BFNA Title: Bryaceae, family description
Author: J. R. Spence 
Date: November 9, 2007
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Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden

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XX. BRYACEAE Schwägrichen

John R. Spence

 

Plants acrocarpous, but sporophytes occasionally appearing lateral due to rapid innovating growth; tiny to robust, as scattered individuals or forming open to dense turfs or cushions, green, silver, white, golden or red, plants often more than one color. Stems short to long, 0.1--10 cm, sometimes julaceous, unbranched to sparsely branching by subfloral innovations, stolons occasionally present; rhizoids sparse to abundant, variously colored, smooth to papillose, micronemata and/or macronemata often present. Leaves imbricate to variously contorted or twisted when dry, erect to erect-spreading when wet, broadly lanceolate, ovate, ovate-lanceolate, obovate to spathulate, 0.03--6 cm, base straight or curved at insertion, decurrent or not; margins plane or revolute, 1- or 2-stratose, rarely multistratose, limbidium present or absent, apex broadly rounded to acute or acuminate, apiculus sometimes present, costa percurrent or subpercurrent, or excurrent as a short- to long-excurrent awn, transverse section with stereid band single, usually well developed, occasionally greatly reduced, with or without guide cells; adaxial supracostal cells irregularly to regularly quadrate or short- to long-rectangular proximally, laminal cells relatively uniform throughout lamina or obscurely to distinctly heterogenous, proximal cells usually quadrate, short- or long-rectangular, often distinctly different in shape or occasionally similar to median and distal, medial cells generally similar to distal cells, distal cells short to very long, hexagonal to rhomboidal, sometimes vermicular, 2--10:1, sometimes occurring in rows oblique to the costa, thin- to thick-walled, walls sometimes pitted, alar cells usually similar to juxtacostal cells, sometimes differentiated into a small group of quadrate cells. Specialized asexual reproduction common, of seven distinct types; spherical to ovoid rhizoidal tubers, filiform rhizoidal gemmae, axillary filiform gemmae, leaf axil bulbils, stem tubers, slender leafless terminal shoots, or leaf axil deciduous brood branchlets. Sexual condition dioicous or monoicous, sometimes variable within species (polyoicous); perigonia and perichaetia terminal or lateral, perichaetial leaves the same size as vegetative leaves or typically larger, sometimes forming a rosette, inner leaves usually highly differentiated, often narrower with a weaker costa. Seta usually 1, sometimes polysetous, variously colored, long, straight, twisted or geniculate. Capsule erect, inclined to nutant, long-exserted, 1--8 mm, ovate, spherical, cylindrical, or pyriform, occasionally zygomorphic, hypophysis well-differentiated or not, sometimes expanded and rugose, exothecial cells near mouth quadrate or short-rectangular, thick-walled, often reddish, in 1--3 or more rows, medial cells longer, short- to long-rectangular with straight or sinuose walls; annulus usually present, revoluble; operculum convex, short to tall-conic, sometimes rostrate; peristome diplolepidous-alternate, rarely reduced to one layer or absent, exostome white to pale yellow or tan, sometimes reddish, teeth triangular to lanceolate, trabeculate, sometimes with small pores along fissural line, endostome hyaline to pale yellow, separate or sometimes adherent to exostome, segments narrow to wide, usually broadly perforate, basal membrane low to high, cilia present, 1--3, usually appendiculate, to variously reduced in number or length or sometimes absent. Calyptra fugacious, cucullate, small, smooth. Spores shed singly or as tetrads, rarely germinating in capsule, 8--60 \um, smooth to papillose, pale yellow, tan or nearly hyaline, rarely darker.

 

Genera 15, species ca. 500 (12 genera, 94 species in flora): worldwide.

 

The Bryaceae is a large family of acrocarpous mosses with a global distribution. Many species are adapted to disturbed soil and are somewhat weedy.  Species exhibit a remarkable array of specialized asexual reproductive structures, perhaps more than in any other bryophyte family. The genus Bryum in the broad sense has a reputation for being taxonomically difficult.  Part of this is because the genus is highly polyphyletic, and also because, traditionally, many species have been distinguished by minor differences in peristome features.  Previous classifications have over-emphasized the peristome, placing taxa with strikingly different gametophytes in the same genus. The gametophytes of Bryum vary widely morphologically, and this has been used as a basis to re-classify the genus and its relatives (J. R. Spence 2005; J. R. Spence and H. P. Ramsay 2005). 

 

Recent genetic research has radically changed understanding of relationships in the family. These studies have shown that Pohlia and related genera, traditionally considered part of the Bryaceae, are more closely related to members of the Mniaceae. Further, Orthodontium is only distantly related to the Bryaceae, while Leptobryum appears to be closest to the Meesiaceae (C. J. Cox and T. A. J. Hedderson 2003). Roellia may also be more closely related to the Mniaceae, and for the flora has been removed to its own family. Within the re-circumscribed Bryaceae, results based on morphology and genetics do not always agree (N. Pedersen et al. 2003). Recent attempts to revise generic limits in the family using DNA evidence have produced large unwieldy clades that defy coherent morphological description. However, to date most phylogenetic research has focused on the chloroplast genome, without taking into account the many critical assumptions underlying the use of DNA sequence data. Recent studies of portions of the chloroplast genome of land plants have indicated that lineage sorting, gene transfer, and paralogy may be fairly common, thus potentially obscuring phylogenetic relationships. Because of these uncertainties, this treatment is based primarily on the morphology of the gametophyte. Differences between phylogenetic and morphological approaches are discussed under each genus.

 

SELECTED REFERENCES: Andrews, A. L. 1935. Bryaceae. In: Grout, A. J. Moss flora of North America, Vol. 2. Newfane: Vermont.   Cox, C. J. and T. A. J. Hedderson. 2003. Phylogenetic relationships within the moss family Bryaceae based on chloroplast DNA evidence. J. Bryol. 25: 31--40.  Nyholm, E. 1993. Illustrated Flora of Nordic mosses, Fasc. 3. Copenhagen and Lund: Nordic Bryological Society.  Ochi, H. 1980. A revision of the Neotropical Bryoideae, Part 1. J. Faculty Educ. Tottori Univ., Nat. Sci. 29: 49--154.  Ochi, H. 1981. A revision of the Neotropical Bryoideae, Part 2. J. Faculty Educ. Tottori Univ., Nat. Sci. 30: 21--55.  Ochi, H. 1992. A revised infrageneric classification of the genus Bryum and related genera (Bryaceae, Musci). Bryobrothera 1: 231--244.  Pedersen, N., C. J. Cox and L. Hedenäs. 2003. Phylogeny of the moss family Bryaceae inferred from chloroplast DNA sequences and morphology. Syst. Bot. 28: 471--482.  Smith, A. J. E. 2004. The moss flora of Britain and Ireland, Ed. 2, Cambridge.  Spence, J. R. 1988. Bryum Hedw. (Bryaceae) in western North America. Bryologist 91: 73--85.  Spence, J. R. 2005. New genera and combinations in Bryaceae (Bryales, Musci) for North America. Phytologia 87: 15--28.  Spence, J. R. 2006. New combinations in the Bryaceae (Bryophyta) for North America. II. Phytologia 89: 110--114.  Spence, J. R. and H. P. Ramsay. 2005. New genera and combinations in the Bryaceae (Bryales, Musci) for Australia. Phytologia 87: 61--72.  Spence, J. R. and H. P. Ramsay. 2006. Bryaceae. In: Flora of Australia, Vol. 51, Mosses 1: 274--348.

 

Sizes of stems and leaves in the keys are as follows; Stems: short (less than 10 mm), medium (10--30 mm), long (more than 30 mm); leaves: small (less than 1.5 mm), medium (1.5--3 mm), large (3-4 mm), robust (more than 4 mm).

 

1.Stems short, mostly less than 10 mm, julaceous; plants green, yellow-green to silver-white; leaves typically less than 1 mm; distal lamina cells long, (2--)3--10:1, proximal cells quadrate to short-rectangular, transition often abrupt; bulbils and rhizoidal tubers sometimes present; dioicous

2. Plants pale yellow or yellow-green, older plants consisting of long very slender and string-like branches; distal laminal cells elongate (6--)8--10:1, often thick-walled; capsules elongate cylindrical   …………………………………….……. 1. Anomobryum, p. XX

2. Plants white, silver-green, green, or brown-green, older plants with short, rounded

strongly julaceous stems or stems gemmiform; distal laminal cells short to moderately

long, 2--6(--8):1; capsules short, ovate, with thick neck ……….……… 3. Bryum, p. XX

1. Stems short to long, rarely julaceous (if so then plants reddish); plants green, yellow-green, red, pink-silver, or rarely hyaline above; leaves (0.5--)1--10 mm; distal lamina cells short to long, mostly 2--6:1, proximal cells variously quadrate, short- to long-rectangular, transition abrupt or not; asexual reproduction of all types present; dioicous or monoicous.

3. Plants rosulate with obovate to spathulate leaves, margins distally serrate.

4. Stolons present; leaves large, often greater than 5 mm; costal stereid band reduced; gemmae lacking ………………11. Rhodobryum, p. XX

4. Stolons absent; leaves typically less than 5 mm, if longer then filiform gemmae present; costal stereid band well developed; rhizoidal tubers and leaf axil filiform gemmae often present

5. Proximal cells of rosette leaves quadrate to short-rectangular, 1--2:1; filiform leaf axil gemmae lacking; capsules erect, peristome reduced

……..…………………………………………..2. Brachymenium, p. XX

5. Proximal laminal cells of rosette leaves rectangular, 2--4:1; filiform gemmae sometimes present in leaf axils; capsules inclined to nodding,  peristome well developed ………… 12. Rosulabryum (in part), p. XX

3. Plants comose to gemmiform, bulbiform or evenly foliate, never distinctly rosulate; leaves ovate, ovate-lanceolate to triangular; margins distally smooth to serrulate

6. Plants typically in dense cushions, golden-silver to pink-silver; leaves with long, spinose, hyaline awn; distal lamina cells rhomboidal, 3--4:1, proximal cells quadrate; capsule long-cylindrical, erect; peristome reduced, cilia absent; epiphytic, corticolous, saxicolous, rarely terricolous  ………………………………………………….….……7. Leptostomopsis, p. XX

6. Plants usually not in dense cushions, of various colors; awn present or absent, rarely hyaline or spinose; distal and proximal lamina cells various; capsule inclined to nutant, if erect then short ovoid; terricolous or saxicolous, rarely corticolous.

7. Plants green, yellow, or red, rarely leaves somewhat hyaline distally; laminal areolation typically heterogenous, distal cells elongate, vermicular, hexagonal, or rhomboidal, thin- to thick-walled, proximal cells quadrate to regularly short- to long-rectangular, alar cells not differentiated; sporophyte terminal; peristome double; seta rarely somewhat twisted, not geniculate.

8. Distal lamina cells 3--6:1, longer than the quadrate or short-rectangular proximal cells; stems gemmiform or elongate and evenly foliate; leaves imbricate, not contorted or twisted when dry, or if somewhat twisted then rhizoidal tubers present; limbidium absent or weak, 1-stratose;  rhizoidal tubers and leaf axil bulbils often present.

9. Plants small, stems mostly less than 1 cm, gemmiform to evenly foliate; leaves 0.5--2.5 mm; leaf axil bulbils often present, tubers if present on long rhizoids in substratum or at base of stem, often abundant; capsules ovate to pyriform, apophysis sometimes inflated and rugose 

…………………………………... 4. Gemmabryum, p. XX

9. Plants medium-sized, stems 1--3 cm, evenly foliate; leaves 1--3.5 mm; tubers if present on micronemata or macronemata on stem, scarce, sometimes absent; capsules pyriform, neck slender …….………6. Imbribryum, p. XX

8. Distal lamina cells mostly 2--4:1, the same length or shorter than short- to long-rectangular proximal cells; stems comose, to evenly foliate; leaves twisted to strongly contorted when dry; limbidium usually present, often strong, 1- or 2-stratose; rhizoidal tubers and leaf axil filiform gemmae, sometimes present, bulbils absent.

10. Leaves ovate to obovate, distal margins serrulate to serrate or rarely nearly smooth, limbidium 1-stratose; rhizoidal tubers usually present, filiform leaf axil gemmae often present; dioicous  ……………………….12. Rosulabryum (in part), p. XX

10. Leaves ovate-lanceolate to ovate; distal leaf margins serrulate to smooth, limbidium 1- or 2-stratose; rhizoidal tubers absent, leaf axil filiform gemmae rare; dioicous or monoicous …………….……10. Ptychostomum, p. XX

7. Plants green to silver, pink-silver, pink-green or red-brown; laminal areolation homogeneous, lax, cells very long, thin walled, 4--8:1, cells near apex sometimes short or rarely irregularly quadrate, alar cells similar to juxtacostal cells or sometimes differentiated, in small quadrate groups; sporophytes sometimes appearing lateral; peristome double, single or absent; seta often twisted or geniculate.

11. Sporophytes appearing lateral; capsules short pyriform to ovate; apophysis short; peristome of exostome teeth only or absent …….…………………………………..… 5. Haplodontium, p. XX

11. Sporophytes terminal; capsules pyriform to distinctly zygomorphic; apophysis often very long;  peristome double.

12. Capsules zygomorphic; seta often geniculate; spores shed as tetrads; plants red-brown to silver or silver-pink; rhizoidal tubers absent …………… 9. Plagiobryum, p. XX

12. Capsules not zygomorphic; seta straight or somewhat twisted but not geniculate; spores shed singly; plants green or pink-green, not silvery; rhizoidal tubers often present.

13. Rhizoidal tubers small, pyriform, brown, 40--60 \um; lamina cells long and narrow, less than 15 \um

wide, alar cells quadrate in small group ……………4. Gemmabryum (in part), p. XX

13. Rhizoidal tubers large, spherical, red, often greater than 200 \um, or absent; lamina cells long and wide, typically greater than 20 \um wide, alar cells not quadrate ……… 8. Plagiobryoides, p. XX

 

1. ANOMOBRYUM Schimper, Syn. Musc. Eur., 382. 1860 * [Greek; anomo, lawless, and bryum, a moss alluding to the somewhat hypnaceous distal laminal cells]

 

John R. Spence

 

Plants acrocarpous; small, gregarious or in thin mats, pale green to yellow-green. Stems short, 0.5--2.5 cm, weakly to strongly julaceous, often appearing string-like, not or weakly branched, stolons absent; rhizoids scarce, micronemata and macronemata absent or present in clusters on proximal stem. Leaves imbricate dry, erect when wet, broadly ovate to ovate-lanceolate, flat to concave, 0.3--1.5 mm, basal insertion straight or slightly curved, not decurrent; margins plane distally, recurved proximally or often plane throughout, 1-stratose, ± nearly smooth, limbidium absent, apex broadly rounded to acute, sometimes slightly hyaline; costa not reaching apex or very short-excurrent in a smooth point, transverse section with single stereid band, usually well developed, guide cells absent; adaxial supracostal cells irregularly to regularly quadrate or short-rectangular at base; laminal cells distinctly heterogenous, proximal cells usually quadrate or short-rectangular, 1--2:1, medial cells generally similar to distal cells, distal cells elongate-hexagonal to vermicular, 6--10:1, not in rows oblique to the costa, thin- to thick-walled, walls not pitted, alar cells not differentiated from juxtacostal cells. Specialized asexual reproduction by small leafy bulbils in leaf axils. Sexual condition dioicous; perigonia and perichaetia terminal, leaves the same size as vegetative leaves or typically larger, not forming a rosette, inner leaves little differentiated. Seta 1, straight to somewhat twisted. Capsule inclined to erect, 2--4 mm, elongate-pyriform, ovate or cylindric, hypophysis somewhat differentiated, exothecial cells near mouth quadrate or short-rectangular, thick-walled, reddish, in 1--2 rows, proximal cells longer, irregularly  long-rectangular with straight or sinuose walls; annulus usually present, revoluble; operculum weakly convex, short-conic, not rostrate; peristome double, single or rarely absent, exostome pale yellow or tan, teeth slender lanceolate, trabeculate, lacking pores along fissural line, endostome hyaline to pale yellow, separate from exostome, or sometimes adherent or absent, segments present or absent, basal membrane low to high, cilia present or sometimes absent. Calyptra fugacious, cucullate, small, smooth. Spores shed singly, not as tetrads, not germinating in capsule, 8--15 \um, finely papillose, pale tan or yellow-tan.

 

Species ca. 30 (2 in the flora). Worldwide.

 

Anomobryum is distributed in temperate to alpine regions of the world, with concentrations in the mountains of Mexico and Central and South America. Although morphologically similar to Bryum, molecular studies consistently show that the types of the two genera, A. julaceum and B. argenteum, are not closely related. Species of Anomobryum can generally be distinguished from Bryum by their elongate, very slender stems, suggestive of pieces of string, and extremely long distal laminal cells. Bryum species tend to have thicker, more rounded stems that are relatively short, and for the most part shorter distal laminal cells  Additional studies are needed with a larger sample of species to confirm the molecular results.

 

SELECTED REFERENCES: Andrews, A. L. 1935. Bryaceae. In: Grout, A. J. Moss flora of North America, Vol. 2. Newfane, Vermont. Holyoak, D.T. and H. Köckinger. A taxonomic revision of some European and Asian bulbiferous species of Anomobryum (Bryophyta: Bryaceae). J. Bryol. 32: 153--169. Ochi, H. 1980. A revision of the Neotropical Bryoideae, Part 1. J. Faculty Educ. Tottori Univ., Nat. Sci. 29: 49--154.  Pedersen, N., C. J. Cox, and L. Hedenäs. 2003. Phylogeny of the moss family Bryaceae inferred from chloroplast DNA sequences and morphology. Syst. Bot. 28: 471--482.  Shaw, J. and A. J. Fife. 1984. The evolutionary and taxonomic significance of peristome morphology in Anomobryum (Bryaceae, Musci). J. Hattori Bot. Lab. 57: 285--298. Spence, J. R. and H. P. Ramsay. 2002. The genus Anomobryum Schimp. (Bryopsida, Bryaceae) in Australia. Telopea 9: 777--792. 

 

 

1. Stems weakly and loosely julaceous; leaves flat to weakly concave, costa variable but in at least some leaves percurrent to short-excurrent; bulbils sometimes present; capsule erect or suberect, peristome reduced, cilia absent …………….………………………….… 1. Anomobryum concinnatum

1. Stems strongly julaceous; leaves strongly concave, costa typically reaching midleaf or 2/3\x lamina length, rarely percurrent; bulbils lacking; capsule nodding, peristome perfect ………………...............................………………..……… 2.  Anomobryum julaceum

 

 

1. Anomobryum concinnatum (Spruce) Lindberg, Öfvers Förh. Kongl. Svenska Vetensk.-Akad. 18: 277. 1861

 

Bryum concinnatum Spruce, Musci Pyren., 121. 1847; Anomobryum julaceum var. concinnatum (Spruce) J. E. Zetterstatt; A. leptostomoides Schimper

 

Plants in thin mats or scattered, pale green to yellow-green. Stems  0.5--1.5 cm, brown to orange, weakly julaceous. Leaves strongly imbricate dry, broadly ovate or ovate-lanceolate, flat to somewhat concave, 0.6--1.4 mm, margins smooth, apex broadly acute, not hyaline, costa percurrent to short-excurrent as smooth point, laminal distal cells elongate-rhomboidal, 40--60 \um, 6:1, thick-walled, not or weakly sinuose. Specialized asexual reproduction of small red-brown leafy bulbils in axils of leaves. Seta 2--3 cm, flexuose. Capsule erect or suberect, red, 1--3 mm,  ovate-cylindric; peristome single, highly reduced, of unknown origin, segments and cilia lacking. Spores 8--15 \um.

 

Capsules very rare, maturing July--September. Locally common on acidic seepy or damp soil or soil over rock, often on ledges; 0--3500 m; Greenland; B.C., N.B., Nfld. & Labr., N.W.T., Que.; Alaska, Calif., Colo., Maine, Mich., Minn., Mo., Mont., N.Y., N.C., Tenn., Utah, Va., Wis.; Mexico; Eurasia.

 

Anomobryum concinnatum is a widespread boreal-temperate species identical to material of the synonym A. leptsostomoides in gametophyte features.  Specimens assigned to the latter name are few and produce erect capsules with highly reduced peristomes.  I have interpreted these as fertile specimens of A. concinnatum, a name with priority.  Because this species is rarely separated from A. julaceum by other workers, the distributions of the two in North America are tentative and need further study. Of the two, A. concinnatum is the more widespread. A recent collection from Utah has erect capsules and leaves with a short-excurrent costa, and in other respects is similar to Mexican material described as A. julaceum var. mexicanun Schimper.  More studies are needed to determine whether this is a good taxon; it is not recognized in this treatment.

 

2. Anomobryum julaceum (Schrader ex P. Gaertner, B. Meyer & Schreber) Schimper, Syn. Musc. Eur., 383. 1860

 

Bryum julaceum (Schrader ex P. Gaertner, B. Meyer & Schreber, Oekon. Fl. Wetterau 3(2): 97. 1802; Pohlia filiformis (Dickson) Andrews

 

Plants in thin mats or scattered, pale green to yellow-green. Stems  0.5--1.5 cm, brown to orange, strongly julaceous. Leaves loosely imbricate dry, broadly ovate or ovate-lanceolate, distinctly concave, 0.6--1.2 mm, margins smooth, apex broadly acute, not hyaline, costa not reaching apex, laminal distal cells elongate-vermicular, 60--100 \um, 8--10:1, thick-walled, sinuose. Specialized asexual reproduction absent. Seta 2--3 cm, flexuose. Capsule inclined to nodding, brown or red-brown, 1--2 mm, ovate-pyriform; peristome double, exostome pale brown basally, hyaline distally, endostome well developed, high, segments broadly perforated, cilia 1--3, appendiculate to nodose. Spores 8--13 \um.

 

Caspules very rare, maturing June--September. Locally common on acidic seepy or damp soil or soil over rock, often on ledges; 0--4000 m; Greenland; Alta., B.C., N.W.T., Nunavut, Yukon; Alaska, Calif., Colo., Conn., Mich., Minn., N.C., N.Y., Pa., Wis.; Mexico; Central America; South America; Eurasia; Africa; Subantarctic Islands.

 

Anomobryum julaceum is a widespread boreal-temperate species. Material from southern California and Mexico has been referred to the var. mexicanum Schimper by past workers, but I do not recognize this variety (see above).

 

2. BRACHYMENIUM Schwägrichen, Sp. Musc. Frond. Suppl. 2(1): 131. 1824 * [Latin brachy, short, and menium, a membrane, alluding to poor development of endostome]

John R. Spence

 

Plants small, forming dense turfs or cushions, green to red-green. Stems short, 0.3--1.5 cm, sometimes rosulate, sparsely to strongly branching by subfloral innovations, rhizoids sparse to abundant, variously colored, smooth to papillose, micronemata often present on stems. Leaves irregularly contorted to spirally twisted around stem when dry, erect-spreading when wet, broadly ovate, 1--2.5 mm, smaller proximally, leaf base not curved at insertion, not or only weakly decurrent; margins plane or revolute proximally, 1-stratose, limbidium absent or present, distal margins serrulate to serrate, apex acute to broadly rounded, differentiated apiculus absent, costa shortly to moderately long-excurrent in a stout, denticulate, colored awn, transverse section with stereid band single and well developed, guide cells present; adaxial supracostal cells irregularly to regularly quadrate or short-rectangular at leaf base, laminal cells heterogeneous, distal and medial cells generally similar, short-rectangular to irregularly hexagonal, 1.5--3(--4):1, not in rows oblique to the costa, thin- to somewhat thick-walled, proximal cells usually quadrate, distinctly different in shape, walls not pitted, alar cells usually similar to juxtacostal cells. Specialized asexual reproduction by spherical rhizoidal tubers and bulbils in leaf axils. Sexual condition dioicous; perigonia and perichaetia terminal, perichaetial leaves same size as vegetative leaves or typically larger, sometimes forming a rosette, inner leaves highly differentiated, often narrower with a weaker costa. Seta single, variously colored, elongate, straight. Capsule erect to suberect, ovate to cylindric, wrinkled when mature, hypophysis slender to well-differentiated, often rugose, operculum short to tall-conic; peristome double, exostome red-brown proximally, pale distally, papillose, teeth linear-lanceolate, separated at base, endostome of basal membrane hyaline to pale yellow, low, segments absent or short and blunt, cilia rudimentary or absent. Spores shed singly, 12--18 \um, smooth or papillose, pale tan to brown.

 

Species ca. 30 (2 in the flora): worldwide in tropical and subtropical regions, primarily Paleotropical.

 

The genus Brachymenium was originally based on the erect capsule with a variously reduced peristome, but recent research indicates the genus to be highly polyphyletic.  Apparently this sporophytic syndrome has evolved independently several times in groups with distinctive gametophytes.  I have used gameophyte morphology as the basis for re-assigning sections and species to other genera. Typical Brachymenium consists of subtropical-tropical epiphytes with gametophytes similar to those of Rosulabryum. Brachymenium andersonii has been re-assigned to Rosulabryum, while the small species B. exile Dozy & Molkenboer is now placed in Gemmabryum near the G. bicolor complex.  Brachymenium niveum and B. systylium have been transferred to the newly erected genus Leptostomopsis, of which the latter is the type. The only remaining North American species, B. macrocarpum and B. mexicanum, are probably not closely related to the type section of Brachymenium.  H. Ochi (1980) placed B. macrocaprum in sect. Peromnium, along with B. regenellii Hampe and B. jamesonii Taylor. This group is similar to some species of Gemmabryum, especially the Paleotropical G. coarctatum (Bosch & Sande Lacoste) J. R. Spence & H. A. Ramsay, but differ in the presence of bulbils and strongly rosulate and twisted leaves.  Because of uncertainty in the phylogenetic relationships of the section, I have tentatively retained B. macrocarpum in Brachymenium pending further study. The species B. mexicanum seems to be related to B. macrocarpum, although also showing similarities to species of the type section. The genus description above is largely based on Brachymenium sect. Peromnium (Mitten) Brotherus.

 

SELECTED REFERENCES: Ochi, H. 1980. A revision of the Neotropical Bryoideae, Part 1. J. Faculty Educ. Tottori Univ., Nat. Sci. 29: 49--154.  Spence, J. R. 2006. New combinations in the Bryaceae (Bryophyta) for North America. II. Phytologia 89: 110--114.

 

 

1. Leaves strongly spirally twisted around stem when dry, limbidium present; large red spherical rhizoidal tubers present in sterile material ......................................................................1. Brachymenium macrocarpum

1. Leaves imbricate to irregularly contorted when dry, limbidium absent; tubers lacking ....................................................................................................................2. Brachymenium mexicanum

 

 

1. Brachymenium macrocarpum Cardot, Rev. Bryol. 38: 6. 1911

 

Plants small, forming dense dark green to olive green cushions, not shiny. Stems 1--2 cm, strongly rosulate, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller.  Leaves ovate, 0.6--1.5 mm,  spirally twisted around stem when dry, forming rosettes when wet, somewhat concave, margins weakly revolute proximally, plane distally, distinctly serrate, limbidium usually present, of 1--2 rows of somewhat elongate incrassate cells, apex broadly rounded-acute; costa short-excurrent into a short stout somewhat yellowish-hyaline denticulate hairpoint; distal and median laminal cells hexagonal to short-rectangular, 20--50 x 15--20 \um, 2--3:1, proximal cells quadrate,  Specialized asexual reproduction of  rhizoidal tubers, spherical, 100--300 \um, red to orange-red, commonly on rhizoids at base of stem. Sexual condition dioicous. Seta 1--2 cm, light brown to red-brown. Capsule 1--3 mm, elongate-ovate to subcylindric, distinctly wrinkled when dry.

 

Capsules mature spring--summer (Apr--Aug). Uncommon on tree trunks, wood and occasionally rock, rarely on mineral soil; 0--1000 m; Ariz., Fla., La., N.Mex., N.C., Tex; Mexico.

 

Brachymenium macrocarpum is a distinctive species that, within its range, should be easy to identify. It forms dense turfs or cushions on trees and rocks.  The very similar B. klotzschii (Schwägrichen) Paris is found to the south in Central and South America.  Many workers lump the two, but there are differences in capsule morphology, leaf shape, lamina areolation, and asexual reproduction, so I have retained the name B. macrocarpum here, based on the Mexican type.  In Mexico, the species is known to produce bulbils in leaf axils, but I have not seen these in material from the flora area. Rosulabryum andicola is similar and its range overlaps with B. macrocarpum, but it has strongly serrate upper margins with very strong limbidium, filiform gemmae in leaf axils, and nodding capsules with a well developed peristome.

 

 

2. Brachymenium mexicanum Montagne, Ann. des Sci. Natur., Bot., sér. 2, 9: 54. 1838

 

Plants in open to dense turfs on soil or rock, green, yellow-green, to golden, often shiny-lustrous. Stems 0.5--1(--2) cm, gemmiform to somewhat rosulate, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller. Leaves 1--2(--3) mm, strongly imbricate, slightly twisted when dry, concave, not decurrent, apex acute, margins plane to weakly recurved proximally, distal margins smooth to serrulate, limbidium absent, costa strong, short to long-excurrent in an awn, distal and mid-laminal cells rhomboidal to elongate-hexagonal, 40--60 x 10--16 \um, mostly 3--4:1, moderately incrassate, proximal laminal cells abruptly quadrate to short-rectangular, 1--2:1, wider and shorter than cells above. Specialized asexual reproduction lacking.  Sexuality dioicous. [Capsule 2--3(--4) mm, erect, pyriform to cylindric with somewhat thickened and rugose apophysis; peristome double, reduced, endsotome low, segments and cilia absent or rudimentary.]

 

Capsules unknown in flora region. Rare on dry rock outcrops or soil covered crevices, 1000 m;  AZ; Mexico, Central America.

 

Brachymenium mexicanum is distinguished from B. macrocarpum by the more or less imbricate  leaves, lack of a limbidium, and absence of rhizoidal tubers. The sporophyte description is based on Mexican material.

 

3. BRYUM Hedwig, Sp. Musc. Frond., 178. 1801 * [Greek bryum, a moss]

John R. Spence

 

Plants small, gregarious or in dense mats or rarely cushions, white, silver-green, olive to pale green, golden, yellow-green, brown or yellow-brown. Stems 0.2--1(--1.5) cm, weakly to strongly rounded julaceous or short-gemmiform, not or weakly branched, innovations smaller than fertile stems, strongly julaceous; stolons absent; rhizoids scarce or common, micronemata and macronemata absent from stems or present in clusters proximally. Leaves dimorphic, fertile stem leaves imbricate dry, erect to erect-spreading when wet, ovate-lanceolate to ovate, flat or concave, 0.2--1(--1.5) mm, base straight or slightly curved at insertion, not decurrent, innovation leaves similar but smaller than fertile stem leaves; margins plane distally, recurved proximally or often plane throughout, 1-stratose, more or less smooth, limbidium absent, apex obtuse to acute or rarely acuminate, sometimes hyaline, costa not reaching apex or excurrent in smooth often hyaline point, apiculus sometimes present if costa not reaching apex, costa in transverse section with stereid band single, usually well developed, guide cells typically absent; adaxial supracostal cells irregularly to regularly quadrate or short-rectangular at base, laminal cells distinctly heterogeneous, distal cells short-rhomboidal to elongate-hexagonal or rarely vermicular, (2--)3--6(--8):1, not in rows oblique to the costa, thin-walled to incrassate, proximal cells usually quadrate or short-rectangular, 1--2(--3):1, wider than more distal cells, medial cells generally similar to distal cells, walls not pitted, alar cells not or somewhat differentiated from juxtacostal cells, usually shorter. Specialized asexual reproduction by small leafy bulbils in axils of leaves, rarely of rhizoidal tubers. Sexual condition dioicous, perigonia and perichaetia terminal, leaves the same size as vegetative leaves or typically larger, not forming a rosette, inner leaves not much differentiated. Seta 1, straight to somewhat twisted. Capsule inclined to erect, small, (1--)2--3 mm, elongate-pyriform or ovate, hypophysis differentiated or not, sometimes thickened and rugose, exothecial cells near mouth quadrate or short-rectangular, thick walled, reddish, in 1--2 rows, more proximal cells longer, irregularly  long-rectangular with straight or sinuose walls; annulus usually present, revoluble; operculum weakly convex, short-conic, not rostrate; peristome double, exostome pale yellow or tan, teeth slender lanceolate, trabeculate, lacking pores along fissural line, endostome hyaline to pale yellow, separate from exostome, or sometimes adherent or absent, segments present, basal membrane low to high, cilia present, appendiculate, or sometimes absent; calpytra fugacious, cucullate, small, smooth. Spores shed singly, not as tetrads, not germinating in capsule, 8--15(--18) \um, finely papillose, pale tan or yellow-tan.

 

Species 40 (7 in the flora): temperate to Arctic-alpine regions worldwide, and in mountains of subtropical or tropical regions.

 

Bryum in the strict sense is a genus of tiny julaceous plants, morphologically similar to Anomobryum. Molecular studies, however, consistently show that the types of the two genera, A. julaceum and B. argenteum, are not closely related. Species of Anomobryum can generally be distinguished from Bryum by their vermicular distal laminal cells, and elongate very slender, string-like stems. Bryum species tend to have shorter laminal cells and thicker more rounded stems that are relatively short.  Additional studies are needed with a larger sample of species to confirm the molecular results. Bryum consists of both silver-white species lacking chlorophyll in the distal portions of the leaves, and green to yellow-green species.  Many collections are difficult to identify as they often consist of very small sterile shoots. There are at least three undescribed species in the flora area, two of which are discussed under related species. The third is highly distinctive and has been collected recently from southeastern Arizona.

