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BFNA Title: Bryaceae Keys |
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Keys to the genera, sections and species of the
Bryaceae in the Flora of North America Region John
R. Spence Size can be of some use in identifying species
of Bryum. Both stem length and leaf length categories are described
below. I have adopted the leaf
lengths from Ochi (1972). Stem length: short
(<10 mm), medium (10-30 mm), long (>30 mm). Leaf length: small (<1.5 mm), medium (1.5-3.0 mm), large
(3.0-4.0 mm), robust (>4.0 mm). [Editor's Note: Several of the generic names were
unpublished (either as new names or raised to generic rank) at the date of
online publication of this draft treatment. For purposes of the FNA, these
names are replaced by "Genus G, Genus P, and Genus L" where the
letter is that of initial letter of the unpublished name. These keys will be
those of the FNA treatment of the Bryaceae, and comments are solicited from
users. These can be sent directly to John Spence John_Spence@nps.gov
(underline between John and Spence) or Spence@thedam.com For a previously BFNA published
preliminary discussion of the Bryaceae, click Preliminary Discussion ] 1. Stems short, julaceous; leaves small, green, yellow-green to silver, if stems not julaceous then plants silver and proximal lamina cells quadrate; lamina areolation Bryum-type, costa Bryum-type …………………………………………………..………………………………..…… Bryum 1. Plants and leaves of
various sizes; fertile stems not julaceous although innovations sometimes so; if plants silver then
proximal lamina cells elongate-rectangular, if julaceous then leaves >2 mm
long and dark red; lamina areolation and costa type various ………………….……………. 2 2. Plants in dense
rhizomatous cushions; leaves mostly imbricate with long spinose partly
hyaline hairpoint, lamina areolation dense, Bryum-type; costa Ptychostomum-type;
capsules erect; epiphytic-corticolous or saxicolous, peristome reduced, cilia
absent ………………………………………………………………………....………….. Genus L 2. Plants not in dense
rhizomatous cushions; areolation dense to lax, hairpoint variable, typically
not spinose, rarely hyaline, often absent, proximal lamina cells short or
long; costa type various capsule erect to nodding, peristome perfect or
reduced; habitats various ……….…………….... 3 3. Inflorescences
appearing lateral; leaves soft with costa not reaching apex to percurrent,
rarely weakly excurrent, lamina areolation Plagiobryum-type, alar cells somewhat differentiated from justacostal
cells, quadrate; costa Bryum-type;
capsule shortly pyriform, lacking peristome or with exostome only
……………………………………….…………………….. Haplodontium 3. Inflorescences
terminal; leaves and costa various, percurrent to excurrent, lamina
areolation various, alar cells rarely differentiated from justacostal cells,
if so then capsule with double peristome; costa type various; capsule shape
various, peristome double ……...………………... 4 4. Leaves strongly
twisted, areolation Bryum-type,
apex broadly obtuse, costa extending into short stout point, Ptychostomum-type, margins serrate
above; capsules erect; saxicolous or epiphytic-corticolous
………………………………………..………………….... Brachymenium 4. Leaves variously
twisted or contorted to imbricate, leaf apex acuminate to acute, if obtuse
then costa percurrent; costa variable, percurrent to long excurrent, upper
margins smooth to serrate, areolation-type various, costa type various;
capsules nodding or erect, terricolous, saxicolous or sometimes rotten wood
but not epiphytic ……………………………………………………….. 5 5. Lamina areolation Bryum-type, or if Plagiobryum-type plants with abundant rhizoidal tubers; leaves
mostly imbricate or rarely slightly twisted at stem apex; stems either budlike
or elongate and evenly foliate; rhizoidal tubers and leaf axis bulbils often
present; costa Ptychostomum-type
……………………………………………..………………………….……….…… Genus
G 5. Lamina areolation
either Rhodobryum-type, or if Plagiobryum-type then plants lacking
gemmae, leaves variously contorted, twisted or imbricate; stems comose,
rosulate or elongate and evenly foliate; rhizoidal tubers or leaf axis
filiform gemmae sometimes present; costa type various
…………………………………….……………………...…………………………….... 6 6. Plants pink, red-green
to pink-silver, innovations sometimes julaceous, lamina areolation Plagiobryum-type, limbidium lacking or
if present bi- to multistratose; capsule distinctly zygomorphic, curved,
gibbous or with well-differentiated furrowed neck, peristome variable, well
developed to reduced, seta curved to geniculate ………………………………………....… 7 6. Plants lacking silvery
tones, with variously shrunken, contorted or twisted leaves, innovations
julaceous or not, laminal areolation Rhodobryum-type,
leaf border of thick elongate cells present or absent, uni- or bi-stratose,
costa Ptychostomum or Rhodobryum-type; capsules not
distinctly zygomorphic or with differentiated neck, peristome various, but
exostome teeth longer than endostome, seta straight
…….……………………………………………………………...……. 8 7. Plants dull red or
pink, stems long and evenly foliate or crowded and comose, innovations not
julaceous, distal lamina cells sometimes very short (1-3:1), broad, with many
irregularly rhomboidal to subquadrate cells; limbidium absent or if present
strong, bi- to multistraose; capsule somewhat zygomorphic, seta slightly
curved to straight; peristome variable, endostome well developed to reduced
and adherent to exostome, teeth longer than endostome segments
…………………………………………………………………………………...... Genus
P 7. Plants silver-pink to
red-green, stems short, leaves crowded, innovations often julaceous, distal
laminal cells long and wide (3-6:1), hexagonal, limbidium absent; capsule
distinctly curved and gibbous, seta short, geniculate or curved, peristome well
developed, endostome longer than exostome ……………………………………………………………….…………… Plagiobryum 8. Stems comose or
elongate with equidistinct leaves, not distinctly rosulate; leaves ovate to
ovate-lanceolate or lanceolate, widest at or below middle of leaf, lamina
margins smooth to finely serrulate but not strongly serrate, border
unistratose or bistratose; rhizoidal tubers lacking, leaf axis filiform
gemmae rarely present; costa Ptychostomum-type;
monosetous; terricolous or saxicolous ……..………………………………………………………………..... Ptychostomum 8. Stems rosulate or
rarely elongate with equidistant leaves; leaves typically obovate or oblong
to spathulate, widest above middle, lamina margins serrulate to distinctly
serrate, unistratose; rhizoidal tubers and leaf axis filiform gemmae often
present, costa Ptychostomum or Rhodobryum-type various; monosetous or
polysetous; habitats various, rarely epiphytic-corticolous
……………………………………………………………………………………….. 9 9. Leaves mostly medium to large, upper
margins usually serrate to serrulate at apex, stolons absent, costa Ptychostomum-type; rhizoidal tubers
generally present, filiform gemmae sometimes present in leaf axils;
micronemata common on stem; on various substrates including bark of living
trees …………………………………………………………………………..
