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BFNA Title: Ptychostomum |
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Edit Level R XX.
PTYCHOSTOMUM Hornschuch 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 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), SELECTED
REFERENCES Andrews, A. L. 1935. Bryaceae. In: A. J. Grout (ed.). Moss flora
of North America north of 1. Stems
evenly foliate to comose with distal leaves distinctly enlarged; leaf base
mostly reddish, 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. Stems
mostly evenly foliate, not distinctly comose; leaf base same color as rest of
leaf, usually green, occasionally red or pink, 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, Phytol. 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. 40
(16 in the flora): widespread soil-inhabiting species in 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; limbidium strong; 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; synoicous ………. 2. Ptychostomum
bimum 6. Leaves comose, leaves enlarged above, not
decurrent, costa excurrent in medium to long awn; 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 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 or pale
orange; 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; Bryum 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, Phytol. 87: 20. 2005 Bryum bimum Schreber, Bot. Zeit. ( 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 incrassate, 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 3. Ptychostomum creberrimum ( Bryum
creberrimum 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.; Bryum 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, Phytol. 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., 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 6. Ptychostomum knowltonii
(Barnes) J. R. Spence, Phytol. 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, Phytol. 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 rows, same color as
lamina; distal and median laminal cells rhomboidal, 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, 10--16 \um. Capsules
mature May--Sept. Locally common on dry to damp soil, 0--3500 m;
boreal-temperate; B.C., Alta.; Alaska, Ariz., Calif., Colo., Idaho, 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 8. Ptychostomum longisetum
(Blandow ex Schwägrichen) J. R. Spence, Phytol. 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 red; 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 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, Phytol. 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 10. Ptychostomum nitidulum
(Lindberg) J. R. Spence, Phytol. 89: 113. 2007 Bryum
nitidulum Lindberg,
Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 23: 545. 1866; Bryum 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; 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, Phytol. 89: 113. 2007 Bryum
ovatum (Hedwig).
Dickson ex Withering, 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., 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, Phytol. 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, Phytol. 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; Ptychostomum
reedii is a highly
distinctive and very rare local endemic, not likely to be confused with any
other species in eastern 16. Ptychostomum salinum
( Bryum
salinum 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., 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. 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. Capsules globose to short-pyriform; leaves
ovate, acute; spores 18--22 \um
...................................................................... 31. Ptychostomum wrightii 4. 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 .……....…. 30. Ptychostomum
warneum 6. Autoicous or polyoicous; capsule
elongate-pyriform to clavate; spores 24--35 \um ................ 7 7. Autoicous; plants green or yellow-green;
costa of rosette leaves percurrent to short percurrent 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 ..............................................
