lacks a central strand, the costa lacks hydroids, Specialized asexual reproduction absent. Sexual and the basal cells in the leaf are relatively narrow.

34. MICROBRYUM Schimper, Syn. Musc. Eur., 10. 1860 · [Greek mikros, small, and bryon, ]

Richard H. Zander ˆ Phascum sect. Microbryum (Schimper) Podpera; Phascum subg. Microbryum (Schimper) Limpricht; Phascum sect. Pottiella (Limpricht) Paris; Phascum subg. Pottiella Limpricht; Pottia sect. Pottiella (Limpricht) Nyholm; Pottia subg. Pottiella (Limpricht) Brotherus

Plants in a low turf, scattered or gregarious, occasionally bulbiform, reddish brown distally, brown proximally. Stem very short, 0.2–0.4 mm, hyalodermis absent, sclerodermis absent or weakly differentiated, central strand present or absent; axillary hairs of 3–6 cells, the proximal 1-2 cells usually with thicker walls. Leaves appressed when dry, weakly spreading, tips occasionally reflexed when moist, lanceolate, elliptical or ovate, adaxial surface weakly concave to broadly channeled, short, 0.6–1.8 mm; base not differentiated, proximal margins not differentiated; distal margins recurved at mid leaf and commonly recurved to near apex, entire or rarely serrulate near apex, margins occasionally less papillose and somewhat thicker walled than medially; apex broadly acute; costa excurrent as an apiculus, mucro or short awn, 628 POTTIACEAE · Microbryum occasionally only percurrent, adaxial outgrowths occasionally present as a pad of enlarged parenchymatous cells, adaxial cells quadrate or short-rectangular or elongate, in 2(4–6) rows; transverse section usually round, adaxial epidermis present, adaxial stereid band absent, guide cells 2(–4) in 1 layer, hydroid strand present, occasionally central, abaxial stereid band present, round to semicircular in sectional shape, abaxial epidermis present; proximal cells differentiated across leaf or higher medially, rectangular, 2–4:1, walls of proximal cells usually thin; distal medial cells quadrate to hexagonal or short-rectangular, occasionally rhomboidal, small to somewhat enlarged, 11–15 µm wide, 1–2:1, 1-stratose; papillae usually simple (seldom 2-fid), hollow, 1–6 per lumen, occasionally branching and tall, cell-walls thin to moderately and evenly thickened, convex on both sides. Specialized asexual reproduction absent. Sexual condition monoicous, usually paroicous, occasionally synoicous. Perichaetia terminal, interior leaves often somewhat enlarged, otherwise little differentiated. Seta very short to elongate (to 4 mm). Capsule stegocarpous or cleistocarpous, theca ovate to short elliptical, apiculate when cleistocarpous, 0.5–1.1 mm, annulus absent or of 1–2 rows of weakly vesiculose cells, persistent; operculum when differentiated low-conic, peristome teeth when present 16, irregular, often rudimentary, ligulate to triangular in shape, untwisted. Calyptra conic-cucullate, occasionally mitrate. Spores large, 20–30 µm. KOH laminal color reaction red. Species 13 (4 in the flora): worldwide in temperate zones. Microbryum is similar to Syntrichia in its broad leaves, single stereid band, and red reaction of the lamina to KOH, but differs in the short stems, distinctive reduction series involving the sporophyte, round to semicircular costal stereid band (reniform in Syntrichia), peristomes (when present) of 16 irregular, often rudimentary, truncate peristome teeth that are large in comparison with the size of the capsule, and operculum (when present) low-conic. Adaxial costal pads of cells are occasionally present (in M. vlassovii). Distinguishing features include variation in ornamentation and size of the spores, and collections of these monoicous taxa usually include sporophytes.

1. Distal laminal cells bulging very strongly medially, projecting as “bottle-shaped” cells on the adaxial surface of the costa ...... 4. Microbryum vlassovii 1. Distal laminal cells weakly convex adaxially, not of distinctive shape. 2. Seta very short, capsules nearly spheric, cleistocarpous ...... 3. Microbryum floerkeanum 2. Seta elongate, capsules elliptical to cylindric, stegocarpous (occasionally operculum not dehiscent) or rarely cleistocarpous. 3. Spores smooth or tuberculate (seldom also weakly papillose), 22–30 µm ...... 1. Microbryum starckeanum 3. Spores papillose or spiculose, 28–39 µm ...... 2.

1. Microbryum starckeanum (Hedwig) R. H. Zander, 1. Capsules cleistocarpous (seldom operculum weakly Bull. Buffalo Soc. Nat. Sci. 32: 240. 1993 differentiated but not dehiscent), spores both tuberculate and papillose, occasionally nearly Weissia starckeana Hedwig, Sp. smooth ...... 1c. Microbryum starckeanum Musc. Frond., 65. 1801; Pottia var. fosbergii starckeana (Hedwig) Müller Hal. 1. Capsules stegocarpous, spores smooth or Distal laminal cells weakly convex tuberculate. superficially, adaxial surface of the 2. Peristome present, usually well-developed but truncate apically, leaves stoutly mucronate to

5 costa not mammillose. Seta

5 short-awned ...... 1a. Microbryum starckeanum elongate. Capsule stegocarpous or var. starckeanum occasionally cleistocarpous, 2. Peristome absent or rudimentary, leaves very cylindrical. Spores smooth or shortly mucronate . . . 1b. Microbryum starckeanum tuberculate (seldom also weakly papillose), 22–30 µm. var. brachyodus Varieties 6 (3 in the flora): North Temperate Zone, disjunct to austral areas. Microbryum · POTTIACEAE 629

1a. Microbryum starckeanum (Hedwig) R. H. Zander The capsule mouth is commonly oblique and constricted var. starckeanum when dry, and the costa is only shortly excurrent as a Pottia arizonica Wareham mucro (M. starckeanum var. starckeanum has short awns). Stem leaves mucronate to short- awned. Capsule stegocarpous, 1c. Microbryum starckeanum var. fosbergii (E. B. peristome present, usually well- Bartram) R. H. Zander, Bull. Buffalo Soc. Nat. Sci. 32: developed but truncate apically. 240. 1993

5 Spores smooth or tuberculate. 5 Capsules mature winter (Jan– Pottia fosbergii E. B. Bartram, Feb). Bare soil, fields; low Bryologist 33: 18. 1930; Phascum elevations; Ariz., Calif.; Mexico fosbergii (E. B. Bartram) J. Guerra (Baja California); Europe; n Africa; Pacific Islands (New Stem leaves mucronate to short- Zealand). awned. Capsule cleistocarpous,