Rosulabryum 9. Leaves mostly robust,
upper margins distinctly serrate, stolons sometimes present, costa Rhodobryum-type; gemmae lacking; stems
lacking micronemata; terricolous ………………... 10 10. Lamina finely rugose, cells large, 90--150
um long by 30--50 um wide, costa not reaching apex; stolons absent ………………………………………………………………………..
Roellia 10. Lamina smooth, cells mostly <100 um
long and <35 um wide; costa not reaching apex, percurrent or excurrent;
stolons present ……………………………………………..
Rhodobryum BRACHYMENIUM 1 Species 1. Brachymenium macrocarpum BRYUM 11 Species 1. Plants silver or white, stems julaceous to gemmiform, upper lamina
hyaline, distal laminal cells hexagonal to rhomboidal, <20 um
……………….……………………………………...…. 2 1. Plants brown, golden, pale yellow, yellow-green to bright green,
stems more or less julaceous, upper lamina not hyaline, distal lamina cells
variable, elongate vermicular to short and rhomboidal, 6--25 um wide
…………………………………………………………………...…. 5 2. Stems elongate, loosely julaceous, leaves somewhat distant, costa
weak, not reaching apex, apiculus lacking, proximal laminal cells
short-rectangular……………………..5. Bryum
gerwigii 2. Stems short, gemmiform to julaceous, leaves crowded, costa
variable, strong and excurrent in distinct recurved hairpoint or not reaching
apex with apiculus present, proximal laminal cells quadrate
………………………………………………………………………………………….. 3 3. Plants hoary white, stems evenly foliate, slender, costa strong,
excurrent in stout hairpoint, recurved when dry ……………………………………………………….….… 7. Bryum lanatum 3. Plants silver to silver-green, stems short, julaceous to
gemmiform, costa not reaching apex, apiculus usually present, irregularly
curved when dry …………..……………………………… 4 4. Plants tiny, stems 1-3 mm, leaves broadly ovate, larger leaves
broader than long, distal laminal cells short rhomboidal, 2--3:1
………………...……..………………… 9. Bryum
nanoargenteum 4. Plants larger, stems 2--10 mm, leaves ovate, longer than wide,
distal laminal cells narrower, 3--5:1 …………...………………………..….…………...……………….…… 1.
Bryum argenteum 5. Distal laminal cells of innovation leaves elongate-vermicular,
>8:1; bulbils sometimes present in leaf axils
……………………………………….……………………………………..…….…. 6 5. Distal laminal cells of innovation leaves not vermicular, shorter
and broader, 2--6:1; bulbils absent …………………………………………………………………………………...……..… 9 6. Fertile stems gemmiform, short, <5 mm long, bulbils sometimes
present in leaf axils of innvations ………………………………………...……………………………… 2. Bryum blindii 6. Fertile stems elongate-julaceous, 3--10 mm long, bulbils present
or absent ……..…………... 7 7. Stems strongly julaceous, leaves concave, costa not reaching apex,
bulbils lacking, capsule nodding, peristome perfect …………………….……………………..………
6. Bryum julaceum 7. Stems weakly and loosely julaceous, leaves flat to weakly concave,
costa variable but often percurrent or very short excurrent, bulbils
sometimes present, capsule nodding to erect, peristome perfect or reduced
………………………………………………………………….... 8 8. Capsules erect, peristome reduced, cilia lacking, bulbils lacking
…….8. Bryum leptostomoides 8. Capsules nodding, peristome perfect, bulbils often present in leaf
axils ………………………………………………………………………….… 4. Bryum
concinnatum 9. Plants brown to golden or yellow-green, laminal cells of fertile
stem leaves elongate hexagonal, 4--6:1 ………………………………………………………...… 10. Bryum oblongum 9. Plants bright green, laminal cells of fertile stem leaves shorter,
2--4:1 irregularly rhomoboidal to hexagonal ……………………………………………………………….……………………
10 10. Distal laminal cells elongate, 3--4:1, 10--16 um wide, main
rhizoids smooth ……………………………………………………………………………..…11. Bryum veronense 10. Distal laminal cells short and irregularly rhomboidal, mostly 2--3:1,
12--25 um wide, main rhizoids coarsely papillose ………………………….…………………….. 3. Bryum calobryoides GENUS G 29 Species Key to Sections 1. Plants with rhizoidal tubers commonly
present, often abundant, at stem base in clusters or on long rhizoids in substrate;
stems evenly foliate, leaves slightly twisted when dry …..…………..
2 1. Plants with leaf axis bulbils, tubers
rarely present, or plants with tubers present, scarce, on macronemata arising
from leaf axils on stem; stems evenly foliate or gemmiform, often in two or
more clumps; leaves imbricate, not twisted when dry ………….………………………………..
4 2. Laminal cells elongate, to 100 um, 5--8:1,
alar cells differentiated, quadrate, justacostal cells elongate, distal
laminal margins mostly entire, costa percurrent or short excurrent; tubers
small (25--60 um), pyriform, brown ………………………………………………………...
Apiculata 2. Laminal cells mostly shorter, <80 um, distal
cells hexagonal (3--5:1), proximal cells quadrate to short-rectangular across
leaf base to costa, distal laminal margins mostly serrulate, costa mostly
distinctly excurrent in short to long hairpoint; tubers various sizes, shapes
and colors …………………………………………………………………………………………..……….. 3 3. Tubers pale tan or pink; leaves folded,
slightly incurved when dry, broadly ovate, distal margins mostly entire;
capsule erect, peristome reduced ………………………….