 

SELECTED REFERENCES  Ochi, H. 1980. A revision of the Neotropical Bryoideae, Part 1. J. Faculty Educ. Tottori Univ., Nat. Sci. 29: 49--154.  Shaw, A. J. 1981. The taxonomy of Bryum oblongum and B. blindii. Canad. J. Bot. 59: 1426--1435.  Frahm, J.-P. 2002. The taxonomic status of Bryum arachnoideum C. Müll. and B. lanatum (P. Beauv.) Brid. Trop. Bryol. 21: 53--56. Pedersen, N., C. J. Cox and L. Hedenäs. 2003. Phylogeny of the moss family Bryaceae inferred from chloroplast DNA sequences and morphology. Syst. Bot. 28: 471--482. Spence, J. R. 1986. Bryum calobryoides, a new species from western North America. Bryologist 89: 215--218. Spence, J. R. 2007. Four species of the Bryaceae new to the U.S.A. Evansia 24: 29--30. Spence, J. R. and H. P. Ramsay. 2002. The genus Anomobryum Schimp. (Bryopsida, Bryaceae) in Australia. Telopea 9: 777--792. Toren, D. and K. Heise. 2009. Bryum chryseum Mitt. (Musci: Bryaceae) new to North America north of Mexico. Evansia 98--101.

 

1. Plants silver-green or white, distal lamina hyaline, including apiculus or awn, distal laminal cells hexagonal to rhomboidal, less than 15 \um wide.

2. Plants hoary white, stems evenly foliate, slender, costa strong, excurrent in stout awn, somewhat recurved when dry; leaf axis bulbils lacking …….…………...… 5. Bryum lanatum

2. Plants silver to silver-green, stems julaceous to gemmiform, costa not reaching apex, apiculus usually present or sometimes absent, irregularly incurved when dry; leaf axis bulbils present ...................................................................................................................... 1. Bryum argenteum

1. Plants yellow-green, golden, olive-green or bright green, distal lamina not hyaline although apiculus if present sometimes hyaline, distal lamina cells variable, elongate vermicular to short and rhomboidal, 6--25 \um wide.

3. Distal laminal cells of innovation leaves elongate-vermicular, more than 6:1, and narrow, less than 8 \um; bulbils sometimes present in leaf axils; leaf base typically red ........…………..…................... 2. Bryum blindii

3. Distal laminal cells of innovation leaves not vermicular, shorter and broader, 2--4:1, mostly more than 8 \um wide; bulbils absent; leaf base red or green.

4. Plants golden or yellow-green; leaves triangular-ovate, costa strong, excurrent in long awn, erect-spreading when dry .............................................................................. 4. Bryum chryseum

4. Plants yellow-green, olive-green or bright green; leaves ovate to ovate-lanceolate, costa not reaching apex or rarely excurrent in a short stout awn.

5. Plants yellow-green, laminal cells of fertile stem leaves elongate-hexagonal, 4--5(--6):1; leaf base red .................…………… 6. Bryum oblongum

5. Plants olive green to bright green, laminal cells of stem leaves shorter, 2--3(--4):1, irregularly rhomoboidal to hexagonal; leaf base green.

6. Leaf apex acute, not cucullate, apiculus lacking or composed of 1--2 cells, distal laminal cells regularly hexagonal, 3--4:1, 10--16 \um wide, basal cells predominantly rectangular in alar region, 2--3:1 ................................ 7. Bryum veronense

6. Leaf apex rounded-obtuse to acute, cucullate, apiculus often present, generally of more than 2 cells, distal laminal cells short and irregularly rhomboidal, mostly 2--3:1, 12--25 \um wide, basal cells predominantly quadrate in alar region ....... 3. Bryum calobryoides

 

 

1. Bryum argenteum Hedwig, Spec. Musc. Frond., 181. 1801

 

Plants gregarious or in dense mats, white-green to silver-green. Stems 0.2--1 cm, strongly rounded julaceous or sometimes gemmiform Leaves imbricate dry, erect when wet, ovate to ovate-lanceolate, somewhat concave, 0.3--1(--1.2) mm; margins plane distally or rarely recurved proximally, apex broadly rounded to acute,  hyaline in distal 1/4--1/2 of leaf or rarely nearly entirely green, costa not reaching apex, hyaline apiculus usually present, sometimes short or nearly absent; distal laminal cells elongate-hexagonal, (30--)40--80 x 10--15, 3--5:1, thin walled or somewhat thickened but not distinctly incrassate, proximal cells usually quadrate or short-rectangular, 1--2:1, 12--18 \um wide. Specialized asexual reproduction by small leafy bulbils in axils of leaves.  Seta elongate, brown to red-brown, 1--2 cm, somewhat twisted when dry. Capsule inclined to pendant, 2--3 mm, ovate, red to red-brown, hypophysis differentiated, somewhat thickened. Spores 8--15 \um.

 

1. Stems loosely julaceous, leaves somewhat distant, green except for tips, apiculus long ....................................................... 1b. Bryum argenteum var. majus

1. Stems densely julaceous or sometimes short and gemmiform, leaves silver-green, hyaline in proximal 1/4--1/2 of leaf, apiculus short or long.

2. Plants small, stems mostly less than 2 mm, sometimes gemmiform, leaves often wider than broad, distal lamina cells relatively short, (2--)3--4:1, apiculus very short or sometimes absent ................................................................................................1c. Bryum argenteum var. muticum

2. Plants larger, stems 2--10 mm, julaceous, leaves usually longer than broad, distal lamina cells elongate, (3--)4--6:1, apiculus present, generally well developed  ...............................................................................................1a. Bryum argenteum var. argenteum

 

Bryum argenteum is perhaps the most widespread and distinctive moss in the world, although in many regions it is found primarily in urban or other heavily disturbed sites, and is mostly absent from native undisturbed habitats in the tropics.  There are about 15 silver-white species described, with species definitions based on a combination of one of  three or four different basic gametophyte morphologies combined with subtle differences in the sporophytes.  However, much of the gametophyte variability in the complex is also exhibited by B. argenteum itself, thus confusing species-level identifications. This complex is urgently in need of world-wide molecular and morphological studies.

 

1a. Bryum argenteum Hedwig var. argenteum

 

Plants silvery-green. Stems distinctly rounded julaceous, 0.5--1.0 cm. Leaves strongly imbricate, ovate to ovate-lanceolate, longer than broad, hyaline in distal 1/3--1/4 of leaf, apiculus well developed; distal laminal cells elongate-hexagonal, 3--4(--5):1.

 

Capsules mature spring--summer (Apr--Jul). Common on soil or soil over rock or in crevices, often in nitrogen enriched sites, usually in disturbed habitats; 0--4000 m; Greenland; St. Pierre and Miquelon; Alta., B.C., Man., N.B., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont., P.E.I., Que., Sask., Yukon; Ala., Alaska, Ariz., Ark., Calif., Colo., Conn., Del., D.C., Fla., Ga., Idaho, Ill., Ind., Iowa, Kans., Ky., La., Maine, Md., Mass., Mich., Minn., Miss., Mo., Mont., Nebr., Nev., N.H., N.J., N.Mex., N.Y., N.C., N.Dak., Ohio, Okla., Oreg., Pa., R.I., S.C., S.Dak., Tenn., Tex., Utah, Vt., Va., Wash., W.Va., Wis., Wyo.; Mexico; West Indies; Central America; South America; Eurasia; Africa; Atlantic Islands (Bermuda); Indian Ocean Islands; Pacific Islands (New Zealand); Australia; Antarctica.

 

1b. Bryum argenteum var. majus Schwägrichen, Sp. Musc. Frond., Suppl. 1, 2: 88. 1816

 

Plants greenish. Stems julaceous, 0.5--1 cm. Leaves loosely imbricate, ovate to ovate-lanceolate, longer than broad, hyaline near the leaf tip, apiculus well developed, long; distal laminal cells elongate-hexagonal, 3--4:1.

 

Capsules unknown. Rare, shaded moist soil, disturbed habitats; 0--500 m; N.H., Maine, Mo; w Europe.

 

Bryum argenteum var. majus is poorly understood, but seems distinctive. It may be more widely distributed.  The pale green loosely imbricate stems with hyaline apiculus are reminiscent of Plagiobrym zierii, but that species has wide elongate laminal cells throughout the leaf, a pinkish coloration, and Arctic-alpine distribution.

 

1c. Bryum argenteum var. muticum Bridel, Bryol. Univ. 1: 846. 1827

 

Plants hoary white to silver-green. Stems gemmiform to shortly julaceous, 0.2--0.8 cm. Leaves ovate to ovate-lanceolate, often broader than long, hyaline in distal 1/2--1/4 of leaf, apiculus absent or very short, composed of 1--2 cells; distal laminal cells short-hexagonal, (1--)2--3:1, sometimes a few quadrate cells present.

 

Capsules unknown. Common, dry soil, soil over rock, crevices, lowland deserts to dry alpine tundra; 0--4,000 m; Ariz., Calif., Colo., Nev., N.Mex., Utah.

 

The var. muticum is widespread in extreme environments, but the world distribution is not well known.  Similar material from Antarctica and Australasia has been called B. subrotundifolium A. Jaeger.

 

2. Bryum blindii Bruch, Schimper & W. Gümbel, Bryol. Eur. 4: 163. 1846

 

Plants gregarious, green to yellow-green. Stems 0.2--1 cm, gemmiform to strongly rounded-julaceous. Leaves imbricate dry, erect when wet, ovate, somewhat concave, 0.3--1(--1.2) mm; margins plane, apex broadly acute, not hyaline in distal portion of leaf, costa not reaching apex,  apiculus absent; distal laminal cells of main (fertile) stem and innovation leaves linear-vermicular to elongate-hexagonal, (60--)70--90(--100) x 6--10(--12), (6--)8--10:1, with yellowish thick walls, proximal cells usually quadrate or short-rectangular, 1--2:1, 8--12 \um wide. Specialized asexual reproduction by small leafy bulbils in axils of leaves.  Seta elongate, red to red-brown, 1--2 cm, somewhat twisted when dry. Capsule inclined to pendant, 1--2 mm, ovate, red to red-brown, hypophysis differentiated, somewhat thickened and rugose when dry. Spores 10--18 \um.

 

Capsules mature summer (June--Aug). Rare, calcareous mineral soil, soil banks, cold-temperate to arctic-alpine regions; 0--3500 m; Greenland; B.C., N.W.T., Nunavut, Ont., Yukon; Cal., Colo.,  Maine, Mont.; n Eurasia.

 

One of the two collections of Bryum blindii  known from Colorado has leaves with a strongly excurrent costa and relatively short distal laminal cells (3--5:1), and is not typical material of the species.

 

3. Bryum calobryoides J. R. Spence, Bryologist 89: 215--218, fig. 1--8. 1986

 

Plants gregarious or in dense mats or cushions, bright green distally, brown proximally. Stems 0.5--1.5(--2) cm, strongly rounded julaceous. Leaves imbricate dry, erect when wet, ovate to suborbicular, concave, 0.2--1(--1.2) mm; margins plane distally, weakly recurved proximally, apex broadly rounded to acute,  not hyaline, costa not reaching apex to rarely percurrent, apiculus typically absent in distal leaves, present in proximal leaves, sometimes hyaline; distal laminal cells irregularly rhomboidal, (25--)30--75 x (8--)12--25, 1--4:1, thin walled or somewhat thickened but not distinctly incrassate, proximal cells usually quadrate or short-rectangular, 1--2:1, 12--18 \um wide. Specialized asexual reproduction lacking.  Sporophytes unknown.

 

Uncommon to locally common, primarily on calcareous damp soil, rocks; 1000--3000 m; Alta., B.C., Que.; Calif., Colo., Idaho, Mont., Oreg., Wash.  Endemic.

 

Bryum calobryoides is easily distinguished by small strongly julaceous stems, strongly concave leaves, and distal leaves broadly rounded and cucullate. An undescribed species from Colorado occurs in similar habitats, but has a short-excurrent costa, acute leaf apices, and unusual, small, irregularly pyriform, rhizoidal tubers with bulging cell walls. Bryum gerwigii (Müller Hal.) Limpricht has been reported from Alberta, but the material is B. calobryoides.  Bryum gerwigii is a local waterfall form of Gemmabryum gemmiparum from central Europe.

 

4. Bryum chryseum Mitten, J. Linn. Soc., Bot. 12: 304. 1869

 

Plants gregarious or in dense mats, shiny golden-yellow to yellow-green or rarely pale green. Stems 0.5--1 cm, slender, weakly julaceous, clusters of red rhizoids often present on stem. Leaves imbricate dry, erect when wet, triangular to ovate, concave, 0.5--1 mm; margins plane distally or rarely recurved proximally, apex acuminate to acute,  not hyaline, costa strong, long-excurrent in slender colored awn, recurved when dry; distal laminal cells elongate-hexagonal to vermicular, (50--)60--90 x 8--12, (4--)6--8:1, thin to somewhat firm walled but not strongly incrassate, proximal cells usually quadrate or short-rectangular, 1--2:1, 12--18 \um wide. Specialized asexual reproduction lacking.  Seta elongate, red, 1--2 cm, somewhat twisted when dry. Capsule inclined to pendant, 3--4 mm, clavate to pyriform, brown, hypophysis slender, not differentiated; exostome sometimes reduced, with cilia sometimes short or nearly absent. Spores (8--)10--15 \um (in Mexican material).

 

Capsules unknown in the flora area. Rare, soil, Mediterranean and other seasonally dry climates; 200--800 m; Calif.; Mexico; Central America; South America (Bolivia, Chile, Columbia, Peru).

 

Bryum chryseum was found in California in two locations in native plant communities by D. Toren and K. Heise. Bryum chryseum is a species characteristic of seasonally dry climates from Mexico to Chile. The small plant size, golden color, and triangular leaves with a strong excurrent costa are diagnostic. It is superficially very similar to B. lanatum, but lacks the hyaline lamina and hoary-white color of that species. Depauperate material from Arizona with somewhat hyaline awns collected by N. G. Miller from calcareous sandy soil near Wilcox may belong to this species rather than to the gametophytically similar Leptostomopsis nivea. The latter differs from B. chryseum in hyaline awns, serrate distal leaf margins, and numerous extremely incrassate and vermicular distal laminal cells.

 

5. Bryum lanatum (P. Beauvois) Bridel, Muscol. Recent. Suppl. 3: 20. 1817

 

Bryum argenteum var. lanatum (P. Beauvois) Hampe, Linnaea 13: 44. 1839

 

Plants gregarious or in dense mats, hoary white. Stems 0.5--1(--1.5) cm, weakly julaceous. Leaves imbricate dry, erect-spreading when wet, ovate to ovate-lanceolate, somewhat concave, 0.3--1(--1.5) mm; margins plane distally or recurved proximally, apex acute,  hyaline in distal 1/2 of leaf, costa excurrent in long slender hyaline awn, recurved when dry; distal laminal cells elongate-hexagonal, 40--60 x 10--16, 4--5:1, walls somewhat thickened to distinctly incrassate, proximal cells usually quadrate or short-rectangular, 1--2:1, 10--16 \um wide. Specialized asexual reproduction absent.  Seta elongate, brown to red-brown, 1--2 cm, somewhat twisted when dry. Capsule inclined to pendant, (1--)2--3 mm, ovate, brown to red-brown, hypophysis differentiated, somewhat thickened and rugose. Spores 8--15 \um.

 

Capsules rare, mature spring--summer (May--Jul). Common to abundant on soil, soil over rock or rock in drier climates; 0--4200 m.; Greenland; Alta., B.C., Man., Nfld. and Labr., N.W.T., N.S., Nunavut, Que., Sask., Yukon; Alaska, Ariz., Calif., Colo., Idaho, Kans., Mo., Mont., Nebr., Nev., N.Mex.,  N.Dak., Okla., Oreg., S.Dak., Tex., Utah, Wash., Wyo; Mexico; West Indies; Central America; South America; Eurasia; Africa; Atlantic Islands (Bermuda); Pacific Islands (Hawaii,  New Zealand); Australia; Antarctica.

 

Although typically considered a variety of Bryum argenteum, B. lanatum from western North America and other parts of the world is quite distinctive.  However, the type of B. lanatum, from Europe, may be a form of B. argenteum, in which case a new name will need to be found.  It is characteristic of undisturbed native plants communities, and is especially common in semi-arid parts of the West on dry soil.  Bryum argenteum generally grows in more moist climates in highly disturbed sites, especially where there is excess nitrogen. The distribution of B. lanatum in the eastern part of North America is not well understood, as this species has been lumped into B. argenteum by all eastern bryologists.

 

6. Bryum oblongum Lindberg, Bot. Not. 1882: 26. 1882

 

Plants gregarious, green to yellow-green. Stems 0.2--1 cm, gemmiform to strongly rounded-julaceous. Leaves imbricate dry, erect when wet, ovate, somewhat concave, 0.3--1(--1.2) mm; margins plane, apex broadly acute, not hyaline in distal portion of leaf, costa not reaching apex,  apiculus absent; distal laminal cells of main (fertile) stem leaves elongate-hexagonal, (30--)35--60(--70) x 6--12 , 4--5(--6):1, thin-walled, not yellowish, distal laminal cells of innovation leaves shorter, proximal cells usually quadrate or short-rectangular, 1--2:1, 8--12 \um wide. Specialized asexual reproduction by small leafy bulbils in axils of leaves.  Seta elongate, red to red-brown, 1--2 cm, somewhat twisted when dry. Capsule inclined to pendant, 1--2 mm, ovate, red to red-brown, hypophysis slender to weakly differentiated, sometimes slightly thickened and weakly rugose when dry. Spores 10--18 \um.

 

Capsules mature summer (June--Aug). Rare, neutral mineral soil or soil banks, boreal to Arctic-alpine regions; 0--1500 m; Greenland; B.C., Yukon; Alaska; n Eurasia.

 

Shorter laminal cells of the innovations leaves distinguish B. oblongum from the closely related B. blindii.  A. J. Shaw (1981) also enumerated other differences, including generally shorter exostome teeth in B. oblongum (300--400 \um) against. longer teeth in B. blindii (400--500 \um). Sterile material from Montana may belong to this species.  Recently, small, sterile, brownish, julaceous plants from alpine areas of New York have been collected that are similar to B. oblongum, but they have a strongly bulging reddish costa and shorter leaf cells.  These collections are likely to represent an undescribed species.

 

7. Bryum veronense De Notaris, Comment. Soc. Crittog. Ital. 2: 212 [112]. 1866

 

Plants in dense mats, bright to dark green or olive-green. Stems 0.4--1 cm, weakly julaceous to string-like. Leaves imbricate when dry, erect when wet, ovate, weakly concave, not cucullate, 0.2--1 mm; margins plane distally or rarely recurved proximally, apex acute, not hyaline, costa not reaching apex, apiculus lacking or a very short apiculus of 1--2 hyaline cells sometimes present; distal laminal cells hexagonal, (25--)30--55 x 10--16, mostly 3:1, thin walled to somewhat thickened but not distinctly incrassate, proximal cells usually short-rectangular, at least in alar region, 2--3:1, 12--18 \um wide. Specialized asexual reproduction lacking. Sporophyte unknown. 

 

Capsules unknown. Very rare, damp to dry soil, soil-filled crevices in sandstone; 1700 m.; Greenland; Ont., Que; Colo., Vt.; Europe.

 

Bryum veronense was first ollected from the Front Range of Colorado near Boulder by W.A. Weber and R. C. Wittmann, in soil-filled crevices in sandstone. The plants agree well with named European material, although they are a dark olive-green rather than the bright green of European material. Since then the species has been collected from Ontario, Quebec and Vermont. This species is similar to B. calobryoides, but lacks the strongly julaceous stems, strongly concave leaves, and cucullate apices of that species.

 

4. GEMMABRYUM J. R. Spence & H. P. Ramsay, Phytologia 87: 63. 2005 * [Latin gemma, bud, and bryum, a moss, alluding to asexual reproduction]

 

John R. Spence

 

Plants small, in open to dense turfs or gregarious on soil or rock, green, yellow-green, pink or red. Stems 0.1--2(--3) cm, gemmiform  to evenly foliate, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller, stems sometimes julaceous; rhizoids variously colored brown, red, violet, yellow or orange. Leaves 0.4--2.5(--3) mm, ovate, ovate-lanceolate to triangular, imbricate to loosely set and slightly twisted when dry, flat or concave, rarely decurrent, apex obtuse to acuminate; margins plane to strongly revolute, distal margins smooth to serrulate, limbidium absent or weak, of 1 layer of elongate incrassate cells; costa strong, percurrent to long-excurrent as an awn, occasionally an apiculus present, costa in cross-section generally with 1 layer of guide cells; distal and mid-laminal cells rhomboidal to elongate-hexagonal, 8--16(--20) \um wide, mostly (3--)4--8:1, thin to moderately incrassate, not porose, proximal laminal cells abruptly quadrate to rectangular, 1--2(--3):1, wider and shorter than more distal cells, inflated group of pink subalar cells rarely present. Specialized asexual reproduction common, of several types, including bulbils or fragile brood branchlets in leaf axils, rhizoidal filiform gemmae and tubers, and stem tubers.  Sexual condition dioicous or synoicous, rarely autoicous; perigonial and perichaetial leaves not much differentiated, outer leaves larger, inner leaves smaller than leaves of innovations. Seta elongate, stout to slender, straight or somewhat flexuose. Capsule 1--3(--5) mm, cylindric, pyriform to ovate, apophysis slender or often thick, sometimes inflated and rugose, erect, inclined or nutant; peristome double, variable, from well developed to reduced, endostome high to low, rarely adherent to exostome, segments generally well developed, with lanceolate to ovate perforations, occasionally reduced, cilia long and appendiculate to short or absent; operculum low conic to tall convex, sometimes apiculate or rarely rostrate. Spores small, smooth to papillose, 8--18(--20) \um.

 

Species ca. 100 (19 in the flora): worldwide in tropical, subtropical, temperate to boreal regions, on all continents; soil or rock.

 

Species of Gemmabryum are small and often sterile, and can be extremely difficult

to identify in the absence of gemmae.  This genus exhibits a remarkable array of asexual reproductive structures, including bulbils, rhizoidal tubers, stem tubers, and filiform rhizoidal gemmae. Sterile collections should be made whenever possible, as fertile material often lacks the diagnostic gemmae. Three major morphological groups exist:  (1) Small gemmiform to evenly foliate species in sect. Gemmabryum (Doliolidium and Dicranobryum sensu H. Ochi 1992) with strongly imbricate leaves and leaf axillary  bulbils; (2) Sect. Tuberibryum (Apalodictyon and Apiculata sensu H. Ochi 1992), with evenly foliate leaves that are often slightly twisted when dry , and which produce rhizoidal tubers; and (3) G. caespiticium and its allies (sect. Caespitibryum), including species with long excurrent awns and concave leaves. Included in the genus is Brachymenium section Dicranobryum, which includes species very similar to the other groups in the genus, especially those in sect. Gemmabryum, but having erect capsules with variously reduced peristomes.  The presence of reduced capsules and erect peristomes has been shown to evolve independently in many separate lineages in the Bryaceae (C. J. Cox and T. A. Hedderson.2003). that appear to converge on Bryum, while other species (e.g., G. subapiculatum) are more similar to Imbribryum. sect. Tuberibryum is intermediate in many ways between Imbribryum and sect. Gemmabryum. There are several undescribed species from California that are discussed under related species.

 

SELECTED REFERENCES Arts, T. 1992. Bryum demaretianum sp. nov., a new species of the B. erythrocarpum complex from Belgium. J. Bryol. 17: 263--267. Arts, T., A. C. Crundwell, and H. L. K. Whitehouse. 1995. Bryum valparaisense Thér. an earlier name for B. pyriferum Crundw. & Whiteh. J. Bryol. 18: 797--801.  Crundwell, A. C. and E. Nyholm. 1964. The European species of the Bryum erythrocarpum complex. Trans. Brit. Bryol. Soc. 4: 597--637. Nyholm, E. 1993. Illustrated flora of Nordic Mosses, Vol. 3. Nordic Bryol. Society, Copenhagen and Lund. Ochi, H. 1992. A revised infrageneric classification of the genus Bryum and related genera (Bryaceae, Musci). Bryobrothera 1: 231--244. Smith, A. J. E. 2004. The moss flora of Britain and Ireland, Ed. 2. Cambridge. Smith, A. J. E. and H. L. K. Whitehouse. 1978. An account of the British species of the Bryum bicolor complex including B. dunense sp. nov. J. Bryol. 10: 29--47. Cox, C. J. and T. A. Hedderson. 2003. Phylogenetic relationships within the moss family Bryaceae based on chloroplast DNA evidence. J. Bryol. 25: 31--40. Spence, J. R. 2005. New genera and combinations in Bryaceae (Bryales, Musci) for North America. Phytologia 87: 15--28. Spence, J. R. 2007. Four species of the Bryaceae new to the U.S.A. Evansia 24: 29--30. Spence, J. R. and H. P. Ramsay. 2005. New genera and combinations in the Bryaceae (Bryales, Musci) for Australia. Phytologia 87: 61--72. Spence, J. R. 2009. Nomenclatural changes in the Bryaceae (Bryopsida) for North America III. Phytologia 91: 493--499. Vanderpoorten, A. and C. E. Zartman. 2002. The Bryum bicolor complex in North America. Bryologist 105: 128--139. Whitehouse, H. L. K. 2001. Bryophytes of arable fields in Québec and Slovakia, including new records of Bryum demaretianum Arts. Lindbergia 26: 29--32. Wilczek, R. and F. Demaert. 1974. Les espèces belges du “complexe Bryum erythrocarpum.” Bull. Jard. Bot. Nat. Belg. 44: 425--438.

 

 

1. Plants with rhizoidal tubers commonly present, often abundant, at stem base in clusters or on long rhizoids in substrate; stems comose to evenly foliate, leaves mostly slightly twisted when dry.

2. Plants pale green to red-green, evenly foliate to strongly comose and caespitose, leaves with long-excurrent awns, sometimes hyaline; subalar group of inflated pink cells often present on leaves of fertile stems, brood branches sometimes present in leaf axils; limbidium often present . . . XXa. Gemmabryum sect. Caespitibryum (in part), p. xx

2. Plants green, pink or red, mostly evenly foliate, with costa not reaching apex to short-excurrent, if longer then rhizoidal tubers common and limbidium absent; subalar group of inflated pink cells absent; brood branchlets absent; limbidium usually absent  . . . XXc. Gemmabryum sect. Tuberibryum, p. xx

1. Plants with leaf axil bulbils, tubers rarely present, or gemmae absent; stems evenly foliate or gemmiform, often in two or more clumps along stem; leaves imbricate, not twisted when dry.

3. Plants lacking bulbils, with somewhat twisted or contorted leaves, costa long-excurrent, sometimes hyaline, subalar group of inflated pink cells often present, brood branchlets sometimes present in leaf axils; limbidium often present . . . XXa Gemmabryum sect. Caespitibryum (in part), p. xx

3. Plants with bulbils, leaves imbricate, costa not reaching apex to short-excurrent, subalar group of inflated pink cells absent, if bulbils lacking then costa percurrent to very short-excurrent as a stout awn, brood branchlets lacking; limbidium usually absent  . . . XXb. Gemmabryum sect. Gemmabryum, p. xx

 

 

4a. GEMMABRYUM sect. CAESPITIBRYUM (Podpĕra) J. R. Spence, Phytologia 91: 497. 2009

 

Bryum subsect. Caespitibryum Podpĕra, Rozpr. České Akad. Vĕd, TY. 2, Vĕdy Mat. PYír.) 10(2): 52. 1901

 

Stems gemmiform to evenly foliate, innovations and fertile stem leaves somewhat differentiated; leaves of innovations smaller, not julaceous; rhizoids brown to red-brown. Leaves imbricate to evenly foliate, costa excurrent in long awn, sometimes hyaline or spinose-denticulate; inflated group of pinkish subalar cells sometimes present on larger distal leaves of fertile stems. Specialized asexual reproduction by deciduous brood branchlets in leaf axils, rarely by rhizoidal tubers.  Sexual condition dioicous. Capsule 2--4(--5) mm, pyriform to subcylindric, apophysis  slender, gradually tapered to seta, not inflated and rugose, nutant; peristome double,  well developed, endostome high, segments well developed, with lanceolate to ovate perforations, cilia long and appendiculate.

 

Species ca. 6 (3 in the flora): primarily Northern Hemisphere boreal-temperate, although also common in Mediterranean climates, all continents except Antarctica.

 

Section Caespitibryum includes G. caespiticium and several narrowly defined segregates.  The taxonomy of the complex is poorly known, and future studies combining morphology and genetic data will be needed to sort out the species. I have placed the group in Gemmabryum, but without strong convictions.  In some respects the species are reminiscent of  species in Ptychostomum section Cladodium, especially to P. creberrimum.  Gemmabryum caespiticium is one of the most widespread mosses in the world.

 

1. Plants small, most leaves less than 1 mm; distal lamina in older leaves becoming hyaline; distal laminal cells short-rectangular to hexagonal, 3--4:1, proximal cells mostly quadrate to short-rectangular, 1--2:1; costa variable but mostly strong, at least some leaves with long-excurrent costa in slender green to hyaline hair-point; rhizoidal tubers  absent

.............................…… 3. Gemmabryum kunzei

1. Plants larger, most leaves 1--2.5 mm; distal lamina green; distal laminal cells long-hexagonal, 4--6:1, proximal cells short- to long-rectangular, 2--4:1 at least along the costa; costa strong, long-excurrent in brown, yellow-brown or sometimes hyaline awn; rhizoidal tubers  present or absent.

2. Leaves strongly concave, hair-point long, more than 1/2 length of lamina, often as long as lamina, denticulate or spinose; tubers absent ........................ 1. Gemmabryum badium

2. Leaves weakly concave, hair-point shorter, typically less than 1/2 length of lamina, smooth, rarely somewhat denticulate; small reddish spherical tubers sometimes present ...............…………… 2. Gemmabryum caespiticium

 

1. Gemmabryum badium (Bruch ex Bridel) J. R. Spence, Phytologia 91: 497. 2009

 

Bryum caespiticium var. badium Bruch ex Bridel, Bryol. Univ.: 850. 1827; B. badium (Bruch ex Bridel) Schimper; Ptychostomum badium (Bruch ex Bridel) J.R. Spence

 

Plants small, in open to dense turfs or gregarious on soil or rock, green, yellow-green, or red-green. Stems 0.1--2(--3) cm, gemmiform to evenly foliate, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller; rhizoids red-brown to red. Leaves 0.5--2(--3) mm, ovate-lanceolate, imbricate, somewhat twisted when dry, strongly concave, not decurrent, apex  acuminate, not becoming hyaline with age, margins plane to strongly revolute, distal margins smooth to serrulate, limbidium distinct to weak, of 1--2 rows of elongate incrassate cells; costa strong, long-excurrent in denticulate colored awn,  often more than 1/2--1/1 lamina length; distal and mid-laminal cells rhomboidal to hexagonal, 12--18(--22) \um wide, mostly (3--)4--5:1, thin to moderately incrassate, proximal laminal cells abruptly quadrate to short-rectangular, 1--2:1 away from costa, 2--4:1 along costa, wider and shorter than more distal cells, inflated group of pink subalar cells present on distal fertile stem leaves. Specialized asexual reproduction by deciduous brood branchlets in axils of distal stem leaves; rhizoidal tubers absent.  Sexual condition dioicous. [Capsule 2--3(--4) mm, pyriform, red-brown, segments bright to pale yellow. Spores small, smooth to papillose, 14--18 \um.]

 

Sporophytes not known in flora area. Rare on dry soil or rock in semi-arid climates; 1000--2000 m; Calif., Nev.; w Eurasia (including Middle East).

 

Gemmabryum badium is similar to G. caespiticium, but distinguished by much longer awns, more strongly concave leaves, and absence of rhizoidal tubers. Gemmabryum badium has only recently been recognized from collections in California and Nevada, and is likely to be found elsewhere in North America.

 

2. Gemmabryum caespiticium (Hedwig) J. R. Spence, Phytologia 91: 497. 2009

 

Bryum caespiticium Hedwig, Sp. Musc. Frond., 180. 1801; Ptychostomum imbricatulum (Müller Hal.) D. T. Holyoak & N. Pedersen

 

Plants small, in open to dense turfs or gregarious on soil or rock, rarely wood, green, yellow-green, or red-green. Stems 0.1--2(--3) cm, gemmiform to evenly foliate, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller; rhizoids red-brown to red. Leaves 0.5--2(--3) mm, ovate-lanceolate, imbricate, somewhat twisted when dry, weakly to moderately concave, not decurrent, apex  acuminate, occasionally becoming hyaline with age, margins plane to strongly revolute, distal margins smooth to serrulate, limbidium distinct to weak, of 1--2 rows of elongate incrassate cells; costa strong, excurrent in short to long, mostly smooth and colored awn,  from 1/4--3/4\x lamina length; distal and mid-laminal cells rhomboidal to hexagonal, 12--18(--20) \um wide, mostly (3--)4--6:1, thin to moderately incrassate, proximal laminal cells abruptly quadrate to rectangular, 1--2:1 away from costa, 2--4:1 along costa, wider and shorter than more distal cells, inflated group of pink subalar cells present on distal fertile stem leaves. Specialized asexual reproduction rare, of small red spherical rhizoidal tubers, 100--2000 \um, cells smooth and of deciduous brood branchlets in axils of distal stem leaves.  Sexual condition dioicous. Capsule 2--3(--4) mm, pyriform, red-brown, segments pale tan, brown, red-brown or rarely pale yellow. Spores small, smooth to papillose, 8--14(--16) \um.

 

Capsules maturing spring-summer (Apr-Aug). Common on disturbed soil, earth banks, rotten wood, rarely on rocks, abundant in disturbed habitats; 0--4,000 m; Greenland; Alta., B.C., Man., N.B., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont., P.E.I., Que., Sask., Yukon; Alaska, Ariz., Ark., Calif., Colo., Conn., Del., D.C., Idaho, Ill., Ind., Iowa, Kans., Ky., Md., Mass., Mich., Minn., Mo., Mont., Nebr., Nev., N.H., N.J., N.Mex., N.Y., N.C., N.Dak., Ohio, Okla., Oreg., Pa., R.I., S.Dak., Tenn., Tex., Utah, Vt., Va., Wash., W.Va., Wis., Wyo.; Mexico:  s South America; Europe; mainland Asia; Africa; Atlantic Islands; Pacific Islands; Australia.

