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BFNA Title: Aulacomniaceae |
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X. Aulacomniaceae
Schimper Norton G. Miller Plants small to large, leaves radially arranged or somewhat complanate; subgametangial branching common. Stems
yellow-green, brown, or reddish brown, irregularly pentagonal in cross
section, hyalodermis present and interrupted or absent, cortex of 2--5 layers
of small thick-walled cells, medulla thick, cells large, usually thick
walled, central strand prominent; rhizoids smooth or roughened.
Leaves lanceolate, linear-lanceolate, oblong-lanceolate,
oblong-elliptical, or oblong-ovate; costa present, shiny abaxially; costa
with two stereid bands, 1 row of guide cells or sometimes 2 at leaf base,
abaxial and adaxial costal epidermis well developed or not; laminal cells collenchymatous-pitted or prominent corner thickenings
absent; axillary hairs usually 5 cells long, terminal cell elongate, hyaline,
lowest one or two cells short, with brown walls. Specialized asexual reproduction common, by propagula terminal
on leafless stem tips. Sexual condition dioicous or autoicous. Perichaetia
terminal, with or without budlike proximal serial perigonia; or perigonia in
terminal, discoidal splash platforms or, budlike
and in a series along stems of male plants; paraphyses present or absent,
isomorphic or weakly differentiated between female and male plants. Seta
usually solitary, upright or slightly inclined, twisted, smooth. Capsule
inclined or horizontal, furrowed, exothecial cells differentially thickened
in longitudinal bands, stomata restricted to inconspicuous apophysis, phaneropore; operculum rostrate, rostrum short or long,
usually blunt; annulus of 2 cell layers, revoluble; peristome diplolepidous,
alternate; exostome papillose;
endostome well developed, cilia 2--4, often nodose or nodulose. Calyptra cucullate.
Spores 8--15 \um, smooth or papillose. Genera
2, species 5 (all in the flora); principally
Northern Hemisphere including the High Arctic region, and sporadically in the
Southern Hemisphere. Leptotheca Schwägrichen had
long been included in the family (V. F. Brotherus
1924), but peristome and gametophytic structure (S. P. Churchill and W. R.
Buck 1982), as well as molecular evidence (N. E. Bell et al. 2007), indicate affinities elsewhere (with Rhizogoniaceae,
S. P. Churchill and W. R. Buck 1982; or Orthodontiaceae,
B. Goffinet et al. 2009). The molecular results of N. E. Bell et al.
(2007) joined Aulacomnium,
Hymenodontopsis
Herzog, Mesochaete Lindberg in a newly recognized Aulacomniales.
These three genera were placed in the Aulacomniaceae
(Rhizogoniales) by B. Goffinet
et al. (2007). In molecular
phylogenies (N. E. Bell and A. E. Newton 2004; T. J. O’Brien 2007), Mesochaete, a
genus of two species in eastern Australia (I. G. Stone 1983), one also on
Lord Howe Island, is sister to Aulacomnium sensu lato (N. E.
Bell and A. E. Newton 2004; T. J. O’Brien 2007). Aulacomniaceae, as treated here, consist of two
genera: Arrhenopterum,
a monotypic genus usually included in Aulacomnium, and Aulacomnium. T. J.
O’Brien (2007) presented molecular evidence that Arrhenopterum (as Aulacomnium heterostichum)
and other species in Aulacomnium he studied were sister groups. Additional molecular and morphological
studies are likely to alter the family circumscription. selected references Blomquist, H. L. and L. L. Robertson. 1941. The development of the peristome in Aulacomnium heterostichum. Bull. Torrey Bot. Cl. 68: 569--584. Holmen, K.
1957. Three West Arctic moss
species in 1.
Leaves lanceolate, oblong-lanceolate, linear-lanceolate, or
oblong-elliptical, spirally arranged, symmetric, width of laminae on either
side of costa equal, leaves tapered toward base, scarcely tapered, or
slightly rounded, insertion broad, apex often irregularly serrulate; papillae
prominent on plane to somewhat domed abaxial and adaxial leaf cell surfaces,
papillae usually one per cell; dioicous………… 1. Aulacomnium, p. XXX 1.