Dicranobryum 3. Tubers typically bright red, red-brown or
yellow; leaves slightly twisted at tips but not folded or incurved when dry,
ovate, ovate-lanceolate to broadly lanceolate; capsule nodding, peristome
perfect ……………………………………………………………………………….
Apalodictyon 4. Leaves small, stems short, fertile stems
gemmiform, innovations slender, gemmiform to evenly foliate, tubers rarely
present, bulbils commonly present in leaf axils of innovations; capsule
cylindrical to short ovate with thick and sometimes wrinkled neck, erect or
nodding, peristome perfect to reduced ……………………………………………………………..………
5 4. Leaves mostly medium, stems evenly foliate
with imbricate leaves, innovations similar to fertile stems; tubers sometimes
present in tomentum on stem, scarce, bulbils absent; capsule pyriform,
nodding, peristome perfect ………………………………………….……
Alpiniformia 5. Capsules erect, peristome reduced; tubers sometimes
present on rhizoids, bulbils lacking; …………….……………………………………………………….…………….…
Dicranobryum 5. Capsules nodding, peristome perfect; tubers
rarely present, bulbils common ……………………………………………………………………..…………….……
Doliolidium ALPINIFORMIA
8
Species
1. Leaves stiff, rigid, strongly imbricate,
distal laminal cells long, incrassate and vermicular, >6:1 …………………………………………………...………………………………………………..
2 1. Leaves rigid to somewhat loosely imbricate,
distal lamina cells shorter, not vermicular, mostly 3-6:1 ……………………………………………...……………………………………………....
3 2. Costa percurrent to short excurrent,
limbidium absent, proximal laminal cells gradually wider proximally,
short-rectangular to quadrate ……………….………..……
1. Genus G alpinum 2. Costa distinctly excurrent in medium-length
point, weak to moderately strong limbidium present, proximal laminal cells
abruptly enlarged, somewhat bulging, rectangular ……………………………………………………………………
3. Genus G microchaeton 3. Stems julaceous; leaves red, purple or
rarely green, strongly concave, apex rounded-obtuse, cucullate, costa not
reaching apex to percurrent, distal lamina cells incrassate, 4--6:1, oblique
to costa, cells across leaf base abruptly enlarged to inflated in 1--2 rows …………………………………………………….…...………………
5. Genus G miniatum 3. Stems not julaceous, leaves red to green or
yellow, concave to flat, apex obtuse, acute or acuminate, not cucullate,
costa not reaching apex to distinctly excurrent, distal lamina cells thin to
somewhat firm-walled, parallel to costa, 3--6:1, cells at leaf base not abruptly
enlarged or inflated …………………………………………………………………………….…………….
4 4. Leaves green or yellow-green, with distinct
limbidium, costa strong, moderately long excurrent ……………………………………………….……
8. Genus G “pseudoriparium” 4. Leaves red, green or yellow-green, limbidium
lacking, costa variable, not reaching apex to short excurrent ………………………………………………………….…………………….….
5 5. Leaves distinctly concave, apex broadly acute to
obtuse, costa not reaching apex or percurrent, hairpoint lacking
………………………………………………………………………………… 6 5. Leaves flat or weakly concave, apex acute
to acuminate, costa short but distinctly excurrent into hairpoint …………………………….……………………………………………………….
7 6. Leaves green to yellow-green, lacking red
tints, loosely set, somewhat distant below, proximal laminal cells
rectangular ...………………………………..………
2. Genus G gemmiparum 6. Leaves red or red-green, red tints usually
present, rigid and imbricate, crowded, proximal laminal cells quadrate ………………………………………...…
6. Genus G muehlenbeckii 7. Plants green or yellow-green, leaves
lacking red tints, distal laminal cells elongate hexagonal, 4--5:1, 10--14 um
wide, proximal cells mostly rectangular, rhizoidal tubers lacking ………………………………………………………………….……
4. Genus G mildeanum 7. Plants green to red, leaves often red, distal
laminal cells short hexagonal, 3--4:1, 14--20 um wide, proximal cells mostly
quadrate, rhizoidal tubers sometimes present ……………………………………………………..….
7. Genus G “pseudomuehlenbeckii” APALODICTYON
10 Species
1. Rhizoidal tubers small, mostly <120 um in
longest axis ……...……...…………………….…
2 1. Tubers larger, mostly 120--250 um …………………………………………………………...
5 2. Rhizoids violet to purple, tubers
red-purple to orange ………..…
18. Genus G violaceum 2. Rhizoids and tubers brown, red-brown or red
……….……..…………………………………
3 3. Tubers spherical, bright red to crimson,
cells protuberant, rhizoids paler than tubers ………………………………………………………………...…
11. Genus G klinggraeffii 3. Tubers spherical to pyriform, brown or
red-brown, cells smooth, rhizoids same color as tubers ….…………………………………………………………………………………………………
4 4. Tubers 40--80 um, spherical to pyriform,
cells 15--20 um, brown or yellow-brown, on long rhizoids in substrate ……………………………………………….
17. Genus G valpariense 4. Tubers 60--120 (rarely to 150) um,
pyriform, cells 40--70 um, bright orange or red-orange, clustered in strings
on side branches of main rhizoids at stem base ………………………………………………………………..…
10. Genus G demaretianum 5. Leaves with distinct limbidium, median
laminal cells wide, 14--20 um; rhizoids red-brown or brown ………………………………………………………………………………….…………
6 5. Leaves lacking distinct limbidium, median
laminal cells narrower, 10--16 um; rhizoids red-brown, brown, violet, purple
or yellow ………..……………………………………….………..
7 6. Tubers 150--250 um, clustered around base
of stem or in axils of proximal leaves, cells distinctly protuberant, 25--50
um wide …………………………………13.
Genus G rubens 6. Tubers 180--300 um wide, on long rhizoids in
substrate, never axillary, cells smooth or weakly protuberant, 40--70 um wide
…….……………………… 9. Genus G bornholmense 7. Rhizoids yellow, tubers yellow to rarely
orange; seta purple with age ……………………………………..…………………………..……
16. Genus G tenuisetum 7. Rhizoids brown, red-brown to purple, tubers
red to red-brown; seta brown or red with age ……...……………………………………………………………………………………………..