A collection from California (Pasadena, “g. 9”, US) 5 peristome rarely differentiated has three variants, including var. starckeanum, var. internally, operculum absent or fosbergii, and what is probably best called var. occasionally weakly differentiated. starckeanum but which has an indehiscent operculum, Spores both tuberculate and even upon heating in KOH and Pohlstoffe. Another, from papillose, occasionally nearly smooth. Arizona (Pima County, E. B. Bartram 990, Musci Acroc. Capsules mature in early spring (Mar). Soil; low to Bor.-Amer. 572, US) includes var. starckeanum, var. moderate elevations; Calif.; Mexico (Baja California). brachyodus, and the indehiscent variant of var. Variety fosbergii has elliptical capsules that lack both starckeanum. Collections with capsules toward the large a differentiated operculum and a peristome (except end of the size scale, previously called Pottia arizonica, California: Ikenberry, CANM, which has a weakly often have entirely smooth spores; commonly, however, differentiated operculum and rudimentary peristome some tuberculate and characteristic of M. starckeanum teeth). This variety apparently has spores that are both can be identified in at least some capsules. Also, D. F. tuberculate and papillose (also true in the type), Chamberlain (1978) described M. starckeanum (as Pottia occasionally nearly smooth. If the spores are unreduced starckeana) as having smooth or tuberculate spores in (though there is no evidence they are), then hybridization his treatment for the British Isles. Capsule contents not would be implicated. R. M. Ros et al. (1994) have differentiating into spores is not uncommon in variants described from Spain a putative hybrid between of M. starckeanum, indicating the possibility of occasional Microbryum starckeanum and Tortula protobryoides (as hybridization. Pottia bryoides), which is evidently much the same as the var. fosbergii. 1b. Microbryum starckeanum var. brachyodus (Bruch & Schimper) R. H. Zander, Bull. Buffalo Soc. 2. Microbryum davallianum (Smith) R. H. Zander, Bull. Nat. Sci. 32: 240. 1993 Buffalo Soc. Nat. Sci. 32: 240. 1993 Anacalypta starckeana var. Gymnostomum davallianum Smith, brachyodus Bruch & Schimper, Ann. Bot. (König & Sims) 1: 577. Bryol. Europ. 2: 47. 1843; Pottia 1805; Pottia davalliana (Smith) arizonica var. mucronulata C. Jensen Wareham; P. starckeana var. 5 Distal laminal cells weakly convex

brachyodus (Bruch & Schimper) 5 5 5 5 5 superficially, adaxial surface of 5 Müller Hal. 5 5 costa not mammillose. Seta Stem leaves very shortly mucro- elongate. Capsule cylindrical, nate. Capsule stegocarpous, stegocarpous but never cleisto- peristome absent or rudimentary. Spores smooth or carpous. Spores granulose, low-papillose or spiculose, tuberculate. large, 28–39 µm. Capsules mature late winter or early spring (Jan–Mar). Varieties 3 (3 in the flora): North America, Mexico, Soil; low to moderate elevations (0–700 m); Ariz., Calif.; Europe, sw Asia, Africa, Pacific Islands (New Zealand). sw Asia; n Africa. Variety brachyodus has a rudimentary peristome (as an irregular proximal membrane) or the peristome absent. 630 POTTIACEAE · Microbryum

1. Capsules commonly wide-mouthed when dehisced, Variety commutatum is known only from California with a single row of thickened cells below mouth, (Berkeley, Howe 79, US), last collected in 1894, from peristome absent, spores spiculose, 31–39 µm which station it is disjunct to the Old World...... 2a. Microbryum davallianum var. davallianum 2c. Microbryum davallianum var. conicum 1. Capsules narrow mouthed when dehisced, with (Schwägrichen) R. H. Zander, Bull. Buffalo Soc. Nat. usually 2 or more rows of thickened cells below mouth, peristome variously absent or present, Sci. 32: 240. 1993 spores merely papillose, 27–34 µm. Gymnostomum conicum 2. Peristome well developed ...... Schwägrichen, Sp. Musc. Frond...... 2b. Microbryum davallianum Suppl. 1(1): 26, plate 9. 1811; var. commutatum Pottia starckeana subsp. conica 2. Peristome rudimentary or absent ...... (Schwägrichen) D. P. Chamberlain; ...... 2c. Microbryum davallianum P. texana Wareham 5 var. conicum 5

5

5 Capsule narrow-mouthed when 2a. Microbryum davallianum (Smith) R. H. Zander var. dehisced, with usually 2 or more davallianum rows of thickened cells below mouth; peristome rudimentary or absent. Spores Pottia starckeana subsp. minutulum papillose, 27–34 µm. (Schwägrichen) D. P. Chamberlain Capsules mature late winter and spring (Feb–Apr). Capsule commonly wide-mouthed Soil; low elevations; Calif., Nebr., Okla., Tex.; Mexico when dehisced, with a single row (Baja California); Europe; sw Asia; w, s Africa; Pacific 5 of thickened cells below mouth; Islands (New Zealand). 5 5 peristome absent. Spores spiculose The San Marcos, Texas specimen (MICH) of var. (finely spinose) and large (31–39 conicum has a flaring mouth like that of var. davallianum. µm.). Another Texas specimen (Orcutt 5565, CANM, WTU) Capsules mature late summer has a weak peristome (a thin line of proximal membrane). to early winter (Sept–Nov). Bare soil, vacant lots, fields; low to moderate elevations; Ont.; Mich., N.Y.; Europe; 3. Microbryum floerkeanum (Weber & D. Mohr) sw Asia; n Africa; Pacific Islands (New Zealand). Schimper, Syn. Musc. Eur., 11. 1860 In var. davallianum the spines of the spores are 2–2.5 µm. Distal laminal papillae are hollow or occasionally Phascum floerkeanum Weber & D. solid, ca. 2 per lumen, usually simple but in some Mohr, Bot. Taschenbuch, 70, fig. specimens (e.g., Ontario, Bruce County, C. Williams 451. 1807 1200, CANM) mostly 2-fid and hollow (with a distinct Distal laminal cells weakly convex saddle between the projections) and distinctly C-shaped 5 superficially, adaxial surface of in optical section. costa not mammillose. Seta extremely short, nearly absent. 2b. Microbryum davallianum var. commutatum Capsule nearly spherical, (Limpricht) R. H. Zander, Bull. Buffalo Soc. Nat. Sci. cleistocarpous. Spores papillose, 32: 240. 1993 20–25 µm. Capsules mature in late fall (Oct–Nov). Soil, fields, Pottia commutata Limpricht, pastures; low elevations; Ont.; Europe; sw Asia; n Africa. Laubm. Deutschl. 1: 537, fig. 160.

