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BFNA Title: Andreaeaceae |
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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. |
