BFNA Title: Rosulabryum
Author: J. Spence
Date: March 21, 2011
Edit Level: R
Version: 1

Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden

BFNA Web site: http://www.mobot.org/plantscience/BFNA/bfnamenu.htm

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XX. ROSULABRYUM J. R. Spence, Bryol. 99: 222. 1996 * [Greek rosula, rosette, and Bryum]

 

Bryum Hedwig sect. Trichophora J. J. Amann, Fl. Mouss. Suisse 2: 241, 1918

 

John R. Spence

 

Plants small to large, in open to dense low turfs or gregarious, green to red-green. Stems short to elongate, mostly strongly rosulate, sometimes in two or more interrupted rosettes, rarely evenly foliate or sub-julaceous, subfloral innovations common; often densely radiculose with brown, red-brown or red papillose rhizoids, micronemata and macronemata present, stolons absent. Leaves ovate, obovate to spathulate, variously contorted to spirally twisted around stem when dry or rarely nearly imbricate, erect to erect-spreading when wet, flat or weakly concave, base straight or slightly curved at insertion, sometimes decurrent; margins plane distally, recurved proximally or sometimes plane, 1-stratose, nearly smooth to distinctly serrate near apex, limbidium present or absent, of 1--6 rows of thick-walled elongate cells, apex broadly rounded to acute, not hyaline, costa excurrent as short- to long-excurrent colored awn, sometimes not reaching apex, variously contorted when dry, in cross-section with a well-developed abaxial stereid band, guide cells present, in 1(--2) layers, adaxial supracostal cells irregularly to regularly short- to long-rectangular at base; laminal areolation heterogeneous, distal and median laminal cells short-rhomboidal (3--5:1) sometimes porose, gradually becoming rectangular and somewhat longer proximally. Specialized asexual reproduction common, of tubers on rhizoids and leaf axis or rarely of rhizoidal unbranched to branched filiform gemmae. Sexual condition dioicous, rarely synoicous, polyoicous or autoicous; perichaetial leaves the same size as vegetative leaves, not forming a rosette, inner leaves differentiated, more acuminate with weaker costa, perigonial leaves often enlarged and distinctly rosulate. Seta usually 1(2--3), elongate, straight. Capsule clavate to cylindric or rarely pyriform, often somewhat arcuate, nutant to inclined; peristome rather uniform, double, well developed; exostome teeth 16, irregularly striate on outer surface; endostome segments 16, same height as exostome, widely perforated; basal membrane high, 1/2--2/3\x length of exostome, papillose; cilia 2--3, appendiculate. Spores small, 8--20 \um, smooth to finely papillose.

 

Species ca. 80 (13 in the flora): worldwide, but concentrated in the Southern Hemisphere and in subtropical to tropical mountains.

 

Rosulabryum is a large, distinctive genus mainly found in tropical mountainous areas and in the Southern Hemisphere in areas of seasonal temperate climates, occurring typically on soil, less commonly on rock or wood, rarely epiphytic.

 

Most Northern Hemisphere bryologists are not familiar with the great morphological diversity of the species of Rosulabryum sect. Rosulabryum, traditionally placed in Bryum sect. Rosulata, as very few extend beyond 20°N.  The most widespread representatives of Rosulabryum in the Northern Hemisphere are the small somewhat atypical species of sect. Trichophora centered around R. capillare.  Recent molecular work suggests that this section of Rosulabryum may be closer to Ptychostomum, and thus convergent on the robust Rosulata clade of the genus. However, very few species have been sampled for molecular work, and it seems unlikely that the complex of characters defining Rosulabryum could have evolved twice in unrelated clades.

 

SELECTED REFERENCES Crundwell, A.C. and H.L.K. Whitehouse. 2001. A revision of Bryum bornholmense Wink. & R. Ruthe. J. Bryol. 23: 171--176. Holyoak, D.T. and N. Pedersen. 2007. Conflicting molecular and morphological evidence within the Bryaceae (Bryopsida) and its implications for generic taxonomy. J. Bryol. 29: 111--124. Mohamed, M.A. Haji. 1979. A taxonomic study of Bryum billardieri Schwaegr. and related species. J. Bryol. 10: 401--465. Spence, J. R. 1996. Rosulabryum genus novum. Bryologist 99: 221--225. Spence, J. R. and H. P. Ramsay. Bryaceae. Flora of Australia. Vol. 51. Mosses I. Canberra. Pp. 274-348. Syed, H. 1973. A taxonomic study of Bryum capillare Hedw. and related species. J. Bryol. 7: 265--326. Wilczek, R. and F. Demaret. 1982. Etude des types de huit espèces du groupe Bryum capillare Hedw. Bull. Jard. Bot. Nat. Belg. 52: 439--462.

