BFNA Title: Hedwigiaceae
38. HEDWIGIACEAE Schimper
P. M. Eckel
Plants acrocarpous, medium to robust, coarse, dull, grayed-opaque when dry, yellow- or grayish green, reddish brown, olive-brown to blackish brown, distal stems and branches may be hoary due to prominent overlapping rigid broad-based awns, or with long, flexuose shining hair-points, in loose or compactly intricate wefts or mats. Stems: primary stems elongate, creeping, with reduced, generally widely spaced leaves; secondary stems variously creeping and erect-ascending or loosely spreading, closely foliate, sparingly and irregularly to subpinnately branched, sometimes bearing thin, radiculose stoloniform or flagelliform-tapered branch or stem tips; cross section with sclerodermis of generally 2--3 rows of small thick-walled cells, cortical cells enlarged, thin to firm-walled, homogeneous, central strand absent; paraphyllia absent; pseudoparaphyllia present, papillose and foliose. Leaves on stems scarcely differentiated from branch leaves, erect to variously secund when dry, spreading when wet, ovate, broadly oblong-ovate, ovate-lanceolate, strongly concave, smooth to shallowly longitudinally undulate-plicate when dry, concolorous throughout and epilose or with whitened, echlorophyllose differentiated tips that are erect to spreading when dry, spreading when wet; leaf base narrowing into the insertion, broadly decurrent to abruptly concave-auriculate; margins erect to incurved, or revolute on one or both sides, differentiated border absent, mostly entire , variously denticulate to irregularly spinose-serrate apically; leaf apex gradually to abruptly acute, or short to long-acuminate, flat to canaliculate or subtubulose, epilose and concolorous, hyaline-whitened apiculate, or hyaline and strongly piliferous with pellucid tips; ecostate; laminal cells 1-stratose throughout, mostly thick-walled and coarsely porose-sinuose, variously papillose on both surfaces; laminal cells near insertion short-rectangular often smooth; mid-basal cells concolorous to deeper-colored; proximal, median, and distal laminal cells subquadrate, short-rectangular or oblong-rhombic 1--2(--3):1, on both surfaces 1- or pluripapillose over lumina and/or walls, papillae simple (unbranched), low and rounded, coarsely tuberculate to regularly or irregularly multifid, columnar, or multifid-coroniform; apical cells chlorophyllose, concolorous, similar to median cells, or cells gradually or abruptly differentiated into an echlorophyllose, broad- or narrow-based, occasionally decurrent, short- or long-triangular apex of translucent, flat-white to hyaline cells with thick and even to strongly porose walls and epapillose and smooth to papillose with papillae simple, low, arranged in lines over the lumina to coarsely and irregularly spinose-papillose, terminating into a variously less papillose pellucid sharp or truncate point or long, smooth, cellularly 1-seriate capillary hair; alar cells differentiated at basal angles, small, oblate, quadrate, or short-rectangular in several marginal rows or somewhat enlarged and thin-walled in concave, gradually differentiated auricles, or thick-walled and porose , concolorous or colored as in mid-basal region, often smooth, becoming papillose distally. Specialized asexual reproduction absent. Sexual condition autoicous, paroicous, rarely synoicous. Perigonia axillary on secondary stems and branches; gemmate. Perichaetia terminal on main stems and branches, often appearing lateral by innovation; perichaetial leaves erect, similar to vegetative leaves or larger, and longer, ovate to narrowly ovate-lanceolate, entire, plane or with a long, deeply channeled, abrupt, epilose acumen, or hyaline-tipped, eciliate to conspicuously ciliate, cilia sparse mid-leaf to dense in the apex, often longer than the leaf length, hyaline, nodose or dentate, flexuose at the toothed ends, lightly papillose. Vaginula with few or dense, long-filiform, yellow-hyaline, smooth paraphyses, not extending onto the developing calyptra, or with long, narrow teeth at the distal ends of some cells, and extending onto the developing calyptra. Seta one per perichaetium, very short or extending beyond the perichaetial leaves, smooth, straight, slender to stout, usually with a sclerotic swelling below the capsule neck. Capsule stegocarpous, erect, immersed to exserted, symmetric, subglobose to obovoid, long ovoid-cylindric, ellipsoid to fusiform (tapering proximally and distally), pyriform to obpyriform, smooth to irregularly wrinkled or regularly furrowed when dry, contracted, sometimes strongly, at base, neck irregularly longitudinally sulcate when dry, contracted distally below a rigid, smooth, incurved, wide or narrow mouth, stomata in neck, cryptoporic or phaneroporic; annulus lacking; operculum plano-convex, sometimes subumbonate-apiculate, or conic and stoutly, bluntly and obliquely short- to long-rostrate; peristome lacking. Calyptra conic-mitrate, undivided at the base or with several lobes, or cucullate, small and covering only the capsule apex to larger and covering one half the capsule length; smooth, naked or pilose. Spores brown, roughly or finely and evenly papillose to vermiculate-papillose, irregularly shaped, spheroid and flattened with an obscure tri-radiate mark on one side.
Genera 4, species 29 (3 genera, 6 species in the flora): nearly worldwide.
