Weissia subg. Hymenostomum (R. Brown) Andrews
Plants in low cushions or turfs or loosely caespitose, green above, brown to tan or yellow below. Stems to 1 cm, hyalodermis weakly differentiated to distinct, seldom absent, sclerodermis weakly differentiated in 1--2 layers, central strand present; axillary hairs with proximal 1--2 cells somewhat thicker walled. Leaves incurved, commonly tubulose and often contorted or spiraled when dry, spreading when moist; long-ligulate, oblong or triangular to long-lanceolate, 1.5--2.5(--4) mm; adaxial surface broadly channeled across leaf; base scarcely differentiated to ovate or rectangular, occasionally half-sheathing; distal margins usually sharply incurved, occasionally merely erect, seldom plane, entire; apex sharply acute to subulate, occasionally broadly acute, obtuse, or weakly cucullate; costa shortly and sharply mucronate, adaxial outgrowths absent, adaxial cells quadrate to short-rectangular except occasionally elongate near apex, in 4--8 rows; transverse section ovate, occasionally circular or semicircular, adaxial epidermis present, adaxial stereid band present, guide cells 4--6(--8) in 1 layer, hydroid strand absent, abaxial stereid band present, crescent in sectional shape, abaxial epidermis present or occasionally absent; basal cells differentiated across leaf, rarely rising higher along margins in a V, rectangular, occasionally rhomboid, 2--5:1; distal medial cells subquadrate to hexagonal, 7--13 μm, papillae 2-fid, 2--6 per lumen, occasionally fused into a large multiplex papilla covering the lumen, occasionally spiculiform and branching. Specialized asexual reproduction rare, as rhizoidal gemmae. Sexual condition monoicous (usually autoicous), occasionally dioicous; perichaetia terminal, interior leaves occasionally sheathing seta, little different from cauline leaves or somewhat larger. Seta 0.1--1.3 cm. Capsule stegocarpic or cleistocarpic; theca elliptical to short-ovate or cylindric, 1--2.2 mm, annulus in ca. 2 rows of persistent, vesiculose cells or not differentiated; operculum when present conic to rostrate; peristome teeth 16 or rudimentary or absent, occasionally removed with operculum, oblong-truncate to long-triangular, often irregularly cleft, straight or twisted weakly counterclockwise, mouth occasionally closed by a remnant of the spore sac. Calyptra cucullate. Spores 14--28 μm. KOH laminal color reaction yellow.
Species ca. 97 (10 in the flora): all continents except Antarctica, found mostly on soil.
A world evaluation of Weissia and Trichostomum (R. H. Zander 1993) at the generic level suggested that species of the two genera might be shuffled into a more natural arrangement if taxa with identical or nearly identical gametophytes were grouped, with differences in sporophytes easily and simply explained by reduction. It is insufficient, however, to simply split Weissia and Trichostomum into two genera, as was implemented by Zander (1993), based on leaf margins sharply incurved or erect to nearly plane, respectively. Weissia jamaicensis and Trichostomum crispulum have similar gametophytes but differ markedly in the leaves of the former being sharply incurved marginally and of the latter nearly plane to merely erect. No major changes were made, however, for this treatment given the need for a full revision. Certain Weissia species with erect leaf margins may have a somewhat cucullate leaf apex similar to that of Trichostomum crispulum, but the lamina of the latter is generally broader, 16--20 cells across on one side of the costa just above mid leaf (except in strongly reduced specimens with short-ovate leaves), while Weissia species with which it may be confused have laminae usually 10-15 cells across a lateral lamina. The present species concepts are largely identical with those of A. Stoneburner (1985) with exceptions as noted.
All species of Weissia with sharply inflexed distal leaf margins have a tendency to show adaxially bulging cells in leaf section, possibly because the inflexion crowds the cells. This differential bulging of cell superfices is probably homoplastic to adaxially bulging distal laminal cells in other pottiaceous genera without sharply inflexed distal margins, such as Hyophila. Thus, the close relationship of Weissia with Hyophila and relatives cladistically demonstrated by R. H. Zander (1993) is probably an artifact. Regarding the hymenium that may be seen closing the newly deoperculate capsules of the eperistomate segregate genus Hymenostomum, such a membrane is present in all species of the genus, being the top of the spore sac, and it is simply more evident in the eperistomate species; see R. H. Zander's (1993) review of the problem. A. Stoneburner summarized evidence that supports synonymy of Weissia and Astomum.
SELECTED REFERENCES
Anderson, L. E. & B. E. Lemmon. 1972. Cytological studies of natural intergeneric hybrids and their parental species in the moss genera Astomum and Weissia. Ann. Missouri Bot. Gard. 59: 382--416. Stoneburner, A.1985 [1986]. Variation and taxonomy of Weissia in the southwestern United States. II. Taxonomic treatment. Bryologist 88: 293--314. Stoneburner, A. and R. Wyatt. 1985.Variation and taxonomy of Weissia in the southwestern United States. I. Biometrical analyses. Mitt. Thüring. Bot. Ges. 11: 175--185. Zander, R. H. 1993. Genera of the Pottiaceae: Mosses of Harsh Environments. Bull. Buffalo Soc. Nat. Sci. 32.
