BFNA Title: Andreaeaceae
Author: R. H. Zander
Date: Dec. 30, 2006
Edit Level: S
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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3. ANDREAEACEAE Dumortier

Richard H. Zander

 

Plants dark green to black, small to large, often in dense turf. Stems erect, irregularly branched, bearing rhizoids at base; central strand absent. Leaves erect or secund, sometimes falcate-secund, short- to long-lanceolate or panduriform; costa absent or single or branched, narrow to broad, percurrent or ending before the apex; in section of uniform cells, lacking lamellae; margins plane to weakly incurved, seldom recurved; laminal cells short throughout or occasionally elongate in leaf base, 1-stratose or 2- to multistratose. Specialized asexual reproduction rare, as filamentous gemmae from laminal cells. Sexual condition usually autoicous, mainly cladautoicous but occasionally gonioautoicous or dioicous; perichaetial leaves commonly differentiated, larger, convolute-sheathing. Sporophytes terminal on an elongate gametophytic stalk, the pseudopodium. Seta  essentially absent. Capsule erect, elliptic, opening by usually 4 lateral longitudinal valves; stomata, annulus, operculum and peristome absent. Calyptra tiny, campanulate-mitrate, often fugacious. Spores spheric, oval or tetrahedral, small to large, 10 to occasionally more than 100 \um, papillose.

 

Genus 1, species ca. 45 (11 in the flora): cosmopolitan.

 

The Andreaeaceae shares with Sphagnaceae the sporophyte raised on a pseudopodium, but the spore sac is derived from the endothecium, not the amphithecium as in the latter family. The spore sac arches over the massive, persistent columella. The longitudinal valves bulge open when the capsules are dry, closing when wet.  Erect, thallose protonemal appendages are common and distinctive.

 

1. ANDREAEA Hedwig, Spec. Musc. Frond. 47. 1801  *  [For J. G. R. Andreae, 1724--1793, apothecary of Hanover, Germany]

 

Plants commonly cemented to substrate. Stems with stalked mucilage hairs in leaf axils, stalks usually brown. Leaves spiraling around stem in several rows, usually brittle, commonly ending in a distinct apiculus of a single cell; costa sometimes poorly differentiated, sometimes not reaching the leaf insertion; laminal cells with thick longitudinal walls and often pitted or sinuose, transverse walls thin. Capsule 0.5--2 mm.

 

Species about 45, varieties about 48 (11 species in the flora): cosmopolitan.

 

This is a genus easily recognized in the field by the dark green to blackish dense turf strongly adherent to a rock habitat, the brittle leaves, and capsule opening by four longitudinal lateral slits. These species are largely temperate montane to Arctic-alpine in distribution, not uncommon on exposed acid rock surfaces. The present treatment follows B. M. Murray’s (1987, 1988a,b) detailed and thorough study closely except that Andreaea alpestris is treated as a synonym of A. rupestris and A. crassinervia is a synonym of A. rothii, following M. V. Corley et al. (1981), while infraspecific taxa are not recognized. The costa may be interpreted as strongly excurrent when it fills the leaf subula.

 

SELECTED REFERENCES   M. F. V. Corley, A. C. Crundwell, R. Düll, M. O. Hill, and A. J. E. Smith. 1981 [1982]. Mosses of Europe and the Azores: an annotated list of species, with synonyms from the recent literature.  J. Bryol.  11: 609--689.   Murray, B. M. 1987. Andreaeaceae. In: G. S. Mogensen, ed. Illustrated Moss Flora of Arctic North America and Greenland. 3. Andreaeobryaceae--Tetraphidaceae. Medd. om Grønland, Biosci. 23: 6--24.   Murray, B. M. 1988a. Systematics of the Andreaeopsida (Bryophyta): two orders with links to Takakia.  Beih. Nova Hedwigia  90: 289--336.   Murray, B. M.  1988b.  The genus Andreaea in Britain and Ireland.  J. Bryol. 15: 17--82. Schultze-Motel, W.  1970.  Monographie der Laubmoosgattung Andreaea. 1. Die costaten Arten. Willdenowia 6: 25--110.