29. Ptychostomum weigelii 8. Leaves not or weakly 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, Phytol. 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, Phytol. 87: 20. 2005 Pohlia
arctica R. Brown,
Chlor. Melvill. 38. 1823; Bryum arcticum (R. Brown) Bruch &
Schimper; Bryum bryoides (R. Brown) Ångstrom in Fries; Bryum
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., 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 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, Phytol. 87: 20. 2005 Bryum
calophyllum R. Brown,
Chlor. Melvill. 38. 1823; Bryum 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., 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 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 resemble Pohlia elongata. 21. Ptychostomum cryophilum
(Mårtensson) J. R. Spence, Phytol. 87: 20. 2005 Bryum
cryophilum Mårtensson,
K.V.A. Afh. Natursk. 15. 1956 Plants in dense swelling turfs, red to red-
green or red-yellow. Stems 2--4(--6) cm, fertile leaves comose,
innovations evenly foliate, leaves somewhat distant. Leaves 0.5--3 mm,
not much enlarged towards stem apex, ovate, concave, younger leaves strongly
contorted to shrunken when dry, yellow-green to pink or red, older leaves
more or less imbricate, pink, red becoming dark red-brown when old, leaf base
green, weakly decurrent; apex obtuse; margins plane or weakly revolute in
lower 1/4 of margin; limbidium strong, of 2--3 rows, partially 2-stratose
from base to mid-leaf; costa red, percurrent or sometimes not reaching apex;
distal and median laminal cells irregularly short-rhomboidal, 15--22(--25)
\um wide, mostly 2--3:1, thin-walled, proximal laminal cells narrower than
more distal cells, rectangular, 3--4:1. Specialized asexual reproduction
absent. Sexual condition dioicous. Seta single, 2--4 cm,
slender, straight to somewhat flexuose, red. Capsule ovate to obovate,
2--4 mm, symmetric, yellow-brown to brown, mouth yellow-brown; peristome well
developed, exostome teeth yellow-brown, lacking pores near base along
mid-line, lamellae straight, basal membrane high, 1/2 to 2/3 height of exostome,
not adherent to exostome, segments with ovate perforations, cilia long,
appendiculate; operculum conic, weakly apiculate. Spores pale yellow,
finely papillose, 16--20 \um. Capsules
mature Jul-Aug. Locally common on wet soil or rocks in streams, wetlands, and
late melting snow beds; Arctic-alpine; 0--2000 m; Greenland; Alta, B.C.,
N.W.T., Nunavut, Yukon; Alaska.; Eurasia (Arctic regions, India, Mongolia,
Nepal). Densely
compact red turfs, ovate concave leaves, and a weak costa are diagnostic of Ptychostomum
cryophilum. 22. Ptychostomum cyclophyllum (Schwägrichen) J. R. Spence, Phytologia
87: 20. 2005 Mnium cyclophyllum Schwägrichen, Sp. Musc. Frond., Supple. 2(2): 160, plate
194. 1827; Bryum cyclophyllum (Schwägrichen) Bruch, Schimper & W.
Gümbel; Bryum tortifolium Bridel Plants in dense or open turfs, green or
yellow-green. Stems 0.5--3(--4) cm, fertile leaves comose, innovations
evenly foliate, leaves somewhat distant. Leaves 0.5--3 mm, not much
enlarged towards stem apex, broadly ovate to orbicular, flat or weakly
concave, strongly contorted to shrunken when dry, green, leaf base green,
weakly decurrent; apex broadly acute to obtuse; margins plane; limbidium
strong, of 2--3 rows, partially 2-stratose from base to mid-leaf; costa not
reaching apex, rarely nearly percurrent; distal and median laminal cells
short-rhomboidal, 18--24 \um wide, mostly 2--3:1, thin-walled, proximal
laminal cells long-rectangular, 3--5:1. Specialized asexual reproduction
rare, of brown finely papillose filiform gemmae in leaf axils. Sexual
condition dioicous. Seta 2--4(--5) cm, slender, straight to
somewhat flexuose, red or brown. Capsule obovate, 2--4 mm, symmetric,
yellow-brown to brown, mouth yellow; peristome well developed, exostome teeth
yellow basally, hyaline distally, lacking pores near base along mid-line,
lamellae straight, basal membrane about 1/2\x height of exostome, not
adherent to exostome, segments with ovate perforations, cilia long,
appendiculate; operculum low-conic, weakly apiculate. Spores pale
yellow, smooth or finely papillose, 14--16 \um. Capsules