EXCLUDED
Weissia condensa (Voit) Lindberg is a name that replaces Weissia tortilis (Schwägrichen) J. K. A. Müller, illegitimate homonym according to M. F. V. Corley et al. (1981). Synonyms of this European species have been applied to specimens of W. controversa in the range of the flora, including Hymenostomum tortile (Schwägrichen) Bruch & Schimper; Weissia euteiches R. H. Zander, superfluous name; and W. tortilis (Schwägrichen) J. K. A. Müller, illegitimate homonym. European specimens of W. condensa usually have cucullate leaf apices, thick costae to 100 µm wide basally, distal laminal cells 8--10 µm wide, and capsules usually eperistomate; the leaves commonly have rather stout and long mucros of (4--)7--10 cells. Weissia sharpii is similar but has large distal laminal cells, 10--13 µm wide (2--)1:1. The combination of a strong mucro, wide costa and generally eperistomate but stegocarpic capsule is not found in the flora area and thus this species is excluded. Although A. Stoneburner (1985) attributed an eperistomate condition to W. condensa, R. H. Zander (1994) pointed out that W. condensa may occasionally have a very rudimentary peristome, one or two segments high, hidden below the capsule stomal rim (e.g. Slovakia, Pilous 139, MO). The thickness of the capsule wall in numbers of cell layers is used by R. H. Zander (1994) to distinguish W. condensa from W. controversa in Mexico, but this is apparently not a good feature because all Weissia capsules examined in the present study are three to four cell layers thick; although W. controversa occasionally has 2--3 layers, with the innermost layer difficult to demonstrate or possibly eroded or resorbed.
Weissia flavescens E. Britton (= Trichostomum brittonianum R. H. Zander) was reported from Florida by W. D. Reese (1991), but these sterile plants proved to be indistinguishable from Trichostomum crispulum.
Weissia perligulata H. A. Crum is a synonym of Trichostomum planifolium (Dixon) R. H. Zander.
Weissia sweetii E. B. Bartram. Although A. Stoneburner placed this species, and W. perligulata, under Trichostomum crispulum. L. R. Stark (1996) correctly recognized it as distinct. It is, however, a synonym of T. planifolium (Dixon) R. H. Zander.
Weissia wimmeriana (Sendtner) Bruch, Schimp. & W. Gümbell is a European species whose report from Minnesota, according to Andrews (1933) is based on W. controversa.
OTHER REFERENCES
Andrews, A. L. 1920. Hymenostomum in North America. I. Delimitation of the genus. Bryologist 23: 28--31.
Andrews, A. L. 1922. Hymenostomum in North America. II. The case of Astomum sullivantii. Bryologist 25: 66--71.
Andrews. A. L. 1924. Hymenostomum in North America. III. Astomum ludovicianum. Bryologist 27: 1--3.
Andrews, A. L. 1933. Hymenostomum in North America. V. Weisia viridula. Bryologist 36: 28--31.
Bartram, E. B. 1927. Weisia and Hymenostomum in southwestern United States. Bryologist 80: 77--83.
Corley, M. F. V., A. C. Crundwell, R. Düll, M. O. Hill and A. J. E. Smith. 1981 [1982]. Mosses of Europe and the Azores; an annotated list of species, with synonyms from the recent literature. J. Bryol. 11: 609--689.
Flowers, S. 1973. Mosses: Utah and the West. Brigham Young Univ. Press, Provo, Utah.
Hill, M. O. 1981. New combinations in European mosses, I. Pottiaceae. J. Bryol. 11: 599--602.
Koponen, T., P. Isoviita and T. Lammes. 1977. The bryophytes of Finland: An annotated checklist. Flora Fennica 6: 1--77.
Redfearn, P. L., Jr. 1976. A Trichostomum from Texas new to the United States. Bryologist 79: 80--82.
Reese, W. D. 1988. Rhizoid gemmae in Weissia controversa. Bryologist 91: 184--185.
Reese, W. D. 1991. Weissia (Hymenostomum) flavescens new to the United States. Bryologist 94: 179--180.
Reese, W. D. and B. E. Lemmon. 1965. A natural hybrid between Weissia and Astomum and notes on the nomenclature of North American species of Astomum. Bryologist 68: 277--283.
Shaw, A. J. 1987. Experimental taxonomy of Weissia controversa and W. sharpii (Musci: Pottiaceae). Syst. Bot. 12: 381--389. cultivation.
Stark, L. R. 1996. The status of Weissia sweetii, a species endemic to the southwestern United States. Bryologist 99: 345--348.
Williams, C. 1966. A natural hybrid in the genus Weissia. Bryologist 69: 361--365.
Zander, R. H. 1994. Weissia. In: A. J. Sharp, H. A. Crum and P. M. Eckel (eds.). Moss Flora of Mexico. Mem. New York Bot. Gard., Vol. 69. 2 vols.