Leaves oblong-ovate, ± complanate, often asymmetric, width of laminae
on either side of costa usually unequal, leaves abruptly rounded to a narrow
insertion, strongly serrate in distal half; papillae weakly developed or
absent, abaxial and adaxial leaf cell walls
bulging, smooth or low papillose-mammillose; autoicous……………..…………
2. Arrhenopterum, p. XXX 1. AULACOMNIUM Schwägrichen,
conserved name. Sp. Musc. Frond.
Suppl. 3 1(1): 215. 1827 “Aulacomnion”
* [Greek, aulax, furrow, alluding to sulcate capsules, and mnion, moss] Orthopixis P. Beauvois;
Sphaerocephalus
Lindberg Stems hyalodermis present or absent, cells of central strand
numerous, thin-walled; rhizoids smooth. Leaves radially
arranged, lanceolate,
oblong-lanceolate, linear-lanceolate, or oblong-elliptical, spirally
arranged, symmetric, width of laminae on either side of costa equal, leaves
tapered toward base, scarcely tapered, or slightly rounded, insertion broad,decurent, margins entire but apex often irregularly
serrulate; papillae prominent on abaxial and adaxial leaf cell surfaces,
usually one per cell Specialized
asexual reproduction by leaf homologs borne
terminally on leafless stem apices or absent. Sexual condition dioicous. Perichaetia
terminal, outer perichaetial leaves sheathing with apices flaring or not,
inner ones narrowly lanceolate, long-acuminate, plicate, apices erect,
sometimes serrulate, paraphyses absent or 9--12 cells, cell length variable,
slightly wider toward tip. Perigonia as terminal discoidal splash platforms, or budlike and arranged in a
series down the stem, outermost leaves narrow or not, apices flaring, inner
ones with a narrowed limb and broad concave base, or conspicuously broader
toward concave base. Seta usually
solitary, red toward base, yellow
distally. Spores 10--15 \um, smooth or slightly papillose. 1. Plants small; fusiform; several-celled
propagules often present in a spherical mass at the apex of leafless stem
tips; leaves lanceolate, apex not prolonged, serrulate, slightly rounded immediately above insertion; rhizoids
conspicuous between leaves near base of
plants………………………1. Aulacomnium androgynum 1. Plants larger; bullet-shaped multicellular
propagules usually present in an elongate series at apex of leafless stem
tips, or propagules absent; leaves linear-lanceolate, oblong-lanceolate, or
oblong-elliptical; leaf apex various, acute, long-acuminate, rounded and
cucullate, insertion broad, not rounded distally; rhizoids conspicuous on
stems between leaves throughout plants or
completely hidden by imbricate leaves.
2. Leaves oblong-elliptical, apices usually
broadly rounded, cucullate; leaf cells
collenchymatous…..……………………………………2. Aulacomnium turgidum 2. Leaves oblong-lanceolate or
linear-lanceolate, apices acute, long-attenuate, or narrowed to a rounded
tip, sometimes irregularly serrulate; leaf cells collenchymatous
or not. 3. Leaves oblong-lanceolate, widest at middle
and tapering both distally and proximally, concave, strongly imbricate when
wet and dry and thus completely hiding rhizoids, leaf apex acute or
prolonged, margins revolute (usually 1.5--2\x) nearly to apex, leaf cells
strongly collenchymatous………… 3. Aulacomnium acuminatum 3. Leaves linear-lanceolate, widest at base
and tapering distally, weakly and irregularly concave, leaves distant and
rhizoids conspicuous when wet or dry, or if imbricate then rhizoids visible
among leaves at base of plants, leaf apex various, usually attenuate to
long-attenuate but sometimes narrowed and rounded, then attenuate and rounded
leaves intermixed, margins revolute (usually 0.75--1.5\x) only in distal 2/3
of leaves, leaf cells collenchymatous or
not….………4. Aulacomnium palustre 1.