mature Jun-Sep. Locally common on wet sandy or organic soil along streams and
in wetlands; 0--3000 m; Greenland; Alta, B.C., Nfld. and Labr., N.W.T.,
Nunavut, Yukon; Alaska, Ariz., Calif., Colo., Mich., Mont., N.J., N.Mex.,
N.Y., Utah, Wash., Wyo.; Eurasia (India and Nepal). Ptychostomum
cyclophyllum is a distinctive Arctic-boreal species with broadly ovate
to orbicular, blunt, more or less flat leaves that are strongly shrunken when
dry. Ptychostomum cryophilum is
almost always reddish, with much more concave leaves that do not shrink much
when dry. Ptychostomum neodamense and P. ovatum are similar,
but have much shorter proximal lamina cells, 1-stratose borders, and red leaf
bases. Ptychostomum cyclophyllum has probably declined in the 23. Ptychostomum marrattii
(Hooker & Wilson) J. R. Spence, Phytol. 87: 21. 2005 Bryum
marratii Hooker &
Wilson, Bryol. Brit., 32b. 1855 Plants in dense or open turfs, yellow-green
to-yellow brown. Stems 0.5--2 cm, fertile leaves comose, innovations
evenly foliate, leaves crowded. Leaves (0.5--)1--2 mm, not much enlarged towards stem apex, ovate, concave,
somewhat contorted to shrunken when dry, yellow-green to green above,
becoming brown in older leaves, leaf base green; apex broadly acute to
obtuse; margins plane; limbidium absent, or rarely a weak single row present,
1-stratose; costa not reaching apex; distal and median laminal cells
short-rhomboidal, 18--25 \um wide, mostly 2--3:1, thin-walled, proximal
laminal cells irregularly rectangular to rhomboidal, somewhat bulging,
3--4:1. Specialized asexual reproduction absent. Sexual condition
autoicous. Seta long, 2--4(--5) cm, stout, somewhat flexuose,
red-purple. Capsule pyriform, 2--3 mm, symmetric, yellow-brown to
red-brown, mouth yellow-brown to red-brown; peristome reduced, exostome teeth
red-brown basally, yellow distally, lacking pores near base along mid-line,
lamellae straight, basal membrane about 1/2\x height of exostome, weakly
adherent to exostome, segments narrowly perforate, cilia absent or
rudimentary; operculum high conic. Spores yellow, finely papillose,
(30--)32--40 \um. Capsules
mature Jul-Sep. Rare and local on moist soil, 0--1000 m; Greenland; Alta;
N.Dak., Ptychostomum
marrattii is a distinctive Arctic-boreal species with oblong, more
or less obtuse green leaves, which lack a limbidium. It is found typically
along the coasts of northern 24. Ptychostomum meesioides
(Kindberg) J. R. Spence, Phytol. 87: 21. 2005 Bryum
meesiodes Kindberg,
Bull. Torr. Bot. Club 16: 95. 1889 Plants in dense or open turfs, green or
yellow-green, sometimes reddish below. Stems 0.5--1(--2) cm, fertile
stems evenly foliate, sometimes comose, innovations evenly foliate, leaves
crowded. Leaves 1--3(--3.5) mm, often gradually enlarged towards stem
apex, ovate-lanceolate, flat, strongly contorted to shrunken when dry, green
to yellow-green or rarely reddish, leaf base green, somewhat to strongly and
narrowly decurrent; apex acuminate; margins revolute proximally, plane
distally; limbidium strong, of 2--3 rows, partially 2-stratose from base to
mid-leaf; costa percurrent to short-excurrent as a stout awn; distal and
median laminal cells short-rhomboidal, 16--25 \um wide, mostly 3--4:1,
thin-walled, proximal laminal cells narrower than more distal cells,
long-rectangular, 3--5:1. Specialized asexual reproduction absent. Sexual
condition dioicous. Seta 2--4 cm, stout, straight to often
flexuose or geniculate, red-brown. Capsule highly variable, elongate-pyriform
to clavate, (2--)3--5 mm, strongly curved and asymmetric, brown, mouth
yellow-brown, median exothecial cells elongate, 40--85 \um long, many cells
3--4:1; peristome well developed, exostome teeth yellow or orange basally,
hyaline distally, lacking pores near base along mid-line, lamellae straight,
basal membrane about 1/2\x height of exostome, not adherent to exostome,
segments with ovate perforations, cilia present, appendiculate; operculum
conic, apiculate. Spores yellow to green, finely papillose, 14--20
\um. Capsules
mature Jun-Sep. Uncommon and scattered on wet soil along streams and in
wetlands, often calcareous, 0--3000 m; BC; Alaska, Idaho, Wash. Ptychostomum
