626 ·POTTIACEAESexual condition dioicous. Seta 6–15 mm, brown.Capsule red, 2.5–3.2 mm, straight or slightly curved, witha distinct neck; operculum 1.5–2mm, red; peristome 0.7–1 mm, the basal membrane about 1/2 the total length.Spores 7–8 µm, lightly papillose.

Soil, deserts and steppe, often forming extensivecarpets; moderate to high elevations; Alta., B.C.; Ariz.,Calif., Colo., Idaho, Mont., Nev., N.Mex., Oreg., Utah,Wash., Wyo.; s, sw Asia (Afghanistan, Iran, Lebanon,Russia, Turkey); n Africa.

Syntrichia caninervis is most common in the colderdeserts and steppes of the flora area, particularly in theMojave and Great Basin deserts and the Columbia Basin.It can be confused in the field with S. ruralis, but goodfield distinctions for S. caninervis include the blackish orolive green color, the imbricate, weakly twisted leaf stancewhen dry, and the back of the costa showing no trace ofred and often having a frosty appearance because of thestellate papillae. Microscopically, S. caninervis is uniquewith its combination of 2-stratose laminae, non-bulgingcell surfaces, and costal cross-sections with sub-stereidcells.

15. Syntrichia papillosissima (Coppey) Loeske,Hedwigia 49: 44. 1910

Barbula papillosissima Coppey, Bull.Séances Soc. Sci. Nancy, sér. 3, 8:314, plate 2, figs. f, g. 1908; B.ruralis var. hirsuta Venturi; Tortulapapillosissima (Coppey) Brotherus;T. ruralis var. hirsuta (Venturi) Paris

Stems 10–25 mm. Leaves claspingat base, infolded and twistedaround the stem when dry,

squarrose-recurved when moist, lingulate-ovate, 2.5–4× 1–1.6 mm, canaliculate to keeled; margins tightlyrevolute in the proximal 3/4–7/8, entire; apices obtuse toacute; costa excurrent into a serrate, hyaline awn that isoften brown, sometimes broadly hyaline at base, stronglypapillose abaxially and serrate because of projecting cellends, yellow-brown; basal cells abruptly differentiated,rectangular, 45–90 × 15–23 µm, quadrate to narrowlyrectangular at the margins; distal cells quadrate topolygonal, 11–18 µm, with tall, bulging mammillae,bearing 1–2 papillae per cell, thick-walled and sometimescollenchymatous. Specialized asexual reproductionabsent. Sexual condition dioicous. Seta brown, 12–18mm. Capsule brown, 3–5 mm, curved, with an abruptneck; operculum ca. 2 mm; peristome ca. 1.8 mm, theupper divisions twisted ca. 2 turns, yellow-brown, thebasal membrane white, ca. 1/2 the total length. Spores10–14 µm, papillose.

Dry soil, rock; moderate to high elevations; B.C.; Ariz.,Calif., Colo., Idaho, Mont., Nev., N.Mex., Oreg., Utah,Wash.; Mexico (Coahuila, Nuevo León); s Europe; c Asia.

Syntrichia papillosissima is primarily a species of theGreat Basin Desert north into the shrub-steppe ecosystemsof the Columbia Basin, where it often occurs as a co-dominant with S. ruralis and S. caninervis. It is similarto a robust S. ruralis, differing most conspicuously in theextremely tall mammillae on the distal leaf cells, eachcrowned by only one or two papillae, unlike the shorterbulging cell surface bearing four or five papillaecharacterizing other species in the S. ruralis complex.Syntrichia papillosissima also has larger distal laminalcells, which are more pellucid than those of S. ruralis.

16. Syntrichia norvegica F. Weber, Arch. Syst.Naturgesch. 1(1): 130, plate 5, fig. 1. 1804

Tortula norvegica (F. Weber)Lindberg

Stems 8–25 mm. Leaves claspingat base, infolded and twistedaround the stem when dry,squarrose-recurved when moist,lingulate-ovate, 2.5–3.5 × 1–1.2mm, canaliculate to keeled;margins tightly revolute in the

proximal 3/4; apices acute to acuminate; costa excurrentinto a serrulate or sometimes serrate, hyaline awn that isoften red at base or throughout but sometimes broadlyhyaline at base, papillose abaxially and sometimesserrulate near the apex because of projecting cell ends,red-brown; basal cells abruptly differentiated, rectangular,45–100 × 16–23 µm, quadrate to narrowly rectangularat the margins; distal cells quadrate, polygonal, orrectangular, 13–18 µm, with 3–6 papillae per cell, bulging,somewhat collenchymatous. Specialized asexualreproduction absent. Sexual condition dioicous. Setabrown, 15–20 mm. Capsule red-brown, 3–4 mm, slightlycurved, with an abrupt neck; operculum ca. 1.8 mm,brown; peristome ca. 1.8 mm, the upper divisions twistedca. 2 turns, red, the basal membrane white, 1/3–1/2 thetotal length. Spores 11–15 µm, papillose.

Soil, rocks; high elevations; Greenland; Alta., B.C.,Ont.; Alaska, Ariz., Calif., Colo., Idaho, Mich., Mont.,Nev., N.Mex., Oreg., Wash.; Mexico; n, c Europe; Asia;Africa (South Africa).

Syntrichia norvegica can be distinguished from S.ruralis by its larger laminal cells, leaf margins less recurveddistally, and apices consistently acute to acuminate. Theawn is often partially to completely red, but the amountof color seems to vary with shade and is not considered

Syntrichia

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rzanderText BoxFlora of North America, Volume 27, 2007

POTTIACEAE 627·definitive. Another potential recognition feature in thefield is that the stereid band in the costa often disappearsnear the apex, making the normally reddish costa appeargreen just before the awn. Note that there are frequentneotenic forms of this species that lack an awn.