The family Hedwigiaceae is characterized by monoicous sexuality, stems with no central strand and capsules with no peristome. The capsules may be immersed or exserted. The generally ovate or oblong leaves are strongly concave, decurrent-auriculate, and abruptly acuminate to acute, , often with striking white apices or piliferous with long, pellucid apical hairs. The completely ecostate leaves and exosporic, globular, non-filamentous protonemal development are distinctive. Leaf cells throughout the family are usually subquadrate, thick-walled, collenchymatous with a diversity of coarse papillae, with patches of elongate cells in the leaf middle near the insertion, an area that is also usually more brightly colored than the rest of the leaf. The cells are generally arranged in longitudinal lines throughout the leaf. The calyptra may be glabrous, sparsely or densely hairy. Without peristomes, the family is difficult to place, but B. H. Allen (2002) gives a good synopsis of the speculative relationships of the Hedwigiaceae with other families. The family has been placed among the pleurocarpous mosses “probably because the leaves are ecostate, a feature extremely rare among acrocarpous arthrodontous mosses” (B. Goffinet and W. R. Buck 2004). The habits are acrocarpous because the archegonia are terminal on the main shoot, and the branching is sympodial (E. DeLuna unpublished) regardless of the plagiotropous habit typical of pleurocarpous mosses. All of the genera in the family possess papillose pseudoparaphyllia that are unique to the family.
Species in the family are usually found in dry, open rock-habitats, and are coarse, rigid plants when dry that are suddenly pliant and soft when moistened. Since the family is monoicous collections usually have sporophytes. All capsules are characterized by a neck region with lax, flaccid cells. When dry the cells collapse making the capsule irregularly and variously pleated at the base., When moist they are swollen rendering the capsule urceolate, strongly fusiform or turbinate when dry, and from globose, broadly oblong or cylindric-elliptic, or pyriform capsules when wet. Although there is no hypophysis, a broadened, sclerotic ring at the apex of the seta forms a kind of knot below the contracted neck, giving the capsule the erroneous appearance of being strumose when dry. Leaf margins with long, multicellular cilia are rare among bryophytes, but the margins of the perichaetial leaves of the widespread Hedwigia ciliata and H. stellata are long-ciliate.
SELECTED REFERENCES Allen, B. H. 2002. Moss Flora of Central America. Part 2. Encalyptaceae--Orthotrichaceae. St. Louis. Biasuso, A. B. 2007. The genus Hedwigia (Hedwigiaceae, Bryophyta) in Argentina. Lindbergia 32: 5--17. Crum, H. and L. E. Anderson. 1981. Mosses of Eastern North America, Vol. 2. Columbia University Press. New York. De Luna, E. 1990. Protonemal development in the Hedwigiaceae (Musci), and its systematic significance. Systematic Botany 15: 192--204. De Luna, E. 1995: The circumscription and phylogenetic relationships of the Hedwigiaceae (Musci). Systematic Botany 20: 347--373. Goffinet, B. and W. R. Buck. 2004. Systematics of the Bryophyta (mosses): from molecules to a revised classification. In: Molecular Systematics of Bryophytes, B. Goffinet, V. Hollowell and R. Magill, eds. Missouri Botanical Garden. Saint Louis. Pp 205--239. Goffinet, B., W. R. Buch and A. J. Shaw. 2008. Morphology, anatomy and classification of the Bryophyta. In: B. Goffinet and A. J. Shaw, eds. Bryophyte Biology, Second Edition. Cambridge. Pp. 55--138. Jones, G. N. 1933. Grimmiaceae In: A. J. Grout, ed. 1928--1940. Moss Flora of North America 2: 1-65. Newfane, Vermont and New York.
1. Capsule immersed , globose, mouth wide when dry; operculum plano-convex to subumbonate; laminal cell papillae usually forked or branched; stem without stoloniform branches; perichaetial bracts ciliate [eciliate in one species]; calyptra small, 0.5--1 mm, pilose to glabrous; stomata cryptoporic. . . . 1. Hedwigia, p. XXX
1. Capsule exserted , long-ellipsoid to pyriform, mouth wide or narrow when dry; operculum conic-apiculate to rostrate; laminal cell papillae always simple; stem with or without stoloniform branches; perichaetial bracts eciliate ; calyptra large, 2.5--3 mm, glabrous; stomata phaneroporic
2. Leaf apices concolorous, acute to abruptly short-acuminate; laminal cells pluripapillose with low, irregularly rounded papillae on both sides; capsules cylindric-fusiform (narrowed at mouth and base), 2--2.2 mm,; operculum medium- to long-rostrate; dry leaves longitudinally sometimes obscurely undulate-plicate; with stoloniform-tapered lateral branches and flagelliform-tapered branch tips; spores finely and evenly papillose. . . 2. Braunia, p. XXX
2. Leaf apices white-hyaline capillary hair-pointed; laminal cells 1-papillose, with a single simple, coarsely tuberculate papilla on both sides; capsules ovoid-pyriform to turbinate (often broadest at the mouth when dry), 1--2 mm,; operculum short-rostrate; dry leaves smooth, not plicate; stems without stoloniform lateral branches ; spores finely vermiculate-papillose . . . . 3. Pseudobraunia, p. XXX
1. HEDWIGIA Palisot de Beauvois, conserved name * [For Johannes Hedwig, 1730--1799, author of the Species Muscorum Frondosorum, 1801]