meesioides is an Arctic-boreal to north temperate species, narrowly
endemic in primarily coastal hyperoceanic areas of western 25. Ptychostomum pallens
(Swartz) J. R. Spence, Phytol. 87: 21. 2005 Bryum pallens Swartz, Monthly Rev. 34: 538. 1801 Plants in dense or open turfs, pale pink, red
or rarely dull green. Stems 1--4(--6) cm, fertile leaves comose,
innovations evenly foliate, leaves crowded. Leaves 1--3.5(--4) mm, often
gradually enlarged towards stem apex, ovate to broadly ovate-lanceolate, flat
or weakly concave, contorted to shrunken when dry, pink, red or rarely green,
leaf base pink or green, somewhat to strongly and narrowly decurrent; apex
acuminate; margins revolute proximally, plane distally; limbidium strong, of
2--3 rows, partially 2-stratose from base to mid-leaf; costa percurrent to
short-excurrent as a stout awn, red to red-brown; distal and median laminal
cells short-rhomboidal, 17--24 \um wide, mostly 2--3:1, thin-walled, not
porose, proximal laminal cells short-rectangular, 2--3(--4):1. Specialized
asexual reproduction rare, of brown-red to green finely papillose
filiform gemmae in leaf axils. Sexual condition dioicous. Seta
2--4(--6) cm, stout, straight to somewhat flexuose, red-brown. Capsule
elongate-pyriform to clavate, (3--)4--5 mm, somewhat curved, yellow-brown,
mouth yellow-brown, median exothecial cells short-rectangular to quadrate,
20--40 \um long, 1--2:1; peristome mostly well-developed, exostome teeth
yellow basally, hyaline distally, lacking pores near base along mid-line,
lamellae straight, basal membrane about 1/2\x height of exostome, not
adherent to exostome, segments with ovate perforations, cilia appendiculate
to nodose, occasionally short; operculum conic, apiculate. Spores
yellow to green, finely papillose, 16--20(--22) \um. Capsules
mature Jun-Sep. Common on wet soil along streams and in wetlands, usually
calcareous, 0--3000 m; Greenland; Alta., B.C., Man., N.B., Nfld. and Labr.,
N.W.T., N.S., Nunavut, Ont., Que., Sask., Yukon,; Alaska, Ariz., Calif.,
Colo., Idaho, Maine, Mich., Minn.,
Mont., Nev., N.Mex., N.Y., Oreg., Utah, Wash., Wyo.; s South America;
Eurasia. Ptychostomum
pallens is a highly variable Arctic-boreal to north temperate
species, related to P. rutilans, P. meesioides and P.
cernuum. The pale pink color is
diagnostic, but is not always present. From P. rutilans it can be
distinguished by its much thinner walled non-porose laminal cells and well
developed peristome; from P. meesiodes by its shorter less arcuate
capsule and short exothecial cells; and from P. cernuum by its
dioicous sexual condition. 26. Ptychostomum rutilans
(Bridel) J. R. Spence, Phytol. 87: 23. 2005 Bryum
rutilans Bridel,
Bryol. Univ. 1: 684. 1826; Bryum aeneum Blytt ex Bruch, Schimper &
W. Gümbel Plants in dense swelling turfs, red to
red-brown. Stems 2--4(--6) cm, fertile leaves comose, innovations
evenly foliate, leaves crowded; often strongly radiculose. Leaves
1--3.5(--4) mm, often gradually enlarged towards stem apex, ovate to broadly
ovate-lanceolate, concave, contorted to shrunken when dry, red, becoming dark
red-brown with age, leaf base uniformly red, not or weakly decurrent; apex acute; margins revolute
proximally, plane distally; limbidium strong, of 2--3 rows, partially
2-stratose from base to mid-leaf; costa percurrent to short-excurrent as a
stout awn, red to red-brown; distal and median laminal cells
short-rhomboidal, 12--18 \um wide, mostly 2--3:1, incrassate and porose,
proximal laminal cells rectangular, 3--4:1. Specialized asexual
reproduction uncommon, of brown-red papillose filiform gemmae in leaf
axils. Sexual condition dioicous. Seta 1--3(--4) cm, stout,
straight to somewhat flexuose, red-brown. Capsule elongate-pyriform, long-necked,
3--5 mm, somewhat curved, brown, mouth yellow-brown; peristome reduced,
exostome teeth yellow basally, hyaline distally, lacking pores near base
along mid-line, lamellae straight, basal membrane low, less than 1/2\x height
of exostome, not adherent to exostome, segments with ovate perforations,
cilia reduced or absent; operculum conic, apiculate. Spores yellow to
green, finely papillose, 16--22(--24) \um.