17. Syntrichia ruralis (Hedwig) F. Weber & D. Mohr,Index Mus. Pl. Crypt., [2]. 1803

Barbula ruralis Hedwig, Sp. Musc.Frond., 121. 1801; Syntrichiaruraliformis (Bescherelle) Dixon;Tortula ruraliformis (Bescherelle)W. Ingham; T. ruralis (Hedwig) P.Gaertner, B. Meyer & Scherbius

Stems 5–15 mm. Leaves claspingat base, infolded and twistedaround the stem when dry, wide-

spreading (in smaller forms) to squarrose-recurved whenmoist, lingulate-ovate, 1.5–3.5 × 0.75–1.25 mm,canaliculate to keeled; margins tightly revolute in theproximal 7/8 or more, entire; apices emarginate to acute;costa excurrent into a serrate (or occasionally only faintlyserrulate), hyaline awn that is often red or sometimesbroadly hyaline at base, weakly to strongly papillose onthe abaxial surface and often serrate near the apexbecause of projecting cell ends, red-brown; basal cellsabruptly differentiated, narrowly rectangular, 35–70(–90) × 11–18 µm, quadrate to narrowly rectangular atthe margins; distal cells quadrate to polygonal, 8–12 µm,with 3–6 papillae per cell, bulging, somewhat obscure.Specialized asexual reproduction absent. Sexual

condition dioicous. Seta red, 5–10 mm. Capsule red-brown, 2–3.5 mm, straight, with an abrupt neck;operculum 1.25–1.75 mm, brown; peristome ca. 1.25mm, the upper divisions twisted ca. 2 turns, red, the basalmembrane white, about 1/3 the total length. Spores 11–15 µm, papillose.

Dry to moist soil and rock; low to high elevations;Greenland; Alta., B.C., Nfld. and Labr. (Nfld.), N.S., Ont.;Alaska, Ariz., Calif., Colo., Idaho, Maine, Mass., Mich.,Mont., Mo., Nev., N.Y., Okla., Oreg., S.Dak., Tex., Utah,Wash., Wyo.; Mexico; s South America; Eurasia; s Africa.

Syntrichia ruralis generally has conspicuouslysquarrose-recurved leaves when wet, with marginsrecurved nearly to the apex, distal portions of the costatoothed abaxially because of projecting cell ends, andrelatively small laminal cells. The decurrent, hyaline baseof the awn sometimes used to distinguish S. ruraliformisis not reliable and can, on occasion, be found in S.princeps, S. papillosissima, and S. norvegica. Specificdistinctions in the S. ruralis complex are subtle, for themost part, requiring cross sections of leaves and stems,and careful measurements. The leaves of S. princeps andS. obtusissima are narrowed near the middle, whereasthose of S. papillosissima, S. norvegica, and S. ruralis arewidest about one-third the way up from the base andthen taper to the apex. The stem of S. princeps and S.obtusissima has a strong central strand of thinner-walledcells, the costa has a group of hydroids just abaxial tothe guide cells, and the basal cells in the leaf are relativelywide. In S. papillosissima, S. norvegica, and S. ruralis,the stem lacks a central strand, the costa lacks hydroids,and the basal cells in the leaf are relatively narrow.

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

Richard H. Zander

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 orweakly differentiated, central strand present or absent; axillary hairs of 3–6 cells, the proximal1-2 cells usually with thicker walls. Leaves appressed when dry, weakly spreading, tipsoccasionally reflexed when moist, lanceolate, elliptical or ovate, adaxial surface weakly concaveto broadly channeled, short, 0.6–1.8 mm; base not differentiated, proximal margins notdifferentiated; distal margins recurved at mid leaf and commonly recurved to near apex, entireor rarely serrulate near apex, margins occasionally less papillose and somewhat thicker walledthan medially; apex broadly acute; costa excurrent as an apiculus, mucro or short awn,

Syntrichia Microbryum·

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628 ·POTTIACEAEoccasionally only percurrent, adaxial outgrowths occasionally present as a pad of enlargedparenchymatous 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, guidecells 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 differentiatedacross leaf or higher medially, rectangular, 2–4:1, walls of proximal cells usually thin; distalmedial cells quadrate to hexagonal or short-rectangular, occasionally rhomboidal, small tosomewhat 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 evenlythickened, convex on both sides. Specialized asexual reproduction absent. Sexual conditionmonoicous, usually paroicous, occasionally synoicous. Perichaetia terminal, interior leavesoften somewhat enlarged, otherwise little differentiated. Seta very short to elongate (to 4 mm).Capsule stegocarpous or cleistocarpous, theca ovate to short elliptical, apiculate whencleistocarpous, 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, oftenrudimentary, ligulate to triangular in shape, untwisted. Calyptra conic-cucullate, occasionallymitrate. 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 thesporophyte, round to semicircular costal stereid band (reniform in Syntrichia), peristomes (whenpresent) of 16 irregular, often rudimentary, truncate peristome teeth that are large in comparisonwith the size of the capsule, and operculum (when present) low-conic. Adaxial costal pads ofcells are occasionally present (in M. vlassovii). Distinguishing features include variation inornamentation and size of the spores, and collections of these monoicous taxa usually includesporophytes.

1. Distal laminal cells bulging very strongly medially, projecting as “bottle-shaped” cells onthe 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 floerkeanum2. 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 starckeanum3. Spores papillose or spiculose, 28–39 µm . . . . . . . . . . . . . . . . . . . . . . . 2. Microbryum davallianum

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1. Microbryum starckeanum (Hedwig) R. H. Zander,Bull. Buffalo Soc. Nat. Sci. 32: 240. 1993

Weissia starckeana Hedwig, Sp.Musc. Frond., 65. 1801; Pottiastarckeana (Hedwig) Müller Hal.

Distal laminal cells weakly convexsuperficially, adaxial surface of thecosta not mammillose. Setaelongate. Capsule stegocarpous oroccasionally cleistocarpous,cylindrical. Spores smooth or

tuberculate (seldom also weakly papillose), 22–30 µm.Varieties 6 (3 in the flora): North Temperate Zone,

disjunct to austral areas.

