BFNA Title: Solenostomataceae

Date: Dec. 6, 2017
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Bryophyte Flora of North America, Provisional Publication

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XX. SOLENOSTOMATACEAE  Stotler & Crandall-Stotler

B. Crandall-Stotler

Plants forming mats or turfs; branches sometimes intercalary from sides of stem in lower part of leaf axil, sometimes replacing ventral half of a leaf; with or without flagella. Leaves alternate, succubous, plane or concave, simple, entire; underleaves absent. Rhizoids scattered over ventral stem. Specialized asexual reproduction absent. Gynoecium terminal on an ordinary leafy branch. Perianth with or without subfloral branches, perianth present, well developed and projecting well beyond the bracts, or reduced and mostly hidden by the bracts, cylindric, usually somewhat plicate distally, mouth contracted or beaked, perigynium low or up to twice the length of the perianth, rarely absent.

Genera 6 (3 in the flora): Cosmopolitan, from the high Arctic to the subantarctic; North America, Central America, South America, Eurasia, Africa, New Zealand, and Australia.

The break-up of the portmanteau Jungermanniaceae Rchb. began in 2007 with the molecular studies of Hentschel et al. (2007), which showed that the family, as traditionally circumscribed (e.g., Crandall-Stotler & Stotler, 2000) was polyphyletic. In Crandall-Stotler et al. (2009), the Solenostomataceae Stotler & Crand.-Stotl.was erected to accommodate the monophyletic lineage of jungermannioid genera that had some level of perigynial development, with other well-supported lineages comprising the Delavayellaceae R. M. Schust. and Jungermanniaceae. Further comprehensive

molecular analyses resulted in the synonomy of Delavayellaceae with Jungermanniaceae and the transfer of Nardia to Gymnomitriaceae H. Klinggr. by Shaw et al. (2015). Currently, Solenostomataceae includes 5 or 6 genera (Söderström et al., 2016), depending on whether Plectocolea is regarded as a distinct genus, as treated here, or as a subgenus of Solenostoma (Shaw et al., 2015).

SELECTED REFERENCES  Crandall-Stotler, B. and R. E. Stotler. 2000. Morphology and classification of the Marchantiophyta, Pp. 21--70 in Shaw, A. J. and B. Goffinet (eds.), Bryophyte Bryology, 1st ed. Cambridge University Press, Cambridge. Crandall-Stotler, B., R. E. Stotler and D. G. Long. 2009. Phylogeny and classification of the Marchantiophyta. Edinburgh J. Bot. 66: 15--198. Hentschel, J., J. A. Paton, H. Schneider and J. Heinrichs. 2007. Acceptance of Liochlaena Nees and Solenostoma Mitt., the systematic position of Eremonotus Pearson and notes on Jungermannia L. s.l. (Jungermanniidae) based on chloroplast DNA sequence data. Pl. Syst. Evol. 268: 147--157. Shaw, B., B. Crandall-Stotler, J. Váňa, R. E. Stotler, M. von Konrat, J. J. Engel, E. C. Davis, et al. 2015. Phylogenetic relationships and morphological evolution in a major clade of leafy liverworts (phylum Marchantiophyta, order Jungermanniales): Suborder Jungermanniineae. Syst. Bot. 40: 27--45. Söderström, L., A. Hagborg, M. von Konrat, S. Bartholomew-Began, D. Bell, L. Briscoe, E. Brown, et al. 2016. World checklist of hornworts and liverworts. PhytoKeys 59: 1--828.

1.  Plants julaceous; leaves imbricate, erect, concave, dorsally appressed, contiguous-connivent, forming bilabiate pairs at the shoot apex; oil bodies irregularly botryoidal; gynoecium appearing as a bilabiate, conical head, subfloral innovations sometimes present; bracts imbricate, dorsally contiguous, bearing conspicuous marginal slime papillae; perianth reduced, included within the bracts; perigynium well developed, bearing the bracts on its surface ..............1. Cryptocolea, p. xxx

1.  Plants not julaceous; leaves distant to imbricate, spreading, not dorsally appressed or contiguous; oil bodies large, spheroidal, finely granular; gynoecium never bilabiate, with the bracts always spreading, without marginal slime papillae; perianth either well-developed or reduced, but never completely hidden by the bracts; perigynium present or absent

2.  Leaves distant to contiguous, never imbricate; rhizoids red to purple; perianth variously reduced, but always partially emergent from the bracts; distally pluriplicate, with the mouth contracted, but not beaked; mid-perianth cells long-rectangular to semi-linear; perigynium well developed, equal to or greater than the length of the perianth ...................2. Plectocolea, p. xxx