Aulacomnium androgynum (Hedwig)
Schwägrichen, Sp. Musc. Frond. Suppl. 3 1:(1):
215. 1827 E F
Bryum androgynum Hedwig, Sp. Musc. Frond.,
178. 1801 Plants small, 1--2 cm, erect, in green or brownish-green tufts or
cushions, rhizoids smooth, basal on plant, absent or greatly reduced
distally; plant apices often without leaves and bearing a tight globose cluster of small fusiform propagules. Stems
yellow or brownish, irregularly pentagonal, hyalodermis absent, cortex 2--4
layers of small, thick-walled cells, medullary cells larger, thick or thin
walled, central strand of small, thin-walled cells. Leaves oblong-lanceolate,
wide to narrow, distant near base of plants, more congested toward apex,
somewhat concave to keeled, acute, irregularly serrulate toward apex, teeth
of 1--3 cells, weakly serrulate proximally; insertion broad, long to
short-decurrent, cells at and near insertion larger, smooth, hyaline,
1-stratose; margin revolute to mid leaf; costa flexuose, percurrent; distal
laminal cells round or ± oblong, walls thick; solitary, blunt, long papilla
on abaxial and adaxial cell walls. Perichaetia with outer perichaetial
leaves sheathing, apices flaring, the inner slender, long-acuminate,
paraphyses 9--12 cells long, cells lengths variable, slightly wider toward
tip.
Perigonia terminal, budlike, perigonia from prior years proximally
in a series, outermost perigonial leaves flaring, inner ones with a narrowed
limb and broad concave base; basal cells large, elongate to rhomboidal, smooth;
paraphyses 7--10 cells long, cells mostly longer than wide, those near tips
slightly wider. Seta 1--1.5 cm. Capsule suberect or horizontal,
2.5--3 mm; operculum
conic, rostrum short, blunt; endostome
cilia nodose. Spores 10--20 \um, smooth.
Organic soil,
rotting logs, mineral soil over rock; low to moderate elevation; Alaska, B.C., N.B., Nfld. and Labrador
(Nfld.), N.S., Ont., P.E.I., Que.; Calif., Colo.,
Idaho, Ill., Maine, Mich., Mont., N.H., N.Y., Ore., S.Dak., Wash., W.Va.,
Wyo. The propagules
of Aulacomnium androgynum,
borne in a spherical cluster at the ends of naked branches, are
distinctive. This is the only Aulacomnium
with small lanceolate leaves that are serrulate mainly beyond mid-leaf and
shortly rounded at the insertion. 2.
Aulacomnium turgidum (Wahlenberg) Schwägrichen, Sp. Musc. Frond. Suppl. 3 1(1)
[Aulacomnium]: 7.
1827 F Mnium
turgidum Wahlenberg,
Fl. Lapp., 351, plate 23. 1812 Plants large, 3--8 cm, julaceous, green, yellow-green, or
fuscous-green, in dense upright-growing cushions or in prostrate mats over
the substratum, branching irregular and often associated with gametangial formation.
Stems yellow, pentagonal in
cross section, cortex of 2--5 irregular layers of small, thick-walled cells,
medullary cells large, walls thick to thin, in radiating groups, central
strand present; rhizoids smooth. Leaves erect, overlapping, concave,
not twisted either wet or dry, mostly covering the rhizoids except in older
parts of plants, oblong, apices rounded-cucullate, not narrowed to the
insertion, decurrent; basal cells oblong, usually in 2 layers, those at leaf
corners sometimes in an excavate group, walls thick, often brown; margins
entire, revolute 1--2\x to just before apex; costa prominent, flexuose toward
apex, laminar cells round to oblong, collenchymatous,
lumens stellate, papillae low, on abaxial and adaxial surfaces. Perichaetia
with outer perichaetial leaves oblong, apices flaring, inner ones
lanceolate, plicate, apices erect, serrulate, cells oblong to rhomboidal,
walls evenly thickened; paraphyses absent.
Perigonia discoidal splash platforms, outer perigonial leaves like
those below, except apices flaring, the inner widening at base, concave;
basal cells irregular, smooth, evenly thickened; paraphyses numerous,
extending above antheridia, 6--9 cells long, middle distal cells wider than
those more distal or proximal. Seta 2--2.5 mm. Capsule
3--5 mm, borne horizontally; operculum conic, rostrum short,
blunt;
endostome nodose. Spores 8--12.5 \um, smooth.