8 8. Rhizoids deep purple, red-purple or violet,
tubers red-purple ...……14. Genus G ruderale 8. Rhizoids brown to red-brown, tubers red to
red-brown ……………………………………….
9 9. Costa strong, long excurrent, yellow-brown,
proximal laminal cells quadrate; on calcareous substrates …………………………………………………………
12. Genus G radiculosum 9. Costa short excurrent, red-brown, proximal
laminal cells short-rectangular; on siliceous substrates ……………………………..…………………..……
15. Genus G subapiculatum APICULATA
1 Species
19. Genus
G apiculatum DICRANOBRYUM
2 Species
1. Stems distinctly gemmiform, often in two or
more interrupted tufts, leaves imbricate wet or dry; tubers absent ……….…………………………………………
22. Genus G mexicanum 1. Stems evenly foliate,
leaves folded and often incurved when dry; tubers present
……………………………………………………………………………. 21. Genus
G exile DOLIOLIDIUM
8 Species
1. Plants with 1--many bulbils in axils of
upper innovation leaves ……...………………………
2 1. Plants lacking bulbils ………………………………………………………………………….
6 2. Capsule with thick, inflated and wrinkled
neck; leaves of fertile stems triangular, laminal margins revolute to above
mid-leaf; bulbils large, single and leafy ………………………………………………………………………
26. Genus G coronatum 2. Capsule with sometwhat thick neck, but not
inflated or distinctly wrinkled; leaves of fertile stems ovate to
ovate-lanceolate, laminal margins recurved proximally; bulbils one to many
per leaf axil, leafy or not ……………………………………………………………………………..
3 3. Bulbils large, 200--750 um, leafy,
primordia rising from near base or mid-bulbil, 1-2 per leaf axil ………………………………………………...………………...…
24. Genus G bicolor 3. Bulbils smaller, 100--350 um, primordia leafy
from upper 1/3 of bulbil to short and peglike or lacking, 5--25 per leaf axil
…………………………………...…………………………………..
4 4. Bulbils small, 100--200 um, round or
cylindric, primordia lacking or very short and peg-like ……………………………………………………………………
28. Genus G gemmilucens 4.Bulbils mostly 150--350 um, pyriform to
conic, distinct primordia present …………..……...
5 5. Young bullbils 150--250 um long, primordia
narrow, acute and tooth-like …………………………………………………………….………
27. Genus G gemmiferum 5. Young bulbils mostly >200 um, primordia
broad, obtuse, leaf-like ………………………………………………………………………….23.
Genus G barnesii 6. Leaves red, distal lamina often hyaline,
costa strong, long excurrent, hairpoint colored, reddish, sometimes
spinulose; capsule cylindrical ……….….…
30. Genus G “rubrifolium” 6. Leaves golden, brown
to yellow-green, distal lamina colored, costa not reaching apex to short
excurrent in stout colored point, not spinulose; capsule cylindrical to ovate
…..………………. 7 7. Capsule ovate, with somewhat thickened
occasionally wrinkled neck, tapered to narrow mouth; leaves flat to weakly
concave, triangular; long slender evenly foliate innovations with red stems
common, leafy to occasionally with leaves reduced or nearly absent ……………………………………………………………………
25. Genus G californicum 7. Capsule cylindrical, mouth wider than urn,
neck not thick or wrinkled; leaves distinctly concave, ovate to
ovate-lanceolate, innovations short, gemmiform with stems not distinctly
reddish ……………………………………..………………..…
29. Genus G “brassicoides” HAPLODONTIUM
2 Species
1. Leaf margins revolute; peristome single,
spores 12-22 um ...… 1. Haplodontium macrocarpum 1. Leaf margins plane; peristome absent,
spores 10-13 um..…..……
2. Haplodontium tehamensis GENUS L
1 Species
1. Genus
L systylium. GENUS P
3 Species
1. Stems long, leaves distant, decurrent, apex
rounded, strong bi- to multi-stratose limbidium present …………………………………….………..……….
3. Genus P incrassatolimbata 1. Stems short, leaves crowded, not decurrent,
apex acute, limbidium weak or absent, unistratose …………………………………………………………………………………………………….
2 2. Synoicous; peristome reduced, exotsome
segments irregular, short, endostome adherent to exostome, fragile, cilia
absent, spores 18-22 um ….……..………
1. Genus P brachyneura 2. Dioicous; peristome well developed,
exostome teeth long, endostome not adherent, cilia present, spores 22-30 um ………………………………...…..……….
2. Genus P vinosula PLAGIOBRYUM
2 Species
1. Plants red-brown to red-green, dark,
innovations leaves erect or erect-spreading, ovate-lanceolate, spores
released as tetrads ……………………………….....
1. Plagiobryum demissum 1. Plants silver-pink, pale, innovations
julaceous, leaves ovate, spores released singly …………………………..………………………………………………….
2. Plagiobryum zierii PTYCHOSTOMUM
38 Species
1. Stems mostly evenly foliate, not comose;
leaf base same color as rest of leaf, usually green, occasionally red or
pink, laminal margins with indistinct, partially bistratose 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 ….…..………………………………
2 1. Stems evenly foliate to comose with leaves
distinctly enlarged above, leaf base mostly reddish, laminal margins with
distinct to indistinct unistratose 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, perichaetial and
inner comal leaves with group of subalar inflated pink cells …………………………………………………………………..…………….3 2. Plants medium to large, dioicous, peristome
mostly perfect, spores small (<22 um) …………………………………………………………………………………….………
Pallentia 2. Plants small to medium, monoicous,
peristome reduced, cilia short or absent, spores generally large (>25 um) …………………………………………………………….……..
Amblyophyllum 3. Plants small, densely caespitose, leaves
ovate-lanceolate, somewhat concave, limbidium indistinct to absent, proximal
lamina cells shorter than median and distal cells; short deciduous branches
sometimes present in axils of distal leaves; dioicous ……..…………….
Caespitibryum 3. Plants medium to large, caespitose, comose
to evenly foliate, leaves broadly ovate, ovate-lanceolate to lanceolate, flat
or concave, proximal laminal cells same length as cells above or somewhat
longer, deciduous branchlets absent; dioicous, autoicous, synoicous or
polyoicous …………………………………………………………………………………………..………..