 

1. Costa absent, leaves mostly lanceolate to ovate or panduriform.

2. Proximal laminal margins denticulate; Greenland . . . 1. Andreaea alpina

2. Proximal laminal margins entire (occasionally crenulate).

3. Cells of basal laminal margins quadrate . . . 2. Andreaea mutabilis

3. Cells of basal laminal margins mostly rectangular.

4. Leaves narrowly lanceolate, medial cell walls pitted and sinuose. . . . 3. Andreaea sinuosa

4. Leaves short-lanceolate to panduriform, medial cell walls variously pitted, straight.

5. Leaves generally curving or secund, short-lanceolate, widest proximally, apices oblique or symmetric. . . 4. Andreaea rupestris

5. Leaves mostly straight, panduriform, widest in distal half, apices usually symmetric. . . 5. Andreaea obovata

1. Costa present, leaves mostly subulate.

6. Leaf margins crenulate to strongly denticulate, laminal papillae present, strong; perichaetial leaves little differentiated . . . . 6. Andreaea nivalis

6. Leaf margins entire or occasionally weakly crenulate, laminal papillae absent; perichaetial leaves differentiated, convolute-sheathing.

7. Spores ca. 10--20 \um; basal laminal cells mostly rectangular, cell walls smooth and straight . . . 7. Andreaea blyttii

7. Spores 20--60 \um; basal laminal cells mostly rounded or quadrate to short-rectangular, walls commonly pitted and sinuose.

8. Costa weak, flattened distally, commonly not reaching the leaf insertion. . . 8. Andreaea heinemannii

8. Costa moderately differentiated to strong, terete, reaching the leaf insertion.

9. Spores usually 50--90 \um . . . 9. Andreaea megistospora

9. Spores usually 35--60 \um.

10. Spores usually 35--60 \um. . . 10. Andreaea rothii

10. Spores usually 20--30 \um . . . 11. Andreaea schofieldiana

 

 

 

1. Andreaea alpina Hedwig, Spec. Musc. Frond. 49, plate 7, fig. 2p. 1801

 

Plants reddish brown to black. Leaves erect-spreading to squarrose, straight, panduriform, widest distally, apex symmetric; costa absent; leaf margins denticulate along leaf base; basal laminal cells rectangular to long-rectangular, marginal cells similar, walls pitted, nodose; medial laminal cells rounded-quadrate to ovate, 1-stratose entirely or sometimes 2-stratose distally, lumens rounded; laminal papillae rare, low. Sexual condition cladautoicous; perichaetial leaves differentiated, convolute-sheathing.  Spores 18--28 \um.

 

Wet acidic or basic rock; s Greenland; n Europe; s and w South America.

 

The spores of Andreaea alpina are of two types, the brown spores generally smaller than the green, and apparently abortive. This comparatively robust species is easily identified by the panduriform leaves with basal marginal denticulations.

 

2. Andreaea mutabilis Hook. f. & Wilson, J. Bot. (London) 3: 536.  1844

 

Plants reddish to black. Leaves erect-spreading, straight or sometimes secund, lanceolate, widest proximally, apex symmetric; costa absent; leaf margins entire; basal laminal cells rectangular to long-rectangular, marginal cells quadrate, walls little pitted; medial laminal cells rounded-quadrate to ovate, 1-stratose entirely or sometimes 2-stratose in patches distally, lumens rounded; laminal papillae rare, low. Sexual condition cladautoicous; perichaetial leaves differentiated, convolute-sheathing.  Spores 12--20(--30) \um.

 

Acidic rock faces, occasionally thin soil; moderate elevations; B.C.; South America (Argentina, Ecuador); nw Europe; Asia (e China); Australia; Atlantic Islands (Kerguelen I.); Pacific Islands (including New Zealand).

