BFNA Title: Bryaceae Keys
Author: John R. Spence 
Date: March 19, 2004
Edit Level: draft of keys
Version: 1

Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden
BFNA Web site: http://www.mobot.org/plantscience/BFNA/bfnamenu.htm

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 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[Editor's Note:  Several of the generic names were unpublished (either as new names or raised to generic rank) at the date of online publication of this draft treatment.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the FNA, these names are replaced by "Genus G, Genus P, and Genus L" where the letter is that of initial letter of the unpublished name.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 1850 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 2530 um, seta 23 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