 

 

1. Filiform gemmae present in leaf axils of distal innovations.

2. Innovations short, rosulate, leaves large, mostly greater than 3 mm, distal lamina margins strongly serrate, limbidium strong ……..……..... 2. Rosulabryum andicola, in part

2. Innovations rosulate or elongate and evenly foliate, leaves small, mostly less than  2 mm, distal lamina margins serrulate to almost smooth, limbidium weak or absent.

3. Innovations rosulate, leaves obovate, flat, rhizoidal tubers red, filiform gemmae brown ………..………………………....10. Rosulabryum laevifilum

3. Innovations evenly foliate, leaves ovate, concave, rhizoidal tubers of various colors, filiform gemmae red, red-brown or brown.

4. Rhizoidal tubers orange to pink-orange, brighter than rhizoids, filiform gemmae red ………....…… 11. Rosulabryum pseudocapillare

4. Rhizoidal tubers and filiform gemmae brown to red-brown, same color as the rhizoids ………………………. 8. Rosulabryum flaccidum

1. Filiform gemmae absent.

5. Plants with elongate evenly foliate stems, leaves large, 2--4 mm, distinctly decurrent, margins sharply serrate distally ….….. 1. Rosulabryum andersonii

5. Plants rosulate or if evenly foliate then leaves small, less than  2 mm and distal margins mostly smooth or only weakly serrulate; leaves decurrent or not.

6. Leaves medium-sized to large, 2--4 mm, often in 2 or more interrupted tufts, margins serrate, limbidium strong or absent distally, costa excurrent as short stout awn, lamina cells incrassate, distinctly porose.

7. Distal margins of leaves lacking limbidium, leaves often in 2 or more interrupted comal tufts, hairpoint recurved when dry

 ……… 4. Rosulabryum canariense

7. Distal margins of leaves with strong limbidium, leaves usually in single tufts, hairpoint variously straight to curved but not distinctly recurved …………......………….…. 2. Rosulabryum andicola, in part

6. Leaves mostly less than  2 mm, usually not in interrupted tufts, variously contorted, twisted or imbricate, margins smooth to serrulate, limbidium present or nearly absent, costa various, not reaching apex to long excurrent in long hairpoint, lamina cells thin to thick-walled, not or weakly porose.

8. Stems more or less evenly foliate, julaceous, leaves not much contorted when dry, distal margins of leaves with weak or absent limbidium, rhizoidal tubers rarely present ..... 6. Rosulabryum elegans

8. Stems rosulate, although innovations sometimes evenly foliate, leaves variously contorted or twisted, rarely innovations leaves imbricate, distal margin of leaf with strong to nearly absent limbidium, rhizoidal tubers commonly present.

9. Plants with strong reddish tints, leaves distinctly decurrent, costa strong in short awn ……..… 7. Rosulabryum erythroloma

9. Plants green, brown-green to red-green, leaves not decurrent, hairpoint excurrent in short, medium or long awn ……….… 10

10. Plants polyoicous (synoicous, some shoots 1-sexual, appearing autoicous, others with 1-sexual lateral gametangia), rhizoidal tubers amber, orange, orange-red to crimson, generally lighter than rhizoids, distal leaf margins distinctly serrate, capsule often strongly nodding, red at maturity.

11. Tubers orange to amber, becoming brown with age, limbidium weak, leaves ovate, distal margins serrulate .........................3. Rosulabryum bornholmense

11. Tubers bright red-orange to crimson, leaves obovate, distal margins serrate, limbidium strong ...................13. Rosulabryum torquescens

10. Plants dioicous, rhizoidal tubers orange, crimson, red, red-brown to brown, mostly same color as rhizoids, if  brighter then either tubers distinctly warty with protuberant cells or distal leaf margins nearly smooth; distal leaf margins serrate, serrulate to almost smooth, capsule inclined to nutant, red-brown to brown.

12. Tubers with distinctly protuberant cells, dark red to red-crimson, clustered at base of stem on short rhizoids, distal lamina margins distinctly serrate, leaves irregularly contorted when dry ...................................12. Rosulabryum rubens

12. Tubers smooth or almost so, orange, red or brown, mostly on long rhizoids away from stem base, distal lamina margins more or less smooth, if serrate then leaves spirally twisted around stem when dry.