The leaves of Microbryum floerkeanum are ovate to 1888; P. davalliana subsp. ˆ ovate-lanceolate and sheath the immersed capsule. The commutata (Limpricht) Podpera papillae occur near the apex of the leaf, one per lumen, Capsule narrow-mouthed when and are large, nearly covering the lumen. The calyptra is

5 dehisced, with usually 2 or more variably papillose, merely rough, or smooth. Variety rows of thickened cells below badium (Bruch & Schimper) Schimper is poorly mouth; peristome well developed distinguishable, but may be segregated by its long- though apically truncate. Spores acuminate leaf apices—the typical variety has short- papillose, 27–34 µm. acuminate apices. Capsules mature late winter (Feb). Soil; low elevations; Calif.; Europe; sw Asia; n Africa. Microbryum · Hilpertia · POTTIACEAE 631

MICROBRYUM ° HILPERTIA

4. Microbryum vlassovii (Lazarenko) R. H. Zander, Substrate and elevational range unknown; B.C.; Calif.; Bull. Buffalo Soc. Nat. Sci. 32: 240. 1993 Asia. Phascum vlassovii Lazarenko, Zhurn Sporophytes were immature in the area of the flora. Inst. Bot. Vseukraïns’k. Akad. Nauk Microbryum vlassovii was reported from British 26/27: 196. 1938 Columbia by T. T. McIntosh (1989) and Spain by M. N. Jiménez et al. (1990), otherwise known from Armenia, Distal laminal cells strongly 5 central Asia and the Ukraine. The long, smooth cell bulging superficially in medial making up the end of the mucro is reminiscent of that of 5 portion of leaf, adaxial surface of Chenia. The rhizoids bear distinctive unicellular costa strongly mammillose, beset hemispherical excrescences that may serve as tubers. with “bottle-shaped” cells. Seta Tortula may have ampullose adaxial costal cells short, nearly absent. Capsule (T. L. Blockeel 1995), but is yellow in KOH. nearly spherical apiculate, cleistocarpous. Spores papillose, ca. 18–24 µm.

35. HILPERTIA R. H. Zander, Phytologia 65: 427. 1989 · [For Friedrich Hilpert, b. 1907, German bryologist]

Richard H. Zander

Plants in loose cushions, greenish brown distally, light brown proximally. Stems to 1 cm, hyalodermis absent, sclerodermis not or weakly differentiated, central strand distinct; axillary hairs to 8 cells long, all hyaline or the proximal one yellow-brown. Stem leaves crowded, larger 632 POTTIACEAE · Hilpertia distally, appressed and tightly spiraled when dry, weakly spreading when moist, 1–1.4 (plus 0.3–0.6 of awn) mm; ovate to circular, adaxial surface flat to quite concave, base not differentiated; distal margins strongly revolute (to 2 ×), entire or broadly dentate above, cells of revolute margin enlarged, strongly chlorophyllose; apex broadly acute, hyaline in an apical triangle; costa narrow but broader above, excurrent as a hyaline awn, lamina inserted laterally, adaxial outgrowths none, superficial adaxial cells long-rectangular, in 2–4 rows; transverse section rounded, adaxial epidermis present, adaxial stereid band absent, guide cells 2 in 1 layer, hydroid strand present, abaxial stereid band present, rounded in cross-sectional shape, abaxial epidermis absent; basal cells weakly differentiated, rectangular, thin-walled; distal medial cells hexagonal to short-rectangular or rhomboid, rather large, (14–)20–25 µm wide, 1–4:1; papillae absent medially, hollow-papillose on revolute margins, cell walls thin to thickened and porose, superficial abaxial cell walls thick, weakly convex superficially on both sides, cells of leaf apex rhomboid to fusiform, smooth. Specialized asexual reproduction by brood bodies borne on proximal rhizoids, brown, spheric to elliptic. Sexual condition polyoicous: synoicous, paroicous, autoicous or perigonia terminal on separate plants. Perichaetia terminal, interior leaves sheathing, usually differentiated, long-oval, margins usually somewhat differentiated, medial laminal cells rectangular, very thin-walled. Seta 0.35–0.4 cm. Capsule stegocarpous, theca elliptic, occasionally weakly ventricose, 1.2–1.5 mm, annulus of 3 rows of smaller, quadrate, highly vesiculose cells; operculum broadly short-conic to long-conic, peristome teeth 32, long, linear, 1 twisted counterclockwise about /2 turn. Calyptra cucullate. Spores 13–16 µm. KOH laminal color reaction red. Species 1: Canada, Europe, Asia. Hilpertia is unique among the Pottioideae in the elaboration of a photosynthetic organ consisting of laminal marginal cells rolled in a spiral tube, thin-walled and hollow-papillose. The species is characteristic of cold steppe habitats.

1. Hilpertia velenovskyi (Schiffner) R. H. Zander, Capsules mature summer. Calcareous silt and shrub Phylologia 65: 429. 1989 steppe; B.C., N.W.T., Nunavut; c Europe; Asia (China). Tortula velenovskyi Schiffner, Nova In the flora area, Hilpertia velenovskyi is a rare but Acta Acad. Caes. Leop.-Carol. distinctive species known only from Northwest German. Nat. Cur. 58: 480, plate Territories, base of a hill on the north bank of the South Nahanni River, 60°33’30”N, 125°23’W, from British 5 17. 1893; Hilpertia scotteri (R. H. 5 Zander & Steere) R. H. Zander; Columbia, E side of Fraser R. at Big Bar, 51°15’N, 122°W, 5 Tortula scotteri R. H. Zander & T. McIntosh 7688, May 13, 1984 (UBC), and from Steere Nunavut, Ellesmere Island, Tanquary Fiord marine forland, 1.5 mi. SW of base camp, 81°23’N, 76°59’W, Stems branching irregularly; G. S. Mogensen 90-134, July 19, 1990 (C). The spiraled rhizoids rare; leaves broadly leaf margin of slightly enlarged, highly photosynthetic toothed at or near the base of the cells is similar to that of Pseudocrossidium replicatum, awn; basal cells ca. 16–18 µm wide, 2–3:1. Specialized but the broad leaf shape and costal anatomy place H. asexual reproduction by brood bodies of (1–)3–4 cells, velenovskii in the Pottioideae. I agree with the appraisal mostly 30–50 µm. Perichaetial leaves to 1.7 mm. Seta of B. C. Tan and J. Zhao (1997) that H. scotteri is a 0.35–0.40 cm, twisted counterclockwise distally, synonym. As with those in Stegonia latifolia, the cells of clockwise proximally. Theca operculum 0.4–1 mm, cells the leaf apex are rhomboid to fusiform and epapillose. twisted 1/2 times counterclockwise; peristome teeth 300– 700 µm, densely branching-spiculose. Calyptra ca. 2.8 mm, smooth. Spores indistinctly papillose, light brown. Chenia · POTTIACEAE 633 36. CHENIA R. H. Zander, Phytologia 65: 424. 1989 · [For Chen Pan Chieh, 1907– 1970, Chinese bryologist]