Dry or moist
Arctic and alpine tundra meadows, heaths, and fellfields,
sometimes on mineral soil or rock, near tree line under spruce, southward on
cirque headwalls; often in pure mats or cushions; low to moderate elevations
in the Arctic, high southward;Greenland; Alta.,
B.C., Nfld. and Labrador, N.W.T., Nunavut, Man.,
Ont., Que., Sask., Yukon; Alaska, N.H., N.Y., Maine, sometimes in deep shaded
canyons; n. Europe, Asia, Africa (high
mountains). The broad cucullate leaf apices and
closely imbricate leaves (hiding the rhizoids) are distinctive for Aulacomnium turgidum. Fossils of this moss have been found well
south of its present range in eastern 3. Aulacomnium
acuminatum (Lindberg & H. Arnell)
Kindberg, Enum. Bryin. Exot.,
suppl. 2. 105. 1891 Sphaerocephalus
acuminatus
Lindberg & H. Arnell, Kongl.
Svenska Vetenskapsakad. Handl.,
n.s. 23(10):
27. 1890 Plants large, to 10 cm, julaceous, sparsely branched, yellow
brown or brown, growing laterally over substratum in mats or upright among
other mosses; propagules absent. Stems yellow or brownish, pentagonal
in cross section, cortical cells in 4 or 5 layers, cells small, thick walled,
medullary cells larger, thick or thin walled, in radial series, central
strand well developed; rhizoids in groups along stem, hidden by leaves.
Leaves erect, tightly overlapping, linear, not twisted when dry,
but sometimes curved, oblong-lanceolate, widest at middle, concave, apex
acuminate to acute, usually long-attenuate in youngest leaves; leaves
narrowed to insertion but insertion broad, decurrent, sometimes slightly auriculate; basal cells in a narrow band, 2-stratose,
cells large, elongate to subquadrate, smooth, walls
thick, brown; margins entire, prominently revolute 1.5--2\x nearly to leaf
apex; costa strong, flexuose in distal leaf, ending before apex, buttressed
at base; laminar cells isodiametric to elongate, corner thickenings massively
collenchymatous, cell lumens thus stellate,
papillose on abaxial and adaxial cell walls.
Perichaetia with subfloral
innovations, outer perichaetial leaves large, flaring, the inner narrow,
long-attenuate, plicate, weakly toothed at apex; paraphyses absent.
Perigonia discoidal, outer perigonial
leaves large, flaring, the inner narrow, expanded concave at base, apices
attenuate; basal cells large, walls evenly thickened; innermost leaves
deltoid, as long as paraphyses;
paraphyses 5--8 cells long, clavate, walls yellow, middle distal cells short
and wide, proximal cells long and narrow. Seta solitary, 1.5--2 cm. Capsule
horizontal or slightly curved, 3 mm; operculum short-conic, rostrum blunt;
endostome cilia in groups of 2--4, weakly nodulose. Spores
10--13 \um, slightly papillose. Dry to moist Arctic tundra
meadows, hummocks, polygon edges, organic or mineral soil, sometimes at
calcareous sites, sometimes in deep canyons southward; low to moderate
elevations; Greenland; B.C., Que.
Man., N.W.T., Nunavut, Yukon; Alaska; Asia (Russia
from w Siberia through the Russian Far East). Aulacomnium acuminatum has an interrupted circumarctic distribution; in North America from the
northern edge of the tree line (and there under white and/or black spruce),
throughout the Arctic islands, with occurrences southward in adjacent alpine
tundra, and as a disjunct north of Lake Superior in 4.
Aulacomnium palustre (Hedwig)
Schwägrichen, Sp. Musc. Frond. Suppl. 3 1(1): 216. 1827
F
Mnium palustre Hedwig, Sp. Musc. Frond.,
188. 1801 Plants large, to 10 cm, in loose, irregular mats or dense pure
cushions, yellow, yellow-green, or brown to blackish when growing suboptimally, often conspicuously tomentose proximally by
abundant matted rhizoids; rhizoids smooth; plant apices often without leaves
and bearing a terminal cluster of bullet- or spearpoint-shaped
propagules, typically flat at attachment.