1. Capsules cleistocarpous (seldom operculum weaklydifferentiated but not dehiscent), spores bothtuberculate and papillose, occasionally nearlysmooth . . . . . . . . . . . . . . 1c. Microbryum starckeanum

var. fosbergii1. Capsules stegocarpous, spores smooth or

tuberculate.2. Peristome present, usually well-developed but

truncate apically, leaves stoutly mucronate toshort-awned . . . . . . . 1a. Microbryum starckeanum

var. starckeanum2. Peristome absent or rudimentary, leaves very

shortly mucronate . . . 1b. Microbryum starckeanumvar. brachyodus

Microbryum

POTTIACEAE 629·1a. Microbryum starckeanum (Hedwig) R. H. Zander

var. starckeanum

Pottia arizonica Wareham

Stem leaves mucronate to short-awned. Capsule stegocarpous,peristome present, usually well-developed but truncate apically.Spores smooth or tuberculate.

Capsules mature winter (Jan–Feb). Bare soil, fields; lowelevations; Ariz., Calif.; Mexico

(Baja California); Europe; n Africa; Pacific Islands (NewZealand).

A collection from California (Pasadena, “g. 9”, US)has three variants, including var. starckeanum, var.fosbergii, and what is probably best called var.starckeanum but which has an indehiscent operculum,even upon heating in KOH and Pohlstoffe. Another, fromArizona (Pima County, E. B. Bartram 990, Musci Acroc.Bor.-Amer. 572, US) includes var. starckeanum, var.brachyodus, and the indehiscent variant of var.starckeanum. Collections with capsules toward the largeend of the size scale, previously called Pottia arizonica,often have entirely smooth spores; commonly, however,some tuberculate and characteristic of M. starckeanumcan be identified in at least some capsules. Also, D. F.Chamberlain (1978) described M. starckeanum (as Pottiastarckeana) as having smooth or tuberculate spores inhis treatment for the British Isles. Capsule contents notdifferentiating into spores is not uncommon in variantsof M. starckeanum, indicating the possibility of occasionalhybridization.

1b. Microbryum starckeanum var. brachyodus(Bruch & Schimper) R. H. Zander, Bull. Buffalo Soc.Nat. Sci. 32: 240. 1993

Anacalypta starckeana var.brachyodus Bruch & Schimper,Bryol. Europ. 2: 47. 1843; Pottiaarizonica var. mucronulataWareham; P. starckeana var.brachyodus (Bruch & Schimper)Müller Hal.

Stem leaves very shortly mucro-nate. Capsule stegocarpous,

peristome absent or rudimentary. Spores smooth ortuberculate.

Capsules mature late winter or early spring (Jan–Mar).Soil; low to moderate elevations (0–700 m); Ariz., Calif.;sw Asia; n Africa.

Variety brachyodus has a rudimentary peristome (asan irregular proximal membrane) or the peristome absent.

The capsule mouth is commonly oblique and constrictedwhen dry, and the costa is only shortly excurrent as amucro (M. starckeanum var. starckeanum has shortawns).

1c. Microbryum starckeanum var. fosbergii (E. B.Bartram) R. H. Zander, Bull. Buffalo Soc. Nat. Sci. 32:240. 1993

Pottia fosbergii E. B. Bartram,Bryologist 33: 18. 1930; Phascumfosbergii (E. B. Bartram) J. Guerra

Stem leaves mucronate to short-awned. Capsule cleistocarpous,peristome rarely differentiatedinternally, operculum absent oroccasionally weakly differentiated.Spores both tuberculate and

papillose, occasionally nearly smooth.Capsules mature in early spring (Mar). Soil; low to

moderate elevations; Calif.; Mexico (Baja California).Variety fosbergii has elliptical capsules that lack both

a differentiated operculum and a peristome (exceptCalifornia: Ikenberry, CANM, which has a weaklydifferentiated operculum and rudimentary peristometeeth). This variety apparently has spores that are bothtuberculate and papillose (also true in the type),occasionally nearly smooth. If the spores are unreduced(though there is no evidence they are), then hybridizationwould be implicated. R. M. Ros et al. (1994) havedescribed from Spain a putative hybrid betweenMicrobryum starckeanum and Tortula protobryoides (asPottia bryoides), which is evidently much the same asthe var. fosbergii.

2. Microbryum davallianum (Smith) R. H. Zander, Bull.Buffalo Soc. Nat. Sci. 32: 240. 1993

Gymnostomum davallianum Smith,Ann. Bot. (König & Sims) 1: 577.1805; Pottia davalliana (Smith)C. Jensen

Distal laminal cells weakly convexsuperficially, adaxial surface ofcosta not mammillose. Setaelongate. Capsule cylindrical,stegocarpous but never cleisto-

carpous. Spores granulose, low-papillose or spiculose,large, 28–39 µm.

Varieties 3 (3 in the flora): North America, Mexico,Europe, sw Asia, Africa, Pacific Islands (New Zealand).

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Microbryum

630 ·POTTIACEAE1. Capsules commonly wide-mouthed when dehisced,

with a single row of thickened cells below mouth,peristome absent, spores spiculose, 31–39 µm. . . . . . . . . . . . . . . . . . . . . 2a. Microbryum davallianum

var. davallianum1. Capsules narrow mouthed when dehisced, with

usually 2 or more rows of thickened cells belowmouth, peristome variously absent or present,spores merely papillose, 27–34 µm.2. Peristome well developed . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . 2b. Microbryum davallianumvar. commutatum

2. Peristome rudimentary or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . 2c. Microbryum davallianum

var. conicum

2a. Microbryum davallianum (Smith) R. H. Zander var.davallianum

Pottia starckeana subsp. minutulum(Schwägrichen) D. P. Chamberlain

Capsule commonly wide-mouthedwhen dehisced, with a single rowof thickened cells below mouth;peristome absent. Spores spiculose(finely spinose) and large (31–39µm.).

