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|Alsia californica (W. J. Hooker & Arnott) Sullivant [Leptodontaceae]|
|photo by Game||drawing by Montalvo of Norris 10414|
Mail a correction to Paul Wilson
Proc. Amer. Acad. Arts Sci. 3: 185. 1855. -- Neckera californica Hook. & Arnott, Bot. Capt. Beechey's Voyage 162. 1833.
Plants forming loose, rather deep wefts; glossy olive-green to yellowish green, often strongly bleached in highly insolated sites. Primary Stems (stolons) to 2 cm long, prostrate and microphyllous. Secondary stems to 8 cm high, ascending or sometimes decumbent ( In areas exposed to salt spray even the stipe is prostrate and only the ultimate branches curl upward.), pinnately or sometimes bipinnately branched in a single plane with branch apices blunt to attenuate or even flagelliform. paraphyllia foliose, deeply and narrowly lobed into narrow segments which are several cells wide at base but uniseriate in the distal 1/2. Stipe Leaves concave, lightly plicate, imbricate with somewhat reflexed acumen when dry, erect-spreading when moist; ovate to ovate-lanceolate, to 2.5 mm, 3–4: 1, decurrent in a triangular segment which may reach to the base of the adjacent leaf. Branch leaves concave and imbricate, not plicate, erect to weakly spreading when moist, ovate, to 1.5 mm, 2–3: 1, narrowly decurrent. Margins reflexed to recurved from base to near mid-leaf and thus appearing weakly plicate, plane above, almost entire to remotely and irregularly serrulate in distal 1/2. apex acute to acuminate, concolorous. Costa thin, extending about 2/5 to 1/2, often forked apically. Median Laminal cells elliptic to rhomboidal, longest in juxtacostal region but becoming shorter toward the apices and margins, 8–12 µm wide, 2–6: 1, thick-walled (lumen/wall ratio about 5:1) but only lightly pitted, smooth. Apical cells often nearly isodiametric. Alar cells thin- to thick-walled, numerous, quadrate to somewhat transversely elongate, to 10 µm wide, only gradually demarcated from laminal cells, extending nearly to costa and an equal distance up the margin. Axillary hairs to 90 µm, 4-celled, with one basal brown cell, not offset from leaf insertion. Rhizoids red-brown, smooth, to 10 µm in diameter, arising fascicled from stem at abaxial costal insertion. Pseudoparaphyllia narrowly lanceolate, 2–3 cells wide at base but uniseriate in distal 1/2, distinguishable from paraphyllia by lack of lobing.
Sexuality dioicous with male plants somewhat smaller and less branched than females, usually located higher on the phorophyte than the females. Perigonia somewhat longer than subtending leaves, nearly spherical, bud-like, densely cloaking the main axis and larger branches. Perigonial bracts deeply concave but abruptly attenuate near apex with attenuation reflexed. Perichaetia on main axis, the bracts to 4 mm long, ecostate or with a rudimentary costa, strongly convolute and investing the seta and the base of the capsule, gradually acute to acuminate from a lanceolate base. Setae smooth, red-brown, 4–6 mm long, mostly geniculate at exit from perichaetia. Capsule exserted, brown, cylindric and symmetrical, to 2 mm, about 2: 1, not sulcate but somewhat wrinkled when dry. Exothecial cells 12–18 µm wide,mostly less than 2: 1 but with scattered longer cells, moderately thin-walled (lumen/wall ratio 7–10:1), with a few phaneroporous stomata on neck. Operculum inclined-rostrate, 1/3–1/2 as long as urn. Annulus poorly developed or occasionally absent, but with suboral cells small, quadrate and thin-walled, in about 10 rows. Exostome teeth 16, erect and pale,linear-lanceolate, 10–15: 1, smooth to inconspicuously papillose. Endostome segments narrow, smooth, about half as long as exostome and with low basal membrane and no cilia. Calyptra cucullate, smooth or with a few hairs. Spores spherical 20–25 µm, smooth to lightly papillose, green and often with premature germination.