 

The combination of small caespitose plants with ovate-lanceolate concave leaves, a strong awn, and inflated subalar cells on fertile stems is diagnostic of Gemmabryum caespiticium.  The last character is also a diagnostic character of section Cladodium of Ptychostomum, and is also known from a few other species in other genera (e.g., Bryum dyffryense Holyoak).

 

 

3. Gemmabryum kunzei (Hornschuch) J.R. Spence, Phytologia 91: 497. 2009

 

Bryum kunzei Hornschuch, Flora 2(1): 90. 1819

 

Plants small, in open to dense turfs or gregarious, green, yellow-green, or red-green. Stems 0.1--0.6(--1) cm, gemmiform to evenly foliate, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller; rhizoids red-brown to red. Leaves 0.3--0.8(--1.2) mm, ovate-lanceolate, imbricate, somewhat twisted when dry, concave, not decurrent, apex  acuminate, proximal third of lamina often becoming hyaline with age, margins plane, distal margins smooth to serrulate, limbidium lacking; costa highly variable, not reaching apex in proximal leaves to long-excurrent in distal leaves, awn green to more often hyaline, slender, smooth to weakly denticulate,  generally 1/3--2/3\x lamina length; distal and mid-laminal cells rhomboidal to hexagonal, 12--18(--20) \um wide, mostly (2--)3--4:1, thin to moderately incrassate, proximal laminal cells abruptly quadrate across leaf base, wider and shorter than more distal cells, inflated group of pink subalar cells present on distal fertile stem leaves. Specialized asexual reproduction not known.  Sexual condition dioicous. Capsule 2--3(--4) mm, pyriform, red-brown, segments pale tan, brown, red-brown or rarely pale yellow. Spores small, smooth to papillose, 8--12(--14) \um.

 

Capsules mature spring--summer (April--July). Common on dry sandy soil and rock, often calcareous, temperate arid to semi-arid regions; 500--2700 m; Ariz., Calif., Colo., N.Mex., Nev., Utah; c Eurasia (including Caucasus).

 

Gemmabryum kunzei is a small long-awned segregate of G. caespiticium, characterized by very small leaves, often hyaline distal lamina and awn, shorter laminal cells, and lack of a limbidium.  It is a nondescript, small, green, often sterile moss, and many collections may represent depauperate material of other small Bryum species. There are unusual very compact cushion-like forms in semi-arid stable environments (under ledges and in alcoves in Nevada and Utah) that may or may not represent G. kunzei. 

 

4b. GEMMABRYUM sect.  GEMMABRYUM

 

Stems gemmiform to evenly foliate, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller, sometimes julaceous; rhizoids brown to red-brown or rarely red. Leaves imbricate, inflated group of pinkish subalar alar cells absent. Specialized asexual reproduction common, by bulbils in leaf axils, very rarely by rhizoidal tubers.  Sexual condition dioicous. Capsule 1--3 mm, ovate, apophysis  thick, abruptly tapered to seta, sometimes inflated and often rugose, erect, inclined or nutant; peristome double,  variable, from well developed to reduced, endostome high to low, rarely adherent to exostome, segments generally well developed, with lanceolate to ovate perforations, occasionally reduced, cilia long and appendiculate to short or sometimes absent.

 

Species ca. 50 (7 in the flora): worldwide, including Antarctica. 

 

The species of sect. Gemmabryum are strongly represented in temperate, subtropical and tropical montane regions of the world, and are especially well represented in seasonally temperate and Mediterranean climates.  They are characterized by gemmiform stems with imbricate leaves, leaf axil bulbils , and variably developed peristomes with erect to nutant capsules.  The capsule is perhaps the most distinctive feature, being short, ovate, and often with a thickened and rugose apophysis.

 

1. Bulbils absent or rarely present; stems slender, string-like; leaves folded along costa when dry; capsules erect, peristome strongly reduced; rhizoidal tubers sometimes present . . . 8. Gemmabryum exile

1. Bulbils usually present; stems evenly foliate to gemmiform but not string-like when dry; leaves not folded inward along costa; capsule inclined to nodding, peristome well developed.

 

2. Capsule with thick, inflated and rugose neck; leaves of fertile stems triangular, laminal margins revolute to above mid-leaf; bulbils large, single and leafy . . . . 6. Gemmabryum coronatum

2. Capsule with somewhat thick neck, but not inflated, weakly rugose; leaves of fertile stems ovate to ovate-lanceolate, laminal margins plane or recurved proximally; bulbils one to many per leaf axil, leafy or not.

 

3. Bulbils large, 200--750 \um, leafy primordia rising from near base or mid-bulbil, 1--2 per leaf axil.

4. Stems slender, string-like; leaves folded along costa; capsule erect . . . . 8. Gemmabryum exile

4. Stems gemmiform to evenly foliate, not string-like when dry; leaves imbricate, not folded; capsule nodding . . . . 7. Gemmabryum dichotomum

 

3. Bulbils smaller, 50--350 \um, leafy primordia from distal 1/3 of bulbil to short and peglike or lacking, 5--25 per leaf axil.

 

5. Bulbils 150--350 \um, pyriform to conic, distinct primordia present.

 

6. Young bulbils 150--250 \um, primordia narrow, acute and tooth-like . . . . 9. Gemmabryum gemmiferum

6. Young bulbils mostly more than 200 \um, primordia broad, obtuse, leaf-like . . . . 4. Gemmabryum barnesii

 

5. Bulbils 50--200 \um, ovate, round or cylindric, primordia lacking or very short and peg-like.

 

7. Leaves yellow to golden or yellow-green, ovate-lanceolate to triangular; bulbils very small, numerous, 50--70 \um, keyhole to obconic in shape . . . . 5. Gemmabryum californicum

7. Leaves bright green, ovate; bulbils larger, green to brown, 100--200 \um, cylindric to pyriform  . . . . 10. Gemmabryum gemmilucens

 

4. Gemmabryum barnesii (J. B. Wood ex Schimper) J. R. Spence, Phytologia 89: 111. 2007

 

Bryum barnesii J. B. Wood ex Schimper, Syn. Musc. Eur. (ed. 2), 471. 1876

 

Plants small, in open to dense turfs or gregarious, green or yellow-green. Stems 0.2--1 cm, gemmiform to evenly foliate, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller. Leaves 0.5--1.5(--2) mm, ovate to ovate-lanceolate, imbricate when dry, erect when wet; weakly concave, not decurrent, margins plane to revolute proximally, apex acute to acuminate, distal margins smooth, limbidium absent; costa strong, percurrent to rarely short in stout awn; distal and mid-laminal cells rhomboidal to elongate-hexagonal, 8--12(--16) \um wide, mostly 3--4:1, thin to moderately incrassate, proximal laminal cells abruptly quadrate to short-rectangular, 1--2:1, wider and shorter than more distal cells. Specialized asexual reproduction of leaf axis bulbils, bulbils (1--)5--25 per axil, green when young, becoming brown with age, cone-shaped, 200--450 \um, with broad obtuse leaf primordia, less than 1/3\x length of bulbil, arising from distal 1/3 of bulbil.  Sexual condition dioicous. Capsule 1--3 mm, purple-red at maturity, ovate, nutant, apophysis thickened, but not or only weakly rugose; peristome double, well developed, endostome high, not adherent to exostome, segments well developed, perforations ovate, cilia long and appendiculate.

 

Capsules mature spring--summer (April-July). Dry to damp soil, often in sandy disturbed sites;

0--1200 m; B.C.; Calif., Colo., Oreg., Wash., Utah; w,s Europe; Asia (Turkey).

 

Gemmabryum barnesii is similar to G. dichotomum, but produces numerous smaller bulbils in the leaf axils. This species is typically found on disturbed soil that dries out following winter rains. I have tentatively accepted the report by A. Vanderpoorten and C. E. Zartman (2002) for Colorado, as it has also been found in Utah.  Reports from Florida need to be confirmed.  The species is most common in Mediterranean climates along the coast of western North America.

 

5. Gemmabryum californicum (Sullivant) J. R. Spence, Phytologia 89: 111. 2007

 

Bryum californicum Sullivant, Expl. Railroad Mississippi Pacific, Desc. Moss. Liverw. 4(5): 188, plate 6. 1856

 

Plants small, in open to dense turfs or gregarious, yellow-green to golden. Stems 0.1--1 cm, gemmiform, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller. Leaves 0.5--1.5(--2) mm, ovate to triangular, imbricate when dry, erect when wet;

strongly concave, not decurrent, margins plane to revolute proximally, apex acute to acuminate, distal margins smooth, limbidium absent; costa strong, percurrent to short-excurrent as a stout awn; distal and mid-laminal cells rhomboidal to elongate-hexagonal, 8--12(--16) \um wide, mostly 3--4:1, thin to moderately incrassate, proximal laminal cells abruptly quadrate to short-rectangular, 1--2:1, wider and shorter than more distal cells. Specialized asexual reproduction of leaf axil bulbils , bulbils tiny, numerous (more than 25), variously shaped but often obconic or keyhole shaped, (40--)50--70 \um, in (2--)3(--4) tiers with a single cell in the first tier, 2 in the second tier, and 4--6 in the distal tier with longitudinal and transverse red or green septa.  Sexual condition dioicous. Capsule 1--2(--3) mm, red to red-brown at maturity, ovate, nutant, apophysis thickened, but not or only weakly rugose; peristome double, well developed, endostome high, not adherent to exostome, segments well developed, perforations ovate, cilia long and appendiculate.

 

Capsules mature spring--summer (April-July). Dry to damp soil or rock, sometimes calcareous, often on sandstone; 0--600 m; Calif.

 

Gemmabryum californicum is a remarkable endemic species producing the smallest known bulbils in the family.  They can be so numerous that they obscure the distal portion of the stem as a dull to bright green powdery mass. The tiny bulbils appear to be released at this stage, and appear to arise from axillary hairs.  If this is confirmed, this would be a new form of specialized asexual reproductive structure based on origin and development for the family. The species is very similar to G. dichotomum, and only separable by the bulbils and the somewhat more triangular-shaped leaves.  In southern and central California there are small strongly imbricate gemmiform plants with a short stout costa, strongly concave leaves, incrassate distal lamina cells, no bulbils, and an elongate-cylindrical capsule that represent an undescribed species. Gemmabryum californicum will probably be found to occur northward in Mediterranean climates to southwestern British Columbia.

 

6. Gemmabryum coronatum (Schwägrichen) J. R. Spence & H. P. Ramsay, Phytologia 87: 66. 2005

 

Bryum coronatum Schwägrichen. Sp. Musc. Frond., Suppl. 1, 2: 103, plate 71. 1816

 

Plants small, in open to dense turfs or gregarious, green or yellow-green. Stems 0.5--1(--2) cm, gemmiform to evenly foliate, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller. Leaves 0.5--1.5(--2) mm, imbricate when dry, ovate-lanceolate to triangular, erect when wet; weakly concave, not decurrent, margins to revolute to mid-leaf or beyond, apex acute to acuminate, distal margins smooth, limbidium absent; costa strong, short-excurrent as a stout awn; distal and mid-laminal cells rhomboidal to elongate-hexagonal, 8--12(--16) \um wide, mostly 3--4:1, thin to moderately incrassate, proximal laminal cells abruptly quadrate to short-rectangular, 1--2:1, wider and shorter than more distal cells. Specialized asexual reproduction of leaf axis bulbils, bulbils 1(--2) per axil, green when young, becoming brown-green with age, elongate ovate, 250--750 \um, with broad, triangular, elongate primordia, more than 1/3\x length of bulbil, with acute tips, arising from just proximal to the middle of bubil or near base.  Sexual condition dioicous. Capsule 1--3(--4) mm, purple-red at maturity, ovate, nutant, apophysis strongly thickened, inflated, strongly rugose; peristome double, well developed, endostome high, not adherent to exostome, segments well developed, perforations ovate, cilia long and appendiculate.

 

Capsules mature spring--summer (April--July). Common on moist soil and soil over rock; 0--500 m; Ala., Fla., Ga., La., Miss., S.C., Tex.; Mexico; West Indies ; Central America; South America; se Asia (including Indonesia); Africa; Pacific Islands; Australia..

 

Gemmabryum coronatum is similar to G. dichotomum, but tropical-subtropical to warm-temperate in distribution with strongly revolute leaf margins and often triangular leaves. Another feature is the strongly thickened apophysis, almost as though inflated and distinctly rugose, which in G. dichotomum is less thickened and more or less smooth.

 

 

7. Gemmabryum dichotomum (Hedwig) J. R. Spence & H. P. Ramsay, Phytologia 87: 66. 2005.

 

Bryum dichotomum Hedwig, Sp. Musc. Frond., 183, plate 42, figs. 8--12. 1801; B. bicolor Dickson; Gemmabryum bicolor (Dickson) J. R. Spence

 

Plants small, in open to dense turfs or gregarious, green or yellow-green. Stems 0.5--1(--2) cm, gemmiform to evenly foliate, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller. Leaves 0.5--1.5(--2) mm, imbricate when dry, ovate-lanceolate to ovate, erect when wet; weakly concave, not decurrent, margins to revolute proximally, apex acute to acuminate, distal margins smooth, limbidium absent; costa strong, percurrent to rarely short-excurrent as a stout awn; distal and mid-laminal cells rhomboidal to elongate-hexagonal, 8--12(--16) \um wide, mostly 3--4:1, thin to moderately incrassate, proximal laminal cells abruptly quadrate to short-rectangular, 1--2:1, wider and shorter than more distal cells. Specialized asexual reproduction by leaf axis bulbils, bulbils 1(--2) per axil, green when young, becoming brown-green with age, elongate-ovate, 250--750 \um long, with broadly triangular, elongate primordia, more than 1/3\x length of bulbil, with acute tips, arising from just proximal to middle of bubil or near base.  Sexual condition dioicous. Capsule 1--3(--4) mm, purple-red at maturity, ovate, nutant, apophysis somewhat thickened, not distinctly inflated, smooth to weakly rugose; peristome double, well developed, endostome high, not adherent to exostome, segments well developed, perforations ovate, cilia long and appendiculate.

 

Capsules mature spring--summer (April--July). Common on dry to moist soil and soil over rock; 0--2000 m; Alta., B.C., N.B., Ont., Que., Sask.; Alaska, Ariz., Ark., Calif., Colo., Conn.,  Kans., Idaho, Ill., Ind., Iowa, Ky., Maine, Mass., Md., Mich., Mo., Nev., N.H., N.J., N.Y., N.C., Ohio,

Oreg., Pa., Vt., Va., Tenn., Wash.; s South America; .Eurasia; n Africa; Atlantic Islands (Macronesia); Pacific Islands (New Zealand); Australia.

 

Gemmabryum dichotomum is a widespread species exhibiting a temperate bi-polar distribution. The Southern Hemisphere material, G. dichotomum in the strict sense, is larger than Northern Hemisphere material that has traditionally gone under the name of Bryum bicolor but is otherwise similar. They may be distinct, but more studies are needed. Ecologically, plants like those in the Southern Hemisphere are much more common and often dominant in Antarctic-subantarctic tundra vegetation, while typical Northern Hemispeher plants are primarily temperate in distribution.

 

 

8. Gemmabryum exile (Dozy & Molkenboer) J. R. Spence & H. P. Ramsay, Phytologia 87: 67. 2005

 

Bryum exile Dozy & Molkenboer, Ann. Sci. Nat. Bot., sér. 3, 2: 300. 1844; Brachymenium exile (Dozy & Molkenboer) Bosch & Sande Lacoste

 

Plants small, in open to dense turfs or gregarious on soil or rock, green to yellow-green. Stems 0.5--1(--2) cm, slender, string-like when dry, evenly foliate, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller, stems not julaceous. Leaves 0.4--1(--1.5) mm, ovate, loosely to closely imbricate along stem, folded adaxially along costa around stem when dry, erect when wet, weakly to strongly concave, not decurrent; apex broadly acute-rounded, margins plane, distal margins smooth, limbidium absent; costa strong, percurrent to short-excurrent in stout awn, occasionally an apiculus present; distal and mid-laminal cells rhomboidal to elongate-hexagonal, (30--)40--60 \um x 8--12(--4) \um, mostly (3--)4--5:1, thin to moderately incrassate, proximal laminal cells abruptly quadrate, wider than more distal cells. Specialized asexual reproduction common, of small leafy bulbils in axils of distal stem leaves, also of small spherical rhizoidal tubers, brown, 100--200 \um, cells smooth.  Sexual condition dioicous. [Capsule 1--2(--3) mm, ovate, erect, red-brown, apophysis thick, rugose; peristome double,  strongly reduced, endostome low, segments short, split, lacking perforations, cilia absent.]

 

Rare on concrete; 0--10 m; Fla.; Mexico; West Indies; Central America; South America; Australia; se Asia; Africa; Pacific Islands.

 

Gemmabryum exile was recently collected from concrete in a parking lot in southern Florida. It is one of the most distinctive and easily identified species in the genus, with its very slender string-like stems and small infolded leaves that are ovate with plane margins. The plants lacked tubers and bulbils.

 

9. Gemmabryum gemmiferum (R. Wilczek & Demaret) J. R. Spence, Phytologia 89: 111. 2007

 

Bryum gemmiferum R. Wilczek & Demaret, Bull. Jard. Bot. Belg. 46: 529, fig. 5. 1976

 

Plants small, in open to dense turfs or gregarious on soil or rock, green or yellow-green. Stems 0.5--1(--2) cm, gemmiform to evenly foliate, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller. Leaves 0.5--1.5(--2) mm, ovate, imbricate when dry, erect when wet, strongly concave, not decurrent; margins plane to revolute proximally, apex acute to acuminate, distal margins smooth, limbidium absent; costa strong, percurrent to rarely short-excurrent as a stout awn; distal and mid-laminal cells rhomboidal to elongate-hexagonal, 8--16(--20) \um wide, mostly 3--4:1, thin to moderately incrassate, proximal laminal cells abruptly quadrate to short-rectangular, 1--2:1, wider and shorter than more distal cells. Specialized asexual reproduction by leaf axis bulbils, bulbils (2--)5--25 per axil, green when young, orange to red when mature, cylindrical to pyriform, 150--250 \um, primordia distinct, narrow, acute to peg-like, arising from upper 1/3--1/4 of bulbil.  Sexual condition dioicous. Capsule 1--2(--3) mm, purple-red at maturity, ovate, nutant, apophysis somewhat thickened, not distinctly inflated, smooth to weakly rugose, peristome double, well developed, endostome high, not adherent to exostome, segments well developed, perforations ovate, cilia long and appendiculate.

 

Occasional on damp soil, often in clearings and fields; 0--500 m; Calif.; w,s Europe; Atlantic Islands (Canary Islands).

 

Gemmabryum gemmiferum is a distinctive Mediterranean climate species. It is typically found on disturbed soil that dries out following winter rains, often in association with ephemeral bryophytes.  It differs from G. gemmilucens and G. dichotomum in bulbil features.

 

10. Gemmabryum gemmilucens (R. Wilczek & Demaret) J. R. Spence, Phytologia 89: 111. 2007

 

Bryum gemmilucens R. Wilczek & Demaret, Bull. Jard. Bot. Belg. 46: 527, fig. 9. 1976

 

Plants small, in open to dense turfs or gregarious on soil or rock, green or yellow-green. Stems 0.5--1(--2) cm, gemmiform to evenly foliate, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller. Leaves 0.5--1.5(--2) mm, ovate, imbricate when dry, erect when wet, strongly concave, not decurrent; margins plane to revolute proximally, apex acute to acuminate, distal margins smooth, limbidium absent; costa strong, percurrent to rarely short in stout awn; distal and mid-laminal cells rhomboidal to elongate-hexagonal, 8--12(--16) \um wide, mostly 3--4:1, thin to moderately incrassate, proximal laminal cells abruptly quadrate to short-rectangular, 1--2:1, wider and shorter than more distal cells. Specialized asexual reproduction by leaf axis bulbils, bulbils (2--)5--25 per axil, yellow to pale orange or brown, cylindric to spheric, 100--200 \um, usually lacking primordia or occasionally short peg-like primordia present.  Sexual condition dioicous. Capsule unknown.

 

Occasional on damp soil, often in clearings and fields; 0--800 m; Calif.; w,s Europe; Atlantic Islands (Canary Islands).  

 

Gemmabryum gemmilucens is, like G. gemmiferum, distinctive Mediterranean climate species, typically found on disturbed soil that dries out following winter rains.  It is often associated with ephemeral bryophytes, and is much more common than G. gemmiferum. A robust form on serpentine rock in central California has much broader leaves and lacks bulbils, and may represent a distinct species.

 

4c. GEMMABRYUM sect.  TUBERIBRYUM J. R. Spence, Phytologia 91: 496. 2009

 

 

Stems evenly foliate, innovations and fertile stem leaves not much different, innovations not julaceous; rhizoids brown, red, or violet. Leaves loosely set, imbricate to somewhat twisted and contorted when dry. Specialized asexual reproduction common, of rhizoidal tubers, spheric to pyriform, variously ornamented, brown, red, crimson, orange, yellow or purple.  Sexual condition dioicous, rarely synoicous. Capsule 1--3(--5) mm, pyriform, apophysis  slender, gradually tapered to seta, never inflated and rugose, inclined or nutant; peristome double,  generally well developed, endostome high, not adherent to exostome, segments generally well developed, with lanceolate to ovate perforations, cilia long and appendiculate.

 

Species about 50 (9 in the flora ). All continents except Antarctica, well represented in temperate regions of the world, with a few species extending to subtropical regions, absent from Arctic-alpine climates.

 

Despite the early work of A. C. Crundwell and E. Nyholm (1964), the species of sect. Tuberibryum are still not recognized by many North American workers, thus the distributions of many remain poorly known.  Many species are weedy, and often occur in agricultural fields.  Several are likely to have been introduced to North America.

 

1. Rhizoidal tubers small, mostly less than 100(--120) \um in longest axis, brown, red, crimson, violet-red, or orange.

2. Plants pink-green or green; distal and median lamina cells very long and narrow, (4--)6--8:1, often more than 80 \um long, alar cells sometimes differentiated in small quadrate groups; tubers mostly pyriform, brown, rhizoids brown; limbidium sometimes present.

3. Lamina cells 6--8:1, 80--100(--120) \um, alar cells quadrate; leafy bulbils occasionally present in leaf axils; limbidium absent . . . . 11. Gemmabryum apiculatum

3. Lamina cells (4--)5--6:1, 40--70--(80) \um, alar cells not differentiated, similar to juxtacostal cells, rectangular; bulbils lacking; a narrow limbidium often present . . . . 18. Gemmabryum valparaisense

2. Plants green to red-green; distal and median lamina cells shorter, mostly 3--4:1, less than 60 \um long,  alar cells similar to adjacent juxtacostal cells; tubers mostly spheric or sometimes pyriform, violet, red, crimson, or orange, rhizoids violet, red, or red-brown; limbidium absent.

4. Rhizoids violet, tubers spherical, red-violet to dark orange . . . . 19. Gemmabryum violaceum

4. Rhizoids brown to red-brown, tubers spherical or pyriform, orange, red to crimson.

5. Tubers spheric, red-brown to crimson, on long rhizoids in substrate . . . .13. Gemmabryum klinggraeffii

5. Tubers pyriform, bright orange or red-orange, clustered in strings on side branches of main rhizoids at stem base . . . . 12. Gemmabryum demaretianum

 

1. Rhizoidal tubers larger, most 120--200(--250) \um, violet, red-violet, yellow, red or red-brown.

6. Rhizoids and tubers red-violet to red . . . . 15. Gemmabryum ruderale

6. Rhizoids yellow, brown or red-brown, tubers yellow or red.

7. Rhizoids yellow, tubers yellow to rarely orange, walls red . . . . 17. Gemmabryum tenuisetum

7. Rhizoids brown to red, tubers red to red-brown, walls red-brown to brown.

8. Costa strong, long-excurrent, yellow-brown, proximal laminal cells quadrate; calcareous substrates . . . . 14. Gemmabryum radiculosum

8. Costa short-excurrent, red-brown, proximal laminal cells short- to long-rectangular; siliceous substrates . . . . 16. Gemmabryum subapiculatum

 

11. Gemmabryum apiculatum (Schwägrichen) J. R. Spence & H. P. Ramsay, Phytologia 87: 65. 2005

 

Bryum apiculatum Schwägrichen, Sp. Musc. Frond., Suppl. 1, 2: 102, plate 72. 1816

 

Plants small, in open to dense turfs or gregarious, green, yellow-green, often with a pinkish tinge; rhizoids brown. Stems 0.4--2(--3) cm, evenly foliate, innovations and fertile stem leaves similar although leaves of innovations smaller. Leaves 0.4--1.5(--2) mm, widely lanceolate to narrowly ovate, imbricate to loosely set and slightly twisted when dry, weakly concave, rarely weakly decurrent, apex acute; margins plane to weakly revolute below, distal margins smooth to serrulate, limbidium absent; costa strong, percurrent to short-excurrent as a slender awn; distal and mid-laminal cells elongate-hexagonal, 60--100(--120) \um long, 8--16(--20) \um wide, mostly 6--8:1, thin to moderately incrassate, proximal laminal cells gradually rectangular, 2--4:1, gradually wider and shorter than more distal cells, with a differentiated group of quadrate cells in alar region. Specialized asexual reproduction common, by small brown pyriform rhizoidal tubers, 40--80 \um, with cells 15--25 \um, smooth, on long rhizoids or sometimes in proximal leaf axils and at stem base; rarely small leafy green bulbils present in axils of distal stem leaves.  Sexual condition dioicous. Capsule 1--3 mm, pyriform, red-brown, inclined or nutant.

 

Locally common on damp soil or soil over rock, especially along shaded streams; 0--300 m; Ala., Fla., Ga., La., Miss., S.C., Tex.; Mexico; West Indies; Central America; South America; Asia; Pacific Islands; Australia.

 

Gemmabryum apiculatum is a nearly pantropical highly variable species. It is known to produce leaf axil bulbils, at least in Paleotropical material, and may represent a microspecies complex. The very long laminal cells, often pinkish tinge to the plants, small brown pyriform tubers, and the small distinct group of quadrate alar cells are diagnostic. Gemmabryum valparaisense is similar, but has shorter laminal cells, lacks the quadrate alar cells, is typically bright green, and has somewhat larger tubers. Another species that may be confused with G. apiculatum is G. sauteri, which has not yet been reported from North America. It is similar to G. valparaisense, but has shorter laminal cells, lacks a limbidium and has much larger tuber cells.

 

12. Gemmabryum demaretianum (Arts) J. R. Spence, Phytologia 89: 111. 2007  I

 

Brym demaretianum Arts, J. Bryol. 17: 263. 1992

 

Plants small, in open to dense turfs or gregarious, green, yellow-green, often with a reddish tinge; rhizoids hyaline, pale yellowish-brown or red-brown. Stems 0.4--1(--1.5) cm, evenly foliate, innovations and fertile stem leaves similar although leaves of innovations smaller, rhizoids brown. Leaves 0.6--1.5(--2) mm, ovate-lanceolate to ovate, loosely set and slightly twisted when dry, weakly concave, not decurrent, apex acuminate, margins plane to weakly revolute basally, distal margins smooth to serrulate, limbidium absent or indistinct, of 1--2 rows of elongate, incrassate cells; costa strong, percurrent to short-excurrent as a slender awn; distal and mid-laminal cells elongate-hexagonal, 50--70(--80) x 12--20(--24) \um, mostly 3--4:1, thin to moderately incrassate, proximal laminal cells gradually rectangular, 2--4:1, gradually wider and shorter than more distal cells, differentiated group of quadrate cells in alar region absent. Specialized asexual reproduction abundant, by small yellow to orange-brown pyriform rhizoidal tubers, 80--150(--170) \um, with cells 25--50 \um, smooth to slightly protuberant but not distinctly so, in clusters of 2--5 on short lateral rhizoids of main rhizoids at base of stem.  Sexual condition dioicous. Capsule unknown.

 

Rare on disturbed soil of agricultural fields, possibly introduced; 0--100 m; Quebec; c,w Europe.

 

The many medium-sized yellow to orange rhizoidal tubers in clusters on short rhizoids at the stem base are diagnostic of Gemmabryum demaretianum.

 

13. Gemmabryum klinggraeffii (Schimper) J. R. Spence & H. P. Ramsay, Phytologia 87: 67. 2005

 

Bryum klinggraeffii Schimper, Höh. Crypt. Preuss., 81. 1858

 

Plants small, in open to dense turfs or gregarious on soil or rock, green to yellow-green. Stems 0.4--2(--3) cm, evenly foliate, innovations and fertile stem leaves similar although leaves of innovations smaller; rhizoids pale tan to brown. Leaves 0.4--1.5(--2) mm, ovate-lanceolate,

loosely set and slightly twisted when dry, weakly concave, not decurrent, apex acute to acuminate, margins plane to weakly revolute below, distal margins serrulate, limbidium absent; costa strong, short-excurrent in slender awn; distal and mid-laminal cells elongate-hexagonal, 40--60 x 10--14(--16) \um, mostly 3--4:1, thin to moderately incrassate, proximal laminal cells short-rectangular, (1--)2:1, gradually wider and shorter than more distal cells, uniform across leaf base. Specialized asexual reproduction common, of irregularly spheric red to crimson rhizoidal tubers, 60--100(--120) \um, with cells 20--30 \um, distinctly protuberant, on long rhizoids in soil.  Sexual condition dioicous. [Capsule 2--4 mm, pyriform, red-brown, inclined or nutant.]

 

Local and scattered on damp disturbed soil in fields, often in agricultural settings, possibly introduced; 0--800 m; Ont., Que.; Calif., Colo., Conn., Ill., Mo., N.Y.; South America (Argentina); w Europe; Asia (China, India, Japan); Atlantic Islands (Iceland); Pacific Islands (New Zealand); Australia.

 

Gemmabryum klinggraeffii is distinguished by its relatively small spherical red to crimson tubers with distinctly protuberant cells and pale rhizoids.

 

14. Gemmabryum radiculosum (Bridel) J. R. Spence & H. P. Ramsay, Phytologia 87: 68. 2005

 

Bryum radiculosum Bridel, Muscol. Recent. Suppl. 3: 18. 1817

 

Plants small, in open to dense turfs or gregarious on soil or rock, green, yellow-green, often with reddish tinge; rhizoids yellow-red to brown or red-brown. Stems 0.5--1(--2) cm, evenly foliate, innovations and fertile stem leaves similar; rhizoids brown. Leaves 0.4--1.5(--2) mm, widely lanceolate to narrowly ovate, loosely set and slightly twisted when dry, weakly concave, not decurrent, apex acute to acuminate, margins plane to revolute below, distal margins serrulate, limbidium absent; costa strong, short to long-excurrent in slender yellow to red awn; distal and mid-laminal cells elongate-hexagonal, 40--60 x 10--12 (14) \um, mostly 3--5:1, thin to moderately incrassate, proximal laminal cells abruptly quadrate to occasionally short-rectangular,  1--2:1,  wider and shorter than more distal cells, uniform across leaf base. Specialized asexual reproduction common, of large spherical red to red-brown rhizoidal tubers, 120--180 \um, with cells 20--40 \um, smooth, on long rhizoids in soil.  Sexual condition dioicous. Capsule 2--3 mm, pyriform, red-brown, inclined or nutant.

 

Capsules mature spring--summer (April--July). Relatively common on dry disturbed calcareous soil and soil over rock; 0--1500 m; Calif., Nev., Oreg.; Mexico; West Indies; Europe; Asia (Japan); n Africa; Atlantic Islands (Canary Islands); Australia.

 

Gemmabryum radiculosum is distinguished by its relatively large reddish spherical tubers with smooth cells, similar in color to the rhizoids, long-excurrent costa, and preference for strongly calcareous substrates. Gemmabryum subapiculatum is similar, but with red tubers that are brighter than the rhizoids, a short-excurrent costa, and a preference for acidic substrates.

 

15. Gemmabryum ruderale (Crundwell & Nyholm) J. R. Spence, Phytologia 89: 111. 2007.

 

Bryum ruderale Crundwell & Nyholm, Bot. Not. 116: 95. 1963

 

Plants small, in open to dense turfs or gregarious on soil or rock, green, yellow-green, often with reddish tinge. Stems 0.4--2(--3) cm, evenly foliate, innovations and fertile stem leaves similar although leaves of innovations smaller; rhizoids red-violet to deep red. Leaves 0.4--1(--1.5) mm, ovate-lanceolate, loosely set and slightly twisted when dry, weakly concave, not decurrent, apex acute, margins plane to weakly revolute below, distal margins smooth to serrulate, limbidium absent; costa strong, short-excurrent in stout awn; distal and mid-laminal cells elongate-hexagonal, (30--)40--60 x 8--14 \um, mostly 3--4:1, thin to moderately incrassate, proximal laminal cells abruptly quadrate to short-rectangular, 2--4:1, wider and shorter than more distal cells, uniform across leaf base. Specialized asexual reproduction common, of large irregularly spherical purple-red or rarely orange rhizoidal tubers, (120--)150--200 \um, with cells 25--50 \um, smooth, on long rhizoids in soil.  Sexual condition dioicous. [Capsule 1--3 mm, pyriform, red-brown, inclined or nutant.]

 

Rare on damp disturbed soil and sand, possibly introduced; 0--1200 m; Ont.; Ariz., Calif., La., Tex.;  Eurasia; n,s Africa; Atlantic Islands (Macaronesia); Pacific Islands (New Zealand).