Capsules mature late summerto early winter (Sept–Nov). Bare soil, vacant lots, fields;low to moderate elevations; Ont.; Mich., N.Y.; Europe;sw Asia; n Africa; Pacific Islands (New Zealand).

In var. davallianum the spines of the spores are 2–2.5µm. Distal laminal papillae are hollow or occasionallysolid, ca. 2 per lumen, usually simple but in somespecimens (e.g., Ontario, Bruce County, C. Williams1200, CANM) mostly 2-fid and hollow (with a distinctsaddle between the projections) and distinctly C-shapedin optical section.

2b. Microbryum davallianum var. commutatum(Limpricht) R. H. Zander, Bull. Buffalo Soc. Nat. Sci.32: 240. 1993

Pottia commutata Limpricht,Laubm. Deutschl. 1: 537, fig. 160.1888; P. davalliana subsp.commutata (Limpricht) Podpera

Capsule narrow-mouthed whendehisced, with usually 2 or morerows of thickened cells belowmouth; peristome well developedthough apically truncate. Sporespapillose, 27–34 µm.

Capsules mature late winter (Feb). Soil; low elevations;Calif.; Europe; sw Asia; n Africa.

Variety commutatum is known only from California(Berkeley, Howe 79, US), last collected in 1894, fromwhich station it is disjunct to the Old World.

2c. Microbryum davallianum var. conicum(Schwägrichen) R. H. Zander, Bull. Buffalo Soc. Nat.Sci. 32: 240. 1993

Gymnostomum conicumSchwägrichen, Sp. Musc. Frond.Suppl. 1(1): 26, plate 9. 1811;Pottia starckeana subsp. conica(Schwägrichen) D. P. Chamberlain;P. texana Wareham

Capsule narrow-mouthed whendehisced, with usually 2 or morerows of thickened cells below

mouth; peristome rudimentary or absent. Sporespapillose, 27–34 µm.

Capsules mature late winter and spring (Feb–Apr).Soil; low elevations; Calif., Nebr., Okla., Tex.; Mexico(Baja California); Europe; sw Asia; w, s Africa; PacificIslands (New Zealand).

The San Marcos, Texas specimen (MICH) of var.conicum has a flaring mouth like that of var. davallianum.Another Texas specimen (Orcutt 5565, CANM, WTU)has a weak peristome (a thin line of proximal membrane).

3. Microbryum floerkeanum (Weber & D. Mohr)Schimper, Syn. Musc. Eur., 11. 1860

Phascum floerkeanum Weber & D.Mohr, Bot. Taschenbuch, 70, fig.451. 1807

Distal laminal cells weakly convexsuperficially, adaxial surface ofcosta not mammillose. Setaextremely short, nearly absent.Capsule nearly spherical,cleistocarpous. Spores papillose,20–25 µm.

Capsules mature in late fall (Oct–Nov). Soil, fields,pastures; low elevations; Ont.; Europe; sw Asia; n Africa.

The leaves of Microbryum floerkeanum are ovate toovate-lanceolate and sheath the immersed capsule. Thepapillae occur near the apex of the leaf, one per lumen,and are large, nearly covering the lumen. The calyptra isvariably papillose, merely rough, or smooth. Varietybadium (Bruch & Schimper) Schimper is poorlydistinguishable, but may be segregated by its long-acuminate leaf apices—the typical variety has short-acuminate apices.

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POTTIACEAE 631·

4. Microbryum vlassovii (Lazarenko) R. H. Zander,Bull. Buffalo Soc. Nat. Sci. 32: 240. 1993

Phascum vlassovii Lazarenko, ZhurnInst. Bot. Vseukraïns’k. Akad. Nauk26/27: 196. 1938

Distal laminal cells stronglybulging superficially in medialportion of leaf, adaxial surface ofcosta strongly mammillose, besetwith “bottle-shaped” cells. Setashort, nearly absent. Capsule

nearly spherical apiculate, cleistocarpous. Sporespapillose, ca. 18–24 µm.

MICROBRYUM ° HILPERTIA

Substrate and elevational range unknown; B.C.; Calif.;Asia.

Sporophytes were immature in the area of the flora.Microbryum vlassovii was reported from BritishColumbia by T. T. McIntosh (1989) and Spain by M. N.Jiménez et al. (1990), otherwise known from Armenia,central Asia and the Ukraine. The long, smooth cellmaking up the end of the mucro is reminiscent of that ofChenia. The rhizoids bear distinctive unicellularhemispherical excrescences that may serve as tubers.Tortula acaulon may have ampullose adaxial costal cells(T. L. Blockeel 1995), but is yellow in KOH.

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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; axillaryhairs to 8 cells long, all hyaline or the proximal one yellow-brown. Stem leaves crowded, larger

Microbryum · Hilpertia

632 ·POTTIACEAEdistally, appressed and tightly spiraled when dry, weakly spreading when moist, 1–1.4 (plus0.3–0.6 of awn) mm; ovate to circular, adaxial surface flat to quite concave, base notdifferentiated; distal margins strongly revolute (to 2 ×), entire or broadly dentate above, cellsof revolute margin enlarged, strongly chlorophyllose; apex broadly acute, hyaline in an apicaltriangle; 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; transversesection 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, abaxialepidermis absent; basal cells weakly differentiated, rectangular, thin-walled; distal medial cellshexagonal to short-rectangular or rhomboid, rather large, (14–)20–25 µm wide, 1–4:1; papillaeabsent 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 apexrhomboid to fusiform, smooth. Specialized asexual reproduction by brood bodies borne onproximal 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 cellsrectangular, 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, highlyvesiculose cells; operculum broadly short-conic to long-conic, peristome teeth 32, long, linear,twisted counterclockwise about 1/2 turn. Calyptra cucullate. Spores 13–16 µm. KOH laminalcolor 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,Phylologia 65: 429. 1989

Tortula velenovskyi Schiffner, NovaActa Acad. Caes. Leop.-Carol.German. Nat. Cur. 58: 480, plate17. 1893; Hilpertia scotteri (R. H.Zander & Steere) R. H. Zander;Tortula scotteri R. H. Zander &Steere

Stems branching irregularly;rhizoids rare; leaves broadlytoothed at or near the base of the

awn; basal cells ca. 16–18 µm wide, 2–3:1. Specializedasexual reproduction by brood bodies of (1–)3–4 cells,mostly 30–50 µm. Perichaetial leaves to 1.7 mm. Seta0.35–0.40 cm, twisted counterclockwise distally,clockwise proximally. Theca operculum 0.4–1 mm, cellstwisted 1/2 times counterclockwise; peristome teeth 300–700 µm, densely branching-spiculose. Calyptra ca. 2.8mm, smooth. Spores indistinctly papillose, light brown.