4 4. Plants evenly foliate to loosely comose; dioicous,
autoicous, synoicous or polyoicous; peristome perfect, basal membrane >1/2
height of exostome, spores small, 10--22 um; filiform gemmae occasionally
present in leaf axils ….……………………………….……
Pseudotriquetra 4. Plants comose to caespitose, monoicous;
peristome reduced, cilia mostly short or absent, basal membrane low, mostly
<1/2 height of exostome, spores 18--50 um; filiform gemmae lacking …….………………………………………………………………………….……..
Penduliformia AMBLYOPYLLUM
10 Species
1. Autoicous; leaf with indistinct unistratose
limbidium, broadly obtuse, costa not reaching apex …………………………………………………………………………
5. Ptychostomum marrattii 1. Autoicous, synoicous or polyoicous; leaf
with distinct partially bistratose limbidium, acute to acuminate, costa not
reaching apex to more commonly percurrent or excurrent …..……...……
2 2. Synoicous; endostome fragile, adherent to
exostome, cilia absent, spores 18--22 um, capsules bright red at maturity,
short and globose ……………………..………
10. Ptychostomum wrightii 2. Autoicous, synoicous or polyoicous; endstome free
from exostome, although reduced, cilia short or absent, capsules brown or
red-brown at maturity, if red then spores larger, clavate to pyriform; spores
>25 um .….……………………………………………………………….…… 3 3. Autoicous; capsule short pyriform to short
ovate, red at maturity …….……………………...
4 3. Autoicous, synoicous or polyoicous; capsule
pyriform to elongate ovate, brown at maturity ………………………………………………….…………………………………………….…..
5 4. Rosette leaves broadly ovate, concave, distal
laminal margins entire, capsule short pyriform ……………………………………………………………………….
3. Ptychostomum axel-blyttii 4. Rosette leaves narrow ovate-lanceolate,
flat, distal laminal margins serrulate, capsule ovate ……………………………………………………………………....
1. Ptychostomum acutiforme 5. Capsule short pyriform to short ovate with
thick neck, seta long …………….…………….…
6 5. Capsule elongate ovate or pyriform, tapered
to slender neck, seta short …………….………..
7 6. Synoicous; leaves concave, broadly ovate,
costa not reaching apex to percurrent, capsule short ovate, with thick neck,
abruptly contracted to seta ……...………..
5. Ptychostomum calophyllum 6. Polyoicous; leaves ovate-lanceolate, flat,
costa excurrent in short hairpoint, capsule short pyriform, neck tapered to seta ………..….………………………….
9. Ptychostomum warneum 7. Polyoicous; median laminal cells wide,
22--35 um, capsule elongate pyriform, symmetric, pendulous, exothecial cells
at mouth in 2--4 rows of transversely rectangular, exostome teeth with small
holes at base along mid-line ………………………..
7. Ptychostomum purpurascens 7. Autoicous or synoicous, median laminal
cells narrower, 18--25 um, capsule elongate ovate to elongate pyriform,
symmetric or curved-gibbous, exothecial cells at mouth mostly quadrate,
exostome teeth lacking small holes …………………………………………………….………..
8 8. Autoicous; costa of rosette leaves
percurrent to short percurrent in stout point, capsule elongate-pyriform,
somewhat curved-gibbous, exostome teeth distally yellow, spores 28--35 um ………………………………………………….……………..……
8. Ptychostomum uliginosum 8. Autoicous or synoicous; costa of rosette
leaves distinctly excurrent in medium-length hairpoint, capsule elongate
pyriform to ovate, symmetric or curved-gibbous, exostome teeth distally
hyaline, spores 24--30 um .……………………………………………..………………..
9 9. Autoicous; hairpoint denticulate, capsule
symmetric, 5--6 rows of orange quadrate exothecial cells at mouth ………………………………………………………….
4. Ptychostomum bryoides 9. Synoicous; hairpoint smooth; capsule
curved-gibbous, 2--3 rows of orange to brown quadrate exothecial cells at
mouth ……………………...………………………
2. Ptychostomum arcticum PALLENTIA
8 Species
1. Leaves longly and broadly decurrent, acute,
costa percurrent …….. 18. Ptychostomum weigelii 1. Leaves not or weakly decurrent, obtuse, acute or
acuminate, costa not reaching apex to excurrent
.………………………………………………………………………………………... 2 2. Leaves broadly ovate to suborbicular, leaf
apex rounded to obtuse, costa not reaching apex to percurrent ……………………………………………………………………………………..….
3 2. Leaves ovate to ovate-lanceolate, leaf apex
acute, costa typically percurrent to short excurrent …………………………………………………………………………………………………….
4 3. Plants red, leaves strongly concave, costa
often percurrent, spores 15--20 um ………………………….……………………………………....….
12. Ptychostomum cryophilum 3. Plants bright green, lacking red tints,
leaves weakly concave to flat, costa not reaching apex, spores 12--16 um ……...…………………………………..……
11. Ptychostomum cyclophyllum 4. Young leaves pink or red-tinged, leaf
margins recurved, filiform gemmae sometimes present in axils of distal
leaves; capsule long clavate, somewhat curved ………………………….….....
5 4. Young leaves yellow, green to brown-green,
leaf margins mostly plane, filiform gemmae lacking; capsule short, pyriform
to turbinate, not curved ………..………………………………
7 5. Laminal cells strongly incrassate, porose,
filiform gemmae common in leaf axils; endostome basal membrane low, cilia
often short …..……………………….…..
15. Ptychostomum rutilans 5. Laminal cells thin walled, not porose,
filiform gemmae rarely present; endostome basal membrane about ½
height of exostome, cilia usually long, well developed …………………….
6 6. Capsule <4 m long, weakly curved to
contorted when mature, exothecial cells short and broad, 1-1.5:1, 20--40 um
by 30--50 um, spores 20--30 um …………………
14. Ptychostomum pallens 6. Capsule often >5 mm, strongly curved to
contorted when mature, exothecial cells elongate, 2--3:1, 14--25 x 40--80 um,
spores 14--20 ……..……….……………
13. Ptychostomum meesioides 7. Leaves green to brown-green, strongly
shrunken and contorted when dry, median laminal cells 15--25 (30) um wide ………..…………………………………….