2.  Leaves contiguous to imbricate; rhizoids hyaline to brownish; perianth well-developed, and long emergent beyond the bracts; distally 4--5 keeled, with the mouth contracted into a short beak; mid-perianth cells isodiametric to shortly rectangular; perigynium absent or less than 0.25 the length of the perianth

            ..........................3. Solenostoma, p. xxx

1. CRYPTOCOLEA R. M. Schuster, Amer. Midland Nat. 49: 417. 1953 • Greek kryptos, cryptic and koleos, female sheath, in reference to the short perianth completely hidden by the perichaetial bracts

Steven L. Jessup

Plants prostrate and creeping with short erect apices, loosely julaceous, appearing laterally compressed below the apex, erect and connivent lateral leaves comprising a dorsal keel; shiny, pellucid green to yellow-brown, often suffused with chestnut-brown. Stems sparsely branching by lateral-intercalary innovations arising below or in the axils of perichaetial bracts, occasionally branching by apical bifurcations, branches sometimes reduced, but otherwise undifferentiated. Lateral leaves broadly and obliquely inserted, succubous-transverse, alternate, antically secund, loosely to tightly imbricate; orbicular to broadly ovate, 0.4--0.8 mm, adaxially concave in the center, margins entire, involute; apical leaves valvate and tightly appressed, weakly recurved to squarrose or broadly reflexed in distal third; leaf cells roughly isodiametric, (20--)25--35(--45) \um, thin walled, trigones small, weakly developed or absent, oil bodies typically 3--5, irregularly botryoidal. Ventral leaves absent. Specialized asexual reproduction absent. Sexual condition dioicous. Androecia terminal as a short spike; antheridia 1--2 per bract, stalk biseriate. Gynoecia terminal, occasionally appearing dorsal by development of subtending branch innovations; forming distinctive conical, laterally compressed, 2-valvate heads, 3--4 mm; gynoecial bracts broadly ovate in 2--4 labiate pairs; perianth entirely concealed by bracts, short tubular, loosely and irregularly plicate, mouth crenulate to distinctly lobed, open to weakly pursed; basal 1/4 to 1/2 of perianth fused with base of gynoecial bracts to form a short thickened perigynium centered on stem axis. Sporophytes rare. Foot embedded in stem tissue below perigynium base. Seta with exterior layer of about 18--25 cells, walls slightly thickened; cortical cells thin-walled, otherwise undifferentiated. Capsule dark red-brown. Elaters contorted, 7--10 \um wide, 2-spiraled, attenuate at tips. Spores 14--16(--19) \um, granulose, red-brown.

Species 1 (1 in the flora); known from a few widely scattered locations in the Arctic and from the disjunct type locality on islands in northern Lake Superior.

Cryptocolea is a phylogenetically isolated member of the Jungermanniaceae, apparently not closely related to any other lineage in the flora. Cryptocolea is probably most closely related to the narrowly endemic and little-known Himalayian genus Diplocolea (T. Amakawa 1963).

SELECTED REFERENCES Schuster, R. M. 1969. Hepaticae and Anthocerotae of North America East of the Hundredth Meridian, Vol. 2. New York.

1. Cryptocolea imbricata R. M. Schuster, Amer. Midland Nat. 49: 417. 1953 F

Plants isolated or clumped in loose patches, 5--10(-- 15) mm. Stems of main sterile shoots and fertile male plants 0.2--0.3 mm wide; fertile shoots of female plants to 0.5 mm wide; rhizoids numerous, long, weakly clumped below the erect apex, clear to brown or faintly purple. Lateral leaf insertion dorsally transverse with short decurrence along the dorsal midline of axis, lateral insertion long-oblique, ventral insertion transverse; margins frequently decolorous in narrow zone. Sexual condition dioicous, male and female plants forming separate, often widely spaced patches, rarely occurring together. Androecia typically comprising several pairs of tightly imbricate androecial bracts, the inflorescence becoming intercalary with apical growth, often forming fertile branches with successive male inflorescences interrupted by a few vegetative leaves. Gynoecia 2-valvate, labial bracts with weak dorsiventral compression at maturity, interior bracts bearing slime papillae along labial contact surfaces; perianth bearing a few scattered marginal slime papillae. Sporophytes (5--)8--10(--15) mm at maturity. Capsule with 12--18 distinct longitudinal ridges.

Capsules mature in summer (July--Aug.). Mature sporophytes are known only from a single collection. Generally restricted to perennially moist basic substrates, peaty soil over basalt, slopes influenced by meltwater from basalt cliffs, calcareous fens, basic mineral soil and calcareous silt deposits, margins of shallow lakes; 0--600 m; Greenland (R. M. Schuster 1969, 1988); Nun. (Ellesmere Island); n Alaska (Brooks Range), Mich., Minn.; Europe (Norway, Sweden); Asia (Russia in Siberia).