13. Limbidium absent or weak, margins almost smooth, innovations elongate, of imbricate, ovate leaves, somewhat reddish or red-brown, rosulate leaves irregularly contorted, tubers orange-red to red ........ 9. Rosulabryum gemmascens

13. Limbidium usually present, margins serrulate, innovations short, rosulate, green with leaves contorted, rosulate leaves spirally twisted around stem, tubers brown-red to brown

......……......… 5. Rosulabryum capillare

 

 

 

1. Rosulabryum andersonii (Crum) J. R. Spence, Novon 19: 398. 2009

 

Brachymenium andersonii Crum, Bryologist 74: 47. 1971

 

Plants large, in tall turfs, bright green. Stems long, 3--6 cm, elongate and evenly foliate, innovations evenly foliate and elongate. Leaves of main stem and innovations similar, spathulate, flat to weakly concave, 3--4.5 mm, distant, spirally twisted when dry, somewhat decurrent, margins plane distally, recurved proximally, serrate from apex to near mid leaf, limbidium strong, of 2--3 rows of hyaline cells, apex broadly acute, costa percurrent in short point, distal and median laminal cells thin-walled, rhomboidal, 3:1, 15--18 \um wide, not prorose, proximal cells longer, narrower and rectangular. Specialized asexual reproduction unkown. Sexual condition dioicous?  Capsules unknown.

 

Rare and local on moist humic soil in broad-leaved forest; 1300 m; N.C. A species of conservation concern.

 

Rosulabryum andersonii was transferred to Rosulabryum because it does not fit in the type section of Brachymenium, which consists of tropical epiphytes with erect capsules. Rosulabryum andersonii is morphologically similar to several robust tropical Rosulabryum species that have elongate evenly foliate stems. Despite repeated searches, the population has never been relocated, and the species may be extinct.

 

2. Rosulabryum andicola (Hooker) Ochyra, Biodiv. Poland 3: 162. 2003

 

Bryum andicola Hooker, Syn. Pl. 1: 58, 1822

 

Plants small to large, in open to dense low turfs or gregarious, green to red-green. Stems short to long, 0.4--5 cm, distinctly and singly rosulate, innovations rosulate. Leaves of main rosette and innovations similar although latter smaller, obovate, 0.6--4 mm, irregularly twisted and spirally twisted around stem when dry, erect-spreading when wet, weakly concave, not decurrent; margins recurved on larger leaves to near tip, distinctly and strongly serrate in distal 1/3 of margin, limbidium strong, of 3--4 layers of pale yellow or whitish cells, apex broadly acute, costa excurrent in short stout awn; distal and median laminal cells thin- to firm-walled, short-rhomboidal (3--5:1), porose, proximal cells long-rectangular in rosette leaves, porose, short-rectangular in smaller proximal leaves. Specialized asexual reproduction by brownish filiform gemmae, densely papillose, and rhizoidal tubers, 300--1000 \um, reddish brown, spheric. Sexual condition dioicous. Capsule cylindric, symmetric, brown, 3--6 mm, nutant to inclined.

 

Capsules mature June--August. Uncommon to locally common on moist to dry soil, soil over rock or rarely wood in mountainous areas; 900--2500 m; se Ariz., s N.Mex., w Tex.; Mexico; Central America; South America; s Africa.

 

When well-developed Rosulabryum andicola is a large, striking species, but at its northern edge of range in the Southwest it is often much smaller.  The strong border and sharply serrate leaves combined with the filiform gemmae serve to distinguish smaller specimens from small species Rosulabryum.  It is most closely related to the southern hemisphere R. billardieri and R. albolimbatum. There is some confusion over the identity of the species, as the type is reported to be similar to R. flaccidum.  Specimens named R. andicola in the sense of M. A. Mohamed (1979) are large, robust plants, very unlike the small R. flaccidum.  If R. andicola is synonymous with R. flaccidum, then the next oldest name for this species is B. ehrenbergianum Müll. Hal.