Richard H. Zander

Phascum sect. Leptophascum Müller Hal., Flora 71: 7. 1888; Phascum subg. Leptophascum (Müller Hal.) G. Roth

Plants forming a low turf, green distally, brownish proximally. Stems short to elongate, to 3 mm; hyalodermis absent, sclerodermis present, central strand weak to strong; axillary hairs small, of 3–4 cells, proximal cell firm-walled. Leaves appressed and contorted when dry, spreading when moist; ligulate to elliptic, adaxial surface occasionally grooved along costa, plane or broadly channeled across leaf. 1.5–2.5 mm; base rectangular or not differentiated in shape; margins distally plane, weakly recurved or plane basally, irregularly denticulate distally with sharp, mid cell projections ending in a simple papilla, marginal cells often smaller; apex rounded to broadly acute, occasionally sharply apiculate; costa weak, ending several (6–9) cells before the apex or percurrent, lamina inserted adaxially to laterally, adaxial cells short-rectangular, narrower than laminal cells, in 2 rows, abaxial cells short-rectangular; transverse section rounded to elliptic, adaxial epidermis present, adaxial stereid band absent, guide cells 2 in 1 layers, hydroid strand present, small to large, abaxial stereid band very weak or occasionally absent, rounded in sectional shape, abaxial epidermis present; proximal cells differentiated across the leaf base except for one or more rows of marginal cells, rectangular; distal medial cells large, (15–)20–25(–30) µm wide, 1:1, bulging-hexagonal, 1-stratose; papillae absent (distal marginal teeth may be interpreted as sharp papillae), cell walls thin, weakly trigonous, convex on both sides. Specialized asexual reproduction by brood bodies that are sometimes present on rhizoids in soil. Sexual condition dioicous. [Seta ca. 1.2 cm. Capsule stegocarpous or cleistocarpous; theca ovate; annulus of 2–3 layers of strongly vesiculose cells, persistent; peristome teeth when present 16, filamentous, somewhat anastomosing, straight or weakly twisted counterclockwise. Calyptra cucullate. Spores ca. 10 µm.] KOH laminal color reaction red. Species 3 (1 in the flora): North America, Mexico, South America, Europe, e Asia, Australia. Chenia includes both cleistocarpous and peristomate taxa. The diagnostic characters are leaves ligulate to spathulate, sharply crenulate to irregularly dentate above with sharp mid-marginal wall projections, often sharply apiculate by a distinctive thick-walled cell or cells, distal laminal cells large, weakly trigonous, epapillose, and red in KOH solution.

SELECTED REFERENCES Neumann, A. J. 1972. Observations on the morphology and biology of the moss Tortula vectensis E. Warb. & Crundw. in Louisiana. Bryologist 75: 580–583. Reese, W. D. 1967. The discovery of Tortula vectensis in North America. Bryologist 70: 112–114. 634 POTTIACEAE · Chenia

CHENIA ° HENNEDIELLA

1. Chenia leptophylla (Müller Hal.) R. H. Zander, Bull. rhizoids in soil, clavate, ca. 100–130 µm. Sporophytes Buffalo Soc. Nat. Sci. 32: 258. 1993 not known from North America. Phascum leptophyllum Müller Hal., Found on bare soil in disturbed, open areas; low to Flora 71: 6. 1888; Chenia high elevations (0–200 m); Ala., La., N.Mex.; Mexico; rhizophylla (Sakurai) R. H. Zander; South America (Bolivia, Brazil); Europe; s Africa; Tortula rhizophylla (Sakurai) Z. Australia; Pacific Islands (Hawaii). Iwatsuki & K. Saito; T. vectensis Chenia leptophylla is one of a mundivagant group that E. F. Warburg & Crundwell is apparently distributed in association with human activities (P. M. Eckel 1986). The small brood bodies 5 5 5 Stems seldom branching; rounded on rhizoids in the soil seem to be the agency for asexual in transverse section; rhizoids reproduction. The very thin-walled distal laminal cells usually few. Leaves ligulate to may appear bright yellow-green in KOH, but elliptic, apex acute or rounded acute and apiculate with examination under the compound microscope reveals an elongate, brownish, thick-walled cell, costa percurrent brick red walls. The elongate, thick-walled, reflexed cell or subpercurrent; proximal laminal cells ca. 20–25 µm forming the leaf apiculus is unique and immediately wide, 2–4:1. Brood bodies sometimes present, borne on diagnostic. Hennediella · POTTIACEAE 635 37. HENNEDIELLA Paris, Actes Soc. Linn. Bordeaux, sér. 5, 9: 232. 1895 · [For Roger Hennedy, 1809–1877, Scottish phycologist]

Richard H. Zander

Hennedia R. Brown bis, Trans. & Proc. New Zealand Inst. 25: 285. 1893, not Hennedya Harvey 1855

