Atrichum

Plants small to robust, not polytrichoid, in loose tufts. Stems simple or rarely sparingly branched. Leaves without a differentiated sheath, membranous, strongly crisped and contorted when dry, widely spreading and plane to ± distinctly concave when moist, with a differentiated border of 1 or more rows of narrow, linear, thick-walled cells, the border usually 2-stratose, the margins toothed to mid-leaf or below with single or paired teeth (teeth and cells of leaf border papillose in A. crispum); costa narrow, abaxially toothed in distal half or rarely smooth, bearing adaxial lamellae; abaxial lamellae none; lamina 1-stratose except for margins, often obliquely transversely undulate and abaxially toothed along the undulations, or almost plane and not abaxially toothed; lamellae ± straight or tall with sinuous margins, rarely rudimentary and almost absent, entire in profile, the marginal cells in section not differentiated, smooth; basal cells of lamina short- to long-rectangular, in longitudinal rows; median cells subquadrate to transversely elongate-hexagonal, in ± regular longitudinal rows, thin- to firm-walled, with or without distinct trigones, slightly convex on both surfaces (adaxial surface bulging-mammillose in A. angustatum), smooth or with minute rounded to striate papillae. Sexual condition dioicous, polygamous, or synoicous; perigonia inconspicuous, the bracts resembling the leaves, or the bracts broadly ovate and overlapping, forming a conspicuous disc, sometimes with more than one male inflorescence in sequence per plant; perichaetial leaves usually narrower and longer than the stem leaves. Seta 1(–3) per perichaetium, straight to flexuose, twisted near the capsule when dry, smooth. Capsule short- to long-cylindric, ± straight to arcuate, terete, finely striate when dry, absent a basal constriction; stomata none; exothecium smooth, the cells elongate-rectangular, not pitted, the longitudinal walls thickened; operculum rostrate, the beak straight to arcuate; peristome teeth 32, simple, linear and crowded. Calyptra cucullate, smooth or merely hispid at tip. Spores finely papillose, appearing smooth.

Distribution

North America, West Indies, Central America, South America, Europe, Asia, Africa, Australia.

Discussion

Species ca. 20 (9 in the flora).

Atrichum is unique in the family in having distinctly bordered, transversely undulate leaves. The peristome, however, is almost identical to that of Psilopilum, suggesting a close relationship. Inflorescence types are important differences between species, but are sometimes difficult to demonstrate. Dioicous (presumably heterothallic) species have antheridia and archegonia on separate male and female plants (A. angustatum, A. crispulum, A. crispum, A. selwynii, A. tenellum). Polygamous (heteroicous) species have both unisexual and bisexual shoots, the latter protandrous, with a male inflorescence followed by a female after an interval of vegetative growth (A. altecristatum, A. cylindricum, A. undulatum). Atrichum flavisetum is synoicous, with antheridia and archegonia within the same involucre, the perigonia hidden among the perichaetial leaves.

T. C. Frye (1937) emphasized the fractions of leaf width obscured by the lamellae, without taking into account many useful characters and important distributional differences. His treatment of distinctive North American taxa as varieties of A. undulatum has delayed recognition of unique aspects of our flora. Three species of the flora are endemic to eastern North America (A. altecristatum, A. crispum, A. cylindricum), and one to the West (A. selwynii). Atrichum crispulum is disjunctive in eastern North America and eastern Asia, whereas true A. undulatum appears to be introduced in North America.

Excluded Species:

Atrichum oerstedianum (Müller Hal.) Mitten

North American specimens named as this species are Atrichum crispulum.

Selected References

Frye, T. C. 1937. Atrichum. In: A. J. Grout. 1928–1940. Moss Flora of North America, North of Mexico. 3 vols. in 12 parts. Newfane, Vt. and New York. Vol. 1, pp. 100–106.

Ireland, R. R. 1969. Taxonomic studies on the genus Atrichum in North America. Canad. J. Bot. 47: 353–368.