 

 

3. Rosulabryum bornholmense (Wink. & R. Ruthe) J. R. Spence, Novon 19: 398. 2009  I

 

Bryum bornholmense Wink. & R. Ruthe, Hedwigia 38 (Beibl. 3): 120. 1899

 

Plants small, in open to dense low turfs or gregarious, red-green. Stems short 0.5--2 cm, fertile stems evenly foliate to weakly rosulate, innovations evenly foliate. Leaves of main rosette and innovations similar, ovate, 1--2 mm, somewhat irregularly twisted to contorted when dry, erect-spreading when wet, weakly concave, not decurrent, margins recurved to mid leaf, weakly but distinctly serrulate in distal 1/3 of margin, limbidium moderately distinct, of 1--2 rows, apex acute, costa slender, excurrent in medium-length colored awn, distal and median laminal cells thick walled, rhomboidal (3--4:1), 14--20 \um wide, not porose, proximal cells short-rectangular, innovation leaves similar but smaller. Specialized asexual reproduction of rhizoidal tubers, on long rhizoids in soil, (120--)180--350 \um, amber, orange, orange-red or dark red, becoming orange-brown, pale brown or dark brown when old, spheric, cells not or only weakly protuberant. Sexual condition dioicous, synoicous or autoicous. Capsule cylindric, symmetric, red to red-brown, 2--3 mm, nutant.

 

Capsules mature April-August. Rare and local on disturbed soil; 100 m; Calif.; Europe.

 

Rosulabryum bornholmense was probably introduced from Europe. A. C. Crundwell and H. L. K. Whitehouse (2001) revised R. bornholmense, providing new criteria to separate it from the closely related R. rubens. In addition to tuber and awn differences, the cells of the tubers in R. bornholmense are 45--60 \um across, while those of R. rubens are 30--35 \um across. Most North American collections can be referred to the latter species.

 

 

4. Rosulabryum canariense (Bridel) Ochyra, Biodiv. Poland 3: 162. 2003

 

Brym canariense Bridel, Muscol. Recent. Suppl. 3: 29. 1817

 

Plants small to large, in open to dense low turfs or gregarious, bright green. Stems short to long, 0.5--2 cm, distinctly rosulate, typically with 2--3 interrupted rosettes on the same stem, innovations rosulate. Leaves of main rosettes and innovations similar although the latter smaller, obovate to spathulate, 2.5--4.5 mm, irregularly contorted when dry, not spirally twisted around stem, erect-spreading when wet, weakly concave, not decurrent; margins recurved on larger leaves to mid leaf, distinctly and strongly serrate in distal 1/3 of margin, limbidium absent or weak, apex acute; costa excurrent in short, slender awn, usually recurved when dry; distal and median laminal cells firm- to thick- walled, rhomboidal (3--5:1), 15--23 \um wide, porose, proximal cells long-rectangular in rosette leaves, porose, short-rectangular in smaller proximal leaves. Specialized asexual reproduction of rhizoidal tubers, 200--400 \um, red, spheric. Sexual condition autoicous or rarely synoicous. Capsule elongate-pyriform, symmetric, red-brown, 3--6 mm, nutant.

 

Capsules mature May--June. Common on moist soil banks and soil over rock in coastal maritime to Mediterranean climates, 0--1600 m, B.C.; Wash., Oreg., Calif.; Mexico; s,w Europe; Asia (Turkey); Atlantic Islands (Macaronesia); reports from East Africa and Central America need to be re-evaluated.

 

Rosulabryum canariense is a large species easily distinguished by the series of interrupted rosettes along the stem, short, recurved hairpoint, weak or absent limbidium, and porose lamina cells.

 

 

5. Rosulabryum capillare (Hedwig) J.R. Spence, Bryologist 99: 223. 1996

 

Bryum capillare Hedwig, Sp. Musc. Frond., 182, 1801

 

Plants small, in open to dense low turfs or gregarious, bright green. Stems short 0.5--1.5 cm, distinctly singly rosulate, innovations short and rosulate. Leaves of main rosette and innovations similar, obovate, 0.5--2.5 mm, regularly spirally twisted around stem, erect-spreading when wet, weakly concave, not decurrent; margins recurved to mid leaf, distinctly serrulate in distal 1/3 of margin, limbidium present, of 1--3 rows, apex acute, costa excurrent in long, sometimes hyaline, slender awn, irregularly twisted when dry; distal and median laminal cells thin-walled, short-rhomboidal (3--4:1), 12--25 \um wide, not porose, proximal cells long-rectangular in rosette leaves. Specialized asexual reproduction of rhizoidal tubers, 200--300 \um, red-brown, same color as rhizoids, spheric. Sexual condition dioicous. Capsule cylindric to elongate-pyriform, symmetric, red-brown, 3--5 mm, inclined.

