BFNA Title: Sanionia
Author: L. Hedenäs
Date: July 2000
Edit Level: Q
Version: 2

Bryophyte Flora of North America, Provisional Publication
Buffalo Museum of Science
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 AMBLYSTEGIACEAE-- Sanionia
Lars Hedenäs

version 2

SANIONIA L. Loeske, Hedwigia 46: 309. 1907 • for C. G. Sanio, German botanist (1832-1891)

Plants medium-sized to large, rarely small, green or yellowish green. Stem pinnately branched ± in one plane or irregularly branched to almost unbranched; with central strand and an outer hyalodermis, cells inside cortex thin-walled; paraphyllia absent; rhizoids smooth, slightly branched, rhizoids or rhizoid initials on stem at or just below leaf insertions; axillary hairs with 1--8 upper hyaline cells. Stem leaves circinate, falcate or almost straight, from rounded-triangular, ovate or broadly ovate base lanceolate and acuminate to very longly acuminate, plicate, rarely smooth, concave; margin plane or recurved above, sometimes incurved below, very finely to strongly denticulate above, finely denticulate or entire below; costa single, ending in acumen; median lamina cells linear, thin-walled or incrassate, not or slightly porose, smooth or some cells in upper leaf sometimes with prorate distal ends on abaxial side; differentiated alar cells few or numerous, shortly to longly rectangular, hyaline, thin-walled and inflated, in transversely triangular or ± isodiametric group, not or hardly decurrent. Branch leaves smaller than stem leaves. Autoicous. Inner perichaetial leaves gradually narrowed to very longly to shortly acuminate apex, plicate; margin unbordered, denticulate or dentate above; costa single and usually ending in acumen; scattered lamina cells in acumen sometimes with prorate distal ends; vaginula with paraphyses. Seta red; capsule cylindrical or shortly cylindrical, curved and horizontal to straight and erect; annulus separating; lid conical. Peristome perfect or specialized; exostome cross-striolate on lower outside, margin dentate above. Spores 10.5--21.0 µm, papillose or finely papillose.

Species belonging to Sanionia are characterized by their well developed hyalodermis, plicate leaves, hyaline, thin-walled, and inflated alar cells that form a transversely triangular or ± isodiametric group, the sometimes prorate distal cell ends on the abaxial leaf lamina, and, except in the most strongly specialised S. georgico--uncinata, very longly acuminate inner perichaetial leaves. Except for S. georgico--uncinata the species mostly occur in drier habitats than members of, e.g., Drepanocladus, Scorpidium, or Warnstorfia. The other members of the traditional, widely circumscribed genus Drepanocladus s. l. that are most closely related to Sanionia are now placed in Drepanocladus and Pseudo--calliergon (see notes after family description). Species of the latter two genera lack a hyalodermis, have 1--3 rather than 1--8 cells in the upper parts of their axillary hairs, and have generally got smooth rather than plicate leaves. However, because some small phenotypes of Sanionia may also have smooth or almost smooth leaves, the last character should be used with caution. The structure of the alar groups are different (see under the respective genera), the upper parts of the axillary hairs are shorter (1--3 celled) than in Sanionia (1--8), and the exostome margins are more weakly dentate than in S. uncinata, which is the only Sanionia species with an unspecialised peristome. Features separating Sanionia from Hamatocaulis, Loeskypnum, Scorpidium, or Warnstorfia are pointed out in the notes after the family description.

A widespread genus with four species in the Holarctic region, all occurring in North America. The genus occurs also in the southern hemisphere. Sanionia is still in need of taxonomic revisions in many geographical areas, and the distributions of the species outside North America, N and W Europe are partly incompletely known.

SELECTED REFERENCES Hedenäs, L. 1989. The genus Sanionia (Musci) in Northwestern Europe, a taxonomic revision. Ann. Bot. Fennici 26: 399--419. Janssens, J. A. 1983. Past and extant distribution of Drepanocladus in North America, with notes on the differentiation of fossil fragments. J. Hattori Bot. Lab. 54: 251--298. Wynne, F. E. 1944. Studies in Drepanocladus. IV. Taxonomy. Bryologist 47: 147--189.