 

Gemmabryum ruderale is characterized by red-purple to violet rhizoids and relatively large spherical purple-red rhizoidal tubers.

 

16. Gemmabryum subapiculatum (Hampe) J. R. Spence & H. P. Ramsay, Phytologia 87: 68. 2005

 

Bryum subapiculatum Hampe, Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 4: 51. 1872

 

Plants small, in open to dense turfs or gregarious on soil or rock, green, yellow-green, often with reddish tinge; rhizoids brown or red-brown. Stems 0.5--1(--2) cm, evenly foliate, innovations and fertile stem leaves similar; rhizoids brown. Leaves 0.4--1.5(--2) mm, broadly lanceolate, ovate-lanceolate to narrowly ovate, loosely set and slightly twisted when dry, weakly concave, not decurrent, apex acute to acuminate, margins plane to revolute below, distal margins serrulate, limbidium absent; costa strong, short-excurrent in slender red-brown awn; distal and mid-laminal cells elongate-hexagonal, 40--60 x 10--12 (14) \um, mostly 3--5:1, thin to moderately incrassate, proximal laminal cells abruptly short-rectangular,  2--3:1,  wider and shorter than more distal cells, uniform across leaf base. Specialized asexual reproduction common, of large spherical bright red rhizoidal tubers, 150--300 \um, with cells 25--50 \um, smooth, on long rhizoids in soil or occasionally at stem base.  Sexual condition dioicous. Capsule 2--3 mm, pyriform, red-brown, inclined or nutant.

 

Capsules mature spring--summer (April--June). Common on disturbed dry to damp soil and soil over rock, often in agricultural fields, 0--2000 m. B.C., N.B., N.S.; Ala., Ariz., Ark., Calif., Colo., Conn., Fla., La., Md., Miss, Mo., Nev., N.J., Wash.; W. Eurasia, Iceland, Israel, Australia, New Zealand.

 

Gemmabryum subapiculatum is the most common member of sect. Tuberibryum in North America. It represents a sort of “catch-all” species for collections that do not fit in the other species, and also vary greatly throughout their distribution.  More work is needed to determine if additional microspecies exist within the species in the broad sense. The large bright red spherical tubers with smooth cells, short-excurrent costa, and rectangular proximal laminal cells are diagnostic.

 

17. Gemmabryum tenuisetum (Limpricht) J. R. Spence & H. P. Ramsay, Phytologia 87: 68. 2005

 

Bryum tenuisetum Limpricht, Jahresber. Schles. Ges. Vaterl. Cult. 74(2): 4. 1897

 

Plants small, in open to dense turfs or gregarious on soil or rock, green, yellow-green, often with reddish tinge. Stems 0.4--1.5(--2) cm, evenly foliate, innovations and fertile stem leaves similar although leaves of innovations smaller, rhizoids yellow. Leaves 0.4--1.5(--2) mm, ovate-lanceolate to narrowly ovate, imbricate to loosely set and slightly twisted when dry, weakly concave, not decurrent, apex acute to acuminate, margins revolute below, distal margins serrulate, limbidium absent; costa strong, excurrent as stout medium to long awn; distal and mid-laminal cells elongate-hexagonal, 50--60(--70) x 12--14(--16) \um, mostly 3--4:1, thin to moderately incrassate, proximal laminal cells quadrate to short-rectangular, 1--2:1, gradually wider and shorter than more distal cells, similar across leaf base. Specialized asexual reproduction common, of yellow, more or less spherical rhizoidal tubers, (100--)120--180(--200) \um, with cells 25--40 \um, smooth, cell walls red, on long rhizoids in soil. Sexual condition dioicous or rarely synoicous. Capsule 2--3(--4) mm, pyriform, red-brown, inclined or nutant.

 

Capsules mature spring--summer (April--June). Rare on damp to drying acidic soil; 0--600 m; possibly introduced; B.C., N.B., N.S., Que.; Calif., Maine, Mass., Nev., N.Y., Wash.; Eurasia;
Atlantic Islands (Tenerife); Australia.

 

Gemmabryum tenuisetum is characterized by relatively large yellow tubers with red cell walls and yellow rhizoids.

 

18. Gemmabryum valparaisense (Thériot) J. R. Spence, Phytologia 89: 112. 2007

 

Bryum valparaisense Thériot, Revista Chilena Hist. Nat. 12: 14, plate 4, fig. 1. 1917; Bryum pyriferum Crundwell & Whitehouse

 

Plants small to medium, in open to dense turfs or gregarious on soil or rock, bright green to yellow-green, rarely with reddish tints. Stems 0.4--1(--2) cm, evenly foliate, innovations and fertile stem leaves similar; rhizoids brown to red-brown. Leaves 0.4--1.5(--2) mm, widely lanceolate to narrowly ovate, densely imbricate, not much twisted when dry, flat to weakly concave, rarely weakly decurrent, apex acute, margins plane to weakly revolute basally, distal margins smooth to serrulate, limbidium absent or present, of 1--2 rows of elongate incrassate cells; costa strong,  percurrent to short-excurrent in slender awn; distal and mid-laminal cells elongate-hexagonal, 60--80 x 8--14(--16) \um, mostly 4--6:1, thin to moderately incrassate, proximal laminal cells gradually rectangular, 2--4:1, gradually wider and shorter than more distal cells, similar across leaf base, differentiated group of quadrate cells in alar region absent. Specialized asexual reproduction common, of small brown pyriform rhizoidal tubers, 40--80 \um, with cells 15--20 \um, smooth to weakly protuberant, on long rhizoids in soil and sometimes in leaf axils of proximal leaves.  Sexual condition dioicous or rarely synoicous. Capsule 2--3(--5) mm, pyriform, red-brown, sub-erect, inclined or nutant; peristome occasionally somewhat reduced, cilia sometimes short or poorly developed.

 

Capsule matures spring (March-May). Locally common on damp soil and soil over rock, usually associated with calcareous springs in arid to semi-arid or Mediterranean climates; 0--1700 m; Ariz., Calif., Colo., Nev., Utah, N.Mex.; Mexico; South America (Chile); Europe (Portugal); n Africa; Atlantic Islands (Canary Islands).

 

The typically bright green plants with small brown pyriform tubers, weak limbidum, and quadrate to short-rectangular cells that are uniform across the leaf base are diagnostic of Gemmabryum valparaisense.  This species seems to be found primarily at calcareous springs or other seepy sites, unlike most of the other species in the section which are found on disturbed soil.

 

19. Gemmabryum violaceum (Crundwell & Nyholm) J. R. Spence, Phytologia 89: 112. 2007

 

Bryum violaceum Crundwell & Nyholm, Bot. Not. 116: 94. 1963

 

Plants small, in open to dense turfs or gregarious on soil or rock, green, yellow-green, often reddish. Stems 0.4--1(--1.5) cm, evenly foliate, innovations and fertile stem leaves similar; rhizoids pale to bright violet, purple or rarely red-purple. Leaves 0.4--1(--1.5) mm, ovate-lanceolate, loosely set and slightly twisted when dry, weakly concave, not decurrent, apex acute, margins plane to weakly revolute basally, distal margins smooth to serrulate, limbidium absent; costa strong, short-excurrent in slender awn; distal and mid-laminal cells elongate-hexagonal, (30--)40--60 x 8--14 \um, mostly 3--4:1, thin to moderately incrassate, proximal laminal cells abruptly quadrate to short-rectangular, 2--4:1, wider and shorter than more distal cells, uniform across leaf base. Specialized asexual reproduction common, of large irregularly spherical purple-red or rarely orange rhizoidal tubers, 60--80(--100) \um, with cells 25--30 \um, smooth, on long rhizoids in soil.  Sexual condition dioicous. [Capsule 1--3 mm, pyriform, red-brown, nutant.]

 

Capsule mature spring--summer (April--July). Damp soil or soil over rock, often in disturbed sites; 0--1000 m; B.C., N.S., Ont., Que.; Ariz., Calif., Idaho, Mass., Mo., Wash., Wis.; South America (Argentina, Chile); Eurasia; Atlantic Islands (Tenerife); Pacific Islands (New Zealand).

 

Gemmabryum violaceum is distinguished by the combination of violet rhizoids and small spherical red to purple-red or orange rhizoidal tubers. Gemmabryum ruderale is similar, but has larger tubers, and European material at least has strongly papillose rhizoids compared to relatively smooth rhizoids of G. violaceum; more collections need to be examined to see if this holds true for North American plants.

 

5. HAPLODONTIUM Hampe, Ann. Sci. Nat., Bot. ser. 5(4): 336. 1865  * [Latin haplo, single, and dontium, tooth, the peristome is single]

 

John Spence

 

Plants small, in open to dense turfs or sometimes as scattered individuals, light green to pink-green, often brown proximally. Stems 0.5--2(--3) cm, evenly foliate, unbranched to freely branched by subfloral innovations, stolons absent; often densely tomentose proximally, macronemata absent, rhizoids sparse to abundant. Leaves somewhat twisted or contorted to subsecund or sometimes imbricate when dry, erect to erect spreading when wet; elliptic, ovate to oblong or sometimes broadly lanceolate, not decurrent; margins plane or narrowly revolute below, 1-stratose, limbidium absent, distal margins smooth or rarely weakly serrulate, apex rounded-obtuse to broadly acute, apiculus absent or occasionally present, costa subpercurrent to percurrent or rarely excurrent, transverse section typically lacking guide cells, adaxial supracostal cells elongate, similar to adjacent laminal cells, laminal cells rather uniform, elongate and thin walled, lax, rhomboidal to elongate-rhomboidal, 40--120 x 10-20 \um, not pitted, distal cells not oblique to costa, alar cells irregularly short- to long-rectangular, but not distinctly differentiated. Specialized asexual reproduction absent. Sexual condition dioicous; perigonia and perichaetia usually appearing lateral due to rapid innovation growth, leaves the same size as or somewhat smaller than vegetative leaves. Seta single, yellow- to red-brown, 3--12 mm, curved to flexuose. Capsule erect, suberect to nutant, subglobose to broadly pyriform, often somewhat obscured by elongate innovations, 1--2.5 mm, sometimes curved or irregularly shaped, hypophysis not enlarged or inflated, exothecial cells irregularly quadrate to short-rectangular; operculum flat to weakly convex, apiculate or umbonate; peristome single, typically exostomial but sometimes of unknown origin, absent, or rarely double, teeth slender, irregularly linear-lanceolate, hyaline, sometimes perforate, smooth or finely papillose. Spores small, 10--20(--24) \um, pale, smooth or finely papillose, not shed in tetrads.

 

Species ca. 40 (2 in the flora): pantropical, subtropical to temperate, centered in the mountains of Africa and South America, but extending to Eurasia, North America, and the Pacific.

 

Because of previous over-reliance on the peristome, the taxonomy of Haplodontium and Mielichhoferia remains confused.  The gametophytes of the two genera are very unlike, with Haplodontium species similar to those of Plagiobryum and Plagiobryoides. Recent genetics research has clearly shown that Mielichhoferia is related to Pohlia, while the species centered around H. macrocarpum are closer to Bryum and relatives.  As our two species are very similar gametophytically to the type of Haplodontium, H. megalocarpum Arnott, I have chosen to place them in Haplodontium. Peristome reduction is common and complex in this genus, from double to single to absent.

 

SELECTED REFERENCES: Brassard, G.R. and T. Hedderson. 1983. The distribution of Mielichhoferia macrocarpa, a North American endemic moss. Bryologist 86: 273--275. Shaw, J. and H. A. Crum. 1984. Peristome homology in Mielichhoferia and a taxonomic account of North American species. J. Hattori Bot. Lab. 57: 363--381. Showers, D. 1980. Mielichhoferia tehamensis, sp. nov., from northern California. Bryologist 83: 365--366.

 

1. Peristome present, operculum plane or weakly convex, sometimes apiculate; leaf margins narrowly revolute, at least proximally, distal lamina cells at tip elongate; leaves somewhat twisted or secund when dry ........... 1. Haplodontium macrocarpum

 

1. Peristome lacking; operculum umbonate; leaf margins plane throughout, distal lamina cells at tip short-rectangular to subquadrate; leaves imbricate, stem julaceous …..…2. Haplodontium tehamense

 

1. Haplodontium macrocarpum (Hooker) Spence, Phytologia 87: 26. 2005

 

Weissia macrocarpa Hooker, Musci Americani; or, Specimens of the Mosses Collected in British North America, 74. 1828; Bryum porsildii (I. Hagen) C. J. Cox & Hedderson; Mielichhoferia macrocarpa (Hooker) Bruch & Schimper ex Jaeger & Sauerbeck

 

Plants pale light green to pink-green, red-brown to brown proximally. Stems 0.5--3 cm, weakly to strongly branched, not julaceous. Leaves  flexuose to weakly secund when dry,  erect when wet, ovate to oblong or broadly lanceolate, 0.6--2 mm; margins narrowly revolute proximally, apex broadly acute to sometimes acuminate, apiculus absent or rarely present, costa subpercurrent to percurrent, pale tan-brown or red, rarely excurrent in short to medium-length hairpoint, laminal cells rather uniform, elongate and thin walled, lax, rhomboidal to elongate-rhomboidal, 40--120 x 10--18 \um, cells near leaf tip usually elongate, 3:1 or more, cells somewhat narrower along margin but not forming a distinct limbidium. Seta single, yellow-brown, 4--12 mm, curved to flexuose. Capsule erect to nutant, subglobse to short-pyriform, 1.3--2.5 mm, operculum flat to weakly convex, weakly apiculate, peristome single, teeth slender, irregularly linear-lanceolate, hyaline, sometimes perforate, smooth or finely papillose. Spores small, 12--20(--24) \um, pale, smooth or finely papillose.

 

Capsules mature summer (Jul--Aug). Rare on damp to seepy rock faces or crevices, typically limestone; 0--3500 m, in southern latitudes restricted to high elevations; widely disjunct, Arctic-alpine; Greenland; Alta, Nfld., Nunavut; Alaska, Colo., Mont., Utah; Eurasia (ne Russia). A species of conservation concern.

 

Although Haplodontium macrocarpum has been thought to be associated with mineral-rich rock, collections come from a wide variety of rock types.  In the Southern and Central Rocky Mountains it is apparently restricted to limestone. Some specimens from the Aleutian Islands have a strongly excurrent costa, and in some respects approach the Asian Mielichhoferia (Haplodontium) himalayana Mitten.

 

2. Haplodontium tehamense (Showers) Spence, Phytologia 87: 26. 2005

 

Mielichhoferia tehamensis Showers, Bryologist 83: 365, figs. 1--3. 1980

 

Plants shiny light green to yellow-green or white-green, light brown below. Stems 0.5--1 cm, stongly branched, somewhat julaceous. Leaves imbricate when dry, erect to erect-spreading when wet; elliptic to  oblong, 0.5--1 mm; margins plane, apex rounded-obtuse to broadly acute, apiculus absent, costa subpercurrent, yellowish, laminal cells rather uniform, elongate and thin walled, lax, rhomboidal to elongate-rhomboidal, 40--80 (--100) x 10--20 \um, distal cells at tip shorter, irregularly short-rectangular to subquadrate. Seta red-brown, 3--7 mm, flexuose. Capsule horizontal to nutant, short and broadly pyriform, 1.2--2.5 mm, operculum convex and umbonate, peristome absent. Spores small, 10--14 \um, pale, more or less smooth.

 

Capsules mature summer (Aug.). Rare in crevices of volcanic rock in montane areas; 2200--2600 m; Calif. A species of conservation concern.

 

Haplodontium tehamense is a distinctive species, with its eperistomate capsule and somewhat julaceous stems. It is still only known from a few locations in Lassen Volcanic National Park.

 

6.. IMBRIBRYUM Pedersen, Bryologist 108: 449. 2005 * [Greek imbri, imbricate, and bryum, a moss, alluding to strongly imbricate leaves]

 

John R. Spence

Plants medium-sized to robust, in open to dense turfs or gregarious on soil or rock, green, yellow-green, red-green or red. Stems 0.5--3(--4) cm, evenly foliate, innovations and fertile stem leaves somewhat differentiated, leaves of innovations smaller, stems sometimes julaceous; rhizoids sparse, at base of stem, occasionally in clusters along stem. Leaves (0.6--)1.5--2.5(--3) mm, strongly imbricate  when dry, erect when wet, flat or concave, rarely decurrent; apex obtuse to acuminate, apiculus absent; margins plane to strongly revolute, smooth to serrulate distally, limbidium absent or occasionally present, of 1--2 rows of more elongate cells; costa percurrent to short-excurrent as an awn, in cross-section generally with 1 layer of guide cells; distal and mid-laminal cells vermicular to rhomboidal or hexagonal, 6--16(--22) \um wide, mostly 3--6(--8):1, thin-walled to incrassate, rarely porose, proximal laminal cells abruptly quadrate or rectangular, 1--2(--3):1, wider and shorter than cells above, 1--2 rows of enlarged colored cells sometimes present across leaf base, inflated group of pink sub-alar cells absent. Specialized asexual reproduction by rhizoidal tubers, spherical, (100--)150--350 \um, red, red-brown, orange, or pink, on rhizoids in proximal leaf axils.  Sexual condition dioicous; perigonial and perichaetial leaves not much differentiated, outer perichaetial leaves larger, inner leaves smaller than leaves of innovations. Seta long-exserted, stout to slender, straight or somewhat flexuose. Capsule 2--6 mm, pyriform to clavate, apophysis slender, not thickened, inflated or rugose; inclined or nutant; peristome double, well developed, exostome teeth yellowish, often separate at base, endostome high, not adherent to exostome, segments well developed, with ovate perforations, cilia long and appendiculate or sometimes nodose; operculum low-conic to convex, sometimes apiculate, rarely rostrate. Spores small, smooth to papillose, 8--18(--20) \um.

 

Species ca. 40 (6 in the flora): Nearly worldwide in tropical, subtropical, temperate to boreal regions, rare in Arctic-alpine climates, absent from Antarctica.

 

Imbribryum consists of relatively large species with elongate, evenly foliate stems and strongly imbricate leaves. Species commonly grow on soil or rock, often associated with waterfalls, seeps and springs, or splashed rock. The genus is especially well represented in the mountains of New Guinea, the Neotropics, southeast Asia, and Africa. Most species produce rhizoidal tubers; the presence of leaf axil bulbils, reported from European material of I. gemmiparum, has not been confirmed for North American collections. Many species are difficult to identify as they are morphologically variable, especially I. alpinum and I. muehlenbeckii. There is one undescribed species in the western U.S., discussed under I. alpinum.

 

SELECTED REFERENCES  Whitehouse, H. L. K. 1963. Bryum riparium Hagen in the British Isles. Trans. Brit. Bryol. Soc. 4: 389--403. Ochi, H. 1980. A revision of the Neotropical Bryoideae, Part 1. J. Faculty Educ. Tottori Univ., Nat. Sci. 29: 49--154. Nyholm, E. 1993. Illustrated flora of Nordic mosses. Fasc. 3. Bryaceae-Rhodobryaceae-Mniaceae-Cinclidiaceae-Plagiomniaceae. Nordic Bryological Society. Pedersen, N. 2005. Validation of Imbribryum (Bryaceae). Bryologist 108: 449. Spence, J. R. 2007. Nomenclatural changes in the Bryaceae (Bryopsida) for North America II. Phytologia 89: 110--114. Spence, J. R. 2007. Four species of the Bryaceae new to the U.S.A. Evansia 24: 29--30. Sagar, T. and P. Wilson. 2009. Niches of common bryophytes in a semi-arid landscape. Bryologist 112: 30--41.

 

 

1. Leaves rigid, strongly imbricate when dry, apex acute; distal laminal cells long, incrassate and vermicular, more than 6:1, stems not julaceous.

2. Costa percurrent to short-excurrent; limbidium absent; proximal laminal cells gradually wider proximally, short-rectangular to quadrate; capsule pyriform with short neck . . .  1. Imbribryum alpinum

2. Costa short to moderately long-excurrent into a slender awn; weak to moderately strong limbidium present; proximal laminal cells abruptly enlarged, somewhat bulging, rectangular; capsule with distinct, long neck. . . 3. Imbribryum microchaeton

1. Leaves rigid to somewhat loosely imbricate; distal lamina cells shorter, not vermicular, mostly 3--5:1; plants sometimes julaceous.

3. Stems strongly julaceous; leaves red or purple, rarely green, strongly concave, apices rounded-obtuse to broadly acute, cucullate; costa percurrent; distal lamina cells incrassate, oblique to costa, proximal laminal cells abruptly enlarged to inflated in 1--2 rows . . . 5. Imbribryum miniatum

3. Stems not or weakly julaceous; leaves red to green or yellow, concave to flat, apices obtuse, acute, or acuminate, not cucullate; costa not reaching apex to excurrent; distal lamina cells thin to somewhat firm-walled, parallel to costa, cells at leaf base not abruptly enlarged or inflated.

4. Leaves flat or weakly concave, ovate-lanceolate, apices acute to acuminate; costa short-excurrent in slender, sometimes denticulate awn  . . . 4. Imbribryum mildeanum

4. Leaves distinctly concave, ovate, apices broadly acute or obtuse, rarely somewhat apiculate; costa not reaching apex or rarely percurrent, awn absent.

5. Leaves green to yellow-green, lacking red tints, loosely set, somewhat distant proximally along stem; proximal laminal cells short-rectangular  . . . 2. Imbribryum gemmiparum

5. Leaves red or red-green, red tints usually present, rigid and imbricate, crowded; proximal laminal cells quadrate  . . . 6. Imbribryum muehlenbeckii

 

 

1. Imbribryum alpinum (Hudson ex Withering) Pedersen, Bryologist 108: 449. 2005

 

Bryum alpinum Hudson ex Withering, Syst. Arrangem. Brit. Pl., Ed. 4, 3: 824. 1801

 

Plants medium-sized, in open to dense turfs, dark red to red-green, rarely entirely green, sometimes becoming black with age. Stems 0.5--2(--3) cm, evenly foliate, somewhat julaceous, with metallic sheen, older portions of stem sometimes densely radiculose. Leaves 1--3 mm, red to red green or sometimes green, strongly imbricate when dry, erect when wet, rigid, broadly lanceolate to narrowly ovate or lingulate or occasionally narrowly triangular, weakly concave, not decurrent; apex acute; margins plane distally, revolute proximally, smooth to finely serrulate distally, limbidium absent; costa reddish, percurrent to short-excurrent as a stout awn; distal and mid-laminal cells elongate-hexagonal to vermicular, (50--)60--80 x (8--)10--12 \um, mostly 6--8:1, strongly incrassate, parallel to costa, proximal laminal cells abruptly quadrate to short-rectangular, 1--2(--3):1, occasionally a single row of colored cell present across leaf base. Specialized asexual reproduction of red to red-brown spherical rhizoidal tubers on rhizoids arising from leaf axils, 100--300 \um, scarce. Seta 1--3 cm, stout, more or less straight, red or red-brown to purple. Capsule 3--5 mm, pyriform, short-necked, inclined to nutant, red-purple; operculum convex, apiculate. Spores smooth to papillose, yellowish, 10--15 \um.

 

Capsules rare, maturing spring--summer. Local and uncommon on moist acidic rock or soil over rock; 0--2500 m; Greenland; Nfld. & Labr.; Alaska (St. Lawrence I.), Calif., Colo., Ill., N.H.; w,n Eurasia; n Africa.

 

Imbribryum alpinum is a relatively rare boreal-temperate species characterized by narrow, shiny-metallic, red leaves and incrassate, elongate distal laminal cells.  Most reddish plants similar to this species in the western U.S. do not conform well to this set of characters.  Instead, most collections have red, broadly ovate leaves, shorter distal lamina cells, and short-excurrent costae.  This material cannot be referred to either I. alpinum or I. muehlenbeckii, and apparently represents an undescribed species. Reports from southern Africa need to be verified as other red-colored species occur in the Southern Hemisphere.

 

2. Imbribryum gemmiparum (De Notaris) J. R. Spence, Phytologia 89: 112. 2007

 

Bryum gemmiparum De Notaris, Comm. Soc. Crittog. Ital.2: 212 [112] [Cronaca Bryol. Ital. 1: 25] 1866

 

Plants medium-sized, in open to dense turfs, bright green to yellow-green near tips, becoming stramineous with age. Stems 1--2(--3) cm, evenly foliate, weakly julaceous, with or without metallic sheen, older portions of stem sometimes densely brown-radiculose. Leaves 1--3 mm, green, younger leaves sometimes yellowish, strongly imbricate when dry, erect when wet, soft, strongly concave, weakly decurrent; apex rounded to acute; margins plane throughout, rarely  revolute proximally, smooth, limbidium absent; costa not reaching apex or percurrent, brown to yellow-brown; distal and mid-laminal cells hexagonal, (30--)40--60 x (14--)16--26 \um wide, mostly 3(--4):1, thin-walled, proximal laminal cells abruptly short-rectangular, 2--3:1, occasionally with quadrate cells intermixed, single row of colored cell across leaf base absent. Specialized asexual reproduction of pink to orange spherical rhizoidal tubers on rhizoids arising from leaf axils, 100--200 \um, rare [leafy bulbils in leaf axils]. Seta 1--3 cm, stout, more or less straight, brown. Capsule 2--3 mm, pyriform, short-necked, inclined to nutant, brown; operculum convex, weakly apiculate. Spores small, smooth to papillose, yellowish, 12--18 \um.

 

Capsules rare, maturing spring--summer. Damp to wet calcareous soil or soil over rock, often associated with springs; 0--1800 m; Alta., B.C., Ont.; Alaska, Ariz., Calif., Colo.,  Idaho, Maine, Mass., Mich., Minn., Mo., Mont., Mo., Nebr., Nev., N.J., N.Mex., N.Y., Okla., Oreg., Pa.,  S.Dak., Utah, Vt, Wash, Wyo.; s Europe; Asia (Turkey); n Africa.

 

European material of Imbribryum gemmiparum is reported to produce small, leafy bulbils in leaf axils, but there is some controversy over this as they are also sometimes described as short innovations with stalks. Molecular studies show that at least some European material appears to be related to the Gemmabryum dichotomum complex.  However, North American material is much larger in size, and is otherwise morphologically closest to other Imbribryum species. In eastern U.S. material, a short leaf apiculus is often present. Future studies may indicate that there are two species present, one in North America and the second in Europe and the Mediterranean. Our plants are typically found in strongly calcareous springs.

 

3. Imbribryum microchaeton (Hampe) J. R. Spence, Phytologia 89: 112. 2007

 

Bryum microchaeton Hampe, Ann. Sci. Nat. Bot., sér. 5, 4: 342. 1865

 

Plants medium-sized, in open to dense turfs, pale shiny green to brown-green. Stems 0.5--2(--3) cm, evenly foliate, not julaceous, with metallic sheen, older portions of stem sometimes densely radiculose. Leaves 1--3 mm, pale shiny green, becoming  brown with age, strongly imbricate when dry, erect when wet, rigid, weakly concave or flat, not decurrent; apex acute to acuminate; margins strongly revolute throughout, sometimes plane distally, smooth to finely serrulate, limbidium present, of 1--2 rows of elongate cells; costa short to moderately long-excurrent into a slender awn; distal and mid-laminal cells vermicular to elongate hexagonal, (8--)10--12 \um wide, mostly 6--8:1, strongly incrassate, proximal laminal cells abruptly enlarged, rectangular, 2--3:1, walls somewhat bulging, row of colored cells lacking across leaf base. Specialized asexual reproduction by spherical rhizoidal tubers, 100--200 \um, red or red-brown, on rhizoids on stem. Seta 1--3 cm, stout, more or less straight, brown to red-brown. Capsule 4--6, mm, elongate-clavate, with long distinct neck, inclined to nutant, brown; operculum convex, apiculate. Spores papillose, yellowish, 16--20 \um.

 

Capsules mature spring--summer. Rare on damp calcareous rock or soil over rock in protected sites; 0--500 m; Calif., Fla.; Mexico; Central America (Guatemala); South America (Argentina, Colombia).

 

Imbribryum microchaeton was recently collected in the Santa Monica Mountains of southern California (T. Sagar and P. Wilson 2009). This species is similar to I. alpinum, differing in the pale green leaves lacking any red tints, stronger costa, presence of a leaf limbidium, and long-necked clavate capsule. There is also a report from southern Florida (H. Ochi 1980); reports from Tahiti need to be verified.

 

 

4. Imbribryum mildeanum (Juratzka) J. R. Spence, Phytologia 89: 112. 2007

 

Bryum mildeanum Juratzka, Verh. Zool.-Bot. Ges. Wien. 12: 967. 1862

 

Plants small to medium-sized, in open to dense turfs, bright green, older portions of stems becoming stramineus. Stems 0.5--2(--3) cm, evenly foliate, not julaceous, lacking metallic sheen, sometimes densely radiculose. Leaves 1--3 mm, strongly imbricate when dry, erect when wet, rigid, ovate-lanceolate, weakly concave or flat, not decurrent; apex acute to acuminate; margins plane distally, revolute proximally, smooth to finely serrulate distally, limbidium absent; costa short-excurrent in slender, sometimes denticulate awn, brown, sometimes red proximally; distal and mid-laminal cells elongate-hexagonal, (30--)40--60 x (8--)10--12 \um, mostly 4--6:1, thin to firm-walled, proximal laminal cells abruptly quadrate, with a few short-rectangular cells intermixed, 1--2:1, row of colored cells across leaf base absent. Specialized asexual reproduction absent. Sporophytes unknown.

 

Damp soil along stream or in wetlands; 100--3500 m; Calif., Colo., Maine, Nev., W.Va.; w Europe, Asia (Turkey); n Africa; Atlantic Islands (Azores, Canary Islands).

 

Imbribryum mildeanum is a small, inconspicuous boreal-montane species that has been found in Maine and subalpine-alpine areas in the West. This species has generally not been recognized by North America workers, but, following the sense of European authors, the species can be confirmed from several boreal to montane sites.  It is probably more widespread than the few records indicate.

 

5. Imbribryum miniatum (Lesquereux) J. R. Spence, Phytologia 89: 112. 2007

 

Bryum miniatum Lesquereux, Mem. Calif. Acad. Sci. 1: 23. 1868

 

Plants medium-sized, in open to dense turfs, dark red to red-green, rarely green, sometimes purple-black. Stems 2--4(--5) cm, evenly foliate, strongly julaceous, with metallic sheen, older portions of stem sometimes densely radiculose. Leaves (1--)2--3 mm, purple-red to brown-red, rarely dull olive-green, strongly imbricate when dry, erect when wet, rigid, ovate, strongly concave, not or weakly decurrent; apex rounded-obtuse to broadly acute, cucullate; margins plane throughout or rarely revolute proximally, smooth to finely serrulate distally, limbidium absent; costa percurrent, red-brown to brown; distal and mid-laminal cells elongate-rhomboidal, (40--)50--60 x (10--)12--14 \um, mostly 4--6:1, strongly incrassate, arranged in rows oblique to the costa at 30--45° angle, proximal laminal cells abruptly enlarged to inflated, quadrate, with a few short-rectangular cells intermixed, 1--2:1, with 2--3 rows of somewhat inflated colored cells present across leaf base. Specialized asexual reproduction absent. Seta 1--3 cm, stout, more or less straight to flexuose, red or red-brown to purple. Capsule 2--4(--5), mm, pyriform, inclined to nutant, red-purple; operculum convex, apiculate. Spores smooth to papillose, yellow-brown, 14--18 \um.

 

Capsules common, maturing spring--summer. Common on damp to wet siliceous rock or soil over rock, often associated with waterfalls or springs; 0--1500 m; B.C., Ont.; Nfld. & Labr.; Ark., Calif., Idaho, Mo., Mont., Oreg., Pa., Wash.; Atlantic Islands (Faroes).

 

Imbribryum miniatum is a distinctive species, with dark red-purple, julaceous stems, oblique and strongly incrassate distal laminal cells, and colored inflated cells across the leaf base. Some collections from Yosemite National Park are remarkably large in stature, with dark green leaves that become nearly black when dry.  They may represent an undescribed variety or species. Imbribryum miniatum appears to be related to several Southern Hemisphere and tropical species, including Bryum crassum (Australasia), B. perconcavifolium (Mexico-Central America), B. recurvulum (southeast Asia), and B. sclerodictyon (New Guinea).

 

6. Imbribryum muehlenbeckii (Bruch & Schimper) Pedersen, Bryologist 108: 449. 2005

 

Bryum muehlenbeckii Bruch & Schimper, Bryol. Eur. 4: 163. pl. 381. 1846

 

Plants small to medium-sized, in open to dense turfs, dark red to red-green, rarely entirely green. Stems 0.5--2(--3) cm, evenly foliate, julaceous, lacking metallic sheen, older portions of stem sometimes densely radiculose. Leaves 1--2(--3) mm, dark red to red-green or sometimes dark green, strongly imbricate when dry, erect when wet, rigid, ovate, distinctly concave, not or weakly decurrent; apex rounded to broadly acute; margins plane distally, revolute proximally, smooth to finely serrulate distally, limbidium absent; costa reddish, not reaching leaf apex or rarely percurrent; distal and mid-laminal cells hexagonal, (40--)50--70 x (16--)18--24 \um, mostly (2--)3--4:1, not incrassate, parallel to costa, proximal laminal cells abruptly quadrate, with a few short-rectangular cells intermixed, 1--2:1, occasionally a single row of colored cells reaching across leaf base. Specialized asexual reproduction by orange-red to red-brown, spherical tubers on rhizoids arising from leaf axils, 100--200 \um, scarce. Seta 1--3 cm, stout, more or less straight, red or red-brown to purple. Capsule 2--3 mm, short pyriform, nutant, dark red-brown; operculum convex, apiculate. Spores small, papillose, yellow-brown, 14--18 \um.

 

Capsules rare, mature summer (June--Aug). Damp shaded siliceous rock or soil in rock crevices, montane; 500--3000 m; Greenland; B.C., Ont., Nfld. & Labr.; Calif., Idaho, Maine, Mich., Mont., Nev., N.H., N.Mex., N.Y., Oreg., Pa., Tenn., Wash.; w Europe; Asia (Caucasus); Atlantic Islands (Madiera).