 

Sporophytes mature May--July. Uncommon to locally common on moist shaded soil, soil banks or rotting wood; 0--2500 m; Greenland;  Alta., B.C., Man., N.B., Nfld. and Labr., N.S., Ont., P.E.I., Que., Sask.; Alaska, Ariz., Ark., Calif., Colo., Conn., Del., Idaho, Ind., Iowa, Ky., Maine, Md., Mass., Mich., Minn., Mo., Mont., Nebr., Nev., N.H., N.J., N.Mex., N.Y., N.Dak., Ohio, Oreg., Pa., R.I., S.C., Tenn., Utah, Vt., Va., Wash., W.Va., Wis., Wyo.; Mexico; West Indies; Central America; South America; Eurasia; Africa; Atlantic Islands (including Macaronesia); Pacific Islands (New Zealand); Australia.

 

Traits diagnostic for the protean Rosulabryum capillare are the leaves spirally twisted around the stem, small size of plant, rosulate innovations, rhizoidal tubers the same color as the rhizoids, leaf with long awn, distinct border, serrulate margins. Rosulabryum gemmascens has a much weaker border with nearly smooth margins, irregularly contorted leaves, and elongate somewhat imbricate innovations of small red-brown ovate leaves. Rosulabryum laevifilum has a much more variable costa that sometimes does not reach the apex, green leaves, and filiform gemmae.  Rosulabryum torquescens has irregularly contorted leaves with strongly serrate margins and red to crimson tubers, brighter than the rhizoids.

 

6. Rosulabryum elegans (Nees) Ochyra, Biodiv. Poland 3: 162. 2003

 

Bryum elegans Nees, Bryol. Univ. 1: 849. 1827; Bryum stirtonii Schimper

 

Plants small, in open to dense low turfs or gregarious, bright to dark green, olive-green or rarely red-green. Stems short 0.5--2 cm, evenly foliate, more or less julaceous. Leaves of main rosette and innovations similar, ovate to obovate, 0.5--1.5 mm, more or less imbricate when dry, sometimes slightly twisted and more loosely set, erect when wet, concave, decurrent or not; margins plane or sometimes recurved to mid leaf, serrulate in distal 1/3 of margin, limbidium present, of 1--2 rows, apex broadly acute to acute, costa excurrent in long, sometimes hyaline, stout awn, straight to somewhat twisted when dry; distal and median laminal cells thin- to thick-walled, short-rhomboidal (3--4:1), 14--30 \um wide, not porose, proximal cells short- to long-rectangular, porose. Specialized asexual reproduction of rhizoidal tubers, very rare, (100--)150--200 \um, brown, same color as rhizoids, spheric. Sexual condition dioicous. [Capsule pyriform, symmetric, red-brown, 2--3 mm, strongly nutant.]

 

Rare on calcareous rock or soil, cool to Arctic-alpine climates, typically in the mountains; 1000--4000 m; Alta., B.C., N.W.T., Nunavut; Alaska, Calif., Maine, Mich., Nev., Oreg., Utah, Wash.; Europe (including European Russia), Asia (Turkey).

 

Capsules of Rosulabryum elegans have not been reported from the flora area. The plants are typically dark green with somewhat julaceous stems, but in shaded moist areas the stems become looser with more widely set and somewhat twisted paler green decurrent leaves.  These latter plants have been named Bryum stirtonii, but they intergrade completely with R. elegans from drier more exposed sites.

 

7. Rosulabryum erythroloma (Kindberg) J. R. Spence, Novon 19: 398. 2009

 

Bryum capillare subsp. erythroloma Kindberg, Eur. N. Amer. Bryin. 2: 358, 1897 [1898];  B. erythroloma (Kindberg) Syed

 

Plants small to medium-sized, open to dense low turfs or gregarious, maroon or red to red-green. Stems short 1--2.5 cm, distinctly singly rosulate, innovations short and rosulate. Leaves of main rosette and innovations similar, obovate, 1--2.5 mm, appressed and not much altered to sometimes spirally twisted around stem, erect-spreading when wet, flat, decurrent; margins recurved to mid leaf, distinctly serrulate in distal 1/3 of margin, limbidium present, of 2--4 rows, red to sometimes yellowish in older leaves, apex acute, costa variable, not reaching apex, percurrent to short excurrent in a stout awn, distal and median laminal cells firm walled, short-rhomboidal (2--3:1), 15--25 \um wide, not porose, proximal cells long-rectangular in rosette leaves. Specialized asexual reproduction of rhizoidal tubers, (180--)200--350 \um, orange, maroon to red, brighter than rhizoids, spheric. Sexual condition dioicous. Capsule cylindric to elongate-pyriform, symmetric, brown to red-brown, 2.5--4 mm, nutant.