 

1. Stem leaves acuminate to longly acuminate, with denticulate to strongly denticulate margins above. Inflated alar cells in ± isodiametric group; transition to supra-alar cells usually gradual; supra-alar cells mostly rectangular, sometimes longly rectangular, and often ± echlorophyllose (These features of the alar and supra-alar cells often give the impression of a longitudinally ovate group of 'alar' cells along the basal leaf margin). Large species, stem leaves 0.48--1.70 mm wide. Endostome reduced, irregularly perforated in all parts. Inner perichaetial leaves ± suddenly narrowed to acute or shortly acuminate apex, with margins strongly denticulate to dentate above. 4. S. georgico--uncinata

1. Stem leaves longly to very longly acuminate, with slightly denticulate to denticulate margins above. Inflated alar cells in transversely triangular group; transition to supra-alar cells sudden; supra-alar cells quadrate to rectangular and mostly chlorophyllose. Small or large species, stem leaves 0.30--1.45 mm wide. Endostome perfect or reduced, perforated only along mid-line of processes. Inner perichaetial leaves gradually narrowed to longly acuminate apex, with margins finely denticulate to denticulate above. 2

2. Supra-alar cells of stem leaves slightly or strongly incrassate, porose, forming a group equal in size to or larger than the alar cell group. Stem leaf costa mostly in the bottom of a deep, in transverse section narrow-angled fold. Larger species (stem leaves 0.62--1.45 mm wide), usually slightly and irregularly branched. Endostome in recently dehisced capsules strongly yellow; basal membrane 20--30% of total endostome height; cilia rudimentary. 3. S. orthothecioides

2. Supra-alar cells of stem leaves thin-walled or slightly incrassate, eporose, usually forming a group equal in size to or smaller than the alar cell group. Stem leaf costa (study cross-section in lower leaf) mostly in the bottom of a shallow, in transverse section wide-angled fold (or not in a fold). Smaller species (stem leaves 0.30--1.05 mm wide), usually ± pinnately branched. Endostome in recently dehisced capsules pale, brownish to yellowish, or strongly yellow; basal membrane 36--45% or 25--33% of total endostome height; cilia well developed or rudimentary. 3

3. Endostome in recently dehisced capsules pale, brownish to yellowish; basal membrane 36--45% of total endostome height; cilia well developed. Capsules horizontal or occasionally erect to inclined 1. S. uncinata

3. Endostome in recently dehisced capsules strongly yellow; basal membrane 25--33% of total endostome height; cilia rudimentary. Capsules erect to slightly inclined. 2. S. symmetrica

1. Sanionia uncinata (J. Hedwig) L. Loeske, Hedwigia 46: 309. 1907.

Hypnum uncinatum J. Hedwig, Spec. Musc. 289. 1801; Drepanocladus uncinatus (J. Hedwig) C. F. E. Warnstorf; Hypnum uncinatum subsp. moseri (N. C. Kindberg) N. C. Kindberg.

Plants slender to medium-sized, usually ± pinnately branched. Stem leaves circinate or falcate, rarely ± straight, from rounded-triangular or ovate base lanceolate, apex longly or very longly acuminate, 0.35--1.05 mm wide, plicate or strongly plicate, rarely smooth; margin plane or rarely partly recurved above, denticulate or finely denticulate above; costa mostly in the bottom of a shallow, wide-angled fold (or not in a fold); cells near leaf apex occasionally with distal ends prorate on back; alar cells in transversely triangular group, transition to supra-alar cells sudden, supra-alar cells quadrate to rectangular and mostly chlorophyllose, thin-walled or slightly incrassate, eporose, usually forming a group equal in size to or smaller than alar cell group. Inner perichaetial leaves gradually narrowed to longly acuminate apex, with margins finely denticulate to denticulate above. Capsules horizontal, rarely erect to inclined; exothecial cells just below capsule mouth in 1--3 rows of ± isodiametric cells; exostome unspecialized, marginal border of tooth widened at zone of transition in pattern of tooth outside; endostome unspecialized, in recently dehisced capsules pale, brownish to yellowish, basal membrane constituting 36--45% of total endostome height, processes perforated only along mid-line, cilia well developed. n = 12, 20, 9+m; outside North America also 10, 11, 30 (Fritsch 1991). The reported chromosome numbers should be considered with caution, since the identity of the investigated material has usually not been checked after recent revisions of the genus.

Sanionia uncinata is the only North American species of the genus with a perfect peristome and usually curved and horizontal capsules. When the shoot is looked at from the dorsal side, the leaves of normally developed shoots appear to be ‘combed’, with leaves pointing stiffly, obliquely forwards-outwards. Weak plants growing in relatively dry situations on rocks or tree stems may have less distinctly ‘combed’ leaves that are sometimes smooth or almost so. From all except S. symmetrica (see note after that species) it is separated by the structure of the alar and supra-alar cells. It is usually somewhat smaller and more regularly pinnately branched than S. orthothecioides and S. georgico--uncinata. From S. georgico--uncinata it is also distinguished by its much more longly acuminate inner perichaetial leaves.