17. Ptychostomum turbinatum 7. Leaves bright yellow-green, weakly
contorted to somewhat imbricate when dry, median laminal cells (20) 30--40
(60) um wide …………..…………….....
16. Ptychostomum schleicheri PENDULIFORMIA
11 Species
1. Endostome strongly adherent to exostome,
giving exostome teeth a chambered look, leaves acuminate, costa excurrent in
long hairpoint, capsules short pyriform, spores 22-35 um ……………………………………………………………………....
26. Ptychostomum pendulum 1. Endostome mostly free from exostome,
exostome teeth not chambered, leaves acute to acuminate, costa not reaching
apex to excurrent in long hairpoint, capsule pyriform, ovate to clavate,
spores 25--50 um …………………………………………………………………….....
2 2. Leaves strongly concave, 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 ………………………....
25. Ptychostomum nitidulum 2. Leaves flat or
concave, twisted or contorted when dry, rarely innovations imbricate, costa
of at least rosette leaves excurrent in short to long hairpoint, limbidium mostly
distinct, spores 18--50 um, capsule pyriform to elongate ovate, straight to
gibbous, cilia present or absent ……….….. 3 3. Autoicous; leaves concave, limbidium
indistinct, costa of lower stem leaves not reaching apex, at least some
rosette leaves with costa short excurrent, filiform gemmae often present in
leaf axils of sterile stems; spores large, 30--40 um; cilia absent ……….……
27. Ptychostomum reedii 3. Synoicous or auoicous; leaves flat to
concave, limbidium distinct, well developed, costa of most leaves short to
long excurrent, filiform gemmae absent, spores 18—50
um, cilia present or absent …………………………………………………………………………………………….
4 4. Mouth of mature capsule red, spores small,
18--28 um, endostome membrane about ½
height of exostome, cilia usually present ………………………………………………………….…….
5 4. Mouth of mature capsule yellow or pale
orange, spores large, 25--50 um, endostome membrane low, <1/2 height of
exostome, cilia rudimentary or absent …………………….…….
7 5. Cilia rudimentary or sometimes 1--2 present;
synoicous; capsule straight, mouth straight, spores 18--28 um ………..……………………………….………..
19. Ptychostomum amblyodon 5. Cilia usually present and well developed,
autoicous or polyoicous; capsule gibbous, mouth oblique, spores 18--24 um ……………………………………………………………..…….…...
6 6. Autoicous, capsule short pyriform …………………...……………….
29. Ptychostomum teres 6. Polyoicous, capsule elongate pyriform ……………………….
22. Ptychostomum intermedium 7. Leaves acute to acuminate, costa of rosette
leaves excurrent into short recurved smooth hairpoint, seta long, 2--6 cm,
spores 25--50 um ….…………..………………………………….
8 7. Leaves acuminate, costa of rosette leaves
excurrent into medium or long straight denticulate hairpoint, seta short,
1--2 cm long, spores 25--30 um …...……....………………………………
9 8. Leaves weakly concave to flat, not keeled,
acuminate, hairpoint distinct, short to medium, spores 40--50 um, seta 2--6
cm long ………………………..…….
24. Ptychostomum longisetum 8. Leaves strongly concave, keeled, acute, hairpoint
percurrent to very short excurrent, spores 25—30
um, seta 2—3 cm ………………………………….…..….
23. Ptychostomum knowltonii 9. Spores dark, blackish, endostome membrane
nearly ½ height of exostome,
sometimes adherent to exostome at base, operculum low-convex, not or weakly
apiculate ……………………………………………….………………...
20. Ptychostomum archangelicum 9. Spores pale, endostome membrane very low,
free from exostome, operculum conical, distinctly apiculate ……………….………………………….…….…………………………...
10 10.
Synoicous, exostome teeth lacking small holes at base, spores 28--32
um, distal laminal cells somewhat incrassate, porose ………….……………………………..21.
Ptychostomum curvatum 10. Polyoicous; exostome teeth with small
holes on midline near base, spores 25--30 um, distal laminal cells thin
walled, not porose ..…………………………..…....
28. Ptychostomum salinum PSEUDOTRIQUETRA
6 Species
1. Autoicous, spores 16-22 um ……………………………….……
34. Ptychostomum pallescens 1. Dioicous, synoicous or polyoicous, spores
mostly 10-16 um .………………………………..
2 2. Dioicous, leaves distinctly decurrent,
limbidium strong, filiform gemmae occasional in axils of leaves ……………………………………………………….
35. Ptychostomum pseudotriquetrum 2. Dioicous, synoicous or polyoicous, border
indistinct to somewhat strong, leaves not or weakly decurrent, filiform
gemmae lacking ……….………………………………………….………….
3 3. Leaves strongly contorted when dry,
concave, apex rounded to obtuse, costa not reaching apex or percurrent;
dioicous …………….……………………….……
33. Ptychostomum neodamense 3. Leaves somewhat twisted or shrunken when
dry, not concave, apex acute to acuminate, costa strong, excurrent in short
to long hairpoint; synoicous or polyoicous …………………..……….
4 4. Stems elongate, leaves evenly foliate, not
much enlarged at apex, somewhat decurrent, costa short-excurrent in stout
hairpoint; synoicous ….………………...…….
30. Ptychostomum bimum 4. Stems short, leaves distinctly comose,
enlarged above, not decurrent, costa excurrent in long hairpoint; synoicous
or polyoicous …...………………………………….………………………
5 5. Synoicous; limbidium narrow, yellowish,
2--3 cells wide ..…. 31. Ptychostomum creberrimum 5. Polyoicous; limbidium wide, 3 or more cells
wide, same color as lamina ………………………..……………………..….………………
32. Ptychostomum lonchocaulon CAESPITIBRYUM
3 Species
1. Plants small, most leaves <1.5 mm, upper
lamina in older leaves becoming hyaline, proximal laminal cells
short-rectangular to quadrate, costa variable, from not reaching apex to
excurrent in slender hyaline hairpoint ….……………………………………………
38. Ptychostomum kunzei 1. Plants larger, most leaves 1.0--2.5 mm,
upper lamina green, proximal laminal cells long rectangular, costa strong,
long excurrent in brown or yellow-brown hairpoint ………….....……
2 2. Leaves strongly concave, hairpoint long,
>1/2 length of lamina, spinose ……………………………………………………………..……..……
36. Ptychostomum badium 2. Leaves weakly concave, hairpoint shorter,
typically <1/2 length of lamina, not spinose ……………….…………………………………………………
37. Ptychostomum angustifolium RHODOBRYUM
2 Species
1. Comal leaves 18--50, mostly >20, costa
reaching apex to short percurrent, margins strongly recurved; costa in
cross-section with small distinct band of stereids; eastern and southern
North America …..……………………………………………….…..……….1.