Plants robust, grayed or glaucous, when dry, generally sordid yellow-green, rarely brightly green, pale buff to red-brown in older parts, or stems dark greenish to reddish black, distal stems and branch tips concolorous, but often matte-white or hoary due to prominent overlapping rigid leaf-awns, fertile branch and stem tips with a tuft of shining flexuose bristles when cilia on margins of perichaetial bracts are well developed. Stems with concolorous leaves or clothed with short to elongate, broad-based, white-hyaline, erect to spreading recurved leaf tips; stoloniform-flagelliform branches absent, pseudoparaphyllia, when present, papillose and foliose. Leaves 1.5--2.5 mm, erect to subsecund and imbricate with erect to spreading tips when dry, erect-spreading to spreading when moist, smooth, occasionally somewhat striolate when dry, not undulate-plicate; leaf margins usually recurved in the proximal 1/2--2/3, sometimes plane, or narrowly to broadly recurved on one side, plane at mid leaf, the margins in the acumen plane to erect to incurved, nearly entire proximally to irregularly papillose with simple to multifid papillae distally, in larger leaves closely and irregularly dentate to spinulose-dentate in the apex; leaf apex acute to acuminate, concolorous when muticous, or with usually broad-based white-hyaline apical areas, often pellucid at the very tip, rather gradually and broadly acute to abruptly and narrowly to longly acuminate to subpiliferous, shallowly to deeply canaliculate-furrowed to tubulose from erect or incurving margins, or between the internal shoulders of the recurving margins, regularly to irregularly papillose proximally to spinose-serrate medially to rather smooth in the extremity, when dry erect to patent, wide-spreading, recurved or reflexed, wide-spreading when moist; proximal laminal cells in mid basal region strongly thick-walled, moderately to strongly porose, long-rectangular, longitudinally pluripapillose with 4--7 simple papillae in one row, strongly yellow- to red-orange across the insertion or extending further distally at the center of the leaf, 0.25--0.33(--0.50) leaf length and often into the alar region in 2--3 rows; median laminal cells subquadrate, short-rectangular or oblong-rhombic, on both surfaces 1- or pluripapillose, papillae simple and also variously regularly or irregularly multifid, thick-walled, moderately to coarsely and irregularly porose-sinuose, becoming progressively more coarsely papillose distally, especially on the abaxial surface; distal laminal cells in the apex chlorophyllose and similar to and concolorous with median cells, or gradually differentiated into an echlorophyllose broad-based, occasionally decurrent, short- or long-triangular apex of translucent, flat-white to hyaline short or elongate cells with thick and even walls, prorulose at the leaf apex and papillose with papillae simple, low, arranged in lines over the lumina to sparsely or coarsely and irregularly spinose-papillose, extending into a less papillose pellucid point; alar cells oblate, quadrate or short-rectangular, even-walled to thickened and porose in poorly delimited ovate or elongate-ovate groups in several rows along the leaf margins, concolorous or colored near the insertion as in mid-basal region. Sexual condition autoicous. Vaginula with sparse to dense long, smooth, occasionally papillose paraphyses, these often also on calyptra. Perichaetial leaves to 4 mm, with hairpoints to 1.6 mm, ovate-lanceolate, margins entire or essentially eciliate to conspicuously long-ciliate on the awns and distal leaf margins, cilia often longer than leaf length, cilia hyaline, loosely to strongly flexuose, smooth or with an occasional long, narrow tooth from the distalmost cell end to low simple-papillose.. Seta very short, 0.5--0.9 mm, hidden by the perichaetial leaves, reddish brown, thick. Capsule about 1 mm, deeply immersed, subglobose to shortly ovoid to obovoid when wet, turbinate-urceolate with a wide mouth when dry and deoperculate, brown proximally, shiny red-brown at the mouth, smooth except the short differentiated neck, which is longitudinally wrinkled to sharply sulcate when dry, capsule contracted below the wide, rigid red-brown, incurved mouth when dry, stomata cryptoporic in the neck; operculum plano-convex, sometimes subumbonate-apiculate. Calyptra 0.5--0.9 mm, conic-mitrate and undivided at the base, or with several lobes, small, and covering only the capsule apex, conic-cucullate, naked or weakly to strongly pilose. Spores ca. 19--30 \um, finely to somewhat coarsely vermiculate-papillose.
Species ca. 6 (3 species in the flora): nearly worldwide.
The genus Hedwigia in North America was recently considered monospecific, but with numerous varieties and forms (G. N. Jones 1933). Leaves of Hedwigia stellata and H. detonsa are generally sharply bicolorous at the often irregular boundary between the hyaline (white) acumen and the green distal lamina, but the apex is relatively more dilute and paler in H. ciliata and its varieties and forms. Pseudobraunia californica also displays strong coloration, usually with the color stronger over the lumina than the cell walls. The apical hyaline cells of vegetative leaves in the key to species are measured from the cell base where the middle lamella meet to the distal end of the cell. Although the lines of the lumen are often indistinct, those of the middle lamellae are usually clear. Note should be made that a “hair-point” is often ascribed to species of Hedwigia in the flora area, but, although the hyaline acumen may be variously elongated, there is never a filiform or hair-like process at the leaf apex as in Pseudobraunia.
The leaf papillae are best developed on the distal abaxial surface, being more reduced on the adaxial surface. The laminar papillae of all species of Hedwigia in the flora are variously branched. Those of Hedwigia ciliata are different from those of their congeners in that they are not columnar in the median abaxial region, but instead sessile. Those of H. stellata and H. detonsa are columnar and are distinctly regularly or irregularly branched from the top of the column, hence the term peltate has been used for this type of papilla. Recent papers recognizing the latter two species often show SEM photos clarifying this feature. Long, multicellular cilia on leaf margins are rare among bryophytes (B. Goffinet et al. 2008), but are present on the perichaetial leaves in the widespread Hedwigia ciliata and the more local H. stellata.
SELECTED REFERENCES Crundwell, A. C. 1995. Hedwigia stellata and H. ciliata in the British Isles. J. Bryol. 18: 807--810. Goffinet, B., W. R. Buch and A. J. Shaw. 2008. Morphology, anatomy and classification of the Bryophyta. In: B. Goffinet and A. J. Shaw, eds. 2008. Bryophyte Biology, Second Edition. Cambridge. Pp. 55--138. Jones, G. N. 1933. Grimmiaceae. In: A. J. Grout, ed., 1928--1940. Moss Flora of North America. 2: 1--65. Newfane, Vermont. Smith, A. J. E. 2004. The Moss Flora of Britain and Ireland, ed. 2. Cambridge.