 

Imbribryum muehlenbeckii is similar to small forms of I. alpinum, but has strongly concave, ovate leaves, short distal laminal cells, and a weak costa.  Most material from western North America can be referred to an undescribed species; see discussion under I. alpinum. Reports from Africa and South America need to be verified.

 

7. LEPTOSTOMOPSIS (Müller Hal.) J. R. Spence & H. A. Ramsay, Phytologia 87: 70, 2005 * [The genus Lepstostomum, Greek lepto, small, and stoma, mouth, alluding to the capsule opening]

 

John R. Spence

Plants small, in dense cushions, turfs or rarely gregarious, pale green-silver, pinkish or yellow-silver. Stems short, 0.4--2 cm, sometimes julaceous, strongly branching by subfloral innovations, stolons absent; rhizoids abundant, micronemata and macronemata present on stems. Leaves imbricate dry, erect to erect-spreading when wet, broadly ovate to spathulate, somewhat concave, 0.4--2.5 mm, base straight or slightly curved at insertion, not decurrent; margins plane distally, recurved proximally or sometimes plane, 1-stratose, nearly smooth to distinctly serrate near apex, limbidium present or absent, apex broadly rounded to acute, sometimes hyaline, costa long-excurrent as a denticulate to spinose hyaline awn, sometimes branched at tip, costa transverse section with one well-developed stereid band, guide cells present; adaxial supracostal cells irregularly to regularly quadrate or short-rectangular at base, laminal cells heterogeneous, proximal cells usually quadrate to short-rectangular, medial cells generally similar to distal cells, distal cells hexagonal to rhomboidal, 3--6:1, occasionally longer, not in rows oblique to the costa, thin- to thick-walled, walls not pitted, alar cells usually distinct in quadrate groups or sometimes transversely elongate in small groups. Specialized asexual reproduction absent. Sexual condition dioicous; perigonia and perichaetia terminal, leaves the same size as vegetative leaves or typically larger, not forming a rosette, inner leaves little differentiated. Seta usually 1, straight to somewhat twisted. Capsule erect, 2--5 mm, cylindrical, hypophysis well-differentiated, somewhat expanded and rugose, exothecial cells near mouth quadrate or short-rectangular, thick-walled, reddish, in 1--2 rows, proximal cells longer, irregularly  long-rectangular with straight or sinuose walls; annulus usually present, revoluble; operculum weakly convex, short-conic, not rostrate; peristome diplolepidous-alternate, exostome pale yellow or tan, sometimes reddish, teeth slender lanceolate, not trabeculate, lacking pores along fissural line, endostome hyaline to pale yellow, separate or sometimes adherent to exostome, segments absent, basal membrane low to high, cilia absent. Calpytra fugacious, cucullate, small, smooth. Spores shed singly, not as tetrads, not germinating in capsule, 12--50 \um, finely papillose, pale tan, yellow-tan or brown.

 

Species 10 (2 in the flora). Southern North America, Mexico, Central and Southern America, Southeast Asia, Java, Africa, India, mostly subtropical to tropical seasonal environments.

 

Leptostomopsis is a distinctive genus superficially similar to Leptostomum.  Recent molecular work suggests that it is basal to the remainder of the Bryaceae. It was originally included in the polyphyletic Brachymenium. Although the molecular work indicates that the genus should be placed in the Bryaceae, morphologically it is close to Leptostomum and a case could be made for moving it to the Leptostomaceae.

 

SELECTED REFERENCES: Andrews, A. L. 1935. Bryaceae. In: A. J. Grout, Moss flora of North America, Vol. 2. Newfane: Vermont. Ochi, H. 1980. A revision of the Neotropical Bryoideae, Part 1. J. Faculty Educ. Tottori Univ., Nat. Sci. 29: 49--154.  Pedersen, N., C. J. Cox and L. Hedenäs. 2003. Phylogeny of the moss family Bryaceae inferred from chloroplast DNA sequences and morphology. Syst. Bot. 28: 471--482. Spence, J. R. and H. P. Ramsay. 2005. New genera and combinations in the Bryaceae (Bryales, Musci) for Australia. Phytologia 87: 61--72.

 

1. Leaves silver-yellow, distal half to third of lamina and awn hyaline, distal lamina cells elongate, 4--6:1, thick walled and usually somewhat sinuose ...........................................................1. Leptostomopsis nivea

1. Leaves pale green or green-silver, lamina green distally, proximal portion of awn colored, usually brown or red-brown, sometimes with pinkish tinge, distal laminal cells shorter and wider, 3--4:1, not thick walled or sinuose ...........................................................................................2. Leptostomopsis systylia

 

1. Leptostomopsis nivea (Bescherelle) J. R. Spence [unpublished as yet]

 

Brachymenium niveum Bescherell, J. de Bot. (Morot) 15: 383. 1901

 

Plants in dense cushions, turfs or gregarious, pale yellow-silver. Stems very short, 0.4--0.8 cm. Leaves imbricate when dry, erect to erect-spreading when wet, broadly ovate, somewhat concave, 0.2--1 mm; margins serrate near apex, limbidium present distally, apex acute, hyaline, costa excurrent as long-excurrent denticulate to spinose hyaline awn, straight to weakly recurved when dry, laminal cells heterogenous, proximal cells usually quadrate to short-rectangular,  distal cells hyaline, rhomboidal to almost vermicular, 30--40 \um, 4--6:1, thick-walled and somewhat sinuose. Sexual condition dioicous. Seta straight to somewhat twisted, reddish brown to yellow. [Capsule erect, 2--3 mm, oblong, hypophysis slender, inconspicuous to somewhat well-differentiated, sometimes expanded and rugose, endostome hyaline to pale yellow, separate or rarely adherent to exostome, segments absent, basal membrane low, cilia absent. Spores 20--23 \um, finely papillose, pale yellow-tan or brown.]

 

Sporophytes unknown in flora area. Rare on dry volcanic rock and sandy soil; 1000--1200 m, Madrean woodland in canyons, rarely on open playa edges; Ariz.; Mexico; Central America.

 

Leptostomopsis nivea was recently collected from two locations in southeastern Arizona, and the plants are very small for the species.  One collection consisted of scattered plants in sandy calcareous soil along the edge of the Wilcox Playa, an unusual habitat for this species. This species occurs on trunks of trees or rarely on dry rock. It is reminiscent of Bryum argenteum, but the yellow cast, thick-walled, somewhat sinuose distal laminal cells, long awn, and serrate margins of the distal portions of the leaves easily distinguish this species.

 

2. Leptostomopsis systylia (Müll. Hal.) J. R. Spence, Phytologia 87: 70, 2005

 

Brachymenium systylium Müll. Hal., Syn. Musc. Frond. 1: 320. 1848

 

Plants in dense cushions or turfs, green-silver to pink-silver. Stems short, 0.4--2 cm, sometimes julaceous. Leaves imbricate when dry, erect when wet, spathulate to somewhat elongate-ovate, concave, 0.3--2 mm, margins plane distally, recurved proximally, serrulate to serrate near apex, limbidium absent, apex broadly rounded to acute, not hyaline, costa percurrent in proximal leaves to long-excurrent as a denticulate to spinose awn distally, weakly to strongly recurved when dry, awn tip hyaline but colored proximally, medial cells generally similar to distal cells, distal cells hexagonal to rhomboidal, 40--60 \um, 3--4:1, occasionally longer, not distinctly thick-walled or sinuose, greenish.  Sexual condition dioicous. Seta straight to somewhat twisted, reddish-brown to yellow. [Capsule erect, 2--4 mm, cylindrical, hypophysis well-differentiated, somewhat expanded and rugose, endostome hyaline to pale yellow, separate or rarely adherent to exostome, segments absent, basal membrane low, cilia absent. Spores 17--25 \um, finely papillose, pale yellow-tan or brown.]

 

Sporophytes unknown in flora area. Uncommon on trunks of trees, especially Quercus, occasionally on rock or soil; 0--2000 m, Ariz., Fla., N.Mex., N.C., Tex.; Mexico, Central America; South America; tropical Africa; se Asia (including India, Java). A pantropical-subtropical species.

The shiny pale yellow-green to pink-tinged plants of Leptostomopsis systylia are found primarily on tree trunks.  This species is much more common and widespread than L. nivea, and the two can be readily distinguished by laminal cell differences.  Recently, material of what appears to be a third Leptostomopsis has been collected from rock in California.  These collections are distinct from our two North American species, and do not match any other known species in the genus.  They are small, with relatively shorter hairpoints, and the plants are pale green.

 

8. PLAGIOBRYOIDES J. R. Spence, Phytologia 87: 24, 2005 * [Greek oides, similarity, and Plagiobryum]

 

John R. Spence

 

Plants small to large, in dense turfs, green, yellow-green, red-brown, pink or red. Stems short to elongate, 0.5--4 cm, evenly foliate, not or rarely weakly julaceous, not or weakly branched, stolons absent; rhizoids scarce, micronemata and macronemata absent from stems or present in clusters on proximal stem. Leaves crowded to distant on stems, somewhat contorted when dry, erect to erect-spreading when wet, broadly ovate to ovate-lanceolate, flat to concave, 0.3--3(--3.5) mm, base straight or slightly curved at insertion, usually red, decurrent or not; margins plane or sometimes recurved proximally, 1--3 stratose, more or less smooth, limbidium present, of elongate cells or absent, apex broadly rounded to acute, not hyaline, costa not reaching apex to very short-excurrent in smooth point, costa transverse section with stereid band single, usually well developed, guide cells absent or sometimes present proximally; adaxial supracostal cells irregularly to regularly elongate-rectangular at base, laminal cells somewhat heterogeneous, wide, mostly more than 16 \um wide, proximal cells usually long-rectangular, sometimes bulging, 4--6:1, medial cells generally similar to distal cells, distal cells highly variable, irregularly rhomboidal, 2--4:1, near tip sometimes more or less quadrate, not in rows oblique to the costa, thin-walled, walls not pitted, alar cells not differentiated from juxtacostal cells. Specialized asexual reproduction of rhizoidal tubers on rhizoids at base of stem and in proximal leaf axils. Sexual condition dioicous or rarely synoicous, perigonia and perichaetia terminal, leaves the same size as vegetative leaves or typically larger, not forming a rosette, inner leaves little differentiated. Seta 1, straight to flexuose or twisted, rarely geniculate. Capsule inclined to suberect, 2--6 mm, elongate-pyriform, apophysis differentiated or not, often elongate, mouth often oblique, exothecial cells near mouth quadrate or short-rectangular, thick walled, reddish, in 1--4 rows, cells below longer, irregularly  long-rectangular with straight or sinuose walls; annulus usually present, revoluble; operculum weakly convex, short-conic, not rostrate; peristome double, exostome pale yellow or tan proximally, hyaline distally, teeth lanceolate, trabeculate, lacking pores along fissural line, endostome hyaline to pale yellow, separate from exostome or sometimes adherent, basal membrane high, segments narrowly to broadly perforate, rarely longer than exostome, rarely absent, cilia present or more commonly absent. Calpytra fugacious, cucullate, small, smooth. Spores shed singly, not as tetrads, not germinating in capsule, 11--30 \um, finely papillose, pale brown, tan or yellow-tan.

 

Species 15 (5 in flora); tropical to warm temperate regions of the world, with concentrations in the Neotropics and southeast Asia.

 

Plagiobryoides is similar to Plagiobryum in gametophyte structure, but has a more or less symmetric capsule with the endostome typically shorter than the exostome, and spores that are shed singly. The two genera are probably closely related, with primary speciation of Plagiobryum in Arctic-alpine regions of the northern hemisphere and Plagiobryoides in the tropics. Molecular studies are not available for these taxa and for the morphologically similar Haplodontium. Although the genus was originally described for the highly unusual Plagiobryoides incrassatolimbata, the overall leaf structure, especially the laminal areolation, is similar for this and the other species, with typically very broad thin-walled cells, becoming very long proximally, while the capsules are also similar in overall shape.  As in Plagiobryum, the endostome segments can sometimes be longer than the exostome teeth, especially in some populations of Plagiobryoides cellularis. Several of our species have only recently been collected in the flora region, and are quite rare. B. Allen (2002) provided valuable information on the Neotropical species.

 

SELECTED REFERENCES Andrews, A. L. 1935. Bryaceae. In: A. J. Grout. Moss flora of North America, vol. 2. Newfane, Vermont. Ochi, H. 1980. A revision of the Neotropical Bryoideae, Part 1. J. Faculty Educ. Tottori Univ., Nat. Sci. 29: 49--154. Shaw, A. J. 1981. Bryum incrassatolimbatum Card., a Mexican species new to the U.S.A. Bryologist 84: 365--367. Ochi, H. and N. Salazar Allen. 1990. Bryum incrassatolimbatum new to Panama, with the first description of its sporophyte. Bryologist 93: 286--287. Allen, B. 2002. Moss Flora of Central America.  Part 2. Encalyptaceae-Orthotrichaceae. Missouri Bot. Gard. Press, St. Louis, Missouri. Spence, J. R. 2005. New genera and combinations in Bryaceae (Bryales, Musci) for North America. Phytologia 87: 15--28. Spence, J. R. 2009. Nomenclatural changes in the Bryaceae (Bryopsida) for North America III. Phytologia 91: 493--499.

 

 

1. Stems elongate, to 4 cm, leaves distant, often red to red-brown when dry, decurrent, apex rounded, strong 2- to multi-stratose limbidium present, rhizoidal tubers absent ..... 3. Plagiobryoides incrassatolimbata

1. Stems short to medium length, 0.5--2 cm, leaves crowded or distant, pale pink, green, yellow-green or yellow-brown, not or only weakly decurrent, apex rounded or acute, limbidium present or absent, 1-stratose, large rhizoidal tubers sometimes present at base of stem.

2. Rhizoidal tubers absent; synoicous; peristome reduced, exostome segments irregular, often short, endostome adherent to exostome, fragile, cilia absent, spores 18--22 \um .............. 1. Plagiobryoides brachyneura

2. Rhizoidal tubers sometimes present; dioicous; peristome well developed, exostome teeth long, endostome not adherent, cilia present or sometimes absent, spores 12--30 \um.

3. Leaf apex broadly rounded, costa not reaching apex, leaves concave, somewhat decurrent .......... 4. Plagiobryoides renauldii

3. Leaf apex acute, costa not reaching apex to short-excurrent, leaves not or weakly concave, not decurrent.

4. Distal lamina cells elongate, (2--)3--4:1, plants pale pinkish green, leaves imbricate when dry, rhizoidal tubers absent; capsule with a distinct elongated apophysis ............ 2. Plagiobryoides cellularis

4. Distal lamina cells often short and broad, some quadrate, 1--2(--3):1, plants bright green, leaves contorted when dry, rhizoidal tubers sometimes present capsule with a short indistinct apophysis .................... 5. Plagiobryoides vinosula

 

 

1. Plagiobryoides brachyneura (Kindberg) J. R. Spence, Phytologia 91: 498. 2009   C E

 

Bryum brachyneuron Kindberg, Ottawa Nat. 5: 179. 1892

 

Plants in dense turfs, green or yellow-green. Stems short to elongate, 0.5--2 cm, not julaceous, somewhat branched, innovations common; rhizoids common on proximal portions of stem. Leaves somewhat contorted when dry, crowded along stem, erect to erect-spreading when wet, narrowly to broadly ovate, flat or weakly concave, 0.5--2(--3.5) mm, not decurrent, base red; margins plane or recurved proximally, 1-stratose, more or less smooth, limbidium variable, absent on younger leaves to present on older leaves, apex broadly acute, not hyaline, costa not reaching apex to very short-excurrent in smooth point, leaves of innovations smaller with more obtuse apex and weaker costa, laminal cells somewhat heterogeneous, distal cells irregular in shape, 35--50 x 18--25 \um, 1--3:1, cells incrassate, proximal cells rectangular, (50--)60--80 x 18--28 \um, 3--5:1,  sometimes bulging. Specialized asexual reproduction absent. Sexual condition synoicous. Seta short, thick, red-brown, (0.5--)1--1.5 cm, flexuose to sometimes geniculate. Capsule inclined to nutant, 2--4 mm, brown, elongate-pyriform, hypophysis strongly differentiated, elongate, operculum weakly convex, conic, apiculate; peristome double, exostome pale yellow or tan proximally, hyaline distally, teeth blunt to lanceolate, irregular in shape, endostome hyaline to pale yellow, adherent to exostome, basal membrane high, segments absent or occasionally a few present, perforations narrow, cilia absent. Spores 18--22 \um, papillose, dark yellow-brown.

 

Capsules mature summer (Jul--Aug.). Rare, damp to seepy rock faces or crevices; 0--100 m, St. Paul, St. George and Agattu Islands, Alaska. A species of conservation concern.

 

Plagiobryoides pachyneura is very distinctive, with elongate-necked capsule, very short twisted seta, and short broad distal laminal cells. A. L. Andrews (1935) suggested that P. brachyneura is similar to species of Plagiobryum.  The combination of gametophyte and sporophyte characters indicates a relationship with P. cellularis and its allies, thus I have transferred the species to Plagiobryoides.  It should be sought for elsewhere in the Bering Sea region, especially in the Aleutian Islands and Siberia.

 

2. Plagiobryoides cellularis (Hooker) J. R. Spence, Phytologia 91: 498. 2009

 

Bryum cellulare Hooker, Sp. Musc. Frond., Suppl. 3: 1(1): 214: a. 1827.

 

Plants in dense short turfs, pale pink-green. Stems short, 0.5--1 cm, weakly julaceous, somewhat branched, innovations common; rhizoids sparse on proximal stem. Leaves mostly imbricate when dry, erect when wet, crowded along stem, narrowly to broadly ovate, flat or weakly concave, 0.4--1(--2.5) mm, not decurrent, base pink; margins plane or recurved proximally, 1-stratose, more or less smooth, limbidium absent or rarely a single indistinct layer of elongate cells present proximally, apex acute, costa reaching apex to very short-excurrent in smooth point, leaves of innovations smaller with more obtuse apex and weaker costa, laminal cells somewhat heterogeneous, distal cells elongate hexagonal, 30--70 x 16--22 \um, 2--4:1, cells thin-walled, proximal cells long rectangular, (60--)80--100 x 18--24 \um, 4--5:1,  sometimes bulging, thin-walled. Specialized asexual reproduction absent. Sexual condition dioicous. [Seta short, thick, red-brown, (1--)2--3 cm, flexuose to twisted. Capsule inclined to nutant, 2--4 mm, brown, elongate-pyriform, hypophysis strongly differentiated, elongate, operculum weakly convex, conic, apiculate; peristome double, exostome pale yellow or tan proximally, hyaline distally, teeth lanceolate,  endostome hyaline to pale yellow, not adherent to exostome, basal membrane high, segments present, sometimes longer than exostome, perforations narrow, cilia usually absent, occasionally 1--2 present. Spores 20--28 \um, papillose, yellow-brown.]

 

Rare on damp to seepy rock; 10 m, Fla., N.C.; Mexico; West Indies; Central America; South America; tropical Africa; se Asia; Australia; Pacific Islands (Indonesia, New Guinea).

.

A widespread pantropical species. Our material is very poor, consisting of a few small sterile shoots from two sites, rocks near the sea on the Florida Keys and wet rock in North Carolina.

 

3. Plagiobryoides incrassatolimbata (Cardot) J. R. Spence, Phytologia 87: 24. 2005

 

Bryum incrassatolimbatum Cardot, Rev. Bryol. 36: 114. 1909

 

Plants in dense turfs, dark green, olive-green, or more commonly dark red, red-brown or brown. Stems short to long, 1--3(--4) cm, not julaceous, somewhat branched, innovations common; rhizoids sparse on proximal stem. Leaves somewhat contorted when dry, erect to erect-spreading when wet, somewhat crowded along stem, narrowly to broadly ovate, concave, 1--2(--3) mm, longly decurrent, base red; margins plane, multistratose, more or less smooth, limbidium of short cells in (1--)2--4 rows, apex broadly acute to obtuse, costa not reaching apex to rarely percurrent, leaves of innovations smaller with more obtuse apex and weaker costa, laminal cells somewhat heterogeneous, distal cells irregularly rhomboidal, 35--60 x 12--24 \um, 1--3:1, cells thin-walled, proximal cells rectangular, (60--)80--100  x 18--28 \um, 3--5:1,  sometimes bulging, thin-walled. Specialized asexual reproduction absent. Sexual condition dioicous. Seta slender, red-brown, 1--1.5 cm, flexuose to sometimes geniculate. Capsule  suberect, 2--3 mm, brown, elongate-pyriform, hypophysis weakly differentiated, relatively short, operculum weakly convex, conic, not apiculate; peristome double, exostome pale yellow proximally, hyaline distally, teeth lanceolate, endostome hyaline, not adherent to exostome, basal membrane high, segments present, perforations narrow, cilia absent. Spores 11--14 \um, smooth or papillose).

 

Capsules mature summer (Jul--Aug.). Rare on damp to seepy rock or soil over rock; 700--2200 m,  Ariz., N.Mex., Okla.; Mexico; Central America.

 

Plagiobryoides incrassatolimbata is characterized by large size of the habit and the unusual multistratose border of short, wide cells. This species is similar in many respects to P. renauldii, from which it be distinguished by its limbidium, generally red-brown colored leaves (sometimes green), and lack of rhizoidal tubers. The sporophyte description is from H. Ochi and N. Salazar Allen (1990).

 

4. Plagiobryoides renauldii (Röll ex Renauld & Cardot) J. R. Spence, Phytologia 91: 498. 2009

 

Bryum renauldii Röll ex Renauld & Cardot, Bull. Soc. Roy. Bot. Belgique 38(1): 13. 1900

 

Plants in dense turfs, dark green to olive-green. Stems short to long, 1--3(--4) cm, not or weakly julaceous, somewhat branched, innovations few; rhizoids sparse on proximal stem. Leaves somewhat contorted when dry, erect when wet, distant along stem, broadly ovate to suborbicular, concave, 0.5--2(--3.5) mm, somewhat decurrent, base red-green; margins plane or recurved proximally, 1-stratose, more or less smooth, limbidium absent, apex broadly acute to obtuse, costa not reaching apex to rarely percurrent , laminal cells heterogeneous, distal cells irregular in shape, 25--50 x 16--25 \um, 1--2(--3):1, cells  thin-walled, proximal cells rectangular, (60--)80--100 x 16--24 \um, 3--5:1, bulging, thin-walled. Specialized asexual reproduction of large tubers on rhizoids at base of stem and in most proximal leaves, spherical, red, (200--)300--500 \um. Sexual condition dioicous?. Sporophytes unknown.

 

Capsules unknown. Rare on wet soil and soil over rock in stream; 2200 m, Ariz.; Mexico; West Indies; Central America; South America.

 

During a bryophyte workshop in the Chiricahua Mountains in 2007, collections of Plagiobryoides renauldii were made on the east side of the mountains along a stream, occurring with P. incrassatolimbata.  This is a robust aquatic species of Plagiobryoides, being overall similar to P. incrassatolimbata, but differing in the lack of a limbidium, the rounded apex, distant leaves and rhizoidal tubers.

 

5. Plagiobryoides vinosula (Cardot) J. R. Spence, Phytologia 91: 498. 2009

 

Brachymenium vinosulum Cardot, Rev. Bryol. 38: 6. 1911

 

Plants in dense turfs, bright green. Stems short to moderately long, 1--2(--3) cm, not julaceous, somewhat branched, innovations common; rhizoids sparse on proximal stem. Leaves somewhat contorted  to imbricate when dry, erect when wet, somewhat crowded along stem, narrowly to broadly ovate, flat or weakly concave, 0.6--2(--3) mm, not decurrent, base red; margins plane or recurved proximally, 1-stratose, more or less smooth, limbidium absent or rarely a single indistinct layer of elongate cells present proximally, apex acute, costa reaching apex to very short-excurrent in smooth point, leaves of innovations smaller with more obtuse apex and weaker costa, laminal cells somewhat heterogeneous, distal cells elongate hexagonal, 25--60 x 16--22 \um, 1--3:1, cells thin-walled, proximal cells long-rectangular, (60--)80--100 x 18--24 \um, 4--5:1,  sometimes bulging, thin-walled. Specialized asexual reproduction by large rhizoidal tubers on rhizoids at base of stem, spherical, red, (200--)250--400 \um. Sexual condition dioicous. Seta short, thick, red-brown, (1--)2--3 cm, flexuose to twisted. Capsule suberect to inclined, 2--3 mm, brown, elongate-pyriform, hypophysis weakly differentiated, short, operculum weakly convex, conic, apiculate; peristome double, exostome pale yellow proximally, hyaline distally, teeth lanceolate,  endostome hyaline to pale yellow, not adherent to exostome, basal membrane high, segments present, sometimes longer than exostome, perforations narrow, cilia usually absent, occasionally 1---2 present. Spores 18--26 \um, papillose, yellow-brown.

 

Capsules mature June-Sept. Locally common, damp calcareous rock at springs, including hot springs; 800--2200 m; Ariz., Calif., Colo., N.Mex., Utah, Wyo.; Mexico.

 

Plagiobryoides vinosula has been synonymized with P. cellularis, but differs in its bright green elongate stems, leaves that are somewhat contorted when dry, typically shorter distal laminal cells, rhizoidal tubers and capsule with a short hypophysis.  This latter feature also occurs in P. cellularis, however, and more work is clearly needed to better delimit the two species. Plagiobryoides vinosula has been found recently in hot springs in Colorado and Wyoming.

 

9. PLAGIOBRYUM Lindberg, Öfv. K. Vet.-Akad. Föhr. 19: 606. 1862 * [Greek plagio, oblique, and bryon, a moss]

 

Terry A. J. Hedderson

 

Plants soft, slender, 0.3--2.5 cm, forming turf or in +/- caespitose clumps, perennial, reddish brown to green or whitish- to silvery green with pink tinges.  Stems 0.3--1.5(--2) cm, highly branched proximal to the apex by sterile innovations, in transverse section with 1--3 layers of cortical cells that are smaller and thicker-walled than the interior  cells, central strand present, red to red-brown rhizoids usually abundant especially near base.  Leaves dimorphic, those of main shoot axes erect-spreading to appressed, larger toward the stem apex, broadly ovate to lanceolate, usually slightly decurrent, apices +/- reflexed, margins plane or recurved, entire, leaves of innovations smaller, more ovate; costa present, single, sub-percurrent to excurrent, in transverse section usually with a few abaxial stereids but these occasionally absent; laminal cells 1-stratose, lax and thin-walled throughout or in older leaves becoming firm-walled, rhombic to rectangular distally, rectangular toward leaf base, toward margins usually 1--3 rows somewhat longer and narrower but not forming distinct border.  Specialised asexual reproduction absent.  Sexual condition dioicous; perichaetia and perigonia terminal; perichaetial leaves usually enlarged; perigonia bud-like with more ovate and abruptly acuminate leaves.  Seta slightly curved to cygneous, often reflexed at base.  Capsule horizontal to pendulous, relatively large, clavate, gibbous, distinctly zygomorphic; neck well-developed, 0.5--2.5 times length of urn, furrowed when dry; urn gibbous, narrowed to an oblique mouth; annulus large, revoluble; operculum small, mammillate to low-conic; peristome well developed with exostome shorter than the endostome, teeth narrow-lanceolate, unbordered, yellowish to reddish yellow;  endostome loosely adherent to exostome, processes narrow with narrow perforations, cilia rudimentary.  Spores adherent in tetrads, at least until maturity, elliptic, densely papillose, yellow- to greenish brown or brown, 26--42 µm.

 

Species 9 (2 in the flora): North America, Central America, Europe, Asia, Africa, Pacific Islands (New Zealand).

 

Plagiobryum is easily distinguished when fruiting by the large, gibbous and distinctly zygomorphic capsules.

 

1.  Plants whitish to silvery-green with pink tinges; innovations julaceous; leaves, especially of innovations, imbricate and broadly ovate with plane margins; exostome only slightly shorter than endostome; spores separate at maturity 1. Plagiobryum zierii

 

1.  Plants reddish brown; innovations not julaceous; leaves erect to spreading, ovate-lanceolate to lanceolate, margins recurved; exostome 0.5--0.7 times height of endostome; spores dispersed in tetrads. …………………... 2. Plagiobryum demissum

 

 

 

1.  Plagiobryum zierii (Hedwig) Lindberg, Öfv. K. Vet.Akad. Förh. 19: 606. 1862

 

Bryum zierii Hedwig, Spec. Musc. Frond., 182. 1801; Pohlia zierii (Hedwig) Schwägrichen

 

Plants 0.8--2.5 cm, whitish or silvery green, usually with pink tinges.  Stems erect, 3--20 mm, soft, usually highly branched by elongate, julaceous sub-perichaetial innovations.  Leaves of innovations broadly ovate to nearly circular, concave, appressed or imbricate, slightly decurrent, 0.7--1.3 x 0.4--0.8 mm, apices acute, ending in a small reflexed point, margins plane, leaves on fertile shoots larger, more narrowly ovate; costa ending in or shortly before leaf point, in transverse section with 2--3 adaxial cells in a single layer, abaxial stereid band usually 2--3 cells thick but occasionally absent; laminal cells lax and thin-walled distally, or in older leaves with +/- firm walls, rhomboidal or elongate-rhomboidal, 2.5--5:1, 13--24 /um wide at mid-leaf, proximal cells rectangular 2--4:1, firmer walled.  Capsule horizontal to pendulous, 4--14 mm, neck 0.9--2.5 times length of urn, exostome yellow to yellow-brown, distinctly papillose in lower 1/2, 0.85--0.9 times length of hyaline endostome.  Spores separate at maturity, 28-40 /um, brown to yellow-brown or greenish brown, papillose.

 

Capsules mature mid to late summer or early autumn.  Ledges, crevices of moist or wet cliff faces, occasionally as scattered plants among other bryophytes, often in the vicinity of waterfalls, most frequently found on basic substrates including basalt, shale, limestone; 0--3,000 m; Greenland; Alta., B.C., Nfld., N.W.T., N.S., Nunavut, Que., Yukon; Alaska, Colo., Idaho, Mont., Oreg., Vt., Wash.; Central America (Guatemala), Europe, Asia, Africa (South Africa).

 

Plagiobryum zierii is a predominantly montane-alpine species that is disjunctively distributed between eastern and western North America.  It differs from P. demissum in the julaceous, whitish or silvery green plants that are usually suffused with tinges of pink.  Plagiobryum demissum is invariably red or reddish brown, and the shoots are non-terete.  The broadly ovate, flat-margined leaves of P. zierii also serve to identify the species.  Sterile collections are more likely to be confused with Bryum argenteum, which is also silvery-green and has a similar leaf shape.  However, plants of B. argenteum are smaller, have laminal cells firm to +/- incrassate distally and quadrate proximally, lack pinkish colouration and occur in drier, often disturbed or nitrogen enriched, habitats.

 

 

2.  Plagiobryum demissum (Hooker) Lindberg, Öfv. K. Vet.-Akad. Förh. 19: 606. 1862

 

Bryum demissum Hooker, Musci Exotica 2: pl. 99. 1819;  Meesia demissa Hoppe & Hornschuch

 

Plants 0.3--1.2 cm, red to reddish brown.  Stems erect, 6--15 mm, soft, usually branched by short sub-perichaetial innovations.  Leaves of innovations ovate-lanceolate to ovate, flat to slightly concave, erect to erect-spreading, 0.9--1.4 mm x 0.25--0.5 mm, ending in a slender, acute apex, margins recurved, on fertile shoots larger, more distinctly lanceolate; costa ending in apex to distinctly excurrent, in transverse section with 2--5 adaxial cells in one or two layers, abaxial stereid band 2--4 cells thick; laminal cells +/- firm-walled distally, narrowly hexagonal or rhomboidal to rectangular, 2.5--4.5:1, 12--20 /um wide at mid-leaf, proximal cells thicker walled, 3--5:1.  Capsule pendulous, 2--5 mm, neck 0.5--1.2 times length of urn, exostome yellowish to brown, nearly smooth to distinctly striate throughout, up to 0.5 times height of hyaline endostome.  Spores released in tetrads, (26--)28--35(--42) /um, brown, strongly papillose.

 

Capsules mature in late summer or autumn. Moist, often base rich, cliffs, or on humusy soil in tundra; 0--4200 m; Greenland; Alta., B.C., Nfld., Nunavut, N.W.T, Yukon; Alaska, Colo.; Europe, Asia. 

 

Plagiobryum demissum is a species of highly disjunctive distribution that is almost always associated with mountainous areas.  It is, surprisingly, little recorded from most areas of the Canadian Arctic.  The species is easily recognised when fertile by the combination of reddish gametophytes and pendulous, zygomorphic and distinctly “hump-backed” capsules.  When sterile the plants are inconspicuous and practically indistinguishable from small reddish Bryum species. 