Plants growing in loose turf or cushions, green distally, reddish brown proximally. Stems ca. 0.5–1 cm; hyalodermis absent or weakly developed, not collapsed, sclerodermis absent or substereid, central strand present; axillary hairs of ca. 5 cells, proximal 1–2 cells occasionally somewhat thick-walled. Stem leaves appressed or incurved when dry, spreading when moist; ovate to long-lanceolate, occasionally ligulate or spathulate, adaxial surface plane or broadly channeled across leaf, (0.5–)3–4(–7) mm; base not differentiated in shape or ovate and somewhat sheathing, proximal margins commonly bordered; distal margins plane, dentate distally or less commonly entire, bordered (sometimes intramarginally) by short-rectangular to elongate cells, usually less papillose, occasionally thick-walled, rarely 2–3-stratose; apex acute or rarely obtuse; costa percurrent or short-excurrent as flat, denticulate mucro, rarely ending 3–4 cells before apex, adaxial cells quadrate to short-rectangular, in 4–5 rows; transverse section round to elliptic, adaxial epidermis present, adaxial stereid band absent, guide cells 2–4 per layer in (1-)2 layers, hydroid strand present, usually large, abaxial stereid band present, of substereid cells, elliptic or round in section, abaxial epidermis present (occasionally only lateral) or absent; proximal cells differentiated across leaf, rectangular, 18–30 µm wide, (2–)4–6:1, walls of proximal cells thin; distal medial cells relatively large, quadrate to hexagonal or short-rectangular, ca. 18–24 µm wide, 1–2:1, often marginally elongate and multistratose; papillae hollow, simple to 2-fid, usually 6–8 or more per lumen, cell walls thin to evenly thickened, flat to weakly convex. Specialized asexual reproduction absent. Sexual condition autoicous, cladautoicous or dioicous. Perichaetia terminal, leaves somewhat larger than the cauline leaves. Seta 0.05–2.5 cm. Capsule stegocarpous, theca ovate to cylindric, occasionally microstomous, 0.8–3.5 mm, annulus of 2– 4 rows of vesiculose cells; operculum conic to rostrate, often narrowly so, occasionally systilius; peristome teeth absent, rudimentary or of 32 long, filamentous teeth, twisted counterclockwise, teeth when well developed to 1.5 mm. Calyptra cucullate, occasionally flaring proximally and not split. Spores 8–30 µm. KOH distal laminal color reaction red. Species 20 (2 in the flora): nearly worldwide, most diverse in austral temperate areas. Hennediella was recently recognized as much larger than previously thought (T. L. Blockeel 1991; R. H. Zander 1989, 1993). The bordered, dentate, plane leaves with flat laminal cells, red in KOH, are distinctive. The sporophyte varies among the species from well developed to much reduced. The operculum is often long, (0.6–)1.2–1.8 mm. The genus is found in soil and on rock, commonly in wet areas.

SELECTED REFERENCE Blockeel, T. L. 1990. The genus Hennediella Par.: A note on the affinities of Tortula brevis Whitehouse & Newton and T. stanfordensis W. C. Steere. J. Bryol. 16: 187–192.

1. Leaves broadly lanceolate to ovate, tubulose and stiff, distal margins 1-stratose, monoicous, commonly fruiting ...... 1. Hennediella heimii 1. Leaves broadly elliptic to obovate, flat and lax, distal margins 2-stratose, dioicous, sporophytes unknown in area of the flora ...... 2. Hennediella stanfordensis 636 POTTIACEAE · Hennediella

1. Hennediella heimii (Hedwig) R. H. Zander, Bull. Yukon; Alaska, Calif., Colo., Idaho, Mont., Nebr., Buffalo Soc. Nat. Sci. 32: 248. 1993 N.Mex., Oreg., Utah, Wash., Wyo.; s South America; Gymnostomum heimii Hedwig, Sp. Europe; Asia; Pacific Islands (New Zealand); Australia; Antarctica. 5 Musc. Frond., 32. 1801; 5 Desmatodon heimii (Hedwig) 5 5 5 Mitten; Pottia heimii (Hedwig) 1b. Hennediella heimii var. arctica (Lindberg) R. H.

5 5 5 Zander, Bull. Buffalo Soc. Nat. Sci. 32: 248. 1993 5 5 5 5 Fürnrohr 5 5 5 5 5 5 5 Pottia heimii var. arctica Lindberg, 5 5 5 Leaves ovate to broadly 5 5 lanceolate, tubulose and stiff; 5 Öfvers. Kongl. Vetensk.-Akad. Förh. 5 23: 551. 1867; Desmatodon heimii

distal margins with a 1-stratose 5 5 border of elongate cells, distal var. arcticus (Lindberg) H. A. Crum; Gymnostomum obtusifolium R. Br. lamina cells 10–16 µm. Sexual condition monoicous, 5 either synoicous or autoicous. Sporophytes often present. bis; P. heimii var. obtusifolia (R. Br. Seta 5–15 mm. Capsule stegocarpous, systilius; theca bis) I. Hagen wide-mouthed, 1–2.4(–3.5) mm; peristome absent. Leaves broadly ovate to broadly Varieties 15 (2 in the flora): North America, s South elliptic, concave and often America, Europe, Asia, Pacific Islands (New Zealand), cucullate, apex usually obtuse, margins usually entire; Australia, Antarctica. costa often ending before the apex. In Hennediella heimii, European infraspecific taxa Capsules mature late summer (Aug). Exposed soil, attributed to the flora area are not distinct in the New silt; low elevations; Greenland; N.W.T., Nunavut, Ont.; World (see R. T. Wareham 1939). S. Flowers (1973) may Alaska; Europe; n Asia. have included H. stanfordensis in his concept of this Variety arctica is a somewhat poorly circumscribed species, judging from the description, illustrations and taxon largely restricted to the Arctic. It is generally much his attribution of dioicy. However, like H. stanfordensis, smaller than var. heimii, with leaves commonly 0.5–0.8 this species may have gemmae borne on proximal mm. rhizoids. This taxon exhibits polymorphisms (discussed by Wareham) with considerable variation in 2. Hennediella stanfordensis (Steere) Blockeel, J. Bryol. differentiation of the leaf border of elongate cells and 16: 191. 1990 marginal dentition, the distal cell ornamentation, and in Tortula stanfordensis Steere, length of capsule. It may be confused with Tortula Bryologist 54: 119, figs. 1–9. 1951; obtusifolia but that species has no trace of marginal Hyophila stanfordensis (Steere) dentition and is yellow in KOH. A. J. E. Smith & H. Whitehouse 1. Leaves ovate to broadly lanceolate, tubulose but Leaves broadly elliptic to obovate, not concave or cucullate, apex acute, margins 5 flat and lax; distal margins with a dentate distally, costa percurrent ...... 2-stratose border of elongate cells, ...... 1a. Hennediella heimii var. heimii distal laminal cells 10–14 µm. 1. Leaves broadly ovate to broadly elliptic, concave and often cucullate, apex usually obtuse, margins Sexual condition dioicous. usually entire, costa often ending before the apex Sporophytes unknown in region of flora...... 1b. Hennediella heimii var. arctica Soil, among grasses, drainage ditches, fields; low to moderate elevations (100–300 m); Calif.; Mexico 1a. Hennediella heimii (Hedwig) R. H. Zander var. (Guerrero); Europe; Australia. heimii Since the description of Hennediella stanfordensis, this mundivagant species has engendered a flurry of Pottia heimioides Kindberg publications on its human-oriented distribution, its 5 Leaves ovate to broadly lanceo- discovery in Europe, and the taxonomic significance of 5