 

Sporophytes mature April--June. Locally common on moist shaded soil, soil banks or rotting wood in lowland forests and other protected sites near the Pacific coast; 0--500 m; B.C.; Calif., Oreg., Wash; Mexico.

 

Rosulabryum erythroloma is a distinctive coastal endemic, characterized by reddish decurrent leaves with a short awn.  The capsules often become strikingly bright red, and nutant as they mature.

 

8. Rosulabryum flaccidum (Bridel) J. R. Spence, Phytologia 89: 114. 2007

 

Bryum flaccidum Bridel, Bryol Univ. 1: 667. 1826

 

Plants small, in open to dense low turfs or gregarious, brown-green to red-green, occasionally bright green. Stems short 0.5--2 cm, fertile stems singly rosulate, rare, plants dominated by numerous slender, evenly foliate, elongate, sterile innovations. Leaves of main rosette and innovations somewhat different, rosette leaves ovate to obovate, 0.6--2 mm, somewhat irregularly twisted to contorted when dry, erect when wet, weakly concave, not decurrent, margins plane or recurved to mid leaf, smooth or weakly serrulate in distal 1/3 of margin, limbidium absent or present, weak, of 1 row, apex acute, costa excurrent in short- to medium-length colored awn, distal and median laminal cells thin-walled, long-rhomboidal (3--5:1), 12--18 \um wide, not porose, proximal cells long-rectangular in rosette leaves, innovation leaves broadly lanceolate to ovate, more or less imbricate when dry, concave, 0.4-1.5 mm, dark green to red-brown, with a short colored awn. Specialized asexual reproduction of filiform gemmae in distal leaf axils or rarely on leaves, brown, finely papillose, and rhizoidal tubers, (100--)150--300 \um, brown to red-brown, spheric. Sexual condition dioicous. Capsule subcylindric, symmetric, brown, 2--3 mm, nutant.

 

Sporophytes rare, mature May--August. Uncommon to locally common, exposed to shaded rock, soil, soil over rock, or rotting wood, rarely on bark; 0--2000 m; Ont.; Ariz., Calif., Colo., Md., Mo., Nev.,  West Indies.

 

In North America, there has been significant confusion over the identity of the small species that produce filiform gemmae in the leaf axils.  H. Syed (1973) named the common widespread species with short, rosulate innovations R. flaccidum, and described a new species, R. laevifilum, for specimens with smooth gemmae. However, these two species completely intergrade, and the name R. flaccidum was incorrectly applied.  The correct name is thus R. laevifilum for what has passed as R. flaccidum in North America. True Rosulabryum flaccidum is a very different species related to R. pseudocapillare, with a type from Hispaniola.

 

9. Rosulabryum gemmascens (Kindberg) J. R. Spence, Novon 19: 399 2009

 

Bryum gemmascens Kindberg, Eur. N. Amer. Bryin. 2: 360. 1897; B. sanguilentum Renauld & Cardot; B. trichophorum Kindberg

 

Plants small, in open to dense low turfs or gregarious, green or brown to red-green. Stems short 0.5--2 cm, distinctly singly rosulate, innovations elongate and evenly foliate. Leaves of fertile rosettes and innovations different, rosette leaves broadly ovate to obovate, 1--2.5 mm, irregularly twisted to contorted when dry, erect-spreading when wet, flat, not decurrent, margins recurved to mid leaf, smooth or weakly serrulate in distal 1/3 of margin, limbidium absent or present, weak, of 1 row, apex acute, costa variable, not reaching apex to excurrent in slender, medium-length, colored or hyaline awn, irregularly twisted when dry; distal and median laminal cells thin-walled, long-rhomboidal (3--5:1), 12--20 \um wide, porose, proximal cells long-rectangular in rosette leaves, innovation leaves broadly lanceolate to ovate, somewhat twisted to weakly imbricate when dry, concave, 0.5--1.5 mm, red-brown, with a short colored to sometimes hyaline awn. Specialized asexual reproduction of filiform gemmae in distal leaf axils, rarely present, brown, finely papillose, and rhizoidal tubers, (60--)100--200 \um, brown, spheric. Sexual condition dioicous. Capsule cylindric, symmetric, brown to red-brown, 2--5 mm, inclined to nutant.