 

10. PTYCHOSTOMUM Hornschuch, Flora 5, 2: syll. 62. 1822 * [Greek ptycho, pleated, and stomum, mouth, alluding to the capsule mouth]

 

John R. Spence

 

Plants small to comparatively very large, in dense or open turfs, red to pink, yellow-green, or brown-green. Stems 0.5--12 cm, brown to red, tufted, comose or evenly foliate, freely branching by sub-floral innovations, innovations elongate and evenly foliate to comose. Leaves of fertile shoots and innovations similar, although innovation leaves smaller, (0.5--)1--4(--5) mm, often gradually enlarged towards stem apex, flat to concave, weakly to strongly contorted or shrunken when dry, erect to erect-spreading when wet, decurrent or not; apex obtuse to acuminate; margins plane or revolute, smooth to denticulate distally, mostly bordered, limbidium 1- or 2-stratose; costa strong, mostly percurrent to long-excurrent in smooth or denticulate awn, in cross-section with abaxial stereid band, 1 layer of guide cells usually present; distal and median laminal cells rhomboidal to elongate hexagonal, mostly 2--4:1, thin to very incrassate, proximal laminal cells gradually short- to long-rectangular, 2--4:1, narrower and generally longer than more distal cells, cells at insertion (subalar) below alar region often inflated, pinkish. Specialized asexual reproduction rare, of filiform gemmae borne in axils of stem leaves and from rhizoids. Sexual condition dioicous, synoicous, autoicous or polyoicous; perigonial and perichaetial leaves somewhat differentiated, outer leaves somewhat enlarged and more acuminate, inner leaves smaller, narrowly ovate-lanceolate to triangular. Seta single, slender to stout, straight to flexuose. Capsule highly variable, from short-ovate to pyriform or clavate, 2--6(--7) mm, suberect to inclined or nutant, peristome double, extremely variable, exostome teeth yellow, orange or brown basally, mostly hyaline distally, lamellae straight or curved in mid-tooth, small pores sometimes present in proximal portion of exostome along mid-line, endostome varying from well-developed with appendiculate cilia and high basal membrane to variously reduced, with cilia variously short or absent, and proximal basal membrane sometimes adherent to exostome, endostome segments narrowly to broadly perforated; operculum conic to low-convex, rounded to apiculate. Spores yellow, brown, black, or green, smooth to distinctly papillose, often variable in size in the same collection and capsule, (8--)10--50 \um.

 

Species ca. 60 (31 in the flora). Worldwide, but concentrated in Northern Hemisphere in Arctic, boreal and alpine regions, also the Southern Hemisphere in cool temperate to subantarctic regions.

 

The species of Ptychostomum exhibit a primary radiation in the Northern Hemisphere, where they form a maze of polyploid Arctic-boreal populations that exhibit high variability and often grade into one another.  They occur on soil, mud, peat, or less commonly on rock or wood. Many species are distinguished by minor differences in capsule shape, operculum development, and peristome structure.  This, more than any other genus, is in urgent need of a worldwide revision.  The two main subgenera of Ptychostomum reflect those species with relatively short proximal lamina cells, inflated subalar cells, 1-stratose limbidium, and comose stems (subg. Cladodium), and those with more elongate foliate stems, not particularly comose, with long-rectangular proximal lamina cells, a thin, partially 2-stratose border, and non-inflated subalar cells (subg. Ptychostomum).  The present treatment does not make use of taxonomic sections since there is considerable confusion over the correct names, with many invalidly published.  A. L. Andrews (1935), E. Nyholm (1993), A. J. E. Smith (2004), and V. I. Zolotov (2000) provide valuable treatments of most of our species. For many species, mature capsules, spores, and sexual condition are needed for proper determination.

 

SELECTED REFERENCES Andrews, A. L. 1935. Bryaceae. In: A. J. Grout (ed.). Moss flora of North America north of Mexico. Newfane, 2: 184--240. Holyoak, D.T. 2004. Taxonomic notes on some European species of Bryum (Bryopsida, Bryaceae). J. Bryol. 26: 247--264. Holyoak, D.T. and L. Hedenäs. 2006. Morphological, ecological and molecular studies of the intergrading Bryum neodamnese and B. pseudotriquetrum. J. Bryol. 28: 299--311. Nyholm, E. 1993. Illustrated flora of Nordic mosses. Fasc. 3. Bryaceae-Rhodobryaceae-Mniaceae-Cinclidiaceae-Plagiomniaceae. Nord. Bryol. Soc. Persson, H. 1952. Critical or otherwise interesting bryophytes from Alaska-Yukon. Bryologist 55: 1--25. Pressel, S., H. W. Matcham, and J. G. Duckett. 2007. Studies of protonemal morphogenesis in mosses. XI. Bryum and allied genera: a plethora of propagules. J. Bryol. 29: 241--258. Smith, A. J. E. 2004. Moss Flora of Britain and Ireland, Ed. 2. Cambridge. Spence, J. R. 2005. New genera and combinations in Bryaceae (Bryales, Musci) for North America. Phytologia 87: 15--28. Spence, J. R. 2007. Nomenclatural changes in the Bryaceae (Bryopsida) for North America II. Phytologia 89: 110--114. Steere, W.C. 1978. The mosses of Arctic Alaska. J. Cramer. Zolotov, V.I. 2000. The genus Bryum (Bryaceae, Musci) in the middle European Russia. Arctoa 9: 155--232.

 

 

1. Leaf base mostly reddish;  stems evenly foliate or comose with distal leaves distinctly enlarged; laminal margins with distinct to indistinct 1-stratose border, distal laminal cells not lax, typically 3--5:1, rhomboidal to hexagonal, proximal lamina cells similar in width and length, but rectangular, occasionally quadrate; gametoecial and inner comal leaves with a group of pink subalar inflated cells  ......................................................................................XXa. Ptychostomum subg. Cladodium

 

1. Leaf base same color as rest of leaf, usually green, occasionally red or pink;  stems mostly evenly foliate, not distinctly comose; laminal margins with indistinct, partially 2-stratose border, distal laminal cells lax, wide and short, 2--4:1, rhomboidal; proximal laminal cells longer and narrower, rectangular; subalar cells not inflated or pinkish .....................XXb Ptychostomum subg. Ptychostomum

 

 

XXa. Ptychostomum subg. Cladodium (Bridel) J. R. Spence, Phytologia 91: 497. 2009

 

Cladodium Bridel, Bryol. Univ. 1: 620. 1826

 

Stems 0.5--4(--6) cm, mostly comose to loosely evenly foliate, leaves usually somewhat enlarged distally; sparsely to densely radiculose. Leaves ovate or ovate-lanceolate, erect-spreading when wet, flat to concave; apex obtuse to acuminate; leaf base red, pink or red sap not present in cell lumens; limbidium usually present, narrow, pale to green or yellow, 1-stratose throughout; costa strong, not reaching apex to long-excurrent, red to red-brown; proximal laminal cells the same length and width as the medial, or sometimes longer, rectangular; leaves of fertile stems and gametoecial leaves with a group of pink or red inflated cells below the alar region. Sexual condition dioicous, synoicous, autoicous or polyoicous. Capsule pyriform to  clavate, sometimes curved; peristome exostome and endostome sometimes adherent, segments with narrowly to broadly ovate perforations, small pores sometimes present at base of exostome teeth. Spores variously yellow, green, brown or black, smooth to coarsely papillose, small to very large, (8--)10--50 \um.

 

Species ca. 20 (16 in the flora): widespread soil-inhabiting species in Arctic, montane and boreal regions of the Northern Hemisphere.

 

Species of subg. Claodium are distinctive in a number of characters, including 1-stratose border, relatively short proximal lamina cells, and tufted or comose habits of many species. Most also have a small group of somewhat inflated pink cells at the leaf insertion in the gametoecial and larger comose leaves, a feature absent from subg. Ptychostomum.  The cells at the leaf base usually have red to red-brown cell walls, but clear sap.

 

1. Plants evenly foliate to loosely comose;  dioicous, autoicous, synoicous or polyoicous; peristome perfect, basal membrane less than or equal to 1/2\x height of exostome; spores generally small, 10--25(--30) \um; filiform gemmae occasionally present in leaf axils.

2. Autoicous; spores 16--20(--22) \um; stems loosely comose to somewhat elongate and ± evenly foliate ................................................ 12. Ptychostomum pallescens

2. Dioicous, synoicous or polyoicous; spores 10--28(--30) \um; stems either evenly foliate or comose.

3. Dioicous; leaves strongly decurrent, stems long and evenly foliate, often densely radiculose; filiform gemmae sometimes present in leaf axils …………. 14. Ptychostomum pseudotriquetrum

3. Dioicous, synoicous or polyoicous; leaves not or weakly decurrent, stems evenly foliate to comose; filiform gemmae lacking.

4. Leaves strongly contorted when dry, concave, apex rounded-acute to obtuse, costa mostly not reaching apex or percurrent; dioicous.

5. Leaves distant, contracted to a narrow base …....… 11. Ptychostomum ovatum

5. Leaves crowded, base wide ………….…9. Ptychostomum neodamense

4. Leaves somewhat twisted or shrunken when dry, not concave, apex acute to acuminate, costa strong, excurrent in short to long awn; synoicous or polyoicous.

6. Stems somewhat elongate, leaves evenly foliate, not much enlarged at apex, somewhat decurrent, costa short-excurrent in stout awn, distal laminal cells often very incrassate, sometimes oblique to costa; synoicous ………. 2. Ptychostomum bimum

6. Leaves comose, leaves enlarged above, not decurrent, costa excurrent in medium to long awn, distal cells thin to moderately thickwalled, never oblique to costa; synoicous or polyoicous.

7. Synoicous; capsule elongate  pyriform, curved and asymmetric; spores (20--)24--28(--30) \um .............. 5. Ptychostomum intermedium (in part)

7. Synoicous or polyoicous; capsule clavate to long-pyriform, symmetric; spores 10--16 \um.

8. Synoicous; limbidium narrow, yellowish, in 2--3 rows wide .…... 3. Ptychostomum creberrimum

8. Polyoicous; limbidium wide, 3 or more rows wide, same color as lamina ....…………………… 7.  Ptychostomum lonchocaulon

1. Plants comose to caespitose, innovations sometimes evenly foliate; autoicous, synoicous or polyoicous; peristome reduced, cilia mostly short or absent, basal membrane low, mostly less than 1/2\x height of exostome; spores (20--)22--50 \um; filiform gemmae absent.

9. Endostome strongly adherent to exostome, giving exostome teeth a chambered appearance; leaves acuminate, costa excurrent in long awn; capsules short-pyriform; spores 22--35 \um

........................…………………………… 13. Ptychostomum pendulum

9. Endostome mostly free from exostome, exostome teeth not chambered; leaves acute to acuminate, costa not reaching apex to excurrent in long awn; capsule pyriform, ovate to clavate; spores (18--)22--50 \um.

11. Leaves strongly concave, innovation leaves imbricate, not much contorted when dry, costa weak, not reaching apex to percurrent, limbidium weak or absent; spores small, 18--23 \um; capsule short-pyriform, somewhat gibbous, cilia generally present ...10. Ptychostomum nitidulum

11. Leaves flat or concave, twisted or contorted when dry, innovations not imbricate, costa of at least rosette leaves excurrent in short to long awn, limbidium mostly distinct; spores (18--)22--50 \um; capsule pyriform to elongate ovate, straight to gibbous, cilia present or absent.

12. Autoicous; leaves concave, limbidium indistinct, costa of  proximal stem leaves not reaching apex, at least some rosette leaves with costa short-excurrent, filiform gemmae often present in leaf axils of innovations; spores large, 30--40 \um; cilia absent …..…… 15. Ptychostomum reedii

12. Synoicous or autoicous; leaves flat to concave, limbidium distinct, well developed, costa of most leaves short to long-excurrent, filiform gemmae absent; spores (18--)22--50 \um; cilia present or absent.

13. Mouth of mature capsule red; spores (18--)22--28(--30) \um; endostome membrane about 1/2\x height of exostome, cilia usually present, sometimes reduced.

14. Cilia rudimentary or sometimes 1--2 and long, appendiculate to nodose cilia present; synoicous; capsule symmetric, mouth straight ………...................... 4. Ptychostomum inclinatum

14. Cilia present and well developed, polyoicous; capsule gibbous, mouth oblique..................... 5. Ptychostomum intermedium (in part)

13. Mouth of mature capsule yellow to pale orange or red; endostome membrane low, less than 1/2\x height of exostome, cilia rudimentary or absent; spores large, 25--50 \um.

15. Leaves acute to acuminate, costa at least of rosette leaves excurrent into short recurved smooth awn; seta long, 2--6 cm; spores 25--50 \um.

16. Leaves weakly concave to flat, not keeled, acuminate, hairpoint distinct, short to medium; spores 40--50 \um; seta 2--6 cm …………………………..…..……. 8. Ptychostomum longisetum

16. Leaves strongly concave, keeled, acute, hairpoint absent to very short-excurrent; spores (22--)25--30 \um; seta 2--3 cm …………………………..….….....…. 6. Ptychostomum knowltonii

15. Leaves acuminate, costa of rosette leaves excurrent into medium or long straight denticulate awn; seta short, 1--2 cm; spores  (22--)25--30 \um.

17. Spores dark, blackish; exostome lacking small pores along midline, endostome membrane nearly 1/2\x height of exostome, sometimes weakly adherent to exostome at base; operculum low-convex, not or weakly apiculate ...…...….........................……... 1. Ptychostomum archangelicum

17. Spores pale; exostome teeth with small pores along midline at base, endostome membrane very low, free from exostome; operculum conical, distinctly apiculate

............................. 16. Ptychostomum salinum

 

 

1. Ptychostomum archangelicum (Bruch, Schimper & W. Gümbel) J. R. Spence, Phytologia 87: 20. 2005

 

Bryum archangelicum Bruch, Schimper & W. Gümbel, Bryol. Eur. 4: 153. 1846; B. curvatum Kaurin & Arnell; Ptychostomum curvatum (Kaurin & Arnell) J. R. Spence

 

Plants small, in dense or open turfs or as scattered individuals, green, red-green or yellow-green. Stems 0.4--1.5 cm, fertile stems tufted, comose, innovations elongate and evenly foliate, often strongly radiculose. Leaves  ovate-lanceolate, (0.4--)0.8--2(--2.5) mm, enlarged towards stem apex, flat to weakly concave, yellow-green to red-green, weakly to strongly contorted or shrunken when dry, erect-spreading when wet, not decurrent; apex acuminate, margins revolute to mid-leaf; limbidium strong, of 2--3 rows; costa red, long-excurrent in awn; distal and median laminal cells rhomboidal, 12--18 \um wide, mostly 3--4:1, mostly thin-walled, proximal laminal cells gradually rectangular, 3--4:1, same width or narrower than more distal cells. Specialized asexual reproduction absent. Sexual condition synoicous. Seta 1--3 cm, straight to flexuose, red or red-brown. Capsule  pyriform, 2--3 mm, symmetric, brown, mouth yellow; peristome mostly well developed to slightly reduced,  exostome teeth yellow basally, hyaline distally, lamellae usually curved in mid-tooth, pores lacking from exostome along mid-line, endostome with high basal membrane, sometimes adherent to exostome, about 1/2\x height of exostome, segments with ovate perforations, cilia reduced or absent; operculum low-convex, apiculate. Spores dark brown to black, strongly papillose, (22--)25--30 \um.

 

Capsules mature Jul--Aug. Locally common on dry calcareous soil in Arctic tundra, occasional in alpine regions to the south; circumpolar Arctic-alpine; Greenland; Alta., B.C., N.W.T., Nunavut, Yukon; Alaska; Eurasia.

 

Ptychostomum archangelicum is a small species found on dry calcareous tundra, and is similar to P. inclinatum.  D. T. Holyoak (2004) placed P. inclinatum and P. curvatum into synonymy under P. archangelicum. Tentatively, I have retained P. inclinatum based on its pale somewhat smaller spores, free endostome, and occasional presence of long cilia. Ptychostomum curvatum is not known from the study area. Capsules are needed for identification.

 

2. Ptychostomum bimum (Schreber) J. R. Spence, Phytologia 87: 20. 2005

 

Bryum bimum Schreber, Bot. Zeit. (Regensburg) 1: 79. 1802.

 

Plants in dense turfs, green or yellow-green. Stems 1.5--3(--4) cm, fertile leaves comose, innovations elongate and evenly foliate; often strongly radiculose. Leaves of fertile shoots and innovations similar, ovate, (1--)2--3(--3.5) mm, not much enlarged towards stem apex, flat to weakly concave, green, weakly twisted to contorted, erect-spreading when wet, weakly decurrent; apex acute, margins revolute to mid-leaf or near apex; costa short-excurrent in stout mostly smooth awn; limbidium strong, in 2--3 rows; distal and median laminal cells rhomboidal, 12--18 \um wide, mostly 3--4:1, mostly firm-walled to sometimes very incrassate, distal cells sometimes at oblique 30--45 degree angle to costa, proximal laminal cells gradually rectangular, 3--4:1, same width or narrower than more distal cells. Specialized asexual reproduction absent. Sexual condition synoicous. Seta 1--3(--4) cm, straight to flexuose, red or red-brown. Capsule  elongate-ovate, 3--5 mm, symmetric, brown, mouth yellow; peristome well developed,  exostome teeth yellow basally, hyaline distally, lamellae usually straight in mid-tooth, pores lacking along exostome mid-line, endostome with high basal membrane, not adherent to exostome, about 1/2\x height of exostome, segments with ovate perforations, cilia long, appendiculate; operculum conic, apiculate. Spores pale yellow or green, finely papillose, (10--)12--16 \um.

 

Capsules mature May--Aug. Locally common on wet soil or soil over rock, occasionally on rock; arctic-boreal-temperate; 0--3000 m; Greenland; Alta., B.C., Man., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont., Que., Sask., Yukon; Alaska, Ariz., Calif., Colo., Conn., Idaho, Ill., Ind., Iowa, Kans., Maine, Mass., Mich., Minn., Mo., Mont., Nebr., Nev., N.H., N.J., N.Dak., Ohio, Oreg., Pa., Utah, Vt., Wash., W.Va., Wis., Wyo.; South America; Eurasia; Africa; Pacific Islands (New Zealand); Australia.

 

Ptychostomum bimum  has in the past been considered a form of P. pseudotriquetrum, but V. I. Zolotov (2002) provided characters that can generally separate the two species.  Ptychostomum bimum is a common species on damp to wet soil or soil over rock, but its distribution is not well understood as it has not little recognized in the past.  It appears to be much more common than P. pseudotriqutrum in eastern North America.

 

3. Ptychostomum creberrimum (Taylor) J. R. Spence & H.P. Ramsay, Phytologia 87: 23. 2005

 

Bryum creberrimum Taylor, Lond. J. Bot., 5: 54. 1846; B. lisae De Notaris var. cuspidatum (Bruch, Schimper & W. Gümbel) Margadant

 

Plants in dense or open turfs, green or yellow-green. Stems 1--2(--3) cm, fertile leaves comose, innovations comose or shortly elongate and evenly foliate; often strongly radiculose. Leaves of fertile shoots and innovations similar, ovate-lanceolate, (1--)2--3(--3.5) mm, somewhat enlarged towards stem apex, flat to weakly concave, green, twisted to contorted, erect-spreading when wet, weakly decurrent; apex acuminate, margins revolute to mid-leaf or near apex; costa long-excurrent in more or less smooth awn; limbidium strong, in 2--3 rows, yellowish; distal and median laminal cells rhomboidal, 14--22 \um wide, mostly 3--4:1, mostly thin to firm-walled, proximal laminal cells gradually rectangular, 3--4:1, same width or sometimes wider than more distal cells. Specialized asexual reproduction absent. Sexual condition synoicous. Seta 1--2(--3) cm, straight to flexuose, red or red-brown. Capsule  elongate-pyriform, 2--4 mm, symmetric, brown, mouth yellow; peristome well developed, exostome teeth yellow basally, hyaline distally, lamellae usually straight in mid-tooth, pores lacking along exostome mid-line, endostome with high basal membrane, not adherent to exostome, about 1/2\x height of exostome, segments with broadly ovate perforations, cilia long, appendiculate; operculum conic, apiculate. Spores pale brown-yellow or green, finely papillose, 10--14(--16) \um.

 

Capsules mature May--Sep. Common on damp to dry soil or soil over rock, 0--3500 m; widespread Arctic-alpine and boreal-temperate; Greenland; Alta., B.C., Man., N.B., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont., P.E.I., Que., Saks., Yukon; Alaska, Ariz., Ark., Calif., Colo., Conn., Del., D.C., Idaho, Ill., Ind., Iowa, Kans., Ky., Maine, Md., Mass., Mich., Minn., Mo., Mont., Nebr., Nev., N.H., N.J., N.Mex., N.Y., N.C., N.Dak., Ohio, Okla., Oreg., Pa., R.I., S.Dak., Tenn., Tex., Utah, Vt., Va., Wash., W.Va., Wyo.; s South America; Eurasia; Pacific Islands (New Zealand); Australia.

 

Ptychostomum creberrimum is one of the most widespread and common species in the genus, occurring throughout the Northern Hemisphere and disjunct in the Southern Hemisphere. It can be distinguished from the closely related P. lonchocoulon by its strictly synoicous sexual condition, and narrower yellowish border. It is similar also to P. pallescens, which is autoicous, has larger spores, and elongate-ovate segment perforations.

 

4. Ptychostomum inclinatum (Swartz ex Bridel) J. R. Spence, Phytologia 87: 21. 2005

 

Cladodium inclinatum (Swartz ex Bridel) Bridel; Bryum amblyodon  Müller Hal.; B. stenotrichum Müller Hal.

 

Plants small, in dense or open turfs or as scattered individuals, green, red-green or yellow-green. Stems 0.5--2(--3) cm, the fertile tufted, comose, the innovations elongate and evenly foliate; often strongly radiculose. Leaves ovate-lanceolate, (0.6--)1--2(--3) mm, enlarged towards stem apex, weakly concave, yellow-green to red-green, weakly to strongly contorted or shrunken when dry, erect-spreading when wet, not decurrent; apex acuminate; margins revolute to mid-leaf, costa red, strong, long-excurrent in smooth to denticulate awn; limbidium strong, in 2--3 rows; distal and median laminal cells rhomboidal, 12--18 \um wide, mostly 3--4:1, mostly thin-walled, proximal laminal cells gradually rectangular, 3--4:1, same width or narrower than more distal cells. Specialized asexual reproduction absent. Sexual condition synoicous. Seta 1--3 cm, straight to flexuose, red or red-brown. Capsule  short to long pyriform, 2--4 mm, brown, symmetric, mouth red to orange-yellow; peristome somewhat reduced,  exostome teeth yellow basally, hyaline distally, lamellae straight to curved in mid-tooth, pores lacking from exostome along mid-line, endostome with high basal membrane, sometimes adherent to exostome, about 1/2\x height of exostome, segments with ovate perforations, cilia 1--2 and appendiculate to reduced; operculum low-convex, apiculate. Spores pale yellow or green, finely  papillose, (18--)20--24 \um.

 

Capsules mature Jun-Aug. Common on dry soil in alpine or Arctic tundra, occasionally at lower elevations; 0--4400 m; circumpolar arctic-alpine; Greenland: Alta., B.C., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont., P.E.I., Que., Yukon; Alaska, Calif., Colo., Idaho, Maine, Mich., Minn., Mont., Nev., N.Y., Oreg., Utah, Wash., Wyo.; South America; Eurasia; Antarctica.

 

Ptychostomum inclinatum is a widespread common Arctic-alpine species, closely related to P. archangelicum, differing in its larger size, occasional presence of cilia, and smaller paler spores.

 

5. Ptychostomum intermedium (Bridel) J. R. Spence, Phytologia 87: 21. 2005

 

Bryum intermedium (Bridel) Blandow, Über. Mecklenb. Moose 6. 1809

 

Plants in dense or open turfs, green or yellow-green. Stems 1--2(--3) cm, fertile leaves comose, innovations comose or shortly elongate and evenly foliate; often strongly radiculose. Leaves of fertile shoots and innovations similar, ovate-lanceolate, (1--)1.5--2(--3) mm, not much enlarged towards stem apex, flat to weakly concave, green, twisted to contorted, erect-spreading when wet, weakly decurrent; apex acuminate; margins revolute to mid-leaf or near apex; costa long-excurrent in colored more or less smooth awn; limbidium weak to strong, 1--2(--3) rows; distal and median laminal cells rhomboidal, 12--20 \um wide, mostly 3--4:1, mostly thin to firm-walled, proximal laminal cells gradually rectangular, 3--4:1, same width or sometimes wider than more distal cells. Specialized asexual reproduction absent. Sexual condition synoicous. Seta 1--2(--3) cm, straight to flexuose, red or red-brown. Capsule elongate-pyriform, 2--4 mm, brown, asymmetric, somewhat curved, mouth oblique, yellow; peristome well developed,  exostome teeth yellow basally, hyaline distally, lamellae usually straight in mid-tooth, pores lacking along exostome mid-line, endostome with high basal membrane, not adherent to exostome, about 1/2\x height of exostome, segments with narrowly ovate perforations, cilia long, nodose; operculum conic, apiculate. Spores pale yellow or green, finely papillose, (18--)20--26(--30) \um.

 

Capsules mature Jun-Sep. Rare on wet soil, boreal-temperate areas; 1000--1500 m; B.C., Greenland; Alta., Man., Ont., Sask.; Eurasia.

 

Although A. L. Andrews (1935) did not confirm material of Ptychostomum intermedium for North America, a few collections from the southern portion of the Boreal zone in Canada can be assigned to this species based on the asymmetric capsules, synoicy, nodulose cilia, and spore size.  Otherwise this species is similar to P. creberrimum and P. lonchocaulon. Mature capsules are needed for identification.

 

6. Ptychostomum knowltonii (Barnes) J. R. Spence, Phytologia 87: 21. 2005

 

Bryum knowltonii Barnes, Bot. Gaz. 14: 44. 1889

 

Plants small, in dense turfs, green or yellow-green. Stems 0.5--2(--3) cm, fertile stems evenly foliate to comose, innovations elongate and evenly foliate; weakly radiculose. Leaves ovate, 1--2(--3) mm, somewhat enlarged towards stem apex, strongly concave, somewhat imbricate to weakly contorted or shrunken when dry, erect-spreading when wet, not decurrent; apex acute to obtuse; margins revolute to mid-leaf; costa red, strong, prominent abaxially, percurrent to short-excurrent in denticulate awn; limbidium strong, (2--)3--4(--5) rows; distal and median laminal cells hexagonal, 14--18 \um wide, (2--)3--4:1, mostly thin-walled, proximal laminal cells gradually rectangular, 3--4:1, same width as cells above. Specialized asexual reproduction absent. Sexual condition synoicous. Seta (2--)3--4 cm, straight to flexuose, slender, red or red-brown. Capsule obovate to short-pyriform, 2--3 mm, yellow-brown, symmetric, mouth red; peristome reduced, exostome teeth yellow to orange proximally, hyaline distally, lamellae straight in mid-tooth, pores lacking from exostome along mid-line, endostome with high basal membrane, not adherent to exostome, less than 1/2\x height of exostome, segments narrow, with ovate perforations, cilia absent or rudimentary; operculum low-convex, apiculate. Spores pale yellow or green, finely papillose, (18--)20--30 \um.

 

Capsules mature Jul-Aug. Locally common to uncommon on wet soil, 0--4000 m; Arctic-alpine; Greenland; B.C., Alta., Nfld. and Labr., N.W.T., Nunavut, Yukon; Alaska, Colo., Wyo.; Arctic-alpine Eurasia.

 

Ptychostomum knowltonii is a distinctive species with its evenly foliate, concave, ovate, keeled leaves, and short capsule on a long seta. In some collections, the capsules are of distinctly different ages in the same tufts.

 

7. Ptychostomum lonchocaulon (Müller Hal.) J. R. Spence, Phytologia 87: 21. 2005

 

Bryum lonchocaulon Müller Hal., Flora 2(6): 90. 1819

 

Plants in dense or open turfs, green or yellow-green. Stems 1--2(--3) cm, fertile leaves comose, innovations comose or shortly elongate and evenly foliate; somewhat radiculose. Leaves of fertile shoots and innovations similar, ovate-lanceolate, (1--)2--3 mm, not much enlarged towards stem apex, flat to weakly concave, green, twisted to contorted, erect-spreading when wet, not decurrent; apex acuminate; margins revolute to mid-leaf or near apex; costa long-excurrent in colored more or less smooth awn; limbidium strong, (2--)3--4(--6) rows, same color as lamina; distal and median laminal cells rhomboidal, (12)14--20 \um wide, mostly 3--4:1, thin to firm-walled, proximal laminal cells gradually rectangular, 3--4:1, same width or sometimes wider than more distal cells. Specialized asexual reproduction absent. Sexual condition polyoicous, mixed synoicous and with male-only gametangia. Seta 1--2(--3) cm, straight to flexuose, red or red-brown. Capsule  elongate-pyriform, 2--4 mm, symmetric, brown, mouth yellow; peristome well developed,  exostome teeth yellow basally, hyaline distally, lamellae usually straight in mid-tooth, pores lacking along exostome mid-line, endostome with high basal membrane, not adherent to exostome, about 1/2\x height of exostome, segments with broadly ovate perforations, cilia long, appendiculate; operculum conic, apiculate. Spores pale brown-yellow or green, finely papillose, (12--)14--18(--20) \um.

 

Capsules mature May--Sept. Locally common on dry to damp soil, 0--3500 m; boreal-temperate; Alta., B.C., , Que.; Alaska, Ariz., Calif., Colo., Idaho, Maine, Minn., Mont., Nev., N.Mex., Oreg., Utah, Wash., Wyo.; Eurasia.

 

Ptychostomum lonchocaulon is very close to P. creberrimum, differing in sexual condition and its stronger non-yellowish limbidium. In North America it is primarily a western species, disjunct in Minnesota. Because P. lonchocaulon has not generally been accepted by most workers, its world distribution is poorly known.  V. I. Zolotov (2002) has a good description of the species and its polyoicous sexual condition.

 

8. Ptychostomum longisetum (Blandow ex Schwägrichen) J. R. Spence, Phytologia 87: 21. 2005

 

Bryum longisteum Blandow ex Schwägrichen, Sp. Frond. Musc. Suppl. 1, 2: 105, plate 74. 1816

 

Plants in dense or open turfs, green or yellow-green. Stems 1--2(--3) cm, fertile leaves comose, innovations comose or shortly elongate and evenly foliate; not strongly radiculose. Leaves of fertile shoots and innovations similar, ovate to broadly ovate-lanceolate, 1--2(--3) mm, enlarged towards stem apex, weakly concave, green or yellow-green, twisted to contorted, erect-spreading when wet, not or weakly decurrent; apex acute; margins revolute to mid-leaf or near apex; costa excurrent in short denticulate awn; limbidium strong, in 2--3 rows; distal and median laminal cells rhomboidal, 12--18 \um wide, mostly 2--3:1, mostly thin to firm-walled, proximal laminal cells gradually rectangular, 3--4:1, same width or sometimes wider than more distal cells. Specialized asexual reproduction absent. Sexual condition synoicous. Seta 3--6(--8) cm, straight to flexuose, red or red-brown. Capsule elongate-pyriform, 2--4 mm, symmetric, brown, mouth yellow; peristome well developed,  exostome teeth yellow basally, hyaline distally, lamellae usually straight in mid-tooth, pores lacking along exostome mid-line, endostome with high basal membrane, not adherent to exostome, less than 1/2\x height of exostome, segments with broadly ovate perforations, cilia short, rudimentary; operculum conic, apiculate. Spores pale yellow-brown or green, finely papillose, variable in size, (38--)40--50 \um.

 

Capsules mature Jun-Sept. Rare and local on damp soil in wetlands; 0--600 m; Greenland; N.W.T., Nfld. and Labr., Nunavut, Yukon; Alaska; Arctic-boreal Eurasia.

 

Ptychostomum longisetum is a circumpolar Arctic-boreal species related to P. inclinatum. Gametophytically it is similar to P. salinum and P. intermedium, but differs in the extremely long seta and very large spores.

 

9. Ptychostomum neodamense (Itzigsohn in Müller Hal.) J. R. Spence, Phytologia 87: 21. 2005

 

Bryum neodamense Itzigsohn in Müller Hal., Syn Musc. Frond. 1: 258. 1848

 

Plants in dense or open turfs, green, red-green or yellow-green. Stems 2--4(--6) cm, fertile stems tufted, comose, innovations elongate and evenly foliate; often strongly radiculose. Leaves of fertile shoots and innovations similar, ovate, (1--)2--3(--4) mm, not much enlarged towards stem apex, strongly concave, yellow-green to green, proximal leaves becoming distinctly black-gray with age, strongly twisted to contorted, erect-spreading when wet, not or weakly decurrent; apex broadly acute to obtuse; margins revolute to mid-leaf or near apex; costa not reaching apex to percurrent, rarely short-excurrent in stout awn; limbidium strong, in 2--3 rows; distal and median laminal cells rhomboidal, 14--20 \um wide, mostly 2--3:1, mostly firm-walled to incrassate, proximal laminal cells gradually rectangular, 3--4:1, same width or wider than more distal cells. Specialized asexual reproduction absent. Sexual condition dioicous. Seta 1--3(--4) cm, straight to flexuose, red or red-brown. Capsule  elongate-ovate, 3--5 mm, symmetric, brown, mouth yellow; peristome well developed,  exostome teeth yellow basally, hyaline distally, lamellae usually straight in mid-tooth, pores lacking along exostome mid-line, endostome with high basal membrane, not adherent to exostome, about 1/2\x height of exostome, segments with ovate perforations, cilia long, appendiculate; operculum conic, apiculate. Spores pale yellow or green, finely papillose, (10--)12--16 \um.

 

Capsules mature Jul-Aug. Uncommon on wet soil or soil over rock, often calcareous, 0--3000 m; primarily Arctic-alpine, but extending into boreal regions; Greenland; B.C., Alta., Nfld. and Labr., N.W.T., Ont., Nunavut, Yukon; Alaska, Calif., Colo.; Eurasia.

 

Ptychostomum neodamense is closely related to P. bimum, P. ovatum and P. pseudotriquetrum, differing in the ovate blunt mostly non-decurrent  leaves that are crowded along the stem.  D. T. Holyoak and L. Hedenäs (2006) did not consider P. neodamense a good species, based on a very limited sampling from Europe, but their results could also be interpreted to suggest that it may be distinct. Clearly more work is needed on the complex of species surrounding P. pseudotriquetrum. Disjunct material from California closely matches European collections, but the Colorado material differs somewhat in overall habit and ecology.