From the sea level to high up in the alpine region, both in forests and in more open habitats, on rocks, logs, stumps, trees, soil or (especially in the north) in mires; Greenland, Alta., B.C., Man., N.B., Nfld., N.W.T., N.S., Ont., P.E.I., Que., Sask., Yukon, Alaska, Ariz., Calif., Colo., Idaho, Maine, Mass., Mich., Minn., Mont., Nev., N.H., N.Mex., N.Y., N.C., Oreg., Pa., S.Dak., Utah, Vt., Wash., Wis., Wyo.; southern Mexico, South America, Eurasia, Africa, Atlantic Islands, Australia, New Zealand, Antarctic.

The identity of Hypnum uncinatum var. plumulosum P. Bruch, W. P. Schimper & W. T. Gümbel (Drepanocladus uncinatus var. plumulosum (P. Bruch, W. P. Schimper & W. T. Gümbel) C. F. E. Warnstorf is somewhat unclear because no type material seems to be extant. However, most of the North American material referred to this name belongs to S. uncinata. Type material of Hypnum (Harpidium) moseri N. C. Kindberg (Hypnum uncinatum subsp. moseri) could not be found among Kindbergs material in S. However, since this taxon should differ from Sanionia uncinata mainly in its non-plicate leaves, and non-plicate leaves occur in some expressions of the latter it is unlikely to represent a separate taxon.

2. Sanionia symmetrica (F. Renauld & J. Cardot) J. A. Wheldon, Lancash. Chesh. Natural. 1921: 13. 1922.

Hypnum uncinatum subsp. symmetricum F. Renauld & J. Cardot, Rev. Bryol. 15: 72. 1888; Drepanocladus uncinatus var. symmetricus (F. Renauld & J. Cardot) A. J. Grout.

Plants medium-sized, usually ± pinnately branched. Stem leaves circinate or falcate, from rounded-triangular base lanceolate, apex longly or very longly acuminate, 0.30--1.05 mm wide, plicate or strongly plicate; margin plane or often narrowly recurved approximately in mid-leaf, denticulate or finely denticulate above; costa mostly in the bottom of a shallow, wide-angled fold (or not in a fold); cells near leaf apex occasionally with distal ends prorate on back; alar cells in transversely triangular group, transition to supra-alar cells sudden, supra-alar cells quadrate to rectangular and mostly chlorophyllose, thin-walled or slightly incrassate, eporose, usually forming a group equal in size to or smaller than alar cell group. Inner perichaetial leaves gradually narrowed to longly acuminate apex, with margins finely denticulate to denticulate above. Capsules erect or slightly inclined; exothecial cells just below capsule mouth in 2--4 rows of ± isodiametric cells; exostome specialized, teeth long and narrow, with border not widened at zone of transition in pattern of tooth outside; endostome specialized, in recently dehisced capsules strongly yellow, basal membrane constituting 25--33% of total endostome height, processes perforated only along mid-line, cilia rudimentary. Chromosome number not known (Fritsch 1991).

The gametophyte of S. symmetrica is impossible to separate with certainty from that of S. uncinata. However, the specialized peristome of S. symmetrica is very different from that of S. uncinata, and there can be no doubt that S. symmetrica is a distinct species. Occasional specimens of S. uncinata may have more or less straight and erect capsules, but in these the capsules are somewhat wider than in S. symmetrica and the peristome is still unspecialized. Sanionia symmetrica differs from S. orthothecioides and S. georgico--uncinata in its smaller size, in being more regularly pinnately branched, and in the structure of the alar and supra-alar cells. Sanionia georgico--uncinata has also got acute or much more shortly acuminate inner perichaetial leaves than S. symmetrica.

Mainly a lowland species, in moist and wet habitats in forests (usually deciduous), on shores of lakes and rivers, and in swamps, on logs, stumps, tree and shrub bases or on the ground; B.C., Idaho, Oreg., Wash.; endemic to western North America.

3. Sanionia orthothecioides (S. O. Lindberg) L. Loeske, Hedwigia 46: 309. 1907.

Hypnum uncinatum subsp. orthothecioides S. O. Lindberg, Oefv. K. Vet. Ak. Foerh. 23: 540. 1867.