 

3. Andreaea sinuosa B. M. Murray, Bryologist 89: 189, figs. 1--17.  1987 

 

Plants reddish brown to black. Leaves weakly spreading, straight or sometimes secund, narrowly lanceolate, widest proximally, apex symmetric; costa absent; leaf margins entire; basal laminal cells rectangular, marginal cells not different in shape, walls thickened, pitted, sinuose; medial laminal cells rounded-quadrate to ovate, 1-stratose entirely or sometimes 2-stratose in patches distally, lumens rounded; laminal papillae low, large, brown to whitish, mainly medial. Sexual condition apparently dioicous, possibly cladautoicous; perichaetial leaves differentiated, convolute-sheathing.  Spores 12--20 \um.

 

Acidic rock in snow beds; low to moderate elevations; B.C., Alaska; nw Europe.

 

Andreaea sinuosa is uncommon, and is distinguished from the similar A. rupestris by the symmetric and often rounded leaf apex and sinuose basal laminal cells.

 

4. Andreaea rupestris Hedwig, Spec. Musc. Frond. 47, plate 7, fig. 2g--o.  1801

 

Andreaea alpestris (Thedenius) Schimper; A. papillosa Lindberg; A. parvifolia J. K. A. Müller; A. rupestris var. acuminata (Schimper) Sharp;  A. rupestris var. alpestris (Thedenius) Sharp; A. rupestris var. papillosa (Lindberg) Podpěra; A. rupestris var. sparsifolia (Zetterstedt) Sharp; A. sparsifolia var. sublaevis Kindberg

 

Plants reddish black, black or greenish brown. Leaves curved or secund to straight, wide-spreading to squarrose, short-lanceolate, widest proximally, apex oblique or symmetric; costa absent; leaf margins entire; basal laminal cells short- to long-rectangular, marginal cells rounded-quadrate to short-rectangular, walls sometimes thickened, usually pitted, straight; medial laminal cells quadrate to short-rectangular, 1-stratose or occasionally 2-stratose in patches, lumens rounded, rectangular or irregularly stellate; laminal papillae usually present, commonly large, whitish. Sexual condition cladautoicous or autoicous; perichaetial leaves differentiated, convolute-sheathing.  Spores 20--32(--50) \um.

 

Neutral to acidic boulders, cliffs and walls, generally wet sites; low to moderate elevations; Greenland; Nunavut, Que.; Alaska; nw Europe.

 

Andreaea rupestris is a species of many morphological variants, more commonly identified by simple elimination. It is similar to A. obovata, but is distinguished by the leaves generally curving or secund, short-lanceolate, widest proximally, apices oblique or symmetric. The degree of expression of laminal papillae is variable on the same plant. Plants identified as A. alpestris, said to differ by straight leaves with low papillae, is probably best considered a high elevation form.

 

5. Andreaea obovata Thedenius, Bot. Not. 5: 78, figs. 27--36.  1849

 

Plants red-brown to purple-black. Leaves straight to secund, panduriform, widest in distal half, apex usually symmetric; costa absent; leaf margins entire; basal laminal cells rectangular to long-rectangular, marginal cells rectangular, walls thick, pitted-nodose; medial laminal cells quadrate, 1-stratose, lumens irregularly stellate to rhombic; laminal papillae low or absent. Sexual condition autoicous; perichaetial leaves differentiated, convolute-sheathing.  Spores 20--35 \um.

 

Rock or soil in streams; low to moderate elevations; Greenland; B.C., Nfld. & Labr. (Labr.), Nunavut, Yukon; Alaska; n Eurasia; c Africa.

 

Like Andreaea alpina, A. obovata has spores in two size classes, the smaller apparently abortive. Andreaea obovata is very rare in the area of the flora, and it can be distinguished from A. rupestris by the panduriform leaves.

 

6. Andreaea nivalis Hooker, Trans. Linn. Soc. London 10: 395. 1811  F

 

Andreaea baileyi (Holzinger) Holzinger; A. macounii Kindberg; A. nivalis var. baileyi Holzinger

 

Plants reddish brown to brownish green. Leaves secund to falcate-secund, mostly subulate, widest in proximal half of leaf, apex symmetric; costa present, percurrent and filling the leaf apices, strong, terete, reaching the leaf insertion; leaf margins crenulate to strongly denticulate; basal and medial laminal cells quadrate to short-rectangular, 1-stratose, walls thin and straight, lumens smoothly rounded, quadrate to short-rectangular on the margins; laminal papillae strong, usually distant. Sexual condition dioicous; perichaetial leaves not differentiated beyond being larger than the cauline.  Spores (18--)20--30(--40) \um.