5 late, tubulose but not concave or 5 the eperistomate sporophyte (e.g., D. G. Long 1979; cucullate, apex acute, margins 5 5 A. J. E. Smith and H. L. K. Whitehouse 1974; Whitehouse 5 5 5 5 5 5 5 5 dentate; costa percurrent. 1975). 5 5 5 5 5 5 Capsules mature Jun–Jul. 5 5 5 Moist, often alkaline soil, banks, frost boils, lake shores, friable shale, near streams and seacoasts; low to high elevations (0–2900 m); Greenland; Alta., B.C., Man., N.B., Nfld. and Labr., Nunavut, Ont., Que., Sask., Acaulon · POTTIACEAE 637

HENNEDIELLA ° ACAULON

38. ACAULON Müller Hal., Bot. Zeitung (Berlin) 5: 99. 1847 · [Greek a-, without, and kaulos, stalk or stem, alluding to stemless habit]

Richard H. Zander

Plants bulbiform, gregarious or scattered, reddish to yellowish brown distally, brown proximally. Stems short, to 0.5 mm; hyalodermis absent, sclerodermis absent, central strand absent; axillary hairs to ca. 5 cells in length, proximal cell walls sometimes thickened. Stem leaves strongly appressed and apices usually reflexed when dry, appressed to weakly spreading when moist; ovate, adaxial surface deeply concave, 0.5–1.75 mm; base not differentiated in shape, proximal margins not differentiated; distal margins plane or very weakly recurved, entire to serrulate or dentate; apex broadly acute, abruptly apiculate; costa excurrent in a stout, sharp apiculus, occasionally as a short, occasionally dentate awn, adaxial outgrowths absent in American species, adaxial cells elongate, in 3–4 rows; transverse section round, adaxial epidermis present, adaxial stereid band absent, guide cells 0–4 in 1 layer, hydroid strand usually present, occasionally central, abaxial stereid band present, usually weak, rounded in sectional shape, abaxial epidermis present; proximal cells not differentiated in shape, rectangular, little wider than distal cells, 3–4:1, walls of proximal cells thin; distal medial cells rounded-quadrate to rhomboid, ca. 13–15 µm wide, 1–4:1, 1-stratose; papillae absent or occasionally large and simple, one per lumen, cell walls evenly thickened, occasionally highly thickened on abaxial walls, convex on both sides of lamina. Specialized asexual reproduction absent. Sexual condition dioicous and perigoniate plants smaller, or monoicous, usually paroicous. Perichaetia terminal, interior leaves somewhat enlarged. Seta very short, to 0.2 mm. Capsule cleistocarpous, spheric, apiculus lacking, ca. 638 POTTIACEAE · Acaulon 0.4–0.7 mm, annulus absent. Calyptra mitrate, often lobed. Spores ca. 25–35(–50) µm. KOH laminal color reaction red. Species 15 (3 in the flora): nearly worldwide, mainly in temperate regions. Acaulon is similar to species of Microbryum in the bulbiform habit, leaves red in KOH, but its species are even smaller in size, and the capsules are spheric, lacking an apiculus. Papillae are absent in two of the three species. Although spore size and ornamentation have figured in in the past, according to H. A. Crum and L. E. Anderson (1981) there is much overlap; but, similarly, their use of differences in margin recurvature and seta length do not hold. Taxa belonging to the old, superficially similar Phascum have been recently placed largely with Tortula and Microbryum (R. H. Zander 1993). I. G. Stone (e.g., 1989) has studied the genus extensively in Australia, and Casas de Puig (e.g., C. Casas de Puig et al. 1990) and her students have done the same for the Iberian Peninsula.

SELECTED REFERENCES Bryan, V. S. 1956b. Cytological and taxonomic studies of some species of Astomum, Acaulon and Phascum. Bryologist 59: 118–129. Grout, A. J. 1922–1940. Acaulon. 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. 194–195.

1. Leaves awned, laminal cells papillose abaxially ...... 2. Acaulon schimperianum 1. Leaves cuspidate or muticus, laminal cells smooth. 2. Plants often three-angled, about 1 mm, leaves keeled, seta about as long as the diameter of the capsule, spores about 30 µm, finely papillose ...... 1. Acaulon triquetrum 2. Plants flattened-globose or three-angled, about 2 mm, leaves broadly channeled, seta short, about 0.3 the diameter of the capsule, spores 30–50 µm, smooth or papillose ...... 3.

1. Acaulon triquetrum (Spruce) Müller Hal., Bot. 2. Acaulon schimperianum (Sullivant) Sullivant in W. S. Zeitung (Berlin) 5: 100. 1847 Sullivant and C. L. Lesquereux, Musc. Bor.-Amer., sched. Phascum triquetrum Spruce, London 8. 1857 J. Bot. 4: 189. 1845 Phascum schimperianum Sullivant in Plants three-angled, about 1 mm. A. Gray, Manual ed. 2, 615. 1856 Stem leaves cuspidate, keeled; Plants elliptic to globose,

5 5 laminal cells, 13–18 µm wide, 1– occasionally three-angled, 1–1.5

5 5 5 3:1, smooth. Seta as long as the mm. Stem leaves awned, broadly 5 5 5 5

5 5 diameter of the capsule. Spores 5 channeled; laminal cells papillose 5

5

25–30(–40) µm, finely papillose. 5 abaxially. Seta as long as the Capsules mature late autumn– diameter of the capsule. Spores spring. Soil, sand, clay, old fields, pastures, roadside spheric, 35-40 µm, papillose. banks, temporarily moist areas; low to moderate Capsules mature winter. Soil, dry washes; low to elevations; Ont., Sask.; Calif., Ill., Iowa, Mass., N.J., Tex., moderate elevations; Ariz., Ill., Iowa, Kans., Tex.; Mexico Va., W.Va.; Europe; Asia; n Africa; Australia. (Nuevo León). Acaulon muticum var. rufescens has reflexed apices Acaulon schimperianum is awned and the distal while A. triquetrum does not always have these, though margins are more strongly dentate than those of the other illustrated as such by H. A. Crum and L. E. Anderson species.