Plants medium-sized to robust, mostly unbranched or sparsely and irregularly branched, rarely ± pinnately branched. Stem leaves falcate or strongly falcate, more rarely straight or almost so, from ovate-triangular to broadly ovate base lanceolate, apex longly or very longly acuminate, 0.62--1.45 mm wide, slightly or usually strongly plicate; margin mostly partly or entirely recurved (most distinct in ventral leaves), sometimes incurved in lower part, finely denticulate or almost entire above; costa mostly in the bottom of a deep narrow-angled fold; cells near leaf apex smooth; alar cells in transversely triangular group transition to supra-alar cells sudden, supra-alar cells quadrate or shortly rectangular, slightly or strongly incrassate, porose, forming a group as large as to much larger than alar cell group. Inner perichaetial leaves gradually narrowed to longly acuminate apex, with margins finely denticulate to denticulate above. Capsules erect or inclined; exothecial cells just below capsule mouth in 3--7 rows of isodiametric or transversely rectangular cells; exostome specialized, teeth long and narrow, with border not widened at zone of transition in pattern of tooth outside; endostome specialized, in recently dehisced capsules strongly yellow, basal membrane constituting 20--30% of total endostome height, processes perforated only along mid-line, cilia rudimentary. Chromosome number not known (Fritsch 1991).

Plants of this and the following species are normally considerably larger than the other two members of the genus. Sanionia orthothecioides is easily separated from S. uncinata and S. georgico--uncinata by its reduced and strongly yellow endostome. From these two species as well as from S. symmetrica it differs by its numerous (3--7 as compared with 1--4) rows of isodiametric or transversely rectangular cells below the capsule mouth, and by the structure of its alar and supra-alar cells. The leaf margin is usually recurved in S. orthothecioides, more rarely so in the other three species, and the costa is usually situated in the bottom of a deep fold in the lower part of the leaves in the first species, which is rare in the other three.

A coastal species that is rarely found in the inland, usually in open habitats, in rock crevices or meadows; Greenland, B.C., N.B., Nfld., N.W.T., Que., Yukon, Alaska; northern Eurasia.

 

4. Sanionia georgico--uncinata (C. Müller) R. Ochyra & L. Hedenäs in R. Ochyra, Moss Flora of King George Island, Antarctica 227. 1998.

Hypnum georgico--uncinatum C. Müller in G. Neumayer, Deutsch. Exp. Int. Polarforsch. 2: 320. 1890; Sanionia nivalis L. Hedenäs, Ann. Bot. Fenn. 26: 411. 1989.

Plants medium-sized to robust, mostly unbranched or sparsely and irregularly branched, rarely ± pinnately branched. Stem leaves falcate or strongly falcate, rarely straight or almost so, from ovate-triangular to broadly ovate base lanceolate, apex acuminate to longly acuminate, 0.48--1.70 mm wide, smooth or plicate, rarely strongly plicate; margin plane or sometimes partly recurved in upper part, usually strongly denticulate above; costa mostly in the bottom of a shallow, wide-angled fold (or not in a fold); cells near leaf apex occasionally with distal ends prorate on back; alar cells in ± isodiametric group, transition to supra-alar cells gradual (alar and supra-alar cells together often forming a ± homogeneous ovate basal marginal group), supra-alar cells rectangular or longly rectangular, thin-walled and eporose (mostly ± echlorophyllose), forming a group as large as to much larger than alar cell group. Inner perichaetial leaves ± suddenly narrowed to acute or shortly acuminate apex, with margins strongly denticulate to dentate above. Capsules ± erect; exothecial cells just below capsule mouth in (1--)2--3 rows of ± isodiametric cells; exostome specialized, teeth narrow in upper part, with border not widened at zone of transition in pattern of tooth outside; endostome specialized, in recently dehisced capsules hyaline or slightly yellowish, basal membrane constituting 17--33% of total endostome height, processes and basal membrane with one or both cell wall layers strongly perforated all over, cilia rudimentary or absent. Chromosome number not known (Fritsch 1991).

Sanionia georgico--uncinata is easily separated from the other three Sanionia species by its reduced and irregularly perforate endostome, its acute to shortly acuminate inner perichaetial leaves with strongly denticulate or dentate upper margins (the other three species have longly acuminate inner perichaetial leaves with finely denticulate or denticulate upper margins) and by the structure of the alar and supra-alar cells. In addition, the stem leaves of S. georgico--uncinata are more shortly acuminate and have more strongly denticulate margins that in the three other species.

Found in mountains in the high- and middle-alpine regions, rarely lower, and in the arctic, in large late snowbeds and on the shores of glacier-fed brooks; Greenland, N.W.T., Yukon, Alaska, Mont.; northern Europe, Antarctic.

OTHER REFERENCE

Fritsch, R. 1991. Index to bryophyte chromosome counts. Bryophytorum Bibliotheca 40: 1--352.

Version History for Sanionia:

Version 2: July 2000

Version 1: November, 1999