Stems pentagonal in cross
section, hyalodermis interrupted, cortex of 4--6 irregular layers of small,
thick-walled cells, medullary cells larger, the outer thicker walled than
inner, corner thickenings present, central strand conspicuous, sometimes
degraded. Leaves broadly to narrowly lanceolate, ± concave, undulate,
weakly plicate, often concave, erect-spreading when wet, twisted when dry or
remaining straight, usually widely spaced and not covering the rhizoids or
imbricate and rhizoids mostly hidden, apices acute or acuminate or narrowed,
rounded, and somewhat cucullate, insertion broad and decurrent, margins
revolute usually 1\x in lower 2/3 and sometimes more but ending well before
leaf apex, irregularly serrulate near leaf tip; costa sinuose, ending just
before leaf apex, epidermis of large hyaline cells ab-
and adaxially; laminar cells with a large central papilla on most abaxial and
adaxial surfaces, cells collenchymatous or corner
thickenings poorly developed, lumens irregularly shaped when collenchymatous; basal cells large, smooth, walls brown,
often 2 cells thick at insertion. Perichaetia terminal, not obvious
splash cups; outer perichaetia leaves similar to those below, inner leaves
narrowly lanceolate; paraphyses absent.
Perigonia terminal, discoidal splash platforms; outer perigonial leaves
narrow, limbs flaring, the inner conspicuously broader toward concave base;
basal cells larger, smooth; paraphyses 8--12 cells long, upper 4--5 cells
short, wider than the slender longer more proximal cells. Seta
3--5 cm. Capsule inclined to
horizontal; operculum short-rostrate, rostrum blunt; endostome segments with
wide gaps, cilia 2--4, nodose. Spores 10--15 \um, smooth. Wet habitats such as fens, marshes,
swamps, often on peat or other organic soil, but sometimes on moist to wet
mineral soil and rock; low to high elevations; Greenland; Alta., B.C., Man., N.B., Nfld. and Labrador, N.S., N.W.T., Nunavut, Ont.,
P.E.I., Que., Sask., Yukon; Ala., Alaska, Ariz., Ark., Calif., Colo., Conn.,
Del., Fla., Ga., Idaho, Ill., Ind., Iowa, Kans., Ky., La., Maine, Md., Mass.,
Mich., Minn., Miss., Mo., Mont., Nebr., Nev., N.H., N.J., N.Mex., N.Y., N.C.,
N.Dak., Ohio, Okla., Oreg., Pa., R.I., S.C., S.Dak., Tenn., Tex., Utah, Vt.,
Va., Wash., W.Va., Wis., Wyo. Mexico;
West Indies (Dominican Republic); n,w South
America; Europe; Asia; Pacific Islands (New Zealand, possibly introduced);
Australia. Aulacomnium
palustre is
essentially a Northern Hemisphere moss, being widespread in North American,
but not necessarily common throughout its range. Numerous varietal and forma
names have been proposed and sometimes accepted to accommodate the great
range of morphological variation present in this moss (e.g., W. Kabiersch 1937; G. Sayre 1935). None of these infraspecific
concepts is adopted here in the absence of thorough studies employing modern
techniques. A distinctive alpine
expression (Aulacomnium
palustre
var. imbricatum
Schwägrichen) has been recognized in 2. ARRHENOPTERUM Hedwig, Sp. Musc. Frond.,
198, plate 46, figs.1--9. 1801 *
[Greek, arrhen,
strong, and pterum,
feather or wing, possibly alluding to the feather-like evenness of leaf
arrangement] Stems hyalodermis absent, cortex two layers of small thick-walled cells, central strand present. Leaves
somewhat complanately arranged, oblong-ovate,
concave, some asymmetrical (laminae on either side of costa at leaf base
unequally wide), abruptly narrowed and rounded to insertion, weakly
decurrent, margins strongly toothed in distal half of leaf but otherwise
undifferentiated, teeth of 1--3 cells or sometimes more, revolute on one side
(or both) in proximal half; costa prominent, flexuose, sometimes with short
lateral spurs, ending before apex, guide cells often in 2 rows near leaf
base, 1 row toward apex; laminal cells round or elongate, walls thick, corner
thickenings absent or small, abaxial and adaxial walls bulging, smooth or
with a single low mammillose papilla. Specialized asexual reproduction by
leaf-like propagules smaller than more proximal leaves, borne on leafless
stem tips. Sexual condition autoicous. Perigonia
budlike, numerous, solitary, borne laterally on stem in a series below
perichaetia, outer and inner
perigonial leaves dimorphic; paraphyses slightly widened distally. Seta short, solitary. 1.