1. Abaxial surface of distal part of leaf pluripapillose with (1--)2--4 simple, sessile or stalked, variously branched papillae per cell; perichaetial leaves ciliate; calyptra pilose or smooth; apical laminal cell of many vegetative leaves obtuse to truncate and pluripapillose, short-rhomboidal, (50--)75--95(--100)\ mu . . . . 1. Hedwigia ciliata
1. Abaxial surface of distal part of leaf 1-papillose, generally with a large, stalked, branched papilla per cell; perichaetial leaves ciliate or not; calyptra smooth ; apical laminal cell of many vegetative leaves sharply pointed, smooth, or distantly, sharply 1-papillate on l margin or pluripapillose long-linear, (80--)120--175(--200) um.
2. Leaf apex wide-spreading to squarrose when dry, with large spinose papillae among shorter papillae, distal cells sharply prorulose; papillae on abaxial leaf surface mostly irregularly, unequally branched; perichaetial leaf margins long-ciliate. . . . 2. Hedwigia stellata
2. Leaf apex erect when dry, with mostly low, rounded papillae in lines, distal cells smooth; papillae on abaxial leaf cells mostly equally branched; perichaetial leaf margins entire . . . . . 3. Hedwigia detonsa
1. Hedwigia ciliata (Hedwig) Palisot de Beauvois, Prodr. Aetheog., 60. 1805
Anictangium ciliatum Hedwig, Sp. Musc. Frond., 40. 1801; Hedwigia ciliata ssp. subnuda Kindberg ex Macoun & Kindberg
Leaves 1.5--3 mm, apices erect, erect-spreading, often variously secund when dry, ending in a short or long acumen, with a narrow-based, small, hyaline area 0.10--0.20 leaf length, or muticous, or with a prominent, broad-based hyaline area to 0.40--0.55 leaf length;
often with a broad or narrow, shallow or deep and sharp canaliculation in the acumen between the shoulders of broadly recurved margins, or by erect to incurved margins, apex of the acumen essentially flat; laminal margins broadly recurved to 1/3 leaf length, narrowly recurved to base of hyaline area on one or both sides or plane mid-leaf and with the distal margins in the acumen variously plane, erect to incurved, or all variations in some form in some leaves on the same plant; medial and distal laminal cells on both sides pluripapillose with (1--)2(--4) simple and sessile and also low-stalked, variously branched papillae per cell; laminal cells in apical hyaline area relatively simple, in distinct or irregular linear rows throughout or sparsely to coarsely and irregularly papillose basally and smooth to minutely rounded-papillose distally, distal cells ending in the hyaline area often irregularly branched to rather broad-based spinose-prorulose; hyaline margins with generally broad teeth, usually irregularly papillose to nearly smooth to the apical cell; apical laminal cell of many cauline leaves obtuse to clearly truncate and also pluripapillose-coronate, the cell relatively short-rhomboidal,(40--)50--75(--80) \um. Perichaetial leaves plane, often with a narrow or broad, shallow or deep canaliculation down the middle of the leaf between broadly recurved margins, or with narrowly recurved margins throughout on one or both sides, long-ciliate on the distal margins . Vaginula sparsely to densely pilose. Calyptra glabrous, sparsely pilose proximally, or densely pilose throughout.
Capsules mature spring to late spring. Dry mostly acidic rocks (granite or sedimentary), occasionally on conglomerates, limestone, rarely soil, cliffs, forming mats on dry, sunny boulders, in dry or wet, open or closed woods, acidic glacial erratic rocks and stones in limestone regions, occasionally tree trunks and branches, asphalt shingles, asphalt road edges; low to high elevations (0--2275 m); Greenland; Alta., B.C., Man., N.B., Nfld. and Labr. (Nfld. and Labr.), N.W.T., N.S., Nunavut, Ont., Que., Sask., Yukon; Ala., Alaska, Ariz., Ark., Calif., Colo., Conn., Del., D.C., 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., Ohio, Okla., Oreg., Pa., R.I., S.C., S.Dak., Tenn., Tex., Vt., Va., Wash., W.Va., Wis., Wyo.; Mexico; West Indies; Central; South America; Eurasia; Africa; Atlantic Islands; Pacific Islands (New Zealand); Australia.
Hedwigia ciliata is common and widespread in the floral area. The pseudoparaphyllia of this species are well illustrated by A. B. Biasuso (2007). The paraphyses of the vaginula and those extending onto the calyptra are smooth, but in forms with dense paraphyses extending onto the calyptra, the paraphyses are sparsely papillose and have sharp lateral teeth on one side at the distal ends of some cells, which are similar to the cells of the cilia on the margins of the perichaetial leaves. In the eastern to north-eastern part of the flora area, the typical facies is generally with the vaginula and calyptra densely pilose, the hyaline apices rarely absent to short or to 0.33 the leaf or longer (to 0.55) in the here excluded variety leucophaea Bruch & Schimper. The papillae at the base of the hyaline apices are coarse and have a chaotic mix of spinose and tall-branched papillae, rather abruptly becoming smooth distally, the papillae only becoming more regular in plants with larger hyaline apices. The laminal margins may be plane, but are usually recurved from the base to mid-leaf or throughout on one or both sides.