Like other members of the Leucodontaceae, Alsia produces archegonia and antheridia almost throughout the year. Most sporophytes are, however, produced in late spring to summer. Alsia californica is an almost strictly coastal lowland (to 300 m) moss with special concentration on the California coast. A collection (Ikenberry 1295a, Columbia, Tuolumne Co, California (UAC) is the only non-coastal site. I have been unable to recollect Alsia from that site but it can be noted that a number of maritime plants are found in the lower foothills of the west face of the Sierra. This may be a reflection of the oceanic embayment in the Central Valley during the high sea levels of interglacial times. The northernmost records cluster around Puget Sound in southern British Columbia and Washington. I hope for recollection to confirm the citation of an Alaska collection (Kellogg, sine loc., FH-JAM). The southernmost records are from Guadelupe Island on the Pacific Coast of Mexico. In coastal forests in California, Alsia is one of the most abundant, often dominant mosses on angiosperms, especially Alnus and Sambucus, In Oregon, Alsia virtually disappears from forested areas except for occasional presence on Sambucus in coastal areas. It occurs more often in Oregon and Washington on man-made substrata such as the trunks of orchard trees. It can reliably be found from California to Washington on rocks on offshore islands which support large colonies of seabirds.
In its typical epiphytic expression, Alsia californica is almost unmistakeable. No other coastal epiphyte within its range has the combination of dioicous plants with plumose stems from a prostrate axis along with very numerous and conspicuous bulfiform perigonia and acicular perichaetia. Microscopically, the abundant lobed paraphyllia may suggest Dendroalsia abietina (Hook.) Britt. but that latter plant has prorations on the median leaf cells so large as to be visible with a handlens. Keying of Alsia may sometimes be difficult because of a tendency for some of the leaves to be almost ecostate. There are epilithic expressions of Alsia, often on offshore islands in colonies of sea birds. These plants are often without perigonia or perichaetia, and the prostrate stems and branches allow field interpretation as a very strange Brachythecium. The bird-island form (repeated on concrete culverts on the mainland) will be identified by the paraphyllia. Another form of Alsia is occasionally found near the tops of large coastal Pseudotsuga. This form has unusually julaceous branchlets, and the paraphyllia are simple and unlobed, lanceolate appendages. Such a form was annotated by Monte Manuel as "not an Alsia". My further study more or less convinces me that this form is referrable to Alsia californica.
It is abundant on the coast of northern California and in similar sites in Washington and British Columbia. Peculiarly, it is quite scarse in coastal Oregon.
|Literature||Bradshaw 1926; Brandegee 1891; Harthill et al. 1979; Holmberg 1969; Jamieson 1969; Kellman 2003; Koch 1950a; Koch and Ikenberry 1954; Lawton 1971; Lesquereux 1868; Manuel 1974; McCleary 1972; Millspaugh and Nuttall 1923; Sayre 1940; Shevock and Toren 2001; Steere 1954; Sullivant 1856; Thomson and Ketchledge 1958; Watson 1880; Whittemore and Sommers 1999; Yurky 1990, 1995. As Alsia californica var. flagellifera Renauld and Cardot 1889.|
|Illustrations||Malcolm et al. 2009 p. 311; Brotherus 1924-25; Lawton 1971; Manuel 1974; Sharp et al. 1994.|
|Bioregions||CaR, CW, NW, SW.|
|Vouchers||Humboldt Co.: Dry Lagoon State Park, Norris 46053; Lake Co.: Hidden Valley at Hartman Creek north of Middletown, Toren & Dearing 5195 (CAS); Monterey Co.: Highway 1, about 10 miles south of Lucia, Los Padres National Forest, R. & I. Duell 1836 (UC); San Luis Obispo Co.: Highway 1 north of Moro Bay about 0.5 mile east of Cambria, Norris 68209; Santa Cruz Co.: Zayante Canyon Road, 4 miles south of Summit Road north of Felton, Norris 86871. Sonoma Co.: Austin Creek near intersection with Russian River, Norris 53138; Yuba Co.: north of County Road 130, 3 miles north of Challenge, Dillingham 989 (CAS).|