Capsules
mature Jul-Aug. Uncommon on wet soil in tundra and along streams and in
wetlands, 0--2000 m; Greenland; N.W.T., Ptychostomum
rutilans is a distinctive Arctic species with dark red shoots,
usually forming dense turfs on wet soil. It is close to P. pallens,
but differs in the incrassate porose lamina cells, and reduced
peristome. Small specimens are similar
to Rosulabryum subelegans, which generally has green-red spirally
twisted leaves and mostly grows on rock in alpine regions of northwestern 27. Ptychostomum schleicheri (Schwägrichen) J. R. Spence, Phytol. 87: 22. 2005 Bryum
schleicheri
Schwägrichen, Sp. Musc., Suppl. 1, 2: 113, plate 73. 1816; Bryum
schleicheri Schwägrichen var. latifolium (Schwägrichen) Schimper Plants in dense or open turfs, pale yellow,
yellow-green or copper. Stems 3--6(--10) cm, fertile leaves comose,
innovations evenly foliate, leaves crowded; weakly radiculose. Leaves
2--4(--5) mm, not much enlarged towards stem apex, broadly ovate, weakly to
strongly concave, upper leaves contorted to shrunken when dry, lower leaves
more or less imbricate, yellow to yellow-copper, leaf base green, not or
weakly decurrent; apex acute; margins plane or revolute proximally, plane
distally; limbidium moderately strong, of 1--3 rows, partially 2-stratose
from base to mid-leaf; costa percurrent to short-excurrent as a slender awn,
pale brown to red-brown; distal and median laminal cells short-rhomboidal,
28--36 \um wide, mostly 2:1, thin-walled, proximal laminal cells rectangular,
3:1. Specialized asexual reproduction none. Sexual condition
dioicous. Seta 2--3(--4) cm, slender, straight to somewhat flexuose,
red-brown. Capsule turbinate to pyriform, 3--5 mm, symmetric,
yellow-brown, mouth yellow; peristome well-developed, exostome teeth yellow
basally, hyaline distally, lacking pores near base along mid-line, lamellae
straight, basal membrane about 1/2\x height of exostome, not adherent to
exostome, segments with ovate perforations, cilia appendiculate to nodose;
operculum conic, rounded. Spores yellow to brown, finely papillose,
16--20(--22) \um. Capsules
mature Jul-Aug. Uncommon on wet soil along streams and on seepy tundra slopes
and late snowmelt areas, 0--4000 m; Greenland; Alta., B.C., N.W.T., Nunavut, Yukon; Alaska, Calif.,
Idaho, Mont., Ore., Wash.; Eurasia. Ptychostomum
schleicheri is an Arctic-alpine species. When well developed it is
distinctively large, with long stems, and crowded yellowish, somewhat
imbricate leaves. Much material named P. schleicheri appears to be
misidentified and can be referred to P. turbinatum. Our material
conforms to the var. latifolium.
The relationships between var. schleicheri and var. latifolium and with P.
turbinatum, have not been worked out yet.
This species is atypical for Ptychostomum, and shows some
morphological similarities to the Mniaceae, especially in the lack of
rhizoidal filiform gemmae, wide laminal cells, and the pronounced shelf-like
leaf base that remains attached to the stem after the leaf is removed. Ptychostomum ovatum is similar but
has a unistratose border and much smaller laminal cells. 28. Ptychostomum turbinatum
(Hedwig) J. R. Spence, Phytol. 87: 22. 2005 Mnium turbinatum Hedwig, Sp. Musc. Frond., 191. 1801; Bryum
turbinatum (Hedwig) Turner Plants in dense or open turfs, dull green or
yellow-green to brownish. Stems (1--)2--4(--6) cm, fertile leaves
comose, innovations evenly foliate, leaves crowded; sparsely radiculose,
sometimes clusters of rhizoids arising from proximal leaf axils. Leaves
1--3 mm, often gradually enlarged towards stem apex, broadly
ovate-lanceolate, somewhat concave, strongly contorted to shrunken when dry,
dull green to yellow-green, base green, not or weakly decurrent; apex acute
to acuminate; margins plane, or occasionally weakly revolute proximally;
limbidium strong, of 2--3 rows, partially 2-stratose from base to mid-leaf;