Cryptocolea imbricata is a rare circumarctic species that is disjunct in a narrow tundra zone along shorelines of a few islands in northern Lake Superior--Susie I., Minnesota, and Apostle I., Michigan (R. M. Schuster 1969), and should be sought in similar refugial habitats elsewhere in the boreal latitudes. It is known outside the flora area from Chukoskiy Peninsula in Siberia (R. N. Schljakov 1975), Spitzbergen, and Swedish Lapland (J. A. Paton 1999). Cryptocolea imbricata is most likely to be confused with Arnellia fennica. Though similar in habit, size, leaf-shape, and color, Arnellia is never shiny, has distinctly opposite lateral leaves, and ventral leaves that are easily revealed. A few misidentified collections are Nardia geoscyphus, which, though also laterally compressed, is distinctly smaller in all respects, and has prominent trigones and well developed ventral leaves.

OTHER REFERENCES  Amakawa, T. 1963. New or little known Asiatic species of the family Jungermanniaceae. I. J. Hattori Bot. Lab. 26: 20--26. Paton, J. A. 1999. The Liverwort Flora of the British Isles. U.K., Colchester.  Schljakov, R. N. 1975. Addimenta ad floram hepaticarum arcticae URSS. Novosti Sist. Nizhsh. Rast. 1975: 318--323.  Schuster, R. M. 1953. Boreal Hepaticae. A Manual of the Liverworts of Minnesota and Adjacent Regions. Amer. Midland Nat. 49: 257-684.  Schuster, R. M. 1988. The Hepaticae of South Greenland. Beiheft zur Nova Hedwigia, Heft 92. Berlin. 

2. PLECTOCOLEA (Mitten) Mitten, Fl. Vit., 405. 1871 * [Greek, pleko, braided basket, and koleos, vagina, alluding to the turbinate distal perianth]

Vadim Bakalin

Solenostoma subg. Plectocolea Mitten, J. Proc. Linn. Soc. Bot. 8: 156 1865; Jungermannia Linnaeus subg. Plectocolea (Mitten) Amakawa

Plants 3--150 x 0.6--5 mm, prostrate to ascending or erect, yellowish brown, brownish, reddish brown, or deep green . Stem transversely elliptic in cross section, branching lateral intercalary or ventral, rarely as postical innovations and scale-like leaved strongly rhizogenous branches; dorsal surface cells rectangular to usually linear, thin- to thick-walled with distinct or indistinct trigones, cuticle mostly striolate; stem cross section mostly differentiated in 2--3 layers, with the outer as a hyalodermis, or if hyalodermis absent a sclerodermis, in some species the differentiation may be unclear to virtually absent. Rhizoids red, red-brown to purple, blackish brown to green, almost always brown with purple or purplish pigmentation at least as traces. Leaves distant to contiguous, never imbricate, obliquely inserted, mostly distinctly decurrent dorsally and sometimes ventrally, midleaf cells subisodiametric, mostly 5--6-gonal, walls colorless to yellowish and brownish, slightly thickened to evidently thickened or thin-walled, trigones distinct in most species, mostly concave, rarely large and convex. Sexual condition dioicous or paroicous. Perianth terminal on main axis, conical to tubular and pyriform, 2-stratose in proximal third, of long-rectangular to semi-linear cells at mid perianth. Perigynium always well-developed, from 1/3 to 2 times the perianth length. Androecia intercalary, with 1--10 pairs of bracts (frequently androecial branch with dead androecia distally), 1--3 antheridia per bract, stalk 2-seriate. Capsule wall 2-stratose. Elaters 2-spiralled, always with homogenous ends, more or less elongate, 1/5--1/10 times the elater length.

Species ca. 50 (7 in the flora): North America, Central America, South America, Eurasia, Africa, Australia, Pacific Islands.

The exact number of Plectocolea species is unknown because many taxa not yet revised have been described from Eastern Asia.

SELECTED REFERENCES Doyle W. T. and R. E. Stotler. 2006. Contribution toward a bryoflora of California III. Keys and Annotated species Catalogue for Liverworts and Hornworts. Madroño.53: 89--197.  Damsholt, K. and J. Váňa. 1977. The genus Jungermannia L. emend. Dumort. (Hepaticae) in  Greenland. Lindbergia. 4: 1--26. Godfrey, J. D. and G. A. Godfrey. 1979. Jungermannia schusteriana, a new hepatic from the Pacific coast of North America. J. Hattori Bot. Lab. 46: 109--117. Hicks, M. L. 1992. Guide to the Liverworts of North Carolina. Durham, North Carolina. Schuster, R. M. 1969. The Hepaticae and Anthocerotae of North America. Vol. 2. New York and London. Schuster, R. M. 1988. The Hepaticae of South Greenland. Beih. Nova Hedwigia 92 . Váňa, J. and  W. S. Hong. 1999. The genus Jungermannia in western North America. Lindbergia 24:133--144.