In the southwestern part of the flora area there is a rather strongly differentiated facies of the species where the vaginula and calyptra are sparsely pilose, the prominent hyaline leaf apical area is usually broad-based, the papillae are simpler in that area and arranged in rather distinct rows, and the leaves display a more concave, non-canaliculate surface. This facies is quite distinct along the Arizona, New Mexico and west-Texas border, but becomes less distinct and merges into what has been identified as Hedwigia ciliata var. leucophaea Bruch & Schimper, which can extend north and east into British Columbia and Michigan. There are two positions on this variation in North America: var. leucophaea and other varieties and forms of H. ciliata are not worthy of recognition;, var. leucophaea is indistinguishable from the southern hemispheric H. nivalis, . It appears, however, that the var. leucophaea of H. ciliata cannot yet be distinguished from H. nivalis.. Because there is essentially no agreement among bryologists regarding the character of Hedwigia ciliata var. leucophaea and H. nivalis, it is impossible to prepare a key differentiating the three (including var. ciliata) implied or proposed taxa in the study area.
Hedwigia ciliata var. ciliata and H. nivalis were lectotypified by L. Hedenäs (1994) and A. B. Biasuso (2007). Although the type of var. leucophaea has apparently been lost, L. Hedenäs (1994) gave formal synonymy for var. leucophaea including citation of types of heterotypic synonyms. W. R. Buck and D. H. Norris (1996) reported that the type of H. nivalis existed at NY, but apparently did not see it as citations of the types of H. stellata and H. detonsa were marked in their publication with ! but not the type of H. nivalis at NY. A. B. Biasuso (2007) reported that “The duplicate specimen of the original material kept in NY was requested but not found (W. Buck, pers. comm.).” W. R. Buck and D. H. Norris (1996) originally proposed the name Hedwigia nivalis for a facies of what was formerly called Hedwigia ciliata in the southwestern US, and for “all tropical American material.” B. Allen (2010) accepted this suggestion of the former authors. M. Lueth and A. Schaefer-Verwimp (2004) reported var. leucophaea as new to South America with the interesting notation that the two specimens of the variety were collected from the type locality of H. nivalis. The differences between Hedwigia ciliata var. leucophaea and H. nivalis according to B. Allen (2010) were the erect (vs. incurved) margins in the channeled hyaline apical portion; that the leaves in var. leucophaea are more oblong (less ovate) than in H. nivalis and the acumina are narrower; and the calyptra is pilose. All of these characters in the flora area are present and variable throughout the range of Hedwigia ciliata, especially in the areas generally west of mid-continent. The description of H. ciliata var. leucophaea given by L. Hedenäs (1994) described both varieties of H. ciliata with “furrowed” or “tubulose” hyaline areas (not “flat” and without recurved margins), noting that var. ciliata may have sparse to dense hairs on the calyptra, but in var. leucophaea, generally with sparse hairs, as can be demonstrated in H. nivalis. The typical facies of H. ciliata var. ciliata, with a very small hyaline area and recurved margins with densely pilose vaginula and calyptra, occurs generally east of mid-continent. That there is a geographically distinct gradation of Hedwigia ciliata var. ciliata, through var. leucophaea to what is proposed as Hedwigia nivalis may also be demonstrated in Europe, with the facies of H. nivalis present in the Mediterranean and Iberian Peninsula regions, but the sample examined by the author was too small for a definite conclusion. Leaves of the typical variety of H. ciliata in Europe seemed always to be longer than those displayed in eastern North America.
SELECTED REFERENCES Allen, B. 2010. Moss Flora of Central America. Part 3. Anomodontaceae--Symphyodontaceae. Missouri Botanical Garden. Saint Louis. Biasuso, A. B. 2007. The genus Hedwigia (Hedwigiaceae, Bryophyta) in Argentina. Lindbergia 32: 5--17. Lueth, M. and A. Schaefer-Verwimp. 2004. Additions to the Bryophyte Flora of the Neotropics. Tropical Bryology 25: 7--17. Hedenäs, L. 1994. The Hedwigia ciliata complex in Sweden, with notes on the occurrence of the taxa in Fennoscandia. J. Bryol. 18: 139--157.
2. Hedwigia stellata Hedenäs, J. Bryol. 18: 144, figs. 1--2. 1994
Leaves 1.5--2.5 mm, with broad, distal hyaline areas wide-spreading to squarrose when dry, ending in a long, broad-based acumen 0.20--0.36(--0.42) leaf length, at the base and middle of the hyaline area broadly shallowly to more deeply channeled due to erect or incurved margins, flat or sometimes twisted at the tips; laminal margins recurved from near the base to midleaf, plane in mid-leaf, erect to incurved in the acumen, medial and distal laminal cells on both sides with 1(--2) large, stalked, strongly and irregularly branched papillae per cell, the branches 3--5, of unequal lengths, especially on the abaxial surface, laminal cells in the hyaline area with relatively simple papillae at the base of the hyaline area arranged in longitudinal lines on the cell lumina, intermixed with scattered, simple, pungent, spinose papillae, the smaller papillae disappear distally generally in mid hyaline area or proximally, the spinose papillae are present to the apex and along margins that become otherwise smooth (epapillose), hyaline margins with smooth, sharply and narrowly spinose teeth throughout; apical laminal cell of many cauline leaves sharply-pointed, the cell relatively long-linear, 120--175(--200) \um, smooth or with an occasional sharp papilla on one margin, or 2-papillate and appearing 2-fid at the tip, Perichaetial leaves plane to weakly concave, margins often narrowly recurved, long-ciliate on the margins of the distal half, especially at the apex. Vaginula sparsely pilose. Calyptra glabrous to sparsely pilose.
Capsules mature spring. Dry boulders, rocks, open areas, outcrops in grassy woodlands, cultivated plum tree, largely coastal areas; low to high elevations (0--1300 m); B.C.; Calif., Oreg., Wash.; Europe (Denmark, Italy, Norway, Portugal, Spain, Sweden, United Kingdom); South America (Chile); Asia (Kazakhstan); Atlantic Islands (Iceland, Faroe Islands).