 

Wet rocks in streams, snow flushes, seeping outcrops; moderate to high elevations; s and sw Greenland; B.C., Nfld and Labrador (Nfld.), Yukon; Alaska, Calif., Oreg., Wash.; Europe; Asia (Japan, Russia).

 

The salient distinguishing traits of Andreaea nivalis are the crenulate to strongly denticulate leaf margins, strong papillae occurring mostly on the abaxial surface of the lamina, and the perichaetial leaves larger, more sharply cordate at the basal angles, but otherwise not much different from the cauline leaves.

 

7. Andreaea blyttii Schimper in P. Bruch, W. P. Schimper & W. Gümbel, Bryol. Eur. 6: 155, plate 13 (635). 1855  F

 

Plants brown to black. Leaves erect, curved or secund, becoming subulate from an ovate or narrow-rectangular base, widest in proximal half of leaf, apex symmetric; costa present, percurrent and filling the leaf apices, strong, terete, usually reaching the leaf insertion; leaf margins entire or crenulate; basal laminal cells rectangular, walls thin and straight; medial laminal cells usually quadrate, 1-stratose or rarely 2-stratose juxtacostally, lumens rounded-quadrate; laminal papillae rare. Sexual condition dioicous; perichaetial leaves differentiated, convolute-sheathing.  Spores (10--)13--15(--20) \um.

 

Rock, alluvium, edges of snow melt areas; low to high elevations; Greenland; B.C., Nfld. & Labr., N.W.T., Nunavut, Que.. Yukon; Alaska, Calif., Oreg., Wash.; n Europe; n Asia; Atlantic Islands (Iceland).

 

Andreaea blyttii is distinguished by leaves with long, narrow subula, nearly filled with the strong costa, and rectangular basal cells, and small spores. Collections commonly lack sporophytes. Distal laminal cells may be minutely mammillose abaxially, and the leaf margins are thus crenulate.

 

8. Andreaea heinemannii Hampe & J. K. A. Müller, Bot. Zeit. 4: 324, plate 2. 1846

 

Andreaea blyttii var. angustata (Limpricht) Schultze-Motel; A. blyttii var. obtusifolia (Berggren) Sharp; A. crassinervia var. obtusifolia Berggren

 

Plants brown to black. Leaves erect to spreading, occasionally secund, broadly subulate from an ovate base, widest in proximal half of leaf, apex symmetric; costa present, percurrent and filling the leaf apices, weak, flattened distally, often not reaching the leaf insertion; leaf margins entire or occasionally weakly crenulate; basal laminal cells quadrate to occasionally short-rectangular, a few marginal cells rectangular, walls usually sinuose; medial laminal cells quadrate, 1-stratose or sometimes 2-stratose in patches, lumens rounded-quadrate; laminal papillae rare, low. Sexual condition apparently autoicous; perichaetial leaves differentiated, convolute-sheathing.  Spores 20--30(--40) \um.

 

Acidic rocks; low to moderate elevations; Greenland; BC, Yukon; Alaska, Calif., Colo., Oreg.; s Europe; se Asia; Atlantic Islands (Canary Is., Madiera I., Kerguelen I.).

 

Andreaea heinemannii is a relatively small species in the genus, having irregularly divergent leaf apices of a flat subula. The weak costa commonly does not reach the leaf insertion, and this species may be mistaken for A. rupestris.