Capsules mature summer. Calcareous silt and shrubsteppe; B.C., N.W.T., Nunavut; c Europe; Asia (China).

In the flora area, Hilpertia velenovskyi is a rare butdistinctive species known only from NorthwestTerritories, base of a hill on the north bank of the SouthNahanni River, 60°33’30”N, 125°23’W, from BritishColumbia, E side of Fraser R. at Big Bar, 51°15’N, 122°W,T. McIntosh 7688, May 13, 1984 (UBC), and fromNunavut, Ellesmere Island, Tanquary Fiord marineforland, 1.5 mi. SW of base camp, 81°23’N, 76°59’W,G. S. Mogensen 90-134, July 19, 1990 (C). The spiraledleaf margin of slightly enlarged, highly photosyntheticcells is similar to that of Pseudocrossidium replicatum,but the broad leaf shape and costal anatomy place H.velenovskii in the Pottioideae. I agree with the appraisalof B. C. Tan and J. Zhao (1997) that H. scotteri is asynonym. As with those in Stegonia latifolia, the cells ofthe leaf apex are rhomboid to fusiform and epapillose.

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Hilpertia

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 3mm; 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, spreadingwhen moist; ligulate to elliptic, adaxial surface occasionally grooved along costa, plane or broadlychanneled across leaf. 1.5–2.5 mm; base rectangular or not differentiated in shape; marginsdistally plane, weakly recurved or plane basally, irregularly denticulate distally with sharp, midcell projections ending in a simple papilla, marginal cells often smaller; apex rounded to broadlyacute, occasionally sharply apiculate; costa weak, ending several (6–9) cells before the apex orpercurrent, lamina inserted adaxially to laterally, adaxial cells short-rectangular, narrower thanlaminal 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 strandpresent, small to large, abaxial stereid band very weak or occasionally absent, rounded insectional shape, abaxial epidermis present; proximal cells differentiated across the leaf base exceptfor 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 beinterpreted as sharp papillae), cell walls thin, weakly trigonous, convex on both sides. Specializedasexual reproduction by brood bodies that are sometimes present on rhizoids in soil. Sexualcondition 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. Calyptracucullate. 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-marginalwall projections, often sharply apiculate by a distinctive thick-walled cell or cells, distal laminalcells 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 NorthAmerica. Bryologist 70: 112–114.

Chenia

634 ·POTTIACEAE

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1. Chenia leptophylla (Müller Hal.) R. H. Zander, Bull.Buffalo Soc. Nat. Sci. 32: 258. 1993

Phascum leptophyllum Müller Hal.,Flora 71: 6. 1888; Cheniarhizophylla (Sakurai) R. H. Zander;Tortula rhizophylla (Sakurai) Z.Iwatsuki & K. Saito; T. vectensisE. F. Warburg & Crundwell

Stems seldom branching; roundedin transverse section; rhizoidsusually few. Leaves ligulate to

elliptic, apex acute or rounded acute and apiculate withan elongate, brownish, thick-walled cell, costa percurrentor subpercurrent; proximal laminal cells ca. 20–25 µmwide, 2–4:1. Brood bodies sometimes present, borne on

rhizoids in soil, clavate, ca. 100–130 µm. Sporophytesnot known from North America.

Found on bare soil in disturbed, open areas; low tohigh elevations (0–200 m); Ala., La., N.Mex.; Mexico;South America (Bolivia, Brazil); Europe; s Africa;Australia; Pacific Islands (Hawaii).

Chenia leptophylla is one of a mundivagant group thatis apparently distributed in association with humanactivities (P. M. Eckel 1986). The small brood bodieson rhizoids in the soil seem to be the agency for asexualreproduction. The very thin-walled distal laminal cellsmay appear bright yellow-green in KOH, butexamination under the compound microscope revealsbrick red walls. The elongate, thick-walled, reflexed cellforming the leaf apiculus is unique and immediatelydiagnostic.

CHENIA ° HENNEDIELLA

Chenia

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 Harvey1855

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 orsubstereid, central strand present; axillary hairs of ca. 5 cells, proximal 1–2 cells occasionallysomewhat thick-walled. Stem leaves appressed or incurved when dry, spreading when moist;ovate to long-lanceolate, occasionally ligulate or spathulate, adaxial surface plane or broadlychanneled across leaf, (0.5–)3–4(–7) mm; base not differentiated in shape or ovate and somewhatsheathing, proximal margins commonly bordered; distal margins plane, dentate distally or lesscommonly 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 beforeapex, 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, ellipticor round in section, abaxial epidermis present (occasionally only lateral) or absent; proximalcells differentiated across leaf, rectangular, 18–30 µm wide, (2–)4–6:1, walls of proximal cellsthin; distal medial cells relatively large, quadrate to hexagonal or short-rectangular, ca. 18–24µm 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. Capsulestegocarpous, 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 andnot 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 tomuch reduced. The operculum is often long, (0.6–)1.2–1.8 mm. The genus is found in soiland 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

Hennediella

636 ·POTTIACEAE1. Hennediella heimii (Hedwig) R. H. Zander, Bull.

