New version Dec. 20, 2011
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 condi
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