costa percurrent to excurrent in a short slender awn; distal and median
laminal cells short-rhomboidal, 18--25(--30) \um wide, mostly 2--3:1,
thin-walled, not porose, proximal laminal cells narrower more distal cells,
rectangular, 3--4:1, thin-walled, not porose. Specialized asexual
reproduction absent. Sexual condition dioicous. Seta
single, 1--3(--4) cm, slender, straight to somewhat flexuose, brown or
red-brown. Capsule turbinate to short-pyriform, 3--5 mm, symmetric,
pale yellow to yellow-brown, becoming black with age, mouth yellow; peristome
well-developed, exostome teeth yellow to orange basally, hyaline distally,
lacking pores near base along mid-line, lamellae straight, basal membrane
about 1/2\x height of exostome, not adherent to exostome, segments with ovate
perforations, cilia appendiculate to nodose; operculum conic, apiculate. Spores
yellow to brown, finely papillose, 18--22 \um. Capsules
maturing Jun-Aug. Locally common on wet soil in calcareous wetlands, 0--3500
m; Greenland; Alta., B.C., Nfld. and
Labr., N.W.T., Yukon; Alaska, Calif., Colo., Idaho, Mont., Nev., Oreg., Utah,
Wash., Wyo.; South America (Argentina, Chile, Ecuador, Peru); Eurasia
(including Turkey), Africa. Ptychostomum
turbinatum is a circumpolar Arctic-boreal to temperate species. The
dull green to yellow-green shoots with strongly shrunken ovate-lanceolate
leaves, and the turbinate capsule are diagnostic. Old capsules often turn
black, and become strongly constricted just below the mouth. Ptychostomum
schleicheri is much larger, pale yellow distally, with wider leaves and
much wider upper lamina cells. Large specimens, 8--12 cm, with long narrow ovate-lanceolate
leaves and elongate-pyriform capsules, represent an undescribed species,
which will be published at a future date. 29. Ptychostomum warneum
(Röhl) J. R. Spence, Phytol. 87: 22. 2005 Mnium caespiticium var. warneum Röhl, Deutschl.
Fl. (ed. 2), Kryptog. Gew. 3: 94. 1813; Bryum
warneum (Röhl) Bridel Plants in dense or open turfs, red, green or
yellow-green. Stems 0.5--2(--3) cm, fertile leaves comose, innovations
evenly foliate, leaves crowded; sparsely radiculose. Leaves 0.5--2 mm,
often gradually enlarged towards stem apex, ovate-lanceolate, flat, strongly
contorted to shrunken when dry, yellow-green to red, base usually green; apex
acuminate; margins revolute proximally; limbidium strong, of 2--3 rows,
partially 2-stratose from base to mid-leaf; costa short-excurrent into a
slender smooth awn; distal and median laminal cells short-rhomboidal, 18--22
\um wide, mostly 2--3:1, thin-walled, proximal laminal cells rectangular,
3--4:1. Specialized asexual reproduction absent. Sexual condition
polyoicous, variable, some populations
mostly autoicous. Seta 2--4(--5), stout, straight to somewhat
flexuose, purple or red. Capsule pyriform, 2--4 mm, symmetric,
yellow-brown or brown, mouth yellow; peristome reduced, exostome teeth yellow
or brown basally, pale yellow to hyaline distally, lacking pores near base
along mid-line, lamellae straight, numerous, often joined by cross-walls,
basal membrane about 1/2\x height of exostome, weakly adherent to exostome,
segments narrowly perforate, cilia absent or rudimentary; operculum tall
conic. Spores pale green or yellow, finely papillose, large, variable
in same capsule (32--)36--48(--50) \um. Capsules
mature Jul-Aug. Rare on damp soil; 0--50 m; Greenland; Que.; n Eurasia (also
Himalayas in Ptychostomum
warneum is an Arctic-boreal species that can be distinguished from
the related P. calophyllum by its pyriform brown capsule with a
slender neck, generally larger spores and acuminate leaves. It is known from
only one location in North America, in the 30. Ptychostomum weigelii
(Sprengel) J. R. Spence, Phytol. 87: 22. 2005 Bryum
weigelii Sprengel,
Mant. Prim. Fl. Hals. 55. 1807 Plants in loose low open turfs, green or