(1981). The exsiccat C. F. Austin, Musci Appalachiani SELECTED REFERENCE Crum, H. A. and L. E. Anderson. 1965. The 52 has a short seta and the is 1.2 mm, and is placed taxonomy and distribution of Acaulon schimperianum. Bryologist 68: here with A. muticum var. rufescens. 208–211. Acaulon· Crumia · POTTIACEAE 639

3. Acaulon muticum (Hedwig) Müller Hal., Bot. Zeitung somewhat elliptic spores. A collection from Texas (Berlin) 5: 99. 1847 (Bastrop County, Bastrop, F. McAllister, Feb. 1934, A. J. Phascum muticum Hedwig, Sp. Grout, North American Musci Perfecti 258, UBC) has Musc. Frond., 23. 1801 spheric, heavily papillose spores, 40–45 µm. Specimens with partially smooth spores but a few large granules Plants elliptic to globose, ca. 2 adherent or scattered through the spore sac (e.g., Oregon, mm. Stem leaves cuspidate, 5 Lane County, Eugene, Alton Baker Park, D. Wagner 5 5 5 broadly channeled; laminal cells 5 1834, Mar. 9, 1978, UBC) are here placed tentatively 5 5 5 5 5 5 5 5 5 5 smooth. Seta short, about 0.3 the 5 5 5 with the typical variety. A specimen from Iowa 5 5 5 5 5 5 5 diameter of the capsule. Spores 5 (Poweshiek County, Conard & Peck v.11.35, MO) has spheric to elliptic, 30–50 µm, leaves blotched red in KOH and, variably among papillose or nearly smooth. capsules, weakly papillose to distinctly crowded-spiculose Varieties 2 (2 in the flora): temperate areas of the spores. Northern Hemisphere, Africa. As H. A. Crum (1969) and A. J. Grout (1939) pointed 3b. Acaulon muticum var. rufescens (A. Jaeger) H. A. out, American specimens reported as Acaulon rubrum Crum, Bryologist 72: 240. 1969 are A. muticum var. rufescens. These plants are often small, rather three-sided and may be confused with A. Acaulon rufescens A. Jaeger, Ber. triquetrum. The leaves of both varieties stain red, at least Thätigk. St. Gallischen Naturwiss. in blotches, in KOH, and they intergrade to some extent. Ges. 1868–1869: 77. 1869 Plants yellow- to red-brown in 5 1. Plants yellow-brown in nature, spores papillose- 5 5 5 nature. Spores variably low punctulose ...... 3a. Acaulon muticum var. muticum 5 5 5 5 5 5 5 5 5 papillose-punctulose or smooth. 1. Plants yellow- to red-brown in nature, spores 5 5 5 5 5 5 5 Capsules mature late fall to smooth or papillose-punctulate ...... 5 ...... 3b. Acaulon muticum var. rufescens early spring. Soil, gravel pit, pasture, lawn; low to moderate 3a. Acaulon muticum (Hedwig) Müller Hal. var. elevations; B.C., Ont., Que., Sask.; Ariz., Calif., Fla., Ill., muticum La., Md., Mich., Miss., Mo., Nebr., N.J., N.Y., Okla., Plants yellow-brown in nature. Oreg., S.C., Tex., Wis.; Mexico. Spores densely low spiculose- A specimen of var. rufescens from New Jersey (Austin papillose. 51, UBC) is reddish and has small (ca. 20 µm), round, Capsules mature late fall to weakly papillose spores; generally, spore size is not a good spring. Soil, among grasses, character. Another specimen, with large (38–40 µm), round, punctulate spores, is piebald with adherent 5 5 pastures; low to moderate 5 5 5 granules similar to the papillae of var. muticum. One 5 5 5 elevations; Calif., Iowa, Kans., 5 Mass., N.Y., N.C., Okla., Tenn., specimen placed here (California, Santa Clara County, Tex.; Europe; Asia; Africa. Stanford University, W. Schofield 5724, Feb. 23, 1955, It may be suspected that young plants of both var. UBC), is rather immature, has a greenish cast and smooth, muticum and var. rufescens sometimes have smooth, elliptic spores about 28-30 µm.

39. CRUMIA W. B. Schofield, Canad. J. Bot. 44: 609, figs. 1–15. 1966 · [For H. A. Crum, 1922–2002, American bryologist]

Richard H. Zander

Plants forming cushions, green to reddish distally, blackish green to iridescent tan proximally. Stems to 3 mm, branching often; hyalodermis absent, sclerodermis absent, central strand absent or very weak; axillary hairs ca. 6 cells in length, proximal 1–2 cells yellowish. Leaves appressed, weakly contorted when dry, spreading when moist; spathulate, adaxial surface flat, shallowly channeled along costa, to 4 mm; base scarcely differentiated in shape, proximal margins bordered by a few rows of long-rectangular cells; margins recurved along 1 or both margins in the proximal 1 /2, entire, bordered by ca. 6 rows of enlarged, rounded rhomboid to rectangular cells, except 640 POTTIACEAE · Crumia the marginal row; apex broadly acute to rounded acute, usually broadly apiculate; costa percurrent or ending 1–2 cells below the apex, adaxial outgrowths absent, adaxial cells elongate in 5–6 rows, transverse section semicircular, adaxial epidermis present, adaxial stereid band absent, guide cells 3–4 in 1 layer, hydroid strand absent, abaxial stereid band present, strong, semicircular in section, abaxial epidermis weakly developed; basal cells differentiated medially, inflated-rectangular, slightly wider than the distal cells, 3–5:1, walls of basal cells thin, brown; distal medial cells hexagonal to shortly rectangular, 13–20 µm wide, 1–2:1, 1-stratose; papillae small, simple to 2-fid, 8–12 per lumen, scattered, cell walls thin, weakly convex on both sides. Specialized asexual reproduction absent. Sexual condition dioicous. Perichaetia terminal, interior leaves not sheathing, little different from cauline leaves. Seta ca. 1.4 mm. Capsule stegocarpous, theca cylindric, 2.5–3 mm, annulus of ca. 2 rows vesiculose cells; operculum conic, 0.8–1.1 mm; peristome teeth 16, cleft to near base into two branches, linear, rami much perforated and anastomosing, twisted very weakly counterclockwise, teeth ca. 200 µm. Calyptra cucullate. Spores ca. 15–18 µm. Laminal KOH color reaction reddish orange. Species 1: w North America. Crumia is a robust moss similar to Scopelophila in the often black or deep brown coloration of the plants, the lack of either a sclerodermis or a hyalodermis in the stem, the spathulate leaf shape and single costal stereid band, but differs in the presence of a stem central strand (though small or occasionally absent), a more strongly differentiated, intramarginal leaf border, and presence of a peristome (though rarely fruiting). Hennediella has bordered leaves but is red in KOH, not reddish orange, and has plane leaf margins with marginal cells not larger than the medial as seen in section. A second species, Crumia deciduidentata, was referred to Tortula by R. H. Zander (1993) on the basis of differences in the peristome, the awned perichaetial leaves, strong stem central strand, and presence of a hydroid strand in the costa.