Ireland, R. R. 1971. Atrichum. In: E. Lawton. Moss Flora of the Pacific Northwest. Nichinan. Pp. 31–33.

Ireland, R. R. 1993. Scanning microscope study of spores of the genus Atrichum (Musci: Polytrichaceae). J. Hattori Bot. Lab. 74: 205–217.

Nyholm, E. 1971. Studies in the genus Atrichum P. Beauv. A short survey of the genus and species. Lindbergia 1: 1–33.

Lower Taxa

Atrichum altecristatum, Atrichum angustatum, Atrichum crispulum, Atrichum crispum, Atrichum cylindricum, Atrichum flavisetum, Atrichum selwynii, Atrichum tenellum, Atrichum undulatum

Key

1	Leaves narrow, less than 1 mm wide at midleaf; median leaf cells 8-17(-19) µm wide, bulging-mammillose on adaxial surface, often distinctly papillose; lamellae 6-9(-12), widely spreading, to 10(-15) cells high.	Atrichum angustatum
1	Leaves broader, mostly 1.5-2 mm wide; median leaf cells 18-24 µm wide or more, merely convex, smooth or finely striate papillose; lamellae 0-2(-6), ± straight, 0-6(-9) cells high	> 2
2	Median leaf cells as much as 40(-52) µm in longest dimension	> 3
2	Median leaf cells averaging 24 µm, rarely exceeding 27 µm in longest dimension	> 4
3	Leaves not undulate, with few or no teeth on abaxial surface; median leaf cells 35-52 µm wide, thin-walled, trigones absent; marginal teeth and border finely striate- papillose; e North America.	Atrichum crispum
3	Leaves strongly undulate, with oblique rows of teeth on abaxial surface; median leaf cells 27-40(-48) µm wide, firm-walled, with small trigones; marginal teeth crowded toward leaf apex, the teeth and border smooth; w North America.	Atrichum selwynii
4	Median leaf cells laxly subquadrate, thin-walled, without trigones; border narrow, with rather distant, small teeth.	Atrichum tenellum
4	Median leaf cells hexagonal, firm-walled, collenchymatous; leaves strongly bordered and sharply toothed	> 5
5	Leaves narrowly lanceolate to linear, subfalcate, not undulate, with few or no abaxial teeth; lamellae (1-)3-4 cells high, strictly parallel; Atlantic and Gulf coastal plain, Mississippi embayment.	Atrichum cylindricum
5	Leaves lanceolate to ovate-lanceolate to broadly elliptic-lanceolate, moderately to strongly undulate, with abaxial teeth on the undulations; lamellae 2-6(-9) cells high, laxly spreading	> 6
6	Plants small to medium, 1-3 cm; leaves narrowly lanceolate to lingulate; lamellae 4-6 cells high or higher	Atrichum altecristatum
6	Plants medium to robust, to 6 cm; leaves ovate-lanceolate to broadly elliptic-lanceolate; lamellae mostly 2-4 cells high	> 7
7	Sporophytes 2-6 per inflorescence, seemingly lateral, persisting from previous years; seta slender, yellowish; capsule short-cylindric and nearly straight; synoicous.	Atrichum flavisetum
7	Sporophytes usually solitary, terminal; seta stout, reddish; capsule somewhat curved and inclined, to arcuate and horizontal; dioicous or polygamous	> 8
8	Plants robust, 2-6 cm, the stems often with matted, whitish rhizoids; capsules 4-7 mm, straight to somewhat curved, erect to inclined; dioicous; soil or humus in shaded habitats, wet banks along streams or margins of fens and swamps.	Atrichum crispulum
8	Plants shorter, 1-5 cm, with reddish rhizoids near base of stem; capsules 2-4(-8) mm, arcuate, horizontal; polygamous; soil in dry weedy habitats, especially roadside ditches	Atrichum undulatum