BFNA Title: Aulacomniaceae
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
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.
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
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.
Brotherus, V. F.
1924. Aulacomniacae. In: Engler, A.,
ed. Die natürlichen Pflanzenfamilien, ed. 2.
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.
G. 1980. Fossil mosses of
J. 2007. The phylogenetic distribution of
pleurocarpous mosses: evidence from cpDNA
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.
Weber, W. A.
and R. C. Wittmann.
2007. Bryophytes of