1. Crumia latifolia (Kindberg) W. B. Schofield, Canad. J. Sporophytes mature spring–summer (May, Jun, Jul). Bot. 44: 610. 1966 Limestone, calcareous shale, siliceous and conglomeritic Merceya latifolia Kindberg, Bull. rock, roadbank, wet areas; low to moderate elevations Torrey Bot. Club 16: 94. 1889; (5–1200 m); B.C.; Ariz., Calif., Mont., Oreg., Utah, Wash. Scopelophila latifolia (Kindberg) Often locally abundant, Crumia latifolia is sometimes Renauld & Cardot reddish orange in exposed situations. The morphology 5 and ecology of this species are discussed at length by S.

5 Stems sparsely radiculose; 5 5 Flowers (1973), W. B. Schofield (1966), and A. J. Grout

5 5 rounded-pentagonal in section. (1928–1940, vol. 1). There is great variation in the 5 Leaf with costal abaxial cells thickness and color of the laminal cell walls, possibly elongate. Sporophytes 1(–2) per correlated with degree of exposure. It has been found in perichaetium. Seta reddish brown fruit in British Columbia and two stations in California. in color, ca. 1.4 mm, twisted clockwise below, counterclockwise above. Capsule with reddish brown theca, peristome teeth of 7 articulations held together by a hyaline membrane. Calyptra ca. 3 mm. Luisierella · POTTIACEAE 641

CRUMIA ° LUISIERELLA

40. LUISIERELLA Thériot & P. de la Varde, Bull. Soc. Bot. France 83: 73, fig. 1. 1936 · [For Alphonse Luisier, 1872–1957, French bryologist] Patricia M. Eckel

Plants scattered or gregarious, dull blackish green. Stems very short, 1.5–2 mm, hyalodermis absent, outer cortex undifferentiated, central strand absent; axillary hairs usually 5 cells in length, proximal cell yellowish. Leaves tubulose and contorted when dry, spreading and recurved from the insertion when moist, ligulate-lanceolate or oblong or long-elliptic, base weakly differentiated to elliptic, margins plane to erect-incurved, entire in proximal region, rounded-crenulate by bulging cells distally, lamina 1-stratose, apex rounded or rounded-obtuse to obtusely acute to occasionally cucullate; costa subpercurrent by 4–5 cells, adaxial outgrowths absent, covered ventrally with a layer of green, rounded-quadrate to short-rectangular bulging cells continuous with those of the lamina but appearing elevated above them, in 4–5 rows; transverse section elliptic to circular, adaxial epidermis strongly differentiated, adaxial stereid band absent, guide cells 2–4(–5) in one row, hydroid strand absent, abaxial stereid band present, 1 strong, semi-elliptic to semi-circular in shape; basal cells sharply differentiated in the lower /6 of the leaf, abruptly inflated but not bulging, smooth, rising higher along the margins and forming a V-shaped area, oblong-rectangular 2–3(–4):1, thick-walled juxtacostally, becoming very thin and lax at the leaf margins; distal laminal cells rounded-quadrate to short-rectangular, 1:1(–2), 1-stratose, essentially smooth, adaxial surface mammillose-bulging, abaxial surface plane, somewhat mammillose near the margins, the walls thickened, firm. Specialized asexual reproduction lacking. Sexual condition gynodioicous, fertile plants synoicous, plants with only 642 POTTIACEAE · Luisierella archegonia not producing sporophytes; perichaetia terminal, interior perichaetial leaves much smaller than the cauline leaves. Seta 0.4–0.5(–0.7) cm, rarely to 2 cm. Capsule stegocarpous, theca narrowly cylindric; annulus compound, large and strongly vesiculose in two rows, persistent; operculum narrowly high-conic; peristome teeth sixteen, linear or narrowly lanceolate, variable in development: absent, rudimentary or to 500 µm, erect. Calyptra cucullate. Spores 8–9 µm. KOH laminal color reaction negative, light yellow to orange. Species 1: se United States, Mexico, West Indies, Central America, South America, Asia. Luisierella is characterized by plants blackish, minute, with ligulate leaves tubulose when dry, laminal cells adaxially mammillose, abaxially plane, adaxial costal surface with bulging rounded cells, and adaxial stereid band absent. The abruptly thin-walled hyaline basal cells extending up the leaf margin in a V shape is distinctive; it is this latter character that quickly distinguishes this genus from similar genera: Gyroweisia, Hyophila, and Plaubelia. Luisierella bears little relation to the genus Tortella, which is also characterized by differentiated basal cells that extend higher on the leaf margin than along the costa. Luisierella has been suggested to be ancestrally related to the genus Quaesticula R. H. Zander (R. H. Zander 1993) through reduction of gametophyte characters and elaboration of reproductive ones.

SELECTED REFERENCES Crum, H. A. and L. E. Anderson. 1961. Luisierella barbula in Georgia. Bryologist 64: 315–320. Deguchi, H. 1987. Luisierella (Pottiaceae, Musci), a moss genus with a disjunctive distribution in Neotropics and Japan. J. Jap. Bot. 62: 7–15. Steere, W. C. 1945. Luisierella, a genus of new to North America. Bryologist 48: 83–85.

1. Luisierella barbula (Schwägrichen) Steere, Bryologist Capsules mature late fall through winter (occasionally 48: 84. 1945 to March). Calcareous rocks, moist to mesic sites, Gymnostomum barbula sinkholes and other limestone substrate (a gravestone); Schwägrichen, Sp. Musc. Frond. low elevations; Ala., Fla., Ga., Tex.; Mexico; West Indies; Suppl. 2(2,1): 77, plate 175. 1826 Central America (Belize); South America (Brazil); Asia (Indonesia, Japan). Plants inconspicuous, nearly stem- Luisierella barbula usually grows in a thin crust less. Stems mostly unbranched, associated with cyanobacteria, Hyophiladelphus agrarius, rhizoids or tomentum not evident. and Weissia jamaicensis, the minute black plants

5 5 Leaves rosulate, 1–1.5(–2) mm, 5 5 contrasting starkly with the usually chalky white rock younger leaves nested in a rosette surface. The somewhat elevated cells on the adaxial of older leaves, which extends to surface of the costa may resemble a differentiated 1 /2 the length of the younger leaves; distal laminal cells structure (a pad of cells), but the cells appear to be deep green in sharp contrast with colorless proximal cells otherwise undifferentiated from those of the lamina. region, irregularly rounded-hexagonal, 8–11 µm. Sexual condition with bulbous vaginula; antheridia tiny, 1/3 the length of the archegonia; plants with only perigonia not seen. Capsule 1–2.5 mm; operculum 0.6–1 mm; peristome teeth short, irregular, sometimes appearing absent.