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FNA | Family List | FNA Vol. 28 | Bryaceae

10a. PTYCHOSTOMUM subg. subg. CLADODIUM (Bridel) J. R. Spence, Phytologia. 91: 498. 2009.

Cladodium Bridel, Bryol. Univ. 1: 620, plate suppl. 3, fig. 9. 1826

Stems 0.5-4(-12) cm, usually comose to evenly foliate; sparsely to densely radiculose. Leaves ovate or ovate-lanceolate, distal leaves usually somewhat enlarged; base red or pink, red sap not present in cell lumina; limbidium distinct to indistinct, narrow, pale to green or yellow, 1-stratose throughout; subalar cells inflated, pink, on gametoecial and inner comal leaves; proximal laminal cells rectangular, (short-rectangular in P. pseudotriquetrum), same length as medial, sometimes longer; distal cells rhomboidal to hexagonal, not lax, usually 3-4:1. Capsule pyriform, ovate, or obovate; endostome sometimes adherent to exostome, segments with narrowly to broadly ovate perforations. Spores 10-50 µm, finely to strongly papillose, yellow, green, brown, or black.

Species ca. 35 (16 in the flora): North America, South America, Eurasia, Africa, Pacific Islands; arctic, montane, and boreal regions.

Species in subg. Claodium are distinctive in a number of characters, including 1-stratose border, relatively short proximal laminal 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 walls but clear sap. The leaves are erect-spreading when moist; the setae are red or red-brown (brown in P. salinum).


1 Sexual condition dioicous, autoicous, synoicous, or polyoicous; peristome well developed; endostome basal membrane 1/2 exostome height; spores 10-26(-30) µm   (2)
+ Sexual condition usually synoicous, rarely autoicous or polyoicous; peristome well developed or reduced; endostome basal membrane usually less than 1/2 exostome height; spores (8-)22-30(-50) µm   (9)
       
2 (1) Sexual condition autoicous; spores 18-22 µm; stems comose, innovations comose or somewhat elongate and evenly foliate.   11 Ptychostomum pallescens
+ Sexual condition dioicous, synoicous, or polyoicous; spores 10-26(-30) µm; stems tufted or comose, rarely evenly foliate, innovations comose or somewhat elongate and evenly foliate   (3)
       
3 (2) Sexual condition dioicous; leaf bases strongly decurrent; stems tufted, weakly comose to evenly foliate, usually strongly radiculose well toward stem apices, innovations elongate and evenly foliate; specialized asexual reproduction occasional, by leaf axil gemmae.   13 Ptychostomum pseudotriquetrum
+ Sexual condition dioicous, synoicous, or polyoicous; leaf bases not or weakly decurrent; stems tufted and/or comose, strongly or weakly radiculose, innovations comose or somewhat elongate and evenly foliate; specialized asexual reproduction absent   (4)
       
4 (3) Leaves strongly twisted to contorted when dry, concave; apices broadly acute to obtuse; costae usually not reaching apex to percurrent; sexual condition dioicous   (5)
+ Leaves somewhat twisted or contorted when dry, flat to weakly concave; apices acute to acuminate; costae short- to long-excurrent; sexual condition synoicous or polyoicous   (6)
       
5 (4) Leaves crowded; bases wide.   9 Ptychostomum neodamense
+ Leaves distant; bases narrow.   16 Ptychostomum subneodamense
       
6 (4) Innovations elongate, evenly foliate; leaves not much enlarged toward stem apex; bases weakly decurrent; costae short-excurrent; distal laminal cell walls sometimes very incrassate, sometimes oblique to costa; sexual condition synoicous.   2 Ptychostomum bimum
+ Innovations comose or somewhat elongate and evenly foliate; leaves usually enlarged toward apex; bases not or weakly decurrent; costae long-excurrent; distal laminal cell walls thin to firm, never oblique to costa; sexual condition synoicous or polyoicous   (7)
       
7 (6) Sexual condition synoicous; spores (18-)20-26(-30) µm.   5 Ptychostomum intermedium (in part)
+ Sexual condition synoicous or polyoicous; spores 10-18(-20) µm   (8)
       
8 (7) Sexual condition synoicous; limbidium in 2 or 3 rows, yellowish.   3 Ptychostomum creberrimum
+ Sexual condition polyoicous; limbidium in (2-)3-4(-6) rows, concolorous.   7 Ptychostomum lonchocaulon
       
9 (1) Endostome strongly adherent to exostome; exostome teeth appearing chambered; leaf apices acuminate; costae long-excurrent; capsules pyriform; spores (22-)25-30 (-36) µm.   12 Ptychostomum pendulum
+ Endostome usually not adherent to exostome; exostome teeth not appearing chambered; leaf apices obtuse, acute, or acuminate; costae not reaching apex to long-excurrent; capsules pyriform or obovate; spores (15-)22-50 µm   (10)
       
10 (9) Leaves strongly concave, ± imbricate to weakly contorted or shrunken when dry; costae not reaching apex to percurrent, rarely short-excurrent; spores 15-22(-25)µm; capsules short-pyriform, asymmetric, curved; endostome cilia usually present.   10 Ptychostomum nitidulum
+ Leaves flat or concave, twisted to contorted or shrunken when dry; costae percurrent to long-excurrent; spores (18-)22-50 µm; capsules obovate or short- to elongate-pyriform, symmetric (asymmetric, somewhat curved in P. intermedium); endostome cilia present or absent   (11)
       
11 (10) Sexual condition autoicous; leaves concave; limbidium indistinct; costae percurrent in proximal leaves to sometimes short-excurrent in distal leaves; specialized asexual reproduction by leaf axil gemmae; spores (28-)30-40 µm.   14 Ptychostomum reedii
+ Sexual condition synoicous; leaves flat to concave; limbidium strong; costae usually short- to long-excurrent; specialized asexual reproduction absent; spores (18-)20-50 µm   (12)
       
12 (11) Capsule mouths orange or yellow; endostome basal membrane 1/2 exostome height, cilia present, sometimes reduced; spores (18-)20-26(-30) µm   (13)
+ Capsule mouths yellow or red; endostome basal membrane usually less than 1/2 exostome height, cilia rudimentary or absent; spores (18-)20-50 µm   (14)
       
13 (12) Cilia rudimentary or sometimes 1 or 2, long, appendiculate to nodose; capsules symmetric, mouths straight.   4 Ptychostomum inclinatum
+ Cilia long, nodose; capsules asymmetric, mouths oblique.   5 Ptychostomum intermedium (in part)
       
14 (12) Leaf apices acute to obtuse; costae percurrent to short-excurrent, awns recurved; setae (2-)3-6(-8) cm; spores (18-)20-50 µm   (15)
+ Leaf apices acuminate; costae short- to long-excurrent, awns straight; setae 1-3 cm; spores (18-)22-30 µm   (16)
       
15 (14) Leaves strongly concave; apices acute to obtuse; costae percurrent to short-excurrent; spores (18-)20-30 µm; setae (2-)3-4 cm.   6 Ptychostomum knowltonii
+ Leaves weakly concave; apices acute; costae short-excurrent; spores (38-)40-50 µm; setae 3-6(-8) cm.   8 Ptychostomum longisetum
       
16 (14) Spores dark brown to black; exostome teeth with pores absent along mid line; endostome sometimes weakly adherent to exostome at base; operculum short-convex, not or weakly apiculate.   1 Ptychostomum archangelicum
+ Spores pale yellow-brown; exostome teeth with small pores present along mid line at base; endostome not adherent to exostome; operculum long-conic, distinctly apiculate.   15 Ptychostomum salinum


 

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