Buffalo Soc. Nat. Sci. 32: 248. 1993

Gymnostomum heimii Hedwig, Sp.Musc. Frond., 32. 1801;Desmatodon heimii (Hedwig)Mitten; Pottia heimii (Hedwig)Fürnrohr

Leaves ovate to broadlylanceolate, tubulose and stiff;distal margins with a 1-stratoseborder of elongate cells, distal

lamina cells 10–16 µm. Sexual condition monoicous,either synoicous or autoicous. Sporophytes often present.Seta 5–15 mm. Capsule stegocarpous, systilius; thecawide-mouthed, 1–2.4(–3.5) mm; peristome absent.

Varieties 15 (2 in the flora): North America, s SouthAmerica, Europe, Asia, Pacific Islands (New Zealand),Australia, Antarctica.

In Hennediella heimii, European infraspecific taxaattributed to the flora area are not distinct in the NewWorld (see R. T. Wareham 1939). S. Flowers (1973) mayhave included H. stanfordensis in his concept of thisspecies, judging from the description, illustrations andhis attribution of dioicy. However, like H. stanfordensis,this species may have gemmae borne on proximalrhizoids. This taxon exhibits polymorphisms (discussedby Wareham) with considerable variation indifferentiation of the leaf border of elongate cells andmarginal dentition, the distal cell ornamentation, and inlength of capsule. It may be confused with Tortulaobtusifolia but that species has no trace of marginaldentition and is yellow in KOH.

1. Leaves ovate to broadly lanceolate, tubulose butnot concave or cucullate, apex acute, marginsdentate distally, costa percurrent . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. Hennediella heimii var. heimii

1. Leaves broadly ovate to broadly elliptic, concaveand often cucullate, apex usually obtuse, marginsusually entire, costa often ending before the apex. . . . . . . . . . . . . . . . . 1b. Hennediella heimii var. arctica

1a. Hennediella heimii (Hedwig) R. H. Zander var.heimii

Pottia heimioides Kindberg

Leaves ovate to broadly lanceo-late, tubulose but not concave orcucullate, apex acute, marginsdentate; costa percurrent.

Capsules mature Jun–Jul.Moist, often alkaline soil, banks,frost boils, lake shores, friableshale, 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.,

Yukon; Alaska, Calif., Colo., Idaho, Mont., Nebr.,N.Mex., Oreg., Utah, Wash., Wyo.; s South America;Europe; Asia; Pacific Islands (New Zealand); Australia;Antarctica.

1b. Hennediella heimii var. arctica (Lindberg) R. H.Zander, Bull. Buffalo Soc. Nat. Sci. 32: 248. 1993

Pottia heimii var. arctica Lindberg,Öfvers. Kongl. Vetensk.-Akad. Förh.23: 551. 1867; Desmatodon heimiivar. arcticus (Lindberg) H. A. Crum;Gymnostomum obtusifolium R. Br.bis; P. heimii var. obtusifolia (R. Br.bis) I. Hagen

Leaves broadly ovate to broadlyelliptic, concave and often

cucullate, apex usually obtuse, margins usually entire;costa often ending before the apex.

Capsules mature late summer (Aug). Exposed soil,silt; low elevations; Greenland; N.W.T., Nunavut, Ont.;Alaska; Europe; n Asia.

Variety arctica is a somewhat poorly circumscribedtaxon largely restricted to the Arctic. It is generally muchsmaller than var. heimii, with leaves commonly 0.5–0.8mm.

2. Hennediella stanfordensis (Steere) Blockeel, J. Bryol.16: 191. 1990

Tortula stanfordensis Steere,Bryologist 54: 119, figs. 1–9. 1951;Hyophila stanfordensis (Steere)A. J. E. Smith & H. Whitehouse

Leaves broadly elliptic to obovate,flat and lax; distal margins with a2-stratose border of elongate cells,distal laminal cells 10–14 µm.Sexual condition dioicous.

Sporophytes unknown in region of flora.Soil, among grasses, drainage ditches, fields; low to

moderate elevations (100–300 m); Calif.; Mexico(Guerrero); Europe; Australia.

Since the description of Hennediella stanfordensis, thismundivagant species has engendered a flurry ofpublications on its human-oriented distribution, itsdiscovery in Europe, and the taxonomic significance ofthe eperistomate sporophyte (e.g., D. G. Long 1979;A. J. E. Smith and H. L. K. Whitehouse 1974; Whitehouse1975).

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Hennediella

POTTIACEAE 637·

38. ACAULON Müller Hal., Bot. Zeitung (Berlin) 5: 99. 1847 · [Greek a-, without, andkaulos, 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; axillaryhairs to ca. 5 cells in length, proximal cell walls sometimes thickened. Stem leaves stronglyappressed 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, proximalmargins not differentiated; distal margins plane or very weakly recurved, entire to serrulate ordentate; 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, adaxialstereid 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, wallsof 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 wallsevenly thickened, occasionally highly thickened on abaxial walls, convex on both sides of lamina.Specialized asexual reproduction absent. Sexual condition dioicous and perigoniate plantssmaller, or monoicous, usually paroicous. Perichaetia terminal, interior leaves somewhatenlarged. Seta very short, to 0.2 mm. Capsule cleistocarpous, spheric, apiculus lacking, ca.

HENNEDIELLA ° ACAULON

Acaulon

638 ·POTTIACEAE0.4–0.7 mm, annulus absent. Calyptra mitrate, often lobed. Spores ca. 25–35(–50) µm. KOHlaminal 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. Papillaeare absent in two of the three species. Although spore size and ornamentation have figured intaxonomy 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. Taxabelonging to the old, superficially similar Phascum have been recently placed largely with Tortulaand Microbryum (R. H. Zander 1993). I. G. Stone (e.g., 1989) has studied the genus extensivelyin Australia, and Casas de Puig (e.g., C. Casas de Puig et al. 1990) and her students have donethe 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 schimperianum1. Leaves cuspidate or muticus, laminal cells smooth.