Rhodobryum ontariense 1. Comal leaves 15--22, mostly <20, costa
not reaching apex, margins weakly recurved below; costa in cross-section with
very small or nearly absent stereid band; coastal Alaska …………………………………………………………………………….
2. Rhodobryum roseum ROELLIA
1 Species
1. Roellia
roellii ROSULABRYUM
11 Species
1. Filiform gemmae
present in axils of upper innovation leaves ……...………………………… 2 1. Filiform gemmae absent
……………………………………………………………………… 5 2. Innovations short,
rosulate, leaves large, mostly >2 mm long, distal lamina margins strongly
serrate, limbidium strong ………..…………………………………....... 2. Rosulabryum andicola 2. Innovations rosulate
or elongate and evenly foliate, leaves small, mostly < 2 mm long, distal
lamina margins serrulate to almost smooth, limbidium weak or absent ………………………....
3 3. Innovations rosulate,
leaves obovate, flat, rhizoidal tubers red, filiform gemmae brown
…………………………...……………………………………………. 9. Rosulabryum
laevifilum 3. Innovations evenly foliate,
leaves ovate, concave, rhizoidal tubers various colors, filiform gemmae red,
red-brown or brown ….…….……………………………………………………… 4 4. Rhizoidal tubers
orange to pink-orange, brighter than rhizoids, filiform gemmae red
………..…………………………………………………..…… 10. Rosulabryum
pseudocapillare 4. Rhizoidal tubers and
filiform gemmae brown ……….…………….. 7. Rosulabryum
flaccidum 5. Plants with elongate
evenly foliate stems, leaves large, 2--4 mm long, distinctly decurrent,
margins sharply serrate distally …………….…………………….….. 1. Rosulabryum andersonii 5. Plants rosulate or if
evenly foliate then leaves small, < 2.0 mm long and distal margins not
sharply serrate, leaves decurrent or not ……………………………………………...….….……. 6 6. Leaves medium-sized to
large, 2--4 mm long, often in 2 or more interrupted tufts, margins serrate,
limbidium strong or absent distally, costa excurrent in short stout
hairpoint, lamina cells incrassate, distinctly porose
..……………………………………………………..………..……. 7 6. Leaves mostly < 2
mm long, usually not in interrupted tufts, variously contorted, twisted or
imbricate, margins smooth to serrulate, limbidium present or nearly absent,
costa various, not reaching apex to long excurrent in long hairpoint, lamina
cells thin to firm-walled, not or weakly porose …...………………………………………………………………………………………..
8 7. Distal margins of
leaves lacking limbidium, leaves often in 2 or more interrupted comal tufts,
hairpoint recurved when dry …………………………………....…… 3. Rosulabryum canariense 7. Distal margins of
leaves with strong limbidium, leaves usually in single tufts, hairpoint
variously straight to curved but not distinctly recurved ……………….. 2. Rosulabryum andicola 8. Stems more or less
evenly foliate, leaves somewhat contorted to julaceous, distal margins of
leaves with weak or absent limbidium, rhizoidal tubers absent ……….... 5. Rosulabryum elegans 8. Stems rosulate,
although innovations sometimes evenly foliate, leaves variously contorted or
twisted, rarely innovations leaves imbricate, distal margin of leaf with
strong to nearly absent limbidium, rhizoidal tubers present
….………………………………………………………….. 9 9. Plants with reddish
tints, leaves distinctly decurrent, costa strong in short hairpoint
……………………………………………………………………… 6. Rosulabryum
erythroloma 9. Plants green,
brown-green to red-green, leaves not decurrent, hairpoint mostly excurrent in
medium to long hairpoint …………………………………..………………….…………..…… 10 10. Plants polyoicous
(synoicous, some shoots unisexual, others with unisexual lateral gametangia),
rhizoidal tubers bright orange-red to crimson, lighter than rhizoids, distal
leaf margins distinctly serrulate, capsule often strongly nodding, red at
maturity …………………………………………………………………...… 11. Rosulabryum torquescens 10. Plants dioicous,
rhizoidal tubers red-brown to brown, same color as rhizoids, distal leaf
margins finely serrulate to almost smooth, capsule often inclined, brown
..…..……………..… 11 11. Limbidium of leaves
weak or absent, distal margins more or less smooth, innovations evenly
foliate, often red-brown and somewhat imbricate, rosulate leaves irregular
twisted …………………………………………………………………….... 8. Rosulabryum
gemmascens 11. Limbidium usually
present, margins finely serrulate, innovations short, rosulate, green with
leaves contorted, rosulate leaves strongly spirally twisted around stem
…………………………………………………………………..……… 4. Rosulabryum
capillare List of Genera and Species –
11 genera, 101 species (sp. nov? –
tentative as of 3/19) Brachymenium 1.
Brachymenium macrocarpum Card.
Bryum
1. Bryum argenteum
Hedw. 2. Bryum blindii
Bruch & Schimp. 3. Bryum calobryoides
Spence 4. Bryum concinnatum
(Spruce) Lindb. 5. Bryum gerwigii
(C. Mueller) Limpr. 6.
Bryum lanatum
P. Beauv.
7.
Bryum leptostomoides Schimp.
8.
Bryum julaceum (Gaert., Meyer & Scherb.) Schimp.
9. Bryum nanoargenteum
sp. nov? 10. Bryum
oblongum Lindb.
11.
Bryum veronense De Not.