rarely pinkish-red. Stems 2--4(--6) cm, fertile leaves comose,
innovations evenly foliate, leaves distant; sparsely radiculose. Leaves
1--3 mm, not much enlarged towards stem apex, ovate-lanceolate, flat,
strongly contorted to shrunken when dry, green, yellow-green, rarely red or
pink, base usually green, strongly and broadly decurrent, decurrencies almost
reaching next more proximal leaf; apex acute; margins revolute proximally,
plane distally; limbidium weak, of 1(--2) rows, partially 2-stratose from
base to mid-leaf; costa not reaching apex to short-excurrent as a slender
awn, green or brown; distal and median laminal cells short-rhomboidal,
18--25(--30) \um wide, mostly 2--3:1, thin-walled, proximal laminal cells
narrower more distal cells, hexagonal to rectangular, 3--4:1. Specialized
asexual reproduction absent. Sexual condition dioicous. Seta
single, 2--4 cm, slender, straight to somewhat flexuose, red-brown. Capsule
turbinate to pyriform, 3--4 mm, symmetric, brown, mouth yellow; peristome
well developed, exostome teeth yellow basally, hyaline distally, lacking
pores near base along mid-line, lamellae straight, basal membrane about 1/2\x
height of exostome, not adherent to exostome, cilia well developed, long,
appendiculate; operculum convex, apiculate. Spores yellow or green,
finely papillose, (12)14--18 \um. Capsules
mature Jun-Aug. Common and widespread on wet soil in wetlands and along
streams, 0--3500 m; Greenland; Alta., B.C., Man., Nfld. and Labr., N.W.T.,
N.S., Nunavut, Ont., P.E.I., Que., Sask., Yukon; Alaska, Ariz., Calif.,
Colo., Conn., Idaho, Maine, Mass.,
Mich., Minn., Mont., Nev., N.H., N.Dak., Ohio, Oreg., Pa.,
Utah, Vt., Va., Wash., W.Va., Wis., Wyo.; South America; Eurasia;
Africa; Pacific Islands (New Zealand); Australia. Ptychostomum
weigelii is a common Arctic-boreal to north temperate species in
wet sites, easily identified by the broad, long marginal decurrencies of the
leaves that nearly reach to the next leaf.
Leaves of the similar P. cyclophyllum and P. turbinatum
are either not decurrent or only produce short very slender decurrencies. 31. Ptychostomum wrightii
(Sullivant & Lesquereux) J. R. Spence, Phytol. 87: 22. 2005 Bryum
wrightii Sullivant
& Lesquereux, Proc. Amer. Acad. Arts. Sci. 4: 278. 1860 Plants in dense or open turfs, red- green or
red-brown. Stems 0.5--1(--2) cm, fertile leaves comose, innovations
evenly foliate; sparsely radiculose. Leaves 0.5--2 mm, not enlarged
towards stem apex, ovate, concave, somewhat contorted to shrunken when dry,
yellow-green to red-green, base green, not decurrent; apex acute; margins
revolute proximally; limbidium strong, of 2--3 rows, partially 2-stratose
from base to mid-leaf; costa strong, percurrent to excurrent in short smooth
awn; distal and median laminal cells short-rhomboidal, 18--25 \um wide,
mostly 2--3:1, thin-walled, proximal laminal cells narrower more distal
cells, long-rectangular, 3--5:1. Specialized asexual reproduction
absent. Sexual condition synoicous. Seta 1--3 cm, slender, straight to somewhat
flexuose. Capsule broadly ovate to pyriform, somewhat inflated, 2--3
mm, symmetric 2--3 mm, symmetric, yellow-red, red or red-brown, often shiny,
mouth yellow-red; peristome reduced, exostome teeth yellow proximally,
hyaline distally, lamellae somewhat curved below, basal membrane low to rudimentary, not or
weakly adherent to exostome, cilia absent; operculum low-conic, smooth or
weakly apiculate. Spores yellow, finely papillose, 18--20(--22)
\um. Capsules
mature Jul-Aug. Locally common on damp calcareous soil and silt in tundra,
0--1500 m; Greenland; Labr., N.W.T., Nunavut, Que., Yukon; Alaska; Arctic
Eurasia. Ptychostomum
wrightii is a small Arctic species with distinctive capsules that
are somewhat inflated, shiny red, and ovate to pyriform, and a poorly
developed fragile endostome. Capsules
are needed for identification. |