The marginal teeth in the hyaline leaf apices of Hedwigia stellata display the same elements as in the apical cell: generally sharply pointed and narrow, with an occasional papilla. In H. ciliata the teeth are broader and often irregularly covered with scattered papillae, especially on their tips. Hedwigia stellata is common in California only in the San Francisco and Los Angeles areas (W. R. Buck and D. H. Norris 1996). It tends to occupy habitats associated with the Pacific coast, especially Vancouver, B.C., and the inlets of islands around the Georgia and Juan de Fuca Straits, such as Vancouver Island, and northward on the Queen Charlotte Islands, leading the authors to speculate that H. stellata may be a recent introduction to North America. Specimens cited by the authors are often associated with trails and roads in public parks. The chlorophyllose, generally yellow-brown boundary between the body of the leaf and the hyaline apex is often irregular in this species with short, longitudinal lines of colored and non-colored cells juxtaposed, resulting in a jagged transverse line. Actually, in many leaves, the ultimate laminal cells are somewhat darker in coloration just at this boundary. This is also true in specimens of H. detonsa.
SELECTED REFERENCE Crundwell, A. C. 1995. Hedwigia stellata and H. ciliata in the British Isles. J. Bryol. 18:807--810.
3. Hedwigia detonsa (M. Howe) W. R. Buck & D. H. Norris, Nova Hedwigia 62: 368. 1996
Hedwigia albicans var. detonsa M. Howe, Erythea 5: 91. 1897
Leaves 2--2.5 mm, with the broad, distal hyaline areas erect when dry, ending in a long, broad-based acumen 0.20--0.40(--0.50) leaf length, at base and middle of the hyaline area channeled to tubulose due to erect to incurved margins, flat or sometimes twisted at the tips, laminal margins broadly or also narrowly recurved in the proximal 1/4 or plane on one or both sides, erect to incurved in the acumen; median and distal laminal cells on both sides with 1(--2) large, stalked, strongly and regularly branched papillae per cell, the short branches 3--5, mostly of equal lengths, especially on the abaxial surface, laminal cells in the hyaline area with relatively low, simple papillae at the base, arranged in longitudinal lines on the cell lumina, usually lacking large, spinose papillae, becoming sparse to absent in the smooth distalmost hyaline portion, margins in the hyaline area with low, simple papillae, with low, rounded teeth from distal cell ends, these sometimes with a small, sharp apical papilla, mostly without spinose papillae, apical laminal cell of many cauline leaves sharply pointed, relatively long-linear, (80--)110--125(--150) \um, smooth or usually with scattered, simple and minute papillae on the lumina and margins. Perichaetial leaves broadly channeled to involute, essentially entire (not long-ciliate) on the margins. Vaginula sparsely pilose. Calyptra glabrous to sparsely pilose.
Capsules mature spring. River canyons and open forested lands, dry or moist acid rocks, including sandstone, boulders and cliff faces near creeks, in open, diffusely lighted mixed forests of Quercus chrysolepis, Calocedrus, Torreya, Umbellularia, Pinus coulteri and Aesculus californica; low to moderate elevations (200--1500 m); Calif.
Unlike the irregular papillae of Hedwigia stellata and H. ciliata, those of H. detonsa as seen in cross section are nearly identical in regularity of form, being moderately stalked and evenly branched. Current studies suggest the species is endemic to Califormia, but it may also be expected in Baja California (W. R. Buck and D. H. Norris 1996). These authors find H. detonsa characteristic of river canyons and open, forested lands, being more inland than H. stellata, and avoiding grasslands and savannas.
SELECTED REFERENCES Buck, W. R. and D. H. Norris. 1996. Hedwigia stellata and H. detonsa (Hedwigiaceae) in North America. Nova Hedwigia 62: 361--370. Hedenäs, L. 1994. The Hedwigia ciliata complex in Sweden, with notes on the occurrence of the taxa in Fennoscandia. J. Bryol. 18: 139--157.
2. BRAUNIA Bruch & Schimper in P. Bruch, W. P. Schimper and W. Gümbel, Bryol. Eur. 3: 159 (fasc. 29--30. Mon. 1). 1846 * [For Alexander Carl Heinrich Braun, 1805--1877, Director of the Berlin Botanic Garden]
Plants robust, deep yellow-green or reddish brown to blackish brown in older parts, uniformly without differentiated white awns or hairs. Stems terminating in acuminate, non piliferous, concolorous leaves, producing flagelliform radiculose stolons laterally and copiously at the base of the stem thence distantly and in some branch and stem apices, the stolons with small, distantly spaced squarrose-recurved abruptly setaceous-acuminate leaves; rhizoids sparse, red-brown, especially evident on stoloniform branches. Leaves usually longitudinally plicate, usually loosely imbricate, with erect to somewhat secund spreading tips when dry, erect-spreading to widely spreading when moist, concave, moderately to distinctly plicate-sulcate when dry, with an acute to abruptly short-acuminate, concolorous apex, strongly channeled between erect margins, the larger leaves often with a small apiculus of several, usually eroded pellucid cells; laminal margins doubly recurved, broadly, and/or narrowly recurved at the base to mid leaf or to the apex on one or both sides, recurved to erect to somewhat incurved in the flat to channeled to subtubulose apex, entire, erose-denticulate at the tip; proximal laminal cells mid-basally