 

Sporophytes mature April--June. Uncommon to locally common, exposed to shaded soil, soil over rock, rotting wood; 0--1000 m; B.C.; Calif., Oreg., Wash.

 

Rosulabryum gemmascens is closely related to R. capillare, but differs in the unusual evenly foliate red-brown innovations, leaves not spirally twisted around stem, very weak or absent limbidium and almost smooth distal margins. It is endemic to coastal areas, and is especially common in Mediterranean climate areas of California.

 

10. Rosulabryum laevifilum (Syed) Ochyra, Biodiv. Poland 3: 162. 2003

 

Bryum laevifilum Syed, J. Bryol. 7: 293. 1973

 

Plants variable, very small to medium, in open to dense low turfs or more often gregarious, bright green. Stems short, 0.3--1.5 cm, distinctly singly rosulate, innovations short and rosulate. Leaves of main rosette and innovations similar, obovate, 0.4--3 mm, irregularly shrunken or contorted but not spirally twisted around stem, erect-spreading when wet, flat, decurrent; margins plane or recurved to mid leaf, serrulate in distal 1/3 of margin, limbidium present but often weak, of 1--2 rows, apex acute, costa highly variable, not reaching apex to excurrent in short slender awn in same rosette, irregularly twisted when dry; distal and median laminal cells thin-walled, short-rhomboidal (3--4:1), 12--20 \um wide, not porose, proximal cells long-rectangular in rosette leaves. Specialized asexual reproduction of filiform gemmae in axils of distal leaves or sometimes arising from leaves, green when young, brown when mature, finely papillose to more or less smooth, and rhizoidal tubers, (70--)100--200 \um, brown, same color as rhizoids, spheric. Sexual condition dioicous. Capsule subcylindric, symmetric, brown to red-brown, 2--4 mm, inclined to nutant.

 

Sporophytes rare, mature April--August. Uncommon to locally common and widely scattered, bark, rotten wood, rarely rock or soil; 0--2500 m; Alta., B.C., Man., Ont., Que., Sask.; Alaska, Ariz., Ark., Calif., Colo., Idaho., Ill., Ind., Iowa, Kans., Ky., Maine, Md., Mass., Mich., Minn., Mo., Mon., Nebr., Nev., N.H., N.J., N. Mex., N.Y., N.C., N.Dak., Ohio, Oreg., Pa., Tenn., Utah, Vt., Va., Wash., Wis., Wyo.; Europe.

 

 

Rosulabryum laevifilum is the flora’s most widespread species of the genus with filiform gemmae, and is the most common corticolous species.  Specimens on bark are often very small compared with those on other substrates. See also the discussion under R. flaccidum.

 

11. Rosulabryum pseudocapillare (Bescherelle) Ochyra, Biodiv. Poland 3: 162. 2003

 

Bryum pseudocapillare Bescherelle, Ann. Sci. Nat. Bot. sér. 6, 3: 205. 1876

 

Plants small, in open to dense low turfs or gregarious, green to red-green. Stems short 0.5--2 cm, fertile stems singly rosulate, rare, plants dominated by numerous slender evenly foliate elongate sterile innovations. Leaves of main rosette and innovations somewhat different, rosette leaves ovate to obovate, 0.6--2 mm, somewhat irregularly twisted to contorted when dry, erect-spreading when wet, flat, not decurrent, margins recurved to mid leaf, smooth or weakly serrulate in distal 1/3 of margin, limbidium absent or present, weak, of 1 row, apex acute, costa variable, not reaching apex to excurrent in slender medium-length colored or hyaline awn, irregularly twisted when dry; distal and median laminal cells thin-walled, long-rhomboidal (3--5:1), 12--18 \um wide, not porose, proximal cells long-rectangular in rosette leaves, innovation leaves broadly lanceolate to ovate, more or less imbricate when dry, concave, 0.4--1.5 mm, red-brown, with a short colored awn. Specialized asexual reproduction of filiform gemmae in distal leaf axils or rarely on leaves, red, finely papillose, and rhizoidal tubers, (100--)150--400 \um, orange, red to pink, spheric. Sexual condition dioicous. Capsule subcylindric, symmetric, brown, 2--3 mm, nutant.