 

10. Ptychostomum nitidulum (Lindberg) J. R. Spence, Phytologia 89: 113. 2007

 

Bryum nitidulum Lindberg, Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 23: 545. 1866; B. teres Lindberg

 

Plants small, in dense or open turfs, yellow-green, yellow-brown, red-green or brown. Stems 0.5--2(--3) cm, fertile stems evenly foliate to comose, innovations elongate and evenly foliate; weakly radiculose. Leaves ovate, 1--2 mm, somewhat enlarged towards stem apex, strongly concave, more or less imbricate to weakly contorted or shrunken when dry, erect-spreading when wet, not decurrent; apex acute to obtuse; margins revolute to mid-leaf; costa red, strong, not reaching apex to percurrent to rarely short-excurrent in denticulate awn; limbidium strong, in (2--)3--4(--5) rows; distal and median laminal cells hexagonal, 12--18 \um wide, 2--3:1, somewhat to distinctly incrassate, proximal laminal cells gradually rectangular, 3--4:1, same width as more distal cells. Specialized asexual reproduction absent. Sexual condition synoicous or sometimes polyoicous, with female-only gametangia. Seta (1--)2--3 cm, straight to flexuose, slender, red or red-brown. Capsule pyriform, 2--3 mm, yellow-brown, asymmetric, curved, mouth yellow; peristome reduced, exostome teeth yellow-brown basally, hyaline distally, lamellae straight in mid-tooth, pores lacking from exostome along mid-line, endostome with low basal membrane, not adherent to exostome, 1/3\x height of exostome, segments narrow, with ovate perforations, cilia usually present, appendiculate, sometimes reduced in length; operculum low-convex, apiculate. Spores brown-yellow, finely papillose, 15--22(--25) \um.

 

Capsules mature Jul-Aug. Locally common on wet soil, 0--1000 m; arctic; Greenland; Yukon, N.W.T., Nunavut; Alaska; Arctic Eurasia.

 

Ptychostomum nitidulum is a small species close to P. intermedium, but differs in ecology, leaf structure and sexual condition. The Arctic P. teres is very close to P. nitidulum, differing in minor characters of the capsule only. D. T. Holyoak (2004) synonymized P. nitidulum with P. intermedium, but I prefer to keep them distinct, based primarily on the leaf differences, until more detailed studies are completed.

 

11. Ptychostomum ovatum (Hedwig) J. R. Spence, Phytologia 89: 113. 2007

 

Bryum ovatum (Hedwig). Dickson ex Withering, Syst. Arr. Brit. Pl. (ed. 4): 3, 795. 1801; B. subneodamense Kindberg

 

Plants in dense or open turfs, olive green, red-green or yellow-green. Stems 4--8(--12) cm, fertile stems tufted, comose, innovations elongate and evenly foliate; weakly radiculose. Leaves of fertile shoots and innovations similar, ovate, (1--)2--3 mm, not much enlarged towards stem apex, concave, yellow-green to green, proximal leaves becoming distinctly black-gray with age, strongly twisted to contorted when dry, erect-spreading when wet, weakly decurrent; apex broadly acute to obtuse; margins revolute to mid-leaf or near apex; costa not reaching apex to percurrent; limbidium strong, in 2--3 rows; distal and median laminal cells rhomboidal, 15--20 \um wide, mostly 2--3:1, mostly thin-walled, proximal laminal cells gradually rectangular, 3:1, same width or wider than more distal cells. Specialized asexual reproduction absent. Sexual condition dioicous. Sporophytes unknown.

 

Rare on wet soil in bogs and wetlands; 0--800 m; Greenland; Alta., N.W.T., Ont., Yukon; n Eurasia.

 

Ptychostomum ovatum is a distinctive species that can only be confused with P. cyclophyllum and P. neodamense.  Diagnostic characters include very distant, strongly shrunken leaves when dry, a rather narrow reddish leaf base, and 1-stratose border. When leaves are stripped off the stem, a long strip of the stem often remains attached to the leaf base.  Ptychostomum cyclophyllum has broadly ovate to orbicular green leaves with very long proximal lamina cells, base not red, and 2-stratose margins. Ptychostomum neodamense has much more crowded leaves with a fairly broad attachment to the stem, but is otherwise similar.  More studies are needed to determine whether they represent extremes of the same species.

 

12. Ptychostomum pallescens (Schleicher ex Schwägrichen) J. R. Spence, Phytologia 87: 21. 2005

 

Bryum pallescens Schleicher ex Schwägrichen, Sp. Musc. Suppl., 2: 107, plate 75. 1816

 

Plants in dense or open turfs, green or yellow-green. Stems 1--3(--4) cm, fertile leaves comose, innovations comose or shortly elongate and evenly foliate; somewhat radiculose. Leaves of fertile shoots and innovations similar, ovate-lanceolate, (1--)2--3(--3.5) mm, somewhat enlarged towards stem apex, flat to weakly concave, green, twisted to contorted, erect-spreading when wet, weakly decurrent; apex acuminate; margins revolute to mid-leaf or near apex; costa long-excurrent in a colored, more or less smooth awn; limbidium strong, in 2--3 rows; distal and median laminal cells rhomboidal, 14--22 \um wide, mostly 3--4:1, mostly thin to firm-walled, proximal laminal cells gradually rectangular, 3--4:1, same width or sometimes wider than more distal cells. Specialized asexual reproduction absent. Sexual condition autoicous. Seta 1--2(--3) cm, straight to flexuose, red or red-brown. Capsule elongate-pyriform, 2--4 mm, symmetric, brown, mouth yellow; peristome well developed,  exostome teeth yellow basally, hyaline distally, lamellae usually straight in mid-tooth, pores lacking along exostome mid-line, endostome with high basal membrane, not adherent to exostome, about 1/2\x height of exostome, segments with elongate oval perforations, 1.5--2:1, cilia long, appendiculate to nodose; operculum conic, apiculate. Spores pale brown-yellow or green, finely papillose, 18--22 \um.

 

Capsules mature May--Aug. Common on damp to wet soil; 0--3300 m; Greenland; Alta., B.C., Man., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont., P.E.I., Que., Sask., Yukon; Alaska, Ariz., Calif., Colo.,  Idaho, Maine, Mass., Mich., Minn., Mo., Mont., Nev., N.H., N.Dak., Ohio, Oreg., Pa.,  S.Dak., Utah, Vt., Va., Wash., W.Va., Wis., Wyo.; South America; Eurasia; Pacific Islands (New Zealand).

 

Ptychostomum pallescens is a common widespread boreal-temperate species favoring wet soil in wetlands, along streams, and in springs. Ptychostomum creberrimum is similar, but has smaller spores, synoicous sexual condition, and ovate segment perforations.

 

13. Ptychostomum pendulum Hornschuch, Flora 5, 2: syll. 62. 1882

 

Bryum algovicum Sendtner ex Müller Hal.

 

Plants small, in dense or open turfs, green, red-green or yellow-green. Stems 0.5--1(--1.5) cm, fertile stems tufted, comose, innovations elongate and evenly foliate, often strongly radiculose. Leaves  ovate-lanceolate, (0.4--)0.8--1.5(--2.5) mm, enlarged towards stem apex, concave, yellow-green to red-green, weakly to strongly contorted or shrunken when dry, erect-spreading when wet, not decurrent; apex acuminate; margins revolute to mid-leaf; limbidium strong, in 2--3 rows; costa red, long-excurrent in denticulate awn; distal and median laminal cells rhomboidal, 12--18 \um wide, mostly 3--4:1, mostly thin-walled, proximal laminal cells gradually rectangular, 3--4:1, same width or narrower than more distal cells. Specialized asexual reproduction absent. Sexual condition synoicous. Seta 1--3 cm, straight to flexuose, red or red-brown. Capsule  pyriform, 2--3 mm, symmetric, brown, mouth red; peristome reduced,  exostome teeth yellow basally, hyaline distally, lamellae indistinct, chambered due to strong adherence of endostome, pores lacking from exostome along mid-line, endostome with basal membrane about 1/3--1/2\x height of exostome,  adherent to exostome, segments with ovate perforations, cilia reduced to nearly absent; operculum low-convex, apiculate. Spores pale brown, strongly papillose, variable in the same collections and capsules, (22--)25--30(--36) \um.

 

Capsules mature May--Aug. Locally common on dry calcareous soil or rock, Arctic-alpine and boreal; 0--4000 m; Greenland; Alta., B.C., Man., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont., Que., Sask., Yukon; Alaska, Ariz., Calif., Colo., Conn., D.C., Idaho, Ill., Ind., Kans., Maine, Mass., Mich., Minn., Mont., Nebr., Nev., N.H., N.J., N.Dak., Ohio, Oreg., Pa., S.Dak., Tenn., Tex., Utah, Vt.,  Wash., W.Va., Wis., Wyo.; South America; Eurasia; n Africa; Pacific Islands (New Zealand); subantarctic islands.

 

Ptychostomum pendulum is a small species found in dry sites, similar overall to P. archangelicum and P. inclinatum.  The larger spores, and unusual chambered appearance of the exostome teeth are diagnostic. Capsules are needed for identification.

 

14. Ptychostomum pseudotriquetrum (Hedwig) J. R. Spence & H. P. Ramsay ex D. T. Holyoak & N. Pedersen, J. Bryol. 29: 120. 2007

 

Mnium pseudotriquetrum Hedwig, Sp. Musc. Frond., 190. 1801; Bryum pseudotriquetrum (Hedwig) Gaertner, Meyer & Scherbius

 

Plants in dense turfs, green, red-green or yellow-green, older shoots becoming red-brown. Stems 2--4(--6) cm, fertile stems tufted, weakly comose to evenly foliate, innovations elongate and evenly foliate; usually strongly radiculose well towards stem tips. Leaves of fertile shoots and innovations similar, ovate, (1--)2--3(--4) mm, not much enlarged towards stem apex, flat to weakly concave, green, red-green or yellow-green, older leaves becoming dull brown-red or brick colored, somewhat twisted to contorted, erect-spreading when wet, strongly and narrowly decurrent; apex acute; margins revolute to mid-leaf or near apex; costa short-excurrent in stout awn; limbidium strong, in 2--3 rows; distal and median laminal cells rhomboidal, 14--22 \um wide, mostly (2--)3:1, firm-walled to often incrassate, proximal laminal cells gradually short-rectangular, 3:1, same width or somewhat wider than more distal cells. Specialized asexual reproduction occasional, of brown filiform papillose gemmae in axils of leaves. Sexual condition dioicous. Seta 1--3(--4) cm, straight to flexuose, red or red-brown. Capsule  elongate-ovate, 3--5 mm, symmetric, brown, mouth yellow; peristome well developed,  exostome teeth yellow basally, hyaline distally, lamellae usually straight in mid-tooth, pores along exostome midline lacking, endostome with high basal membrane, not adherent to exostome, about 1/2\x height of exostome, segments with ovate perforations, cilia long, appendiculate; operculum conic, apiculate. Spores pale yellow or green, finely papillose, 12--18 \um.

 

Capsules mature Apr--Oct. Common to abundant on wet soil, soil over rock or rock, often in fens; widespread arctic-boreal-temperate; 0--4000 m; Greenland; Alta., B.C., Man., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont., P.E.I., Que., Sask., Yukon; Alaska, Ariz., Ark., Calif., Colo., Conn., Del., D.C., Ga., Idaho, Ill., Ind., Iowa, Kans., Ky., Maine, Md., Mass., Mich., Minn., Mo., Mont., Nebr., Nev., N.H., N.J., N.C., N.Dak., Ohio, Okla., Oreg., Pa., R.I., S.Dak., Tenn., Tex., Utah, Vt., Va., Wash., W.Va., Wis., Wyo.; South America; Eurasia; Africa; Pacific Islands (New Zealand); Australia.

 

Ptychostomum pseudotriquetrum is one of the most common and widespread species in the Bryaceae, absent only from the subtropics, tropics, and central Pacific islands. The ovate decurrent leaves, short awn, dense areolation, dioicous sexual condition, and long radiculose stems are characteristic.  Ptychostomum bimum is smaller, with much weaker decurrencies, and is synoicous.

 

15. Ptychostomum reedii (Robinson) J. R. Spence, Phytologia 87: 22. 2005

 

Bryum reedii Robinson, Bryol. 69: 107, 1966

 

Plants in dense turfs, dark green. Stems 1--2 cm, fertile leaves comose, innovations comose or shortly elongate and evenly foliate; not strongly radiculose. Leaves of fertile shoots and innovations similar, oblong or ovate, (0.5--)1--2 mm, concave, not much enlarged towards stem apex, flat to weakly concave, green, twisted to contorted, erect-spreading when wet, not decurrent; apex acute, sometimes apiculate; margins plane or revolute to mid-leaf; costa percurrent in proximal leaves to sometimes excurrent in short slender awn in distal leaves; limbidium somewhat indistinct, 1--2 rows; distal and median laminal cells rhomboidal, 12--16 \um wide, mostly 3--4:1, thin-walled, proximal laminal cells gradually rectangular, 3--4:1, same width or sometimes wider than more distal cells. Specialized asexual reproduction of filiform green or green-brown gemmae in leaf axils. Sexual condition autoicous. Seta 2--3 cm, straight to flexuose, brown. Capsule  pyriform, 2--3 mm, symmetric, brown, mouth yellow; peristome reduced,  exostome teeth yellow basally, hyaline distally, lamellae usually straight in mid-tooth, pores lacking along exostome mid-line, endostome variable, basal membrane low, not adherent to exostome, segments with narrowly ovate perforations, sometimes reduced, cilia variable, absent or occasionally present and well developed, appendiculate or nodose; operculum low-conic, apiculate. Spores green, finely papillose, (28--)30--40 \um.

 

Capsules mature Jun-Aug. Rare on soil over serpentine rock or in serpentine rock crevices, of conservation concern; 0--100 m; Del., Md.

 

Ptychostomum reedii is a highly distinctive and very rare local endemic, not likely to be confused with any other species in eastern North America. Material from Delaware has abundant filiform gemmae in the leaf axils.

 

16. Ptychostomum salinum (Hagen ex Limpricht) J. R. Spence, Phytologia 87: 22. 2005

 

Bryum salinum Hagen ex Limpricht, Laubm. Deutschl. 2: 334. 1892

 

Plants in dense or open turfs, green or yellow-green. Stems 1--2(--3) cm, fertile leaves comose, innovations comose or shortly elongate and evenly foliate; not strongly radiculose. Leaves of fertile shoots and innovations similar, ovate-lanceolate, (0.5--)1--2(--3) mm, enlarged towards stem apex, flat to weakly concave, green, twisted to contorted, erect-spreading when wet, weakly decurrent; apex acuminate; margins revolute to mid-leaf or near apex; costa excurrent in short to long colored more or less smooth awn; limbidium strong, in 2--3 rows; distal and median laminal cells rhomboidal, 16--20 \um wide, mostly 2--3:1, mostly thin to firm-walled, proximal laminal cells gradually rectangular, 3--4:1, same width or sometimes wider than more distal cells. Specialized asexual reproduction absent. Sexual condition synoicous. Seta 1--2(--3) cm, straight to flexuose, red or red-brown. Capsule  elongate-pyriform, 2--4 mm, symmetric, brown, mouth yellow; peristome well developed,  exostome teeth yellow basally, hyaline distally, lamellae usually straight in mid-tooth, 1--2(--3) small pores present along exostome mid-line at base of tooth, endostome with high basal membrane, not adherent to exostome, about 1/2\x height of exostome, segments with broadly ovate perforations, cilia short, rudimentary; operculum tall conic, apiculate. Spores pale yellow-brown, finely papillose, variable in size, (18--)22--30 \um.

 

Capsules mature Jul-Sept. Rare and local on damp soil near the ocean in Arctic tundra; 0--500 m; Greenland; N.W.T., Man., Nfld. and Labr., Nunavut, Yukon; Alaska; Eurasia.

 

Ptychostomum salinum is a circumpolar Arctic-boreal species similar to P. creberrimum and P. pallescens in habit, but is smaller and has a strongly reduced peristome.  The small pores at the base of the mid-line of the exostome teeth and the large spores are diagnostic.  Spore size varies considerably in the some capsules for unknown reasons.

 

XXb. Ptychostomum subg. Ptychostomum

 

Stems mostly evenly foliate, leaves sometimes enlarged distally; sparsely to moderately radiculose. Leaves ovate, ovate-lanceolate or orbicular, leaf base green, or if red then same color as rest of leaf, purple, pink or red sap often present in cell lumens; limbidium usually present, narrow, yellowish, partly 2-stratose in proximal half; proximal laminal cells longer than the medial, rectangular, cells below alar region not inflated or pink. Sexual condition dioicous, synoicous, autoicous or polyoicous, often variable within species. Capsule shape variable, ovate, pyriform, clavate, sometimes curved; peristome endostome occasionally fragile and poorly developed, segments with narrow linear to ovate perforations. Spores variously colored yellow, green or brown, smooth to finely papillose, small to large, 10--40 \um.

 

Species ca. 20 (15 in the flora): common on wet soil in fens and wetlands or along streams and around lakes in Arctic and boreal regions of the Northern Hemisphere.

 

Species of subg. Ptychostomum are distinctive in a number of characters, especially the 2-stratose margin and the unusual color of the sap in cells of the leaf base.  These cells tend to be green, rarely evenly pink or red, but the cell sap is colored, often a bright purple, pink, orange or red.  Usually this occurs as irregular and randomly distributed patches of cells with colored sap. Subgenus Cladodium does not exhibit this feature. There are two groups of species in subg. Ptychostomum, those with moniocous sexual condition, reduced peristomes and large spores, and those that are dioicous, have more or less well-developed peristomes, and small spores.

 

1. Stems mostly less than 3 cm; monoicous; peristome reduced, cilia short or absent; spores generally large, mostly greater than 24 \um.

2. Autoicous; leaf with indistinct 1-stratose limbidium, apex broadly obtuse, costa not reaching apex ……………………… 23. Ptychostomum marrattii

2. Autoicous, synoicous or polyoicous; leaf with distinct partially 2-stratose limbidium, apex acute to acuminate, costa not reaching apex to more commonly percurrent or excurrent.

3. Autoicous or synoicous; endostome often fragile, sometimes adherent to exostome, cilia absent; spores 18--30 \um; capsules bright red at maturity, short-ovate, short pyriform to globose.

4. Synoicous; Capsules globose to short-pyriform; leaves ovate, acute; spores 18--22 \um ...................................................................... 31. Ptychostomum wrightii

4. Autoicous; Capsule short-ovate, leaves ovate-lanceolate, long-acuminate; spores 25--30 \um ...................................................................... 17. Ptychostomum acutiforme

3. Autoicous or polyoicous; endostome free to weakly adherent to exostome at base, cilia short or absent; capsules brown or red-brown at maturity or if red then spores greater than 30 \um; capsules clavate to pyriform; spores greater than 25 \um.

5. Autoicous; capsule short-ovate, red, red-brown or brown at maturity, apophysis thickened ....................................…............ 19. Ptychostomum calophyllum

5. Polyoicous; capsule pyriform to elongate ovate, yellow-brown to brown at maturity, apophysis slender.

6. Polyoicous; capsule short-pyriform; spores 38--45 \um .……....…. 29. Ptychostomum warneum

6. Autoicous or polyoicous; capsule elongate-pyriform to clavate; spores 24--35 \um.

7. Autoicous; plants green or yellow-green; costa of rosette leaves percurrent to short excurrent in stout awn; capsule elongate-pyriform, somewhat curved-gibbous, exostome teeth distally yellow; spores 28--35 \um …………………..….………………..… 20. Ptychostomum cernuum

7. Polyoicous, many gametangia either male or female; plants often red or pink; costa of rosette leaves distinctly excurrent in medium-length awn; capsule elongate-pyriform to clavate, more or less symmetric to somewhat curved, exostome teeth distally hyaline; spores 24--30 \um ............................. 18. Ptychostomum arcticum

1. Stems mostly (1--)2--10 cm; dioicous; peristome mostly perfect; spores small, less than 20 \um.

8. Leaves longly and broadly decurrent, apex acute, costa percurrent

.............................................. 30. Ptychostomum weigelii

8. Leaves not or somewhat decurrent, apex obtuse, acute or acuminate, costa not reaching apex to excurrent.

9. Leaves broadly ovate to suborbicular, apex of young leaves rounded to obtuse, costa not reaching apex to percurrent.

10. Plants red, leaves strongly concave, costa often percurrent; spores 15--20 \um ………… 21. Ptychostomum cryophilum

10. Plants bright green, lacking red tints, leaves weakly concave to flat, costa not reaching apex; spores 12--16 \um ……...…………………………………......… 22. Ptychostomum cyclophyllum

9. Leaves narrowly ovate to ovate-lanceolate, apex of young leaves acute, costa typically percurrent to short-excurrent.

11. Leaves yellow, green to brown-green, not decurrent, leaf margins mostly plane; filiform gemmae lacking; capsule pyriform or turbinate, not curved.

14. Leaves green to brown-green, strongly shrunken and contorted when dry, narrowly ovate to broadly ovate-lanceolate, median laminal cells 15--25(--30) \um wide; stems 1--3(--4) cm ............................................................ 28. Ptychostomum  turbinatum

14. Leaves bright yellow-green, somewhat imbricate when dry, broadly ovate, median laminal cells (20--)30--40(--50) \um wide; stems 3--6(--8) cm ………....…..... 27. Ptychostomum schleicheri

11. At least young leaves pink or red-tinged, somewhat decurrent, leaf margins recurved; filiform gemmae sometimes present in axils of distal leaves; capsule long-clavate, somewhat curved.

12. Laminal cells strongly incrassate; porose; filiform gemmae common in leaf axils; endostome basal membrane low, cilia short …............… 26. Ptychostomum rutilans

12. Laminal cells thin-walled, not porose; filiform gemmae rarely present; endostome basal membrane about 1/2\x height of exostome, cilia usually long, well developed …13

13. Capsule less than 4 mm, weakly curved to contorted when mature, exothecial cells short and broad, 1--1.5:1, 30--50  x 20--40 \um; spores 20--30 \um …...…….. 25. Ptychostomum pallens

13. Capsule often greater than 5 mm, strongly curved to contorted when mature, exothecial cells elongate, 2--3:1, 40--80 x 14--25 \um; spores 14--20 \um ….....……….…....…… 24. Ptychostomum meesioides

 

 

17. Ptychostomum acutiforme (Limpricht) J. R. Spence, Phytologia 89: 112. 2007

 

Bryum acutiforme Limpricht, Tromsø Mus. Aarsheft 21--22: 156. 1901

 

Plants in dense or open turfs, green, yellow-green to brown. Stems 1--2 cm, fertile leaves comose, innovations evenly foliate, leaves somewhat distant. Leaves 0.5--2(--3) mm, ovate-lanceolate, acuminate, flat to more often concave, weakly contorted to somewhat imbricate when dry, erect-spreading when wet, yellow-green to yellow-brown, leaf base red near costa, green otherwise, not decurrent; apex acute to acuminate; margins plane; limbidium moderately strong, (1--)2--3 rows, partially 2-stratose from base to mid-leaf; costa red to red-brown, not reaching apex to percurrent, short, slender apiculus sometimes present; distal laminal cells short-rhomboidal, 18--25 \um wide, mostly 2--3:1, thin-walled to incrassate, median cells somewhat longer, incrassate, proximal laminal cells narrower more distal cells, long-rectangular, (3--)4--5:1. Specialized asexual reproduction absent. Sexual condition autoicous. Seta moderately long, 2--4 cm, stout, straight to somewhat flexuose, red-brown. Capsule short, ovate, with thick apophysis abruptly contracted into seta, 2--3 mm, symmetric, red-brown or red, mouth yellow to yellow-orange; peristome reduced, exostome teeth yellow basally, hyaline distally, lacking pores along mid-line, lamellae more or less straight in middle of tooth, basal membrane low, less than 1/2\x height of exostome, not adherent to exostome, segments with narrow, lanceolate to slit-like perforations, cilia absent or rudimentary; operculum conic, apiculate. Spores yellow or brown, finely papillose, size often variable in same capsule, 28--32 \um. 

 

Capsules mature Jul-Aug. Rare on moist gravelly to sandy soil, near the sea or along streams and around lakes, Arctic-alpine; 0--600 m; Greenland; Nfld. and Labr., N.W.T., Nunavut; Alaska; n Eurasia.

 

Ptychostomum acutiforme is related to P. calophyllum, but differs in its ovate-lanceolate leaves with acuminate apices.

 

18. Ptychostomum arcticum (R. Brown) J. R. Spence, Phytologia 87: 20. 2005

 

Pohlia arctica R. Brown, Chlor. Melvill. 38. 1823; Bryum arcticum (R. Brown) Bruch & Schimper; B. bryoides (R. Brown) Ångstrom in Fries; B. purpurascens (R. Brown) Bruch, Schimper & W. Gümbel

 

Plants in dense or open turfs, red, green or yellow-green. Stems 0.5--3 cm, fertile leaves comose, innovations evenly foliate, leaves crowded. Leaves 0.5--3 mm, often gradually enlarged towards stem apex, ovate-lanceolate, flat, strongly contorted to shrunken when dry, shiny yellow-green to red or pink, leaf base usually green, not decurrent; apex acute to acuminate; margins revolute proximally; limbidium strong, of 2--3 rows, partially 2-stratose from base to mid-leaf; costa strong, excurrent in a medium to long smooth or denticulate awn; distal and median laminal cells short-rhomboidal, 15--25 \um wide, mostly 2--3:1, thin to moderately thick walled, proximal laminal cells narrower more distal cells, long-rectangular, 3--5:1. Specialized asexual reproduction absent. Sexual condition polyoicous, variable, often populations either synoicous or only archegoniate. Seta 1--3 cm, slender, straight to somewhat flexuose, brown or red-brown. Capsule highly variable, from elongate-pyriform to clavate, 3--5 mm, symmetric to somewhat curved, brown, mouth orange-brown to red; peristome variable, reduced, exostome teeth yellow or brown, hyaline distally, sometimes with small pores near base along mid-line, lamellae straight, basal membrane low to about 1/2\x height of exostome, weakly adherent to exostome, segments with narrow slit-like perforations, cilia absent or rudimentary; operculum low-conic, weakly apiculate. Spores yellow, finely papillose, size often variable in same capsule, 24--30(--32) \um. 

 

Capsules mature Jul-Aug. Common on moist soil banks and wet soil; Arctic-alpine; 0--4500 m; Greenland; Alta., B.C., Man., Nfld. and Labr., Que.; Alaska, Calif., Mont.,  Nev., Oreg., Wash., Wyo.; Eurasia

 

Ptychostomum arcticum is a widespread, circumpolar Arctic-alpine species, common in Arctic tundra. It can be confused with other pink-red species, including P. cryophilum, P. pallens, and P. rutilans.  Its smaller more or less flat and often shiny red leaves, polyoicous sexual condition and large spores distinguish it from these other species.  Ptychostomum arcticum is one of the most variable and confusing species in the genus. Much of this variation has been described as a series of very closely related species, including P. bryoides and P. purpurascens, differing in minor details of the leaves and capsule. D. T. Holyoak (2004) synonymized several of these species with P. arcticum, which I have tentatively accepted. More work is needed, especially a combination of quantitative morphology and molecular studies, to determine the true status of some of these segregates. The segregates that have been reported from the Flora region are keyed out below based on E. Nyholm (1993).  Their distribution and ecology remain poorly understood, but P. purpurascens is reported from Arctic North America, while P. bryoides is reported from Greenland. Capsules are needed for identification.

 

1. Distal lamina cells wide, 22--35 \um; capsule elongate-pyriform, symmetric, exothecial cells below mouth in 2--4 rows, transversely rectangular, exostome teeth with small holes along mid-line at base ........................................................... Ptychostomum purpurascens (R. Brown ) J. R. Spence

1. Distal lamina cells narrower, 17--24 \um; capsule clavate to ovate, symmetric or curved, exothecial cells quadrate, in 2--6 rows, exostome teeth lacking small holes.

2. Autoicous; hairpoint denticulate; capsule symmetric, exothecial cells orange, in 5--6 rows below mouth

................................................................................. Ptychostomum bryoides (R. Brown) J. R. Spence

2. Synoicous; hairpoint smooth; capsule curved, exothecial cells brown to orange, in 2--3 rows below

mouth ...................................................................... Ptychostomum arcticum (R. Brown) J. R. Spence

 

 

19. Ptychostomum calophyllum (R. Brown) J. R. Spence, Phytologia 87: 20. 2005

 

Bryum calophyllum R. Brown, Chlor. Melvill. 38. 1823; B. axel-blyttii H. Philibert

 

Plants in dense or open turfs, green, yellow-green to brown. Stems 1--3 cm, fertile leaves comose, innovations evenly foliate, leaves somewhat distant. Leaves 0.5--2.5(--3) mm, broadly ovate, flat to more often concave, weakly contorted to somewhat imbricate when dry, erect-spreading when wet, yellow-green to yellow-brown, leaf base red near costa, green otherwise, not decurrent; apex obtuse to broadly acute; margins mostly plane; limbidium moderately strong, (1--)2--3 rows, partially 2-stratose from base to mid-leaf; costa red to red-brown, not reaching apex to percurrent, short blunt apiculus sometimes present; distal laminal cells short-rhomboidal, 12--25 \um wide, mostly 2--3:1, incrassate, median cells somewhat longer, incrassate, proximal laminal cells narrower than more distal cells, long-rectangular, (3--)4--5:1. Specialized asexual reproduction absent. Sexual condition autoicous. Seta  moderately long, 2--4 cm, stout, straight to somewhat flexuose, red-brown. Capsule short, ovate, with thick apophysis abruptly contracted into seta, or rarely short-pyriform with a slender neck, 2--3 mm, symmetric, yellow-brown, mouth yellow to yellow-orange; peristome reduced, exostome teeth red to red-brown or yellow-brown basally, hyaline distally, mostly lacking pores or rarely a few present at base of tooth, lamellae curved in middle of tooth, basal membrane low, less than 1/2\x height of exostome, not adherent to exostome, segments with narrow lanceolate to slit-like perforations, cilia absent or rudimentary; operculum low-conic and apiculate. Spores yellow or brown, finely papillose, size often variable in same capsule, (28--)30--40 \um. 

 

Capsules mature Jul-Aug. Rare, moist gravelly to sandy soil, near the sea or along streams and around lakes, Arctic-alpine; 0--1000 m, Greenland; N.W.T., Nunavut, Que., Labr.; Alaska; n Eurasia.

 

Ptychostomum calophyllum is a distinctive species, with the short-ovate capsule and ovate leaves with a blunt apex and a strong limbidium. Sterile material from alpine regions of Colorado matches Arctic material and may belong here. This is a variable species with several segregates synonymized under P. calophyllum by D. T. Holyoak (2004).   Although I have tentatively accepted part of Holyoak's analysis, P. acutiforme seems quite distinct and is therefore retained in this treatment.  A key to the segregates reported from the Flora region, based on the work of E. Nyholm (1993), is included below. The variation, ecology and distribution of these species is poorly understood. Capsules are needed for identification. Ptychostomum axel-blytti is only known from Greenland.

 

1. Capsule yellow-brown to brown when mature, short-ovate with a thick neck ......... Ptychostomum calophyllum (R. Brown) J.R. Spence

1. Capsule red when mature, short-ovate with a thick neck to short-pyriform with a slender neck.

2. Leaves of fertile stems broadly ovate, more or less obtuse, concave, distal margins smooth; capsule short-pyriform, neck slender .......................................... Ptychostomum axel-blyttii (H. Philibert) J.R. Spence

2. Leaves of fertile stems ovate-lanceolate, acute, flat, distal margins denticulate; capsule short-ovate with a thick neck ......................................................................... Ptychostomum acutiforme (Limpricht) J.R. Spence

 

 

20. Ptychostomum cernuum Hornschuch, Flora 5, 2: syll. 62, 1822

 

Bryum uliginosum (Bridel) Bruch, Schimper & W. Gümbel

 

Plants in dense or open turfs, green or yellow-green. Stems 0.5--3 cm, fertile leaves comose, innovations evenly foliate, leaves crowded. Leaves 1--3.5(--4) mm, often gradually enlarged towards stem apex, ovate-lanceolate, flat, strongly contorted to shrunken when dry, green to yellow-green, leaf base green, not decurrent; apex acuminate; margins revolute proximally, plane distally; limbidium strong, of 2--3 rows, partially 2-stratose from base to mid-leaf; costa percurrent to short-excurrent as a stout awn; distal and median laminal cells short-rhomboidal, 18--22 \um wide, mostly 3--4:1, thin-walled, proximal laminal cells narrower than more distal cells, long-rectangular, 3--5:1. Specialized asexual reproduction absent. Sexual condition autoicous. Seta 2--4 cm, stout, straight to somewhat flexuose, yellow-brown or brown. Capsule highly variable, elongate-pyriform to clavate, (3--)4--6(--7) mm, somewhat to strongly curved, brown, mouth yellow-brown; peristome reduced, exostome teeth yellow throughout or rarely hyaline distally, lacking pores near base along mid-line, lamellae straight, basal membrane about 1/2\x height of exostome, not adherent to exostome, segments with ovate perforations, cilia absent or rudimentary; operculum conic, apiculate. Spores yellow to green, finely papillose, 28--32(--35) \um. 

 

Capsules mature Jun-Sep. Uncommon and scattered on wet soil along streams and in wetlands, often calcareous habitats, 0--3000 m; Greenland; Alta, BC, Labr. and Nfld., Man., N.W.T., Nunavut, Ont., Que., Sask., Yukon; Colo., Ill., N.Y., Iowa, Mich., Minn., Nebr., N.Mex, Colo, Pa., Ohio, N.Dak, S.Dak; s South America; Eurasia.

 

Ptychostomum cernuum is a circumpolar Arctic-boreal to north-temperate species and is characterized by its long ovate-lanceolate leaves with a short awn, green leaf base, autoicous sexual condition, and extremely long, curved capsule. It is related to P. pallens, but differs in the longer more strongly curved capsule, autoicous sexual condition, larger spores, and  yellow-green color.  Ptychostomum turbinatum is similar, and generally capsules are needed to separate these species.  Ptychostomum cernuum is autoicous, smaller and has longer narrower ovate-lanceolate leaves, while P. turbinatum is dioicous, often very large in stature, and has more broader, more ovate leaves. The species superficially resembles Pohlia elongata.