2. Plants often three-angled, about 1 mm, leaves keeled, seta about as long as the diameterof the capsule, spores about 30 µm, finely papillose . . . . . . . . . . . . . . . . . . . . . . . 1. Acaulon triquetrum

2. Plants flattened-globose or three-angled, about 2 mm, leaves broadly channeled, setashort, about 0.3 the diameter of the capsule, spores 30–50 µm, smooth or papillose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. Acaulon muticum

1. Acaulon triquetrum (Spruce) Müller Hal., Bot.Zeitung (Berlin) 5: 100. 1847

Phascum triquetrum Spruce, LondonJ. Bot. 4: 189. 1845

Plants three-angled, about 1 mm.Stem leaves cuspidate, keeled;laminal cells, 13–18 µm wide, 1–3:1, smooth. Seta as long as thediameter of the capsule. Spores25–30(–40) µm, finely papillose.

Capsules mature late autumn–spring. Soil, sand, clay, old fields, pastures, roadsidebanks, temporarily moist areas; low to moderateelevations; Ont., Sask.; Calif., Ill., Iowa, Mass., N.J., Tex.,Va., W.Va.; Europe; Asia; n Africa; Australia.

Acaulon muticum var. rufescens has reflexed apiceswhile A. triquetrum does not always have these, thoughillustrated as such by H. A. Crum and L. E. Anderson(1981). The exsiccat C. F. Austin, Musci Appalachiani52 has a short seta and the plant is 1.2 mm, and is placedhere with A. muticum var. rufescens.

2. Acaulon schimperianum (Sullivant) Sullivant in W. S.Sullivant and C. L. Lesquereux, Musc. Bor.-Amer., sched.8. 1857

Phascum schimperianum Sullivant inA. Gray, Manual ed. 2, 615. 1856

Plants elliptic to globose,occasionally three-angled, 1–1.5mm. Stem leaves awned, broadlychanneled; laminal cells papilloseabaxially. Seta as long as thediameter of the capsule. Sporesspheric, 35-40 µm, papillose.

Capsules mature winter. Soil, dry washes; low tomoderate elevations; Ariz., Ill., Iowa, Kans., Tex.; Mexico(Nuevo León).

Acaulon schimperianum is awned and the distalmargins are more strongly dentate than those of the otherspecies.

SELECTED REFERENCE Crum, H. A. and L. E. Anderson. 1965. Thetaxonomy and distribution of Acaulon schimperianum. Bryologist 68:208–211.

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Acaulon

POTTIACEAE 639·3. Acaulon muticum (Hedwig) Müller Hal., Bot. Zeitung

(Berlin) 5: 99. 1847

Phascum muticum Hedwig, Sp.Musc. Frond., 23. 1801

Plants elliptic to globose, ca. 2mm. Stem leaves cuspidate,broadly channeled; laminal cellssmooth. Seta short, about 0.3 thediameter of the capsule. Sporesspheric to elliptic, 30–50 µm,papillose or nearly smooth.

Varieties 2 (2 in the flora): temperate areas of theNorthern Hemisphere, Africa.

As H. A. Crum (1969) and A. J. Grout (1939) pointedout, American specimens reported as Acaulon rubrumare A. muticum var. rufescens. These plants are oftensmall, rather three-sided and may be confused with A.triquetrum. The leaves of both varieties stain red, at leastin blotches, in KOH, and they intergrade to some extent.

1. Plants yellow-brown in nature, spores papillose-punctulose . . . . . . . 3a. Acaulon muticum var. muticum

1. Plants yellow- to red-brown in nature, sporessmooth or papillose-punctulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3b. Acaulon muticum var. rufescens

3a. Acaulon muticum (Hedwig) Müller Hal. var.muticum

Plants yellow-brown in nature.Spores densely low spiculose-papillose.

Capsules mature late fall tospring. Soil, among grasses,pastures; low to moderateelevations; Calif., Iowa, Kans.,Mass., N.Y., N.C., Okla., Tenn.,Tex.; Europe; Asia; Africa.

It may be suspected that young plants of both var.muticum and var. rufescens sometimes have smooth,

somewhat elliptic spores. A collection from Texas(Bastrop County, Bastrop, F. McAllister, Feb. 1934, A. J.Grout, North American Musci Perfecti 258, UBC) hasspheric, heavily papillose spores, 40–45 µm. Specimenswith partially smooth spores but a few large granulesadherent or scattered through the spore sac (e.g., Oregon,Lane County, Eugene, Alton Baker Park, D. Wagner1834, Mar. 9, 1978, UBC) are here placed tentativelywith the typical variety. A specimen from Iowa(Poweshiek County, Conard & Peck v.11.35, MO) hasleaves blotched red in KOH and, variably amongcapsules, weakly papillose to distinctly crowded-spiculosespores.

3b. Acaulon muticum var. rufescens (A. Jaeger) H. A.Crum, Bryologist 72: 240. 1969

Acaulon rufescens A. Jaeger, Ber.Thätigk. St. Gallischen Naturwiss.Ges. 1868–1869: 77. 1869

Plants yellow- to red-brown innature. Spores variably lowpapillose-punctulose or smooth.

Capsules mature late fall toearly spring. Soil, gravel pit,pasture, lawn; low to moderate

elevations; B.C., Ont., Que., Sask.; Ariz., Calif., Fla., Ill.,La., Md., Mich., Miss., Mo., Nebr., N.J., N.Y., Okla.,Oreg., S.C., Tex., Wis.; Mexico.

A specimen of var. rufescens from New Jersey (Austin51, UBC) is reddish and has small (ca. 20 µm), round,weakly papillose spores; generally, spore size is not a goodcharacter. Another specimen, with large (38–40 µm),round, punctulate spores, is piebald with adherentgranules similar to the papillae of var. muticum. Onespecimen placed here (California, Santa Clara County,Stanford University, W. Schofield 5724, Feb. 23, 1955,UBC), is rather immature, has a greenish cast and smooth,elliptic spores about 28-30 µm.