 

Sporophytes rare, mature April--July. Uncommon to locally common, damp rotting wood, bark, rarely soil in subtropical regions; 0--500 m; Ala., Fla., Ga., La., Miss., S.C.; Mexico; West Indies; Central America; Pacific Islands (Hawaii).

 

Rosulabryum pseudocapillare is very closely related to R. flaccidum, but differs in ecology, distribution, and color of the filiform gemmae and tubers. See the discussion under R. flaccidum.

 

12. Rosulabryum rubens (Mitten) J. R. Spence, Novon 19: 399. 2009  I

 

Bryum rubens Mitten, Hooker's J. Bot. Kew Gard. Misc. 8: 232. 1856

 

Plants small, in open to dense low turfs or gregarious, red-green. Stems short 0.5--2 cm, fertile stems evenly foliate to weakly rosulate, innovations evenly foliate. Leaves of main rosette and innovations similar, ovate, 1--2.5 mm, somewhat irregularly twisted to contorted when dry, erect-spreading when wet, weakly concave, not decurrent, margins recurved to mid leaf, distinctly serrulate in distal 1/3 of margin, limbidium moderately distinct, of 1--2 rows, apex acute, costa slender, excurrent in short colored awn, distal and median laminal cells thin-walled, rhomboidal (3--4:1), 15--20 \um wide, not porose, proximal cells short-rectangular, innovation leaves similar but smaller. Specialized asexual reproduction by rhizoidal tubers, on short rhizoids at base of stem, (120--)150--300 \um, crimson, red to dark red, spheric, cells strongly protuberant. Sexual condition dioicous. Capsule cylindric, symmetric, red to red-brown, 2--3 mm, nutant.

 

Capsules mature April-August. Rare and local, disturbed soil, occasionally concrete; 0--1000 m; B.C., Que.; Calif., Md., N.J., Okla., Tenn.; Europe; Asia (India); Australia.

 

Rosulabryum rubens was most probably introduced from Europe, and is likely to be more widely distributed in temperate North America than the records indicate. Although originally considered related to Gemmabryum subapiculatum, morphology as well as recent molecular work support a position near R. capillare.

 

13. Rosulabryum torquescens (Bruch ex De Not.) J. R. Spence, Bryologist 99: 223. 199

 

Bryum torquescens Bruch ex De Notaris, Syllab. Musc., 163. 1838; B. capillare subsp. torquescens (Bruch ex De Notaris) Kindberg

 

Plants small to more often medium-sized, in open to dense low turfs or gregarious, green or red-green. Stems short 1.5--2 cm, distinctly singly rosulate, innovations short and rosulate. Leaves of main rosette and innovations similar, broadly ovate to obovate, 1.5--3 mm, slightly twisted to contorted when dry, erect-spreading when wet, flat, not decurrent; margins recurved from mid leaf to near apex, distinctly and strongly serrate in distal 1/3 of margin, limbidium present, strong, of 2--4 rows, apex acute, costa excurrent in short to medium stout or slender awn, irregularly twisted when dry; distal and median laminal cells firm-walled, long-rhomboidal (3--5:1), 12--20 \um wide, not porose, proximal cells long-rectangular in rosette leaves. Specialized asexual reproduction of rhizoidal tubers, (100--)200--300 \um, scarlet, crimson to red, brighter than rhizoids, spheric. Sexual condition dioicous or polyoicous. Capsule elongate-pyriform, symmetric, red, 3--6 mm, strongly nutant.

 

Sporophytes mature April--June. Uncommon to locally common, soil or rock over soil, rarely on rotting wood; 0--1500 m; B.C.; Ariz., Calif., Nev., Oreg., Wash.; South America; Eurasia; Africa; Atlantic Islands (including Macaronesia); Pacific Islands (New Zealand); Australia.

 

Rosulabryum torquescens is found on all continents except Antarctica, particularly in seasonal Mediterranean, warm-temperate to subtropical regions, in climates that are warmer and more seasonal than for R. capillare. This is a distinctive species, characterized by irregularly contorted leaves with a short to medium-length awn, strong limbidium, strongly serrate distal margins, mixed sexuality, bright red to crimson tubers, and strongly nutant (at maturity), red capsules.  In the flora region it is most common in Mediterranean climates along the coast, but extends inland in semi-arid to arid regions of the Southwest.