 

21. Ptychostomum cryophilum (Mårtensson) J. R. Spence, Phytologia 87: 20. 2005

 

Bryum cryophilum Mårtensson, K.V.A. Afh. Natursk. 15. 1956

 

Plants in dense swelling turfs, red to red- green or red-yellow. Stems 2--4(--6) cm, fertile leaves comose, innovations evenly foliate, leaves somewhat distant. Leaves 0.5--3 mm, not much enlarged towards stem apex, ovate, concave, younger leaves strongly contorted to shrunken when dry, yellow-green to pink or red, older leaves more or less imbricate, pink, red becoming dark red-brown when old, leaf base green, weakly decurrent; apex obtuse; margins plane or weakly revolute in lower 1/4 of margin; limbidium strong, of 2--3 rows, partially 2-stratose from base to mid-leaf; costa red, percurrent or sometimes not reaching apex; distal and median laminal cells irregularly short-rhomboidal, 15--22(--25) \um wide, mostly 2--3:1, thin-walled, proximal laminal cells narrower than more distal cells, rectangular, 3--4:1. Specialized asexual reproduction absent. Sexual condition dioicous. Seta single, 2--4 cm, slender, straight to somewhat flexuose, red. Capsule ovate to obovate, 2--4 mm, symmetric, yellow-brown to brown, mouth yellow-brown; peristome well developed, exostome teeth yellow-brown, lacking pores near base along mid-line, lamellae straight, basal membrane high, 1/2 to 2/3 height of exostome, not adherent to exostome, segments with ovate perforations, cilia long, appendiculate; operculum conic, weakly apiculate. Spores pale yellow, finely papillose, 16--20 \um. 

 

Capsules mature Jul-Aug. Locally common on wet soil or rocks in streams, wetlands, and late melting snow beds; Arctic-alpine; 0--2000 m; Greenland; Alta, B.C., N.W.T., Nunavut, Yukon; Alaska.; Eurasia (Arctic regions, India, Mongolia, Nepal).

 

Densely compact red turfs, ovate concave leaves, and a weak costa are diagnostic of Ptychostomum cryophilum.

 

22. Ptychostomum cyclophyllum (Schwägrichen) J. R. Spence, Phytologia 87: 20. 2005

 

Mnium cyclophyllum Schwägrichen,  Sp. Musc. Frond., Supple. 2(2): 160, plate 194. 1827; Bryum cyclophyllum (Schwägrichen) Bruch, Schimper & W. Gümbel; B. tortifolium Bridel

 

Plants in dense or open turfs, green or yellow-green. Stems 0.5--3(--4) cm, fertile leaves comose, innovations evenly foliate, leaves somewhat distant. Leaves 0.5--3 mm, not much enlarged towards stem apex, broadly ovate to orbicular, flat or weakly concave, strongly contorted to shrunken when dry, green, leaf base green, weakly decurrent; apex broadly acute to obtuse; margins plane; limbidium strong, of 2--3 rows, partially 2-stratose from base to mid-leaf; costa not reaching apex, rarely nearly percurrent; distal and median laminal cells short-rhomboidal, 18--24 \um wide, mostly 2--3:1, thin-walled, proximal laminal cells long-rectangular, 3--5:1. Specialized asexual reproduction rare, of brown finely papillose filiform gemmae in leaf axils. Sexual condition dioicous. Seta 2--4(--5) cm, slender, straight to somewhat flexuose, red or brown. Capsule obovate, 2--4 mm, symmetric, yellow-brown to brown, mouth yellow; peristome well developed, exostome teeth yellow basally, hyaline distally, lacking pores near base along mid-line, lamellae straight, basal membrane about 1/2\x height of exostome, not adherent to exostome, segments with ovate perforations, cilia long, appendiculate; operculum low-conic, weakly apiculate. Spores pale yellow, smooth or finely papillose, 14--16 \um.

 

Capsules mature Jun-Sep. Locally common on wet sandy or organic soil along streams and in wetlands; 0--3000 m; Greenland; Alta, B.C., Nfld. and Labr., N.W.T., Nunavut, Yukon; Alaska, Ariz., Calif., Colo., Mich., Mont., N.J., N.Mex., N.Y., Utah, Wash., Wyo.; Eurasia (India and Nepal).

 

Ptychostomum cyclophyllum is a distinctive Arctic-boreal species with broadly ovate to orbicular, blunt, more or less flat leaves that are strongly shrunken when dry.  Ptychostomum cryophilum is almost always reddish, with much more concave leaves that do not shrink much when dry. Ptychostomum neodamense and P. ovatum are similar, but have much shorter proximal lamina cells, 1-stratose borders, and red leaf bases. Ptychostomum cyclophyllum has probably declined in the U.S., as it grows in wetlands and along streams in mid-elevation regions, areas where human development is often concentrated.

 

23. Ptychostomum marrattii (Hooker & Wilson) J. R. Spence, Phytologia 87: 21. 2005

 

Bryum marratii Hooker & Wilson, Bryol. Brit., 32b. 1855

 

Plants in dense or open turfs, yellow-green to-yellow brown. Stems 0.5--2 cm, fertile leaves comose, innovations evenly foliate, leaves crowded. Leaves (0.5--)1--2 mm, not much  enlarged towards stem apex, ovate, concave, somewhat contorted to shrunken when dry, yellow-green to green above, becoming brown in older leaves, leaf base green; apex broadly acute to obtuse; margins plane; limbidium absent, or rarely a weak single row present, 1-stratose; costa not reaching apex; distal and median laminal cells short-rhomboidal, 18--25 \um wide, mostly 2--3:1, thin-walled, proximal laminal cells irregularly rectangular to rhomboidal, somewhat bulging, 3--4:1. Specialized asexual reproduction absent. Sexual condition autoicous. Seta long, 2--4(--5) cm, stout, somewhat flexuose, red-purple. Capsule pyriform, 2--3 mm, symmetric, yellow-brown to red-brown, mouth yellow-brown to red-brown; peristome reduced, exostome teeth red-brown basally, yellow distally, lacking pores near base along mid-line, lamellae straight, basal membrane about 1/2\x height of exostome, weakly adherent to exostome, segments narrowly perforate, cilia absent or rudimentary; operculum high conic. Spores yellow, finely papillose, (30--)32--40 \um. 

 

Capsules mature Jul-Sep. Rare and local on moist soil, 0--1000 m; Greenland; Alta; N.Dak., Wash.; nw Europe.

 

Ptychostomum marrattii is a distinctive Arctic-boreal species with oblong, more or less obtuse green leaves, which lack a limbidium. It is found typically along the coasts of northern Europe. Two of the three North American collections are from interior regions.

 

24. Ptychostomum meesioides (Kindberg) J. R. Spence, Phytologia 87: 21. 2005

 

Bryum meesiodes Kindberg, Bull. Torr. Bot. Club 16: 95. 1889

 

Plants in dense or open turfs, green or yellow-green, sometimes reddish below. Stems 0.5--1(--2) cm, fertile stems evenly foliate, sometimes comose, innovations evenly foliate, leaves crowded. Leaves 1--3(--3.5) mm, often gradually enlarged towards stem apex, ovate-lanceolate, flat, strongly contorted to shrunken when dry, green to yellow-green or rarely reddish, leaf base green, somewhat to strongly and narrowly decurrent; apex acuminate; margins revolute proximally, plane distally; limbidium strong, of 2--3 rows, partially 2-stratose from base to mid-leaf; costa percurrent to short-excurrent as a stout awn; distal and median laminal cells short-rhomboidal, 16--25 \um wide, mostly 3--4:1, thin-walled, proximal laminal cells narrower than more distal cells, long-rectangular, 3--5:1. Specialized asexual reproduction absent. Sexual condition dioicous. Seta 2--4 cm, stout, straight to often flexuose or geniculate, red-brown. Capsule highly variable, elongate-pyriform to clavate, (2--)3--5 mm, strongly curved and asymmetric, brown, mouth yellow-brown, median exothecial cells elongate, 40--85 \um long, many cells 3--4:1; peristome well developed, exostome teeth yellow or orange basally, hyaline distally, lacking pores near base along mid-line, lamellae straight, basal membrane about 1/2\x height of exostome, not adherent to exostome, segments with ovate perforations, cilia present, appendiculate; operculum conic, apiculate. Spores yellow to green, finely papillose, 14--20 \um. 

 

Capsules mature Jun-Sep. Uncommon and scattered on wet soil along streams and in wetlands, often calcareous, 0--3000 m; BC; Alaska, Idaho, Wash.

 

Ptychostomum meesioides is an Arctic-boreal to north temperate species, narrowly endemic in primarily coastal hyperoceanic areas of western North America. It is very similar to P. pallens, although somewhat smaller and green-colored rather than the typical pink coloration of the latter. The diagnostic characters are those of the sporophyte, primarily the strongly curved capsules and elongate median exothecial cells. The median exothecial cells of P. pallens are shorter, 1--2:1.  Capsules are needed for identification.

 

25. Ptychostomum pallens (Swartz) J. R. Spence, Phytologia 87: 21. 2005

 

Bryum pallens Swartz, Monthly Rev. 34: 538. 1801

 

Plants in dense or open turfs, pale pink, red or rarely dull green. Stems 1--4(--6) cm, fertile leaves comose, innovations evenly foliate, leaves crowded. Leaves 1--3.5(--4) mm, often gradually enlarged towards stem apex, ovate to broadly ovate-lanceolate, flat or weakly concave, contorted to shrunken when dry, pink, red or rarely green, leaf base pink or green, somewhat to strongly and narrowly decurrent; apex acuminate; margins revolute proximally, plane distally; limbidium strong, of 2--3 rows, partially 2-stratose from base to mid-leaf; costa percurrent to short-excurrent as a stout awn, red to red-brown; distal and median laminal cells short-rhomboidal, 17--24 \um wide, mostly 2--3:1, thin-walled, not porose, proximal laminal cells short-rectangular, 2--3(--4):1. Specialized asexual reproduction rare, of brown-red to green finely papillose filiform gemmae in leaf axils. Sexual condition dioicous. Seta 2--4(--6) cm, stout, straight to somewhat flexuose, red-brown. Capsule elongate-pyriform to clavate, (3--)4--5 mm, somewhat curved, yellow-brown, mouth yellow-brown, median exothecial cells short-rectangular to quadrate, 20--40 \um long, 1--2:1; peristome mostly well-developed, exostome teeth yellow basally, hyaline distally, lacking pores near base along mid-line, lamellae straight, basal membrane about 1/2\x height of exostome, not adherent to exostome, segments with ovate perforations, cilia appendiculate to nodose, occasionally short; operculum conic, apiculate. Spores yellow to green, finely papillose, 16--20(--22) \um. 

 

Capsules mature Jun-Sep. Common on wet soil along streams and in wetlands, usually calcareous, 0--3000 m; Greenland; Alta., B.C., Man., N.B., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont., Que., Sask., Yukon,; Alaska, Ariz., Calif., Colo., Idaho, Maine, Mich., Minn.,  Mont., Nev., N.Mex., N.Y., Oreg., Utah, Wash., Wyo.; s South America; Eurasia.

 

Ptychostomum pallens is a highly variable Arctic-boreal to north temperate species, related to P. rutilans, P. meesioides and P. cernuum.  The pale pink color is diagnostic, but is not always present. From P. rutilans it can be distinguished by its much thinner walled non-porose laminal cells and well developed peristome; from P. meesiodes by its shorter less arcuate capsule and short exothecial cells; and from P. cernuum by its dioicous sexual condition.

 

26. Ptychostomum rutilans (Bridel) J. R. Spence, Phytologia 87: 23. 2005

 

Bryum rutilans Bridel, Bryol. Univ. 1: 684. 1826; B. aeneum Blytt ex Bruch, Schimper & W. Gümbel

 

Plants in dense swelling turfs, red to red-brown. Stems 2--4(--6) cm, fertile leaves comose, innovations evenly foliate, leaves crowded; often strongly radiculose. Leaves 1--3.5(--4) mm, often gradually enlarged towards stem apex, ovate to broadly ovate-lanceolate, concave, contorted to shrunken when dry, red, becoming dark red-brown with age, leaf base uniformly red, not or weakly  decurrent; apex acute; margins revolute proximally, plane distally; limbidium strong, of 2--3 rows, partially 2-stratose from base to mid-leaf; costa percurrent to short-excurrent as a stout awn, red to red-brown; distal and median laminal cells short-rhomboidal, 12--18 \um wide, mostly 2--3:1, incrassate and porose, proximal laminal cells rectangular, 3--4:1. Specialized asexual reproduction uncommon, of brown-red papillose filiform gemmae in leaf axils. Sexual condition dioicous. Seta 1--3(--4) cm, stout, straight to somewhat flexuose, red-brown. Capsule elongate-pyriform, long-necked, 3--5 mm, somewhat curved, brown, mouth yellow-brown; peristome reduced, exostome teeth yellow basally, hyaline distally, lacking pores near base along mid-line, lamellae straight, basal membrane low, less than 1/2\x height of exostome, not adherent to exostome, segments with ovate perforations, cilia reduced or absent; operculum conic, apiculate. Spores yellow to green, finely papillose, 16--22(--24) \um. 

 

Capsules mature Jul-Aug. Uncommon on wet soil in tundra and along streams and in wetlands, 0--2000 m; Greenland; N.W.T., Nunavut; Alaska; n Eurasia.

 

Ptychostomum rutilans is a distinctive Arctic species with dark red shoots, usually forming dense turfs on wet soil. It is close to P. pallens, but differs in the incrassate porose lamina cells, and reduced peristome.  Small specimens are similar to Rosulabryum subelegans, which generally has green-red spirally twisted leaves and mostly grows on rock in alpine regions of northwestern Europe.  D. T. Holyoak (2004) considered them the same species because the type of R. subelegans also has bistratose borders.  I prefer to keep them separate until more detailed studies are completed. If they are conspecific, then the name Bryum (Ptychostomum) aeneum Blytt ex Bruch, Schimper & W. Gümbel should be used for the plants in North America.

 

27. Ptychostomum schleicheri (Schwägrichen) J. R. Spence, Phytologia 87: 22. 2005

 

Bryum schleicheri Schwägrichen, Sp. Musc., Suppl. 1, 2: 113, plate 73. 1816; B. schleicheri Schwägrichen var. latifolium (Schwägrichen) Schimper

 

Plants in dense or open turfs, pale yellow, yellow-green or copper. Stems 3--6(--10) cm, fertile leaves comose, innovations evenly foliate, leaves crowded; weakly radiculose. Leaves 2--4(--5) mm, not much enlarged towards stem apex, broadly ovate, weakly to strongly concave, upper leaves contorted to shrunken when dry, lower leaves more or less imbricate, yellow to yellow-copper, leaf base green, not or weakly decurrent; apex acute; margins plane or revolute proximally, plane distally; limbidium moderately strong, of 1--3 rows, partially 2-stratose from base to mid-leaf; costa percurrent to short-excurrent as a slender awn, pale brown to red-brown; distal and median laminal cells short-rhomboidal, 28--36 \um wide, mostly 2:1, thin-walled, proximal laminal cells rectangular, 3:1. Specialized asexual reproduction none. Sexual condition dioicous. Seta 2--3(--4) cm, slender, straight to somewhat flexuose, red-brown. Capsule turbinate to pyriform, 3--5 mm, symmetric, yellow-brown, mouth yellow; peristome well-developed, exostome teeth yellow basally, hyaline distally, lacking pores near base along mid-line, lamellae straight, basal membrane about 1/2\x height of exostome, not adherent to exostome, segments with ovate perforations, cilia appendiculate to nodose; operculum conic, rounded. Spores yellow to brown, finely papillose, 16--20(--22) \um. 

 

Capsules mature Jul-Aug. Uncommon on wet soil along streams and on seepy tundra slopes and late snowmelt areas, 0--4000 m; Greenland; Alta., B.C.,  N.W.T., Nunavut, Yukon; Alaska, Calif., Idaho, Mont., Ore., Wash.; Eurasia.

 

Ptychostomum schleicheri is an Arctic-alpine species. When well developed it is distinctively large, with long stems, and crowded yellowish, somewhat imbricate leaves. Much material named P. schleicheri appears to be misidentified and can be referred to P. turbinatum. Our material conforms to the var. latifolium.  The relationships between var. schleicheri  and var. latifolium and with P. turbinatum, have not been worked out yet.  This species is atypical for Ptychostomum, and shows some morphological similarities to the Mniaceae, especially in the lack of tmema cells in the rhizoidal filiform gemmae, extremely wide laminal cells, and the pronounced shelf-like leaf base that remains attached to the stem after the leaf is removed.  Ptychostomum ovatum is similar but has a unistratose border, usually contorted and shrunken leaves when dry, and much smaller laminal cells.

 

28. Ptychostomum turbinatum (Hedwig) J. R. Spence, Phytologia 87: 22. 2005

 

Mnium turbinatum Hedwig, Sp. Musc. Frond., 191. 1801; Bryum turbinatum (Hedwig) Turner

 

Plants in dense or open turfs, dull green or yellow-green to brownish. Stems (1--)2--4(--6) cm, fertile leaves comose, innovations evenly foliate, leaves crowded; sparsely radiculose, sometimes clusters of rhizoids arising from proximal leaf axils. Leaves 1--3 mm, often gradually enlarged towards stem apex, broadly ovate-lanceolate, somewhat concave, strongly contorted to shrunken when dry, dull green to yellow-green, base green, not or weakly decurrent; apex acute to acuminate; margins plane, or occasionally weakly revolute proximally; limbidium strong, of 2--3 rows, partially 2-stratose from base to mid-leaf; costa percurrent to excurrent in a short slender awn; distal and median laminal cells short-rhomboidal, 18--25(--30) \um wide, mostly 2--3:1, thin-walled, not porose, proximal laminal cells narrower more distal cells, rectangular, 3--4:1, thin-walled, not porose. Specialized asexual reproduction absent. Sexual condition dioicous. Seta single, 1--3(--4) cm, slender, straight to somewhat flexuose, brown or red-brown. Capsule turbinate to short-pyriform, 3--5 mm, symmetric, pale yellow to yellow-brown, becoming black with age, mouth yellow; peristome well-developed, exostome teeth yellow to orange basally, hyaline distally, lacking pores near base along mid-line, lamellae straight, basal membrane about 1/2\x height of exostome, not adherent to exostome, segments with ovate perforations, cilia appendiculate to nodose; operculum conic, apiculate. Spores yellow to brown, finely papillose, 18--22 \um.

 

Capsules maturing Jun-Aug. Locally common on wet soil in calcareous wetlands, 0--3500 m; Greenland; Alta., B.C.,  Nfld. and Labr., N.W.T., Yukon; Alaska, Calif., Colo., Idaho, Mont., Nev., Oreg., Utah, Wash., Wyo.; South America (Argentina, Chile, Ecuador, Peru); Eurasia (including Turkey), Africa.

 

Ptychostomum turbinatum is a circumpolar Arctic-boreal to temperate species. The dull green to yellow-green shoots with strongly shrunken ovate-lanceolate leaves, and the turbinate capsule are diagnostic. Old capsules often turn black, and become strongly constricted just below the mouth. Ptychostomum schleicheri is much larger, pale yellow distally, with wider leaves and much wider upper lamina cells. Large specimens, 8--12 cm, with long narrow ovate-lanceolate leaves and elongate-pyriform capsules, represent an undescribed species, which will be published at a future date.

 

29. Ptychostomum warneum (Röhl) J. R. Spence, Phytologia 87: 22. 2005

Mnium caespiticium var. warneum Röhl,  Deutschl. Fl. (ed. 2), Kryptog. Gew. 3: 94. 1813;

Bryum warneum (Röhl) Bridel

 

Plants in dense or open turfs, red, green or yellow-green. Stems 0.5--2(--3) cm, fertile leaves comose, innovations evenly foliate, leaves crowded; sparsely radiculose. Leaves 0.5--2 mm, often gradually enlarged towards stem apex, ovate-lanceolate, flat, strongly contorted to shrunken when dry, yellow-green to red, base usually green; apex acuminate; margins revolute proximally; limbidium strong, of 2--3 rows, partially 2-stratose from base to mid-leaf; costa short-excurrent into a slender smooth awn; distal and median laminal cells short-rhomboidal, 18--22 \um wide, mostly 2--3:1, thin-walled, proximal laminal cells rectangular, 3--4:1. Specialized asexual reproduction absent. Sexual condition polyoicous, variable, some populations  mostly autoicous. Seta 2--4(--5), stout, straight to somewhat flexuose, purple or red. Capsule pyriform, 2--4 mm, symmetric, yellow-brown or brown, mouth yellow; peristome reduced, exostome teeth yellow or brown basally, pale yellow to hyaline distally, lacking pores near base along mid-line, lamellae straight, numerous, often joined by cross-walls, basal membrane about 1/2\x height of exostome, weakly adherent to exostome, segments narrowly perforate, cilia absent or rudimentary; operculum tall conic. Spores pale green or yellow, finely papillose, large, variable in same capsule (32--)36--48(--50) \um.

 

Capsules mature Jul-Aug. Rare on damp soil; 0--50 m; Greenland; Que.; n Eurasia (also Himalayas in Nepal).

 

Ptychostomum warneum is an Arctic-boreal species that can be distinguished from the related P. calophyllum by its pyriform brown capsule with a slender neck, generally larger spores and acuminate leaves. It is known from only one location in North America, in the Mingan Islands. Capsules are needed for identification.

 

30. Ptychostomum weigelii (Sprengel) J. R. Spence, Phytologia 87: 22. 2005

 

Bryum weigelii Sprengel, Mant. Prim. Fl. Hals. 55. 1807

 

Plants in loose low open turfs, green or rarely pinkish-red. Stems 2--4(--6) cm, fertile leaves comose, innovations evenly foliate, leaves distant; sparsely radiculose. Leaves 1--3 mm, not much enlarged towards stem apex, ovate-lanceolate, flat, strongly contorted to shrunken when dry, green, yellow-green, rarely red or pink, base usually green, strongly and broadly decurrent, decurrencies almost reaching next more proximal leaf; apex acute; margins revolute proximally, plane distally; limbidium weak, of 1(--2) rows, partially 2-stratose from base to mid-leaf; costa not reaching apex to short-excurrent as a slender awn, green or brown; distal and median laminal cells short-rhomboidal, 18--25(--30) \um wide, mostly 2--3:1, thin-walled, proximal laminal cells narrower more distal cells, hexagonal to rectangular, 3--4:1. Specialized asexual reproduction absent. Sexual condition dioicous. Seta single, 2--4 cm, slender, straight to somewhat flexuose, red-brown. Capsule turbinate to pyriform, 3--4 mm, symmetric, brown, mouth yellow; peristome well developed, exostome teeth yellow basally, hyaline distally, lacking pores near base along mid-line, lamellae straight, basal membrane about 1/2\x height of exostome, not adherent to exostome, cilia well developed, long, appendiculate; operculum convex, apiculate. Spores yellow or green, finely papillose, (12)14--18 \um. 

 

Capsules mature Jun-Aug. Common and widespread on wet soil in wetlands and along streams, 0--3500 m; Greenland; Alta., B.C., Man., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont., P.E.I., Que., Sask., Yukon; Alaska, Ariz., Calif., Colo., Conn., Idaho,  Maine, Mass., Mich., Minn., Mont.,  Nev., N.H.,  N.Dak., Ohio,  Oreg., Pa.,  Utah, Vt., Va., Wash., W.Va., Wis., Wyo.; South America; Eurasia; Africa; Pacific Islands (New Zealand); Australia.

 

Ptychostomum weigelii is a common Arctic-boreal to north temperate species in wet sites, easily identified by the broad, long marginal decurrencies of the leaves that nearly reach to the next leaf.  Leaves of the similar P. cyclophyllum and P. turbinatum are either not decurrent or only produce short very slender decurrencies. Colorado material is pinkish-red.

 

 

31. Ptychostomum wrightii (Sullivant & Lesquereux) J. R. Spence, Phytologia 87: 22. 2005

 

Bryum wrightii Sullivant & Lesquereux, Proc. Amer. Acad. Arts. Sci. 4: 278. 1860

 

Plants in dense or open turfs, red- green or red-brown. Stems 0.5--1(--2) cm, fertile leaves comose, innovations evenly foliate; sparsely radiculose. Leaves 0.5--2 mm, not enlarged towards stem apex, ovate, concave, somewhat contorted to shrunken when dry, yellow-green to red-green, base green, not decurrent; apex acute; margins revolute proximally; limbidium strong, of 2--3 rows, partially 2-stratose from base to mid-leaf; costa strong, percurrent to excurrent in short smooth awn; distal and median laminal cells short-rhomboidal, 18--25 \um wide, mostly 2--3:1, thin-walled, proximal laminal cells narrower more distal cells, long-rectangular, 3--5:1. Specialized asexual reproduction absent. Sexual condition synoicous. Seta  1--3 cm, slender, straight to somewhat flexuose. Capsule broadly ovate to pyriform, somewhat inflated, 2--3 mm, symmetric 2--3 mm, symmetric, yellow-red, red or red-brown, often shiny, mouth yellow-red; peristome reduced, exostome teeth yellow proximally, hyaline distally, lamellae somewhat curved below,  basal membrane low to rudimentary, not or weakly adherent to exostome, cilia absent; operculum low-conic, smooth or weakly apiculate. Spores yellow, finely papillose, 18--20(--22) \um. 

 

Capsules mature Jul-Aug. Locally common on damp calcareous soil and silt in tundra, 0--1500 m; Greenland; Labr., N.W.T., Nunavut, Que., Yukon; Alaska; Arctic Eurasia.

 

Ptychostomum wrightii is a small Arctic species with distinctive capsules that are somewhat inflated, shiny red, and ovate to pyriform, and a poorly developed fragile endostome.  Capsules are needed for identification.

 

11.. RHODOBRYUM (Schimper) Limpricht, Die Laubmoose Deutschlands, Oesterreichs und der Schweiz 2: 444. 1892  *  [Greek rhodo, rose, and Bryum, a moss genus]

John R. Spence

 

Plants large for the family, dark green or olive-green, sometimes with red tints. Stems secondary to 6 cm, erect, arising from wiry creeping stoloniferous primary stems, unbranched or sometimes innovations arising from below the terminal rosette; rhizoids sparse to abundant, at base of stem or arising as macronemata in leaf axils, micronemata lacking on leafy stems. Leaves small and scale-like proximally, becoming enlarged distally and crowded in a terminal rosette, 4--12 mm, strongly contorted and shrunken when dry, erect-spreading when moist; margins bordered by elongate thickened cells or sometimes border weak or absent, 1-stratose, strongly serrate from mid leaf to apex; distal and median laminal cells hexagonal to rhomboidal, 3--4:1, proximal laminal cells elongate-rectangular, longer than distal cells, alar cells not differentiated; costa typically strong, subpercurrent to more often percurrent to short excurrent as a short and often recurved hairpoint or apiculus, in cross section with 2--4 layers of enlarged guide cells, stereid band small or absent. Specialized asexual reproduction lacking. Sexual condition dioicous; inner perichaetial leaves somewhat differentiated, smaller and narrower than surrounding rosette leaves; inner perigonial leaves small, broad, over-arching enlarged disc-like pale perigonia with abundant paraphyses. Seta 1--8 per perichaetium, red or brown, long-exserted, 2--5 cm, straight to slightly flexuose. Capsule inclined to nutant, brown to red-brown, oblong to cylindric, 3--5 mm, slightly curved and narrowed to mouth, mouth sometimes oblique; operculum low-conic; peristome double, well-developed, exostome teeth lanceolate, acuminate, brown or yellow-brown proximally, hyaline near apex; endostome with high basal membrane, segments lanceolate to subulate, keeled and perforate, cilia 2-4, nodose to appendiculate. Spores 10--22(--25) \um, finely papillose.

 

Species ca. 25 (2 in the flora): worldwide in temperate to tropical regions, all continents except Antarctica.

 

Rhodobryum is a genus characterized by relatively large plants with leaves in a distinct rosette (at least in the region of the flora), stoloniferous primary stems, and costa with guide cells in two or more layers and stereid band reduced.  The chromosomes of Rhodobryum are distinctly different from those of Bryum and Rosulabryum (H. P. Ramsay and J. R. Spence 1996).  Like Roellia and Rosulabryum, more than one sporophyte can mature from the same perichaetium.  Roellia differs from Rhodobryum by its lack of stolons, less contorted shiny pale green leaves that are finely rugose, and much larger laminal cells.  Robust specimens of Rosulabryum andicola and R. canariense differ in their strongly developed stereid band with a single layer of guide cells, smaller leaves, lack of stolons, and presence of rhizoidal tubers and filiform leaf-axis gemmae.  Most species of Rhodobryum occur in the tropics, especially in montane regions.  Recent DNA research suggests that the genus, excepting only Leptostomopsis, is basal to and sister to the remainder of the Bryaceae.

 

SELECTED REFERENCES  Iwatsuki, Z. and T. Koponen. 1972. On the taxonomy and distribution of Rhodobryum roseum and its related species (Bryophyta). Acta Bot. Fennica 96: 1--22. Koponen, T., X. Li and M. Zang. 1982. A synopsis of Rhodobryum (Musci, Bryaceae) in China. Ann. Bot. Fennici 19: 75--80. Mohamed, M.A. Haji. 1984. A synopsis of the genus Rhodobryum in Asia. J. Hattori Bot. Lab. 55: 281--293. Ramsay, H. P. and J. R. Spence. 1996. Chromosome data on Australasian Bryaceae. J. Hattori Bot. Lab. 80: 251--270.

 

1. Rosette leaves typically 18--55 in number, leaf margins strongly revolute to mid leaf or well beyond, stereid band relatively well developed, reaching dorsal epidermal cells, without layer of thin-walled cells between.         1. Rhodobryum ontariense

 

1. Rosette leaves typically 15--21 in number, leaf margins weakly recurved to about mid-leaf, stereid band small, with at least one distinct layer of thin-walled cells before dorsal epidermal cells.     2. Rhodobryum roseum

 

1. Rhodobryum ontariense (Kindberg) Kindberg, Spec. Eur. N. Amer. Bryin. 2: 346. 1897

 

Plants 1--5 cm tall, mostly unbranched or rarely with slender sub-apical innovations. Stem leaves 4--10 mm, numerous in rosettes, from 18--55, typically more than 20; margins strongly  revolute to above mid leaf, often nearly to apex, apex broadly acute to cuspidate; costa percurrent to short-excurrent into a slender hairpoint in rosette leaves, in cross section with distinct stereid band, reaching dorsal epidermal layer without intervening thin-walled layer of cells; distal and median laminal cells hexagonal, 25--35 × 50--80 \um, (3--4:1), proximal cells longer, to 100 \um, rectangular. Inner perichaetial leaves with costa strong, long-excurrent in denticulate hairpoint. Spores 16--24 \um.

 

Capsules mature late winter--late fall (Feb.--Nov.). Common on rich soil in forests, along forest edges, on rotten logs, tree bases, soil over rock or rock, often calcareous, sometimes in boggy sites; 0--3000 m, restricted to higher elevations in southern latitudes; Alta, Man., N.B., Nfld., N.S., Ont., Que., Sask.; se Ariz., Ark., Conn., Del., D.C., n Ga., Ill., Ind., Iowa, Kans., Ky., Maine, Md., Mass., Mich., Minn., Mo., Nebr., N.H., N.J., N.Mex., N.Y., N.C., N.Dak., Ohio, Pa., R.I., n S.C., S.Dak., Tenn., w Tex., Vt., Va., W.Va., Wis.; Mexico; Eurasia; Asia (India, Japan, mainland southeast Asia).

 

Rhodobryum ontariense is a common and characteristic species of the eastern deciduous forests, occurring as far south as Arkansas and Georgia, with disjunct populations in the mountains of west Texas, New Mexico and southeastern Arizona.  Rhodobryum ontariense is not found in Arctic tundra and is rare in the northern boreal forests.

 

2. Rhodobryum roseum (Hedwig) Limpricht, Laubm. Deutschl. 3: 444. 1892

 

Plants 1--3 cm tall, commonly branched by slender sub-apical innovations. Stem leaves 3--8 mm, relatively few in rosettes, 18--22; margins revolute to about mid leaf or less, apex acute; costa variable, from subpercurrent to percurrent, with slender apiculus, to short-excurrent into a slender hairpoint on median rosette leaves, in cross section with small stereid band, not reaching dorsal epidermal layer because of a layer of intervening thin-walled cells; distal and median laminal cells hexagonal, 25--35 × 50--80 \um, (3--4:1), proximal cells longer, to 100 \um, rectangular. Inner perichaetial leaves with costa variable, from subpercurrent to percurrent or excurrent into a short, smooth to weakly denticulate hairpoint. Spores 16--20 \um.

 

Capsules mature late summer (Jul.--Sep.). Rare, rich soil, humus and litter in coastal tundra and shrublands or occasionally forests; 0--300 m; B.C.; Alaska; Eurasia; Asia (India, Japan).

 

Rhodobryum roseum is found only in coastal and near-coastal regions of Alaska, principally the Aleutian Islands, and adjacent far northwestern British Columbia, and disjunct in the Queen Charlotte Islands.  Sporophytes are rarely produced, and many populations consist of small plants that are sterile or only female. At least some locations of this species are in areas that remained ice-free during the last glaciation.

 

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

 

 

John R. Spence

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

 

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, especially Africa, 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. The center of diversity appears to be sub-Saharan Africa.

 

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 variable, sometimes strong and ending 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. A specimen reportedly collected in 1988 from Mexico needs to be evaluated.

 

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 rows 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, and 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, and is usually synoicous.

 

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., N.Y., 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.