Arrhenopterum heterostichum Hedwig, Sp. Musc. Frond.,
198, plate 46, figs.1--9. 1801
F Aulacomnium heterostichum (Hedwig)
Bruch & Schimper in P. Bruch, W. P. Schimper & W. Gümbel
Plants small, densely branched, mostly from
subfloral branches, weakly complanate, apices nitid
when dry, plants scarcely altered when wet, in mats; rhizoids conspicuous
along stem. Stems reddish brown or yellow.
Perichaetia terminal, outer
perichaetial leaves symmetrical, flaring, narrowed to insertion, inner ones
narrow, attenuate to long attenuate; paraphyses 8--15 cells long, slightly
widened toward apex. Perigonia sessile; outer perigonial leaves short,
apices slightly flaring, narrowed to insertion, the inner scarcely longer
than antheridia, acute, concave
basally, margins entire; basal cells elongate, larger than upper
cells, smooth. Seta reddish throughout, 1--1.5 cm. Capsule 3.5--5
mm. Spores 10--15 \um, punctate. Soil, mineral or humified, slopes of ravines and similar habitats,
sometimes on bark at tree bases; mostly low elevations; N.S., Ont.; Ala., Ark., Conn., Del.,
Fla., Ga., Ill., Ind., Iowa, Kans., Ky., La., Maine, Mass., Md., Mich.,
Minn., Miss., Mo., N.C., N.J., N.Y., Ohio, Okla., Pa., R.I., S.C., Tenn.,
Tex, Va., Vt., W.Va., Wis.; Asia (China, Japan, Korea, Russian Far East). The range of Arrhenopterum heterostichum in
North American corresponds largely to the region of temperate deciduous or
mixed deciduous-conifer forest, ranging westward into forested portions of
the tall grass prairie. By separating Arrhenopterum and Aulacomnium, the latter is made morphologically more
homogeneous. Leaf stance (±
complanate) and asymmetry (laminae on either side of the costa unequally
wide), prominent marginal teeth, leaf laminal cells with low mammillose-papillae
or cells bulging and smooth, and autoicous sexuality set this moss apart from
species of Aulacomnium. The Southern Hemisphere Australian genus Mesochaete is
sister to Arrhenopterum
in molecular phylogenies. SELECTED
REFERENCES Brotherus, V. F.
1924. Aulacomniacae. In: Engler, A.,
ed. Die natürlichen Pflanzenfamilien, ed. 2.
Vol. 10. Churchill, S.
P. and W. R. Buck. 1982. A taxonomic investigation of Leptotheca (Rhizogoniace). Brittonia 34: 1--11. Goffinet, B. and W. R. Buck. 2004.
Systematics of the Bryophyta (mosses): From
molecules to a revised classification.
Monogr. Syst. Bot. Missouri Bot. Gard. 98: 205--239. Goffinet, B., W. R. Buck, and A. J. Shaw. 2009.
Morphology, anatomy, and classification of the Bryophyta. In: Goffinet, B.
and A. J. Shaw. Bryophyte Biology, edition 2.
Kabiersch, W.
1937. Studien
über die ostasiatischen Arten einiger Laubmoosfamilien II (Rhizogoniaceae,
Bartramiaceae, Aulacomniaceae,
Meeseaceae). Hedwigia 76:
71--136. Miller, N.
G. 1980. Fossil mosses of O’Brien, T.
J. 2007. The phylogenetic distribution of
pleurocarpous mosses: evidence from cpDNA
sequences. In: Sayre, G. 1935.
Aulacomniaceae. In: Grout, A. J., Moss Flora of North
America North of Mexico 2: 149--152,
plate 66. Shaw, A. J.
and L. E. Anderson. 1986. Morphology and homology of the peristome
teeth in Hymenodon
and Hymenodontopsis
(Rhizogoniaceae: Musci). Syst. Bot. 11: 466--454. Stone, Weber, W. A.
and R. C. Wittmann.
2007. Bryophytes of |