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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 absentor 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, shallowlychanneled along costa, to 4 mm; base scarcely differentiated in shape, proximal margins borderedby a few rows of long-rectangular cells; margins recurved along 1 or both margins in the proximal1/2, entire, bordered by ca. 6 rows of enlarged, rounded rhomboid to rectangular cells, except

Acaulon Crumia·

640 ·POTTIACEAEthe marginal row; apex broadly acute to rounded acute, usually broadly apiculate; costapercurrent or ending 1–2 cells below the apex, adaxial outgrowths absent, adaxial cells elongatein 5–6 rows, transverse section semicircular, adaxial epidermis present, adaxial stereid bandabsent, 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; papillaesmall, 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, interiorleaves 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, 0.8–1.1mm; peristome teeth 16, cleft to near base into two branches, linear, rami much perforated andanastomosing, 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 leafshape and single costal stereid band, but differs in the presence of a stem central strand (thoughsmall or occasionally absent), a more strongly differentiated, intramarginal leaf border, andpresence of a peristome (though rarely fruiting). Hennediella has bordered leaves but is red inKOH, not reddish orange, and has plane leaf margins with marginal cells not larger than themedial as seen in section. A second species, Crumia deciduidentata, was referred to Tortula byR. 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.Bot. 44: 610. 1966

Merceya latifolia Kindberg, Bull.Torrey Bot. Club 16: 94. 1889;Scopelophila latifolia (Kindberg)Renauld & Cardot

Stems sparsely radiculose;rounded-pentagonal in section.Leaf with costal abaxial cellselongate. Sporophytes 1(–2) perperichaetium. Seta reddish brown

in color, ca. 1.4 mm, twisted clockwise below,counterclockwise above. Capsule with reddish browntheca, peristome teeth of 7 articulations held together bya hyaline membrane. Calyptra ca. 3 mm.

Sporophytes mature spring–summer (May, Jun, Jul).Limestone, calcareous shale, siliceous and conglomeriticrock, roadbank, wet areas; low to moderate elevations(5–1200 m); B.C.; Ariz., Calif., Mont., Oreg., Utah, Wash.

Often locally abundant, Crumia latifolia is sometimesreddish orange in exposed situations. The morphologyand ecology of this species are discussed at length by S.Flowers (1973), W. B. Schofield (1966), and A. J. Grout(1928–1940, vol. 1). There is great variation in thethickness and color of the laminal cell walls, possiblycorrelated with degree of exposure. It has been found infruit in British Columbia and two stations in California.

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Crumia

POTTIACEAE 641·

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, hyalodermisabsent, outer cortex undifferentiated, central strand absent; axillary hairs usually 5 cells inlength, proximal cell yellowish. Leaves tubulose and contorted when dry, spreading andrecurved from the insertion when moist, ligulate-lanceolate or oblong or long-elliptic, baseweakly 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-obtuseto obtusely acute to occasionally cucullate; costa subpercurrent by 4–5 cells, adaxial outgrowthsabsent, covered ventrally with a layer of green, rounded-quadrate to short-rectangular bulgingcells 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 stereidband absent, guide cells 2–4(–5) in one row, hydroid strand absent, abaxial stereid band present,strong, semi-elliptic to semi-circular in shape; basal cells sharply differentiated in the lower 1/6of the leaf, abruptly inflated but not bulging, smooth, rising higher along the margins andforming a V-shaped area, oblong-rectangular 2–3(–4):1, thick-walled juxtacostally, becomingvery 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 surfaceplane, somewhat mammillose near the margins, the walls thickened, firm. Specialized asexualreproduction lacking. Sexual condition gynodioicous, fertile plants synoicous, plants with only

CRUMIA ° LUISIERELLA

Luisierella

642 ·POTTIACEAEarchegonia not producing sporophytes; perichaetia terminal, interior perichaetial leaves muchsmaller 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. Spores8–9 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 roundedcells, and adaxial stereid band absent. The abruptly thin-walled hyaline basal cells extendingup the leaf margin in a V shape is distinctive; it is this latter character that quickly distinguishesthis genus from similar genera: Gyroweisia, Hyophila, and Plaubelia. Luisierella bears littlerelation to the genus Tortella, which is also characterized by differentiated basal cells that extendhigher on the leaf margin than along the costa. Luisierella has been suggested to be ancestrallyrelated to the genus Quaesticula R. H. Zander (R. H. Zander 1993) through reduction ofgametophyte 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 mosses new to North America. Bryologist 48: 83–85.

1. Luisierella barbula (Schwägrichen) Steere, Bryologist48: 84. 1945

Gymnostomum barbulaSchwägrichen, Sp. Musc. Frond.Suppl. 2(2,1): 77, plate 175. 1826

Plants inconspicuous, nearly stem-less. Stems mostly unbranched,rhizoids or tomentum not evident.Leaves rosulate, 1–1.5(–2) mm,younger leaves nested in a rosetteof older leaves, which extends to

1/2 the length of the younger leaves; distal laminal cellsdeep green in sharp contrast with colorless proximal cellsregion, irregularly rounded-hexagonal, 8–11 µm. Sexualcondition with bulbous vaginula; antheridia tiny, 1/3 thelength of the archegonia; plants with only perigonia notseen. Capsule 1–2.5 mm; operculum 0.6–1 mm;peristome teeth short, irregular, sometimes appearingabsent.

Capsules mature late fall through winter (occasionallyto March). Calcareous rocks, moist to mesic sites,sinkholes and other limestone substrate (a gravestone);low elevations; Ala., Fla., Ga., Tex.; Mexico; West Indies;Central America (Belize); South America (Brazil); Asia(Indonesia, Japan).

Luisierella barbula usually grows in a thin crustassociated with cyanobacteria, Hyophiladelphus agrarius,and Weissia jamaicensis, the minute black plantscontrasting starkly with the usually chalky white rocksurface. The somewhat elevated cells on the adaxialsurface of the costa may resemble a differentiatedstructure (a pad of cells), but the cells appear to beotherwise undifferentiated from those of the lamina.

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Luisierella