Genus
G
Section
Alpiniformia 1. Genus G alpinum
(With.) Spence 2. Genus G gemmiparum
(De Not.) Spence 3. Genus G microchaeton
(Hampe) Spence 4. Genus G mildeanum
(Jur.) Spence 5. Genus G miniatum
(Lesq.) Spence 6. Genus G muehlenbeckii
(Bruch) Spence 7. Genus G pseudomuehlenbeckii
sp. nov? 8.
Genus G pseudoriparium sp. nov?
Section
Apalodictyon 9. Genus G bornholmense (Winkel. &
Ruthe) Spence 10. Genus G demaretianum
(Arts) Spence 11. Genus G klinggraeffii (Schimp.)
Spence & Ramsay
12. Genus G radiculosum (Bridel)
Spence & Ramsay
13. Genus G rubens (Mitt.) Spence & Ramsay
14.
Genus G ruderale (Crundw. & Nyholm) Spence
15. Genus G subapiculatum
(Hampe) Spence & Ramsay 16. Genus G tenuisetum
(Limpr.) Spence & Ramsay 17.
Genus G valpariense (Thér.) Spence
18.
Genus G violaceum (Crundw. & Nyholm)
Spence Section
Apiculata
19. Genus G apiculatum (Schwaegr.) Spence & Ramsay Section
Dicranobryum 20.
Genus G exile (Dozy & Molk.)
Spence & Ramsay 21. Genus G mexicanum (Mont.) Spence Section
Doliolidium 22. Genus G barnesii (Wood ex Schimp.) Spence
23.
Genus G bicolor (Dicks.) Spence 24.
Genus G brassicoides sp. nov. 25.
Genus G californicum (Lesq.) Spence 26.
Genus G coronatum (Schwaegr.)
Spence & Ramsay 27.
Genus G gemmiferum (Wilcz. &
Dem.) Spence 28.
Genus G gemmilucens (Wilcz. & Dem.)
Spence 29.
Genus G rubrifolium sp. nov. Haplodontium
Hampe (Mielichhoferia Nees &
Hornsh. in part)
1. Haplodontium macrocarpum
(Hook.) Spence 2. Haplodontium tehamensis
(Showers) Spence Genus L
(C. Muell.) Spence
1.
Genus L systylium (C. Muell.) Spence
Genus
P Spence
1. Genus P brachyneura
(Kindb.) Spence 2. Genus P incrassatolimbata
(Card.) Spence 3. Genus P vinulosa (Card.) Spence Plagiobryum Lindb.
1.
Plagiobryum demissum (Hook.) Lindb.
2. Plagiobryum zierii
(Hedw.) Lindb. Ptychostomum Hornschuch
Section Amblyophyllum 1.
Ptychostomum acutiforme
(Limpr.) Spence
2. Ptychostomum
arcticum (Brown) Spence 3. Ptychostomum axel-blyttii
(Philib.) Spence 4. Ptychostomum bryoides (R. Brown) Spence
5. Ptychostomum calophyllum
(R. Brown) Spence 6. Ptychostomum cernuum
(Hornsch.) Spence 7. Ptychostomum
marattii (Wilson) Spence 8. Ptychostomum
purpurascens (R. Brown) Spence 9. Ptychostomum
warneum (Brifel) Spence 10. Ptychostomum
wrightii (Sull.) Spence Section Pallentia 11. Ptychostomum cyclophyllum
(Schwaegr.) Spence 12.
Ptychostomum cryophilum (Mart.) Spence 13. Ptychostomum meesioides
(Kindb.) Spence 14. Ptychostomum pallens
(Swartz) Spence 15. Ptychostomum rutilans
(Bridel) Spence 16. Ptychostomum schleicheri
(Schwaegr.) Spence 17. Ptychostomum turbinatum
(Hedwig) Spence 18. Ptychostomum weigelii
(Spreng.) Spence Section
Penduliformia 19. Ptychostomum archangelicum
(Bruch & Schimp.) Spence 20. Ptychostomum amblyodon
(C. Muell.) Spence 21. Ptychostomum curvatum
(Kaurin & Arnell) Spence 22. Ptychostomum intermedium
(Bridel) Spence. 23. Ptychostomum knowltonii
(Barnes) Spence 24. Ptychostomum longisetum
(Schwaegr.) Spence 25. Ptychostomum nitidulum
(Lindb.) Spence 26. Ptychostomum pendulum
Hornsch. 27. Ptychostomum reedii
(Robins.) Spence 28. Ptychostomum salinum
(Limpr.) Spence 29. Ptychostomum teres
(Lindb.) Spence Section
Pseudotriquetra 30. Ptychostomum bimum
(Schreber) Spence 31. Ptychostomum creberrium
(Tayl.) Spence & Ramsay 32. Ptychostomum lonchocaulon
(C. Muell.) Spence 33. Ptychostomum neodamense
(Itzig.) Spence 34. Ptychostomum pallescens
(Schwaegr.) Spence 35. Ptychostomum pseudotriquetrum
(Hedwig) Spence & Ramsay Section Caespitibryum 36.
Ptychostomum
angustifolium (Bridel) Spence & Ramsay 37. Ptychostomum badium (Bridel)
Spence
38. Ptychostomum kunzei (Hoppe et.
Hornsch.) Spence Roellia
Kindb. 1. Roellia roellii
(Broth. ex Ruell) Andrew. ex Crum Rhodobryum (Schimp.) Limpr.
1.
Rhodobryum ontariense (Kindb.) Paris 2. Rhodobryum roseum
(Hedw.) Limpr Rosulabryum Spence
Section
Rosulata 1. Rosulabryum andersonii
(Crum) Spence 2. Rosulabryum andicola
(Hook.) Spence 3. Rosulabbryum canariense
(Bridel) Spence Section
Trichophora 4. Rosulabryum capillare
(Hedw.) Spence 5.
Rosulabryum elegans Nees ex Bridel 6. Rosulabryum erythroloma
(Kindb.) Syed 7. Rosulabryum flaccidum
Bridel 8. Rosulabryum gemmascens
Kindb. 9. Rosulabryum laevifilum
Syed 10. Rosulabryum pseudocapillare
Besch. 11. Rosulabryum torquescens
(De Not.) Spence |