quadrate in one row at the insertion, becoming long-rectangular to linear, more or less evenly thin to thick-walled and not to moderately porose, more deeply yellow to yellow-orange at the insertion usually in only one or two rows, seldom extending as much as 1/5 leaf length, pluripapillose longitudinally with 4--8 small simple papillae in one row, more strongly yellow or orangish yellow, sometimes extending distally to 0.20 lamina; median laminal cells subquadrate, short-rectangular or rhombic, on both surfaces pluripapillose, the papillae obscure, simple, low and irregularly rounded, 3--4(--6) per lumen, thin to thick-walled, the walls even or weakly to strongly porose-sinuose; distal laminal cells in the apex chlorophyllose, similar to and concolorous with median cells, becoming somewhat longer in long-acuminate leaves, 3--4:1, and less papillose, occasionally in larger leaves with several hyaline apical cells or chlorophyllose cells with a pellucid margin; alar cells usually isodiametric, in numerous rows at basal margins, thick-walled or somewhat enlarged and thinner walled near the insertion and gradually forming an excavate auricule, smooth to mostly 3-papillate, concolorous or more strongly colored in 1--2 rows as in the mid-basal region. Sexual condition autoicous, paroicous, rarely synoicous on the same plants or tufts. Vaginula with many long, hyaline, smooth paraphyses, these not extending onto the developing calyptra. Perichaetial leaves 2--2.5(--3) mm, similar to vegetative leaves, longitudinally weakly to strongly plicate, longer, ovate- or oblong-lanceolate with a channeled, abrupt acumen, margins entire (non-ciliate), apex non-pilose with a sharp apical cell. Seta elongate, (5--)7--14(--25) mm, extending beyond the perichaetial leaves, dark reddish yellow, stout, often with a prominent knot of sclerotic tissue at the distal end. Capsule (1.9--)2--2.2 mm, exserted, long ovoid-cylindric, ellipsoid when wet to fusiform (tapering proximally and distally) when dry, smooth when wet to irregularly striate-wrinkled to irregularly furrowed when dry, ovate (broadest at the base) when wet to variously strongly contracted at the short neck, contracted below a narrow, rigid, smooth incurved mouth; stomata phaneroporic in the neck; operculum conic, stoutly, bluntly and obliquely medium- to long-rostrate. Calyptra, 2--4 mm, covering to below the capsule middle, cucullate, naked. Spores 18--23 \um, finely and evenly papillose.
Species 23 (2 in the flora): mostly tropical and subtropical worldwide.
The habit of Braunia may be immediately distinguished from that of the other two genera in the flora area by the presence of flagelliform-stoloniferous branches emerging laterally from the stem or from branch apices. The undersides of the mats are a tangle of wiry flagelliform stolons that are generated laterally from the stem bases. They do not appear to be obviously involved in the proliferation of the plant and may have more of a rhizoidal function, at least effecting to keep the acrocarpous stem closer to the substrate. The essentially concolorous apices of the leaves are without whitened areas or filiform processes and the leaves are plicate-undulate, like the ridges in a scallop shell. The capsules are not immersed, as in Hedwigia, but distinctly exserted. The cells are pluripapillose, the papillae low, irregularly rounded rather than tuberculate or obviously branched.
1. Leaf margins recurved at base, plane above mid leaf; distal medial cell walls straight or weakly sinuose; plants autoicous . . . . 1. Braunia andrieuxii
1. Leaf margins recurved to near the apex; distal medial cell walls strongly sinuose; plants paroicous. . . . 2. Braunia secunda
1. Braunia andrieuxii Lorentz, Moosstudien, 164. 1864 F
Braunia secunda var. andrieuxii (Lorentz) Thériot
Plants with stolons apical on secondary stems and distally on their branches. Leaves with margins recurved proximally; distal medial cells with nearly straight or weakly sinuose walls. Sexual condition autoicous.
Capsules mature spring. Shaded or open cliffs faces and rock outcrops in gorges and canyons, talus, granite boulders, dry rocks; low to high elevations (0--1680 m); Ariz., Tex.; Mexico (Jalisco, Michoacán, Oaxaca).
Leaf apices are shorter and broader in Braunia andrieuxii than in B. secunda. Braunia audrieuxii seems paler yellow than B secunda, the cells relatively thinner-walled, moderately porose and smooth at the base. The contrasts in color on the lamina are less pronounced.
2. Braunia secunda (Hooker) Bruch & Schimper, Bryol. Eur. 3: 161 (fasc. 29--31. Monogr. 3). 1846
Hedwigia secunda Hooker, Musci Exot. 1: 46. 1818
Plants with stolons lateral on primary and secondary stems and branches. Leaves with margins recurved from base to apex; distal medial cells with strongly sinuose walls. Sexual condition paroicous.
Capsules mature spring. Shaded or exposed acidic cliffs, boulders, rocks, igneous rock faces and shaded ravines, in Quercus and Pinus woodlands with Arctostaphylos; moderate to high elevations (1500--1950 m); Ariz., N.Mex., Tex.; Mexico; Central America; South America; Africa; Asia (India).
The apices of the leaves of Braunia secunda are longer and narrower than those of B. andrieuxii and the leaf is more lanceolate and long-acuminate. Stoloniferous secondary stems and branches are also characteristic of various species outside of the floral area in Hedwigia and Hedwigidium, but not in the monotypic California genus Pseudobraunia.