 

9. Andreaea megistospora B. M. Murray, Bryologist 90: 18, figs. 9--23, 25--28.  1987

 

Andreaea megistospora var. epapillosa (B. M. Murray) Crum & L.E. Anderson; A. megistospora subsp. papillosa B. M. Murray

 

Plants brown to black. Leaves erect-spreading or secund, subulate-lanceolate from an ovate base, widest in proximal half of leaf, apex symmetric; costa present, percurrent and sometimes filling the leaf apices, strong but narrow, bulging abaxially, usually reaching the leaf insertion; leaf margins entire or occasionally weakly crenulate; basal laminal cells quadrate to short-rectangular, marginal cells mostly rounded-quadrate, walls weakly pitted-sinuose; medial laminal cells rounded-quadrate, 1-stratose or 2-stratose in patches or completely, lumens rounded; laminal papillae smooth to papillose. Sexual condition cladautoicous or gonioatoicous; perichaetial leaves differentiated, convolute-sheathing.  Spores (40--)50--90(--110) \um.

 

Acidic rock; low to moderate elevations; B.C.; Alaska, Wash.; nw Europe.

 

Andreaea megistospora is distinguished from the similar A. rothii by its large spores, and may simply prove to be a large form. The variety papillosa is a minor variant commonly occurring in mixture with the typical variety, and doubtfully warrants a name.

 

10. Andreaea rothii F. Weber & D. Mohr, Bot. Taschenb. 386, plate 11, fig. 7--8. 1807

 

Andreaea crassinervia Bruch; A. huntii Limpricht; A. rothii var. crassinervia (Bruch) Mönkemeyer

 

Plants brown to black. Leaves erect-spreading, occasionally secund, broadly subulate from an ovate base, widest in proximal half of leaf, apex symmetric; costa present, percurrent and usually filling the leaf apices, moderately differentiated to strong, terete, reaching the leaf insertion; leaf margins entire or occasionally weakly crenulate; basal laminal cells quadrate to occasionally short-rectangular, marginal cells mostly quadrate or rounded, walls usually sinuose; medial laminal cells quadrate, 1-stratose to nearly completely 2-stratose, lumens rounded-quadrate; laminal papillae rare, low. Sexual condition clad- or gonioautoicous; perichaetial leaves differentiated, convolute-sheathing.  Spores 35--60(--70) \um.

 

Siliceous rock, cliffs, boulders; low to high elevations; Greenland; B.C., Ont., N.B., Nfld. & Labr., N.S.; Ala., Alaska, Calif., Conn., Fld., Ga., Maine, Md., Mich., N.Y., N.C., Ohio, Oreg., Pa., Tenn., Vt., Va., Wash., W.Va.; n and c Europe.

 

H. A. Crum and L. E. Anderson (1981) chose not to recognize as Andreaea crassinervia Bruch those eastern North American specimens of A. rothii with excurrent or poorly defined costae (not bordered by laminal cells in the subula). B. M. Murray (1987) excluded this name from the Arctic, while M. F. V. Corley (1981) submerged it with A. rothii.  The essentially European Andreaea rothii var. falcata (Schimper) Lindberg (synonym A. rothii var. papillosa J. K. A. Müller) is only poorly distinguished from the typical variety and does not warrant recognition here, at least on the basis of the single specimen reported for the area of the flora. The previously used traits of spore size and costa filling the acumen or not are here found to intergrade between such taxa.

 

11. Andreaea schofieldiana B. M. Murray, Bryologist 90: 15, fig. 1--7. 1987  E

 

Plants greenish brown to brown-black. Leaves spreading, straight to secund, broadly subulate from an ovate base or lanceolate, widest in proximal half of leaf, apex symmetric; costa present, percurrent and usually filling the leaf apices, moderately differentiated to more often strong, terete, commonly reaching the leaf insertion; leaf margins entire or occasionally weakly crenulate; basal laminal cells rectangular, marginal cells grading to short-rectangular, walls usually sinuose, sometimes slightly pitted; medial laminal cells rounded-quadrate, 2-stratose entirely or in large patches except near costa where commonly 1-stratose, rarely 1-stratose throughout, lumens rounded; laminal papillae rare, low. Sexual condition cladautoicous; perichaetial leaves differentiated, convolute-sheathing.  Spores 20--30 \um.

 

Dry rock outcrops; moderate elevations; sw B.C.; n California.

 

The small spores immediately separate Andreaea schofieldiana from the similar A. rothii.