3. PSEUDOBRAUNIA (Lesquereux & James) Brotherus in A. Engler & K. Prantl, Nat. Pflanzenfam. 1(3): 715. 1905 * [Name based on a close but spurious resemblance to the genus Braunia) E
Braunia subg. Pseudobraunia Lesquereux & James, Man. Mosses N. America, 153. 1884
Plants medium-sized, somewhat pale yellow-green to yellow-brown or usually red-brown, not hoary by conspicuous awns, stem and branches often hoary when sparsely clothed by long, narrowly based flexuose, shining hairs from the leaf tips. Stems with leaves proximally smaller, concolorous, less apiculate, quickly and gradually becoming enlarged and ultimately pilose, terminating in clusters of hyaline, flexuose pilose leaves at the ends of stems and new branches, with tufts of long, flexuose bristles from the apices of vegetative leaves distally or confined to the stem or branch tips; stoloniform-flagelliform branches absent. Leaves to 2 mm, smooth (not plicate) when dry, papillose on both leaf surfaces, more or less closely appressed-imbricate with somewhat to strongly secund, spreading tips when dry, erect-spreading to widely spreading when moist, not plicate; acute to more or less gradually acuminate, to piliferous capillary hair-points gradually absent to short, broad-based and confluent with the narrowing lamina apex, opaque to translucent white, papillose proximally, then smooth, entire to subdenticulate, when long, in one 1-cellular series, appearing jointed at the cell ends, straight to subflexuose; leaf margins doubly recurved, broadly and/or narrowly recurved to just beyond mid leaf, in the broad acumen flat, entire; proximal laminal cells mid-basally short-rectangular at the insertion to long-rectangular, thick-walled, moderately porose, concolorous or slightly more strongly yellow to orange-yellow at the insertion or to 0.20 lamina and into the alar region in 2--3 rows, 1-papillate (not in rows), appearing smooth; median leaf cells subquadrate, short-rectangular or oblong-rhombic, on both surfaces with one or less frequently two stoutly tuberculate simple (unbranched) papilla over the lumen, the walls thick and coarsely and irregularly porose; distal laminal cells in the apex similar to and concolorous with median cells, abruptly epapillose, differentiated into an abruptly hyaline, flat, rather narrow-based short-triangular apex of cells that are thick-walled, elongate, to 7:1, smooth, strongly porose, the tip variously extended into a short or long, cellularly 1-seriate, sometimes flexuose filiform hair ending in a single sharp, smooth cell; alar cells more or less isodiametric, in a concolorous excavate group of somewhat larger, more even and more thin-walled cells, smooth to mostly 1-papillate distally. Sexual condition autoicous. Vaginula with numerous, long, smooth, hyaline paraphyses, these not extending onto the developing calyptra. Perichaetial leaves similar to the vegetative leaves or larger, with apices irregularly rounded to broadly acute, with or without a short apiculus of hyaline cells, margins completely entire (not ciliate). Seta (4.5--)5--9(--10) mm, slender, pale yellow. Capsule ca. 1 mm, exserted, yellow- to red-brown, short and broad, smooth and ovoid-pyriform when wet, longitudinally sulcate-ribbed and obpyriform to turbinate (tapering to the seta and broadest at the mouth) when dry, contracted at the short neck when dry, constricted below a wide mouth when dry, stomata phaneroporic in the neck; operculum conic, stoutly, bluntly and obliquely short-rostrate. Calyptra 2.5--3 mm, covering to below the capsule middle, cucullate, naked. Spores 24--28 \um, finely to coarsely vermiculate-papillose.
Species 1 in the flora; western North America
Often the lumina of Pseudobraunia are a brighter color than the paler, thick cell walls, darkening at the border with the hyaline apex just as they are at the insertion. This is a monotypic genus endemic to western North America. Both Pseudobraunia and Braunia are more yellow, yellow-orange than Hedwigia, which is often a paler or richer if somewhat sordid green. Pseudobraunia has a long, flexuose filiform apical process on its leaves, unlike the rigid, more triangular white apical region of Hedwigia, which has immersed capsules, unlike the exserted capsules of Pseudobraunia. The two species of Braunia in the floral area has essentially no hyaline apical differentiation.
1. Pseudobraunia californica (Lesquereux) Brotherus in A. Engler & K. Prantl, Nat. Pflanzenfam. 1(3): 716. 1905 E F
Braunia californica Lesquereux, Trans. Amer. Philos. Soc. 13: 8. 1865; Hedwigia pilifera Mitten; Pseudobraunia californica var. pilifera (Mitten) Brotherus
Plants stoloniferous. Stems to ca. 8 cm. Leaves 0.75--1.5 mm exclusive of the hair-points, well-developed hair-points to 2 mm. Capsule 1--1.3 mm.
Capsules mature spring. Rocks usually dry during part of the year, metamorphic or igneous outcrops in oak woodlands, river canyons and ravines in the Inner Coast Ranges, granite boulders or vertical rock faces; exposed and open mossy granitic terraces and rock outcrops in a transition zone of chaparral and oak; low to high elevations, 100--1200 m; B.C.; Calif., Idaho, Oreg., Wash.
Pseudobraunia californica is commonly associated with species of Hedwigia on periodically moist rock outcrops. The hairpoints are best developed in the broader branch and stem tips, variously floccose in different populations to sparsely present. They are absent or nearly so at the stem and branch bases and gradually develop apically along the stem. The papillae, one per lumen and stoutly and simply tuberculate, will distinguish this species from many others in the family.
Hedwigia nivalis (Müller Hal.) Mitten
As noted above, the occurrence of this taxon in the flora area was discussed by W. R. Buck and D. H. Norris (1996), but without definitive conclusion.
Hedwigia ciliata var. leucophaea Bruch & Schimper
Only the typical variety of H. ciliata was accepted for the flora of North America by L. E. Anderson et al. (1990). This long-known European variety was accepted as a northern variant of H. ciliata by L. Hedenäs (1994), but it was not reported or recognized in several northern or circum-boreal floras, such as that of Canada (R. R. Ireland et al. 1987).
Anderson, L. E., H. A. Crum. 1990. List of the Mosses of North America North of Mexico. Bryologist 93: 448--499.
Ireland, R. R., G. R. Brassard, W. B. Schofield, and D. H. Vitt. 1987. Checklist of the mosses of Canada II. Lindbergia 13:1--62.