1. Plants dioicous, leaves with distinct rim of swollen cells, far eastern portion of flora … 1. Plectocolea crenuliformis

1. Plants paroicous or dioicous, leaves without distinct rim of swollen cells rim, although marginal cells may be more intensively colored.

2. Plants paroicous.

3. Plants pellucid, merely soft, light green to yellowish green, leaves obliquely oriented, leaf margin (especially in distal part of plant) frequently undulate. Perigynium low, ca. 1/3 times the perianth length or less; rhizoids purplish … 2. Plectocolea fossombronioides

3. Plants not pellucid, merely rigid, deep green to brownish green and (when small) almost constantly with marked traces of purple coloration; leaves not undulate, more or less vertically oriented at least in normally developed shoots. Perigynium more than 1-- 1.5 times perianth length; rhizoids purple to brownish.

4. Plants large, wider than 2.2 mm, rhizoids purple, leaves vertically oriented … 4. Plectocolea obovata

4. Plants small, less than 2 mm in width, rhizoids purplish to brownish, leaves vertically to obliquely oriented … 7. Plectocolea subelliptica

2. Plants dioicous.

5. Plants pale green to pale yellowish brownish to brownish-green, leaves in larger plants frequently undulate at margin, shoots mostly less than 2 mm in width, perigynium ca. 1/2 of perianth length, rhizoids pale purplish to almost colorless and brownish … 3. Plectocolea hyalina

5. Plants deep to dark green, brown green, purplish brown and blackish green, leaves not or slightly undulate at margin (in plants larger than 3 mm in width), shoots mostly wider than 2 mm, perigynium the same size or longer than perianth, rhizoids purple.

6. Plants 1.4--2.1 mm wide, perigynium from 1--1.5 times longer than perianth, oil-bodies granulate  … 5. Plectocolea obscura

6. Plants 2.1--4.8 mm wide, perigynium 2 times longer than perianth, oil-bodies in the midleaf commonly botryoidal … 6. Plectocolea schusteriana

1. Plectocolea crenuliformis (Austin) Mitten, Trans. Linn. Soc. Bot. Ser. 2, 3: 198. 1891 E

Jungermannia crenuliformis Austin, Bull. Torrey Bot. Club 3: 10 1872; Solenostoma crenuliforme (Austin) Stephani

Plants 3--15(--20) x 1.1--2.5 mm, prostrate to ascending, yellowish brown, brownish to brown reddish, with characteristically more deeply colored leaf margin, rarely in shady places pale greenish to whitish and very soft and gentle; female plants larger than the sterile. Stem 105--360(--385) \um in width and 80--250(--305) \um in height, branching lateral intercalary or ventral, the latter rare; dorsal surface cells 63--175 x (12--)15--25 \um, linear, rectangular to obliquely rectangular, walls thin to very thick and brownish (ca. 2.5--3.5 \um in thickness), trigones indistinct, cuticle loosely striolate; in the stem cross section there are at least four variants of structure: (1) three-layered with outer cells (hyaloderm) ca. 17--30 \um in diameter, thin-walled, walls colorless, middle stratum (scleroderm), cells ca. 12--15 \um in diameter, thick-walled, walls yellowish, inner cells ca. 12--23 \um in diameter, slightly thick-walled to thin-walled; (2) in the stem cross section there are 2 strata: outer with slightly thick-walled cells, ca. 25--27 \um in diameter, inner with cell wall thin to slightly thickened, cells mostly hexagonal, ca. 25--37 \um in diameter, cell walls colorless; (3) no distinct differentiation in the stem cross section, walls become thicker to dorsal side, outer cells larger, ca. 25--33 in diameter, inner smaller, ca. 20--25 in diameter, becoming smaller near ventral side; (4) outer cells smaller than inner, ca. 17--23 \um in diameter, thick-walled, walls brownish, but in ventral side outer cells rather thin-walled, walls purplish, inner cells slightly thick-walled, walls flexuous, ca. 25--35 \um in diameter, trigones indistinct to small concave. Rhizoids red, red-brown to purple, blackish brown and brown, sparse to numerous and dense in indistinct fascicles, decurrent along the stem or erect. Leaves distant, inserted at angle 15--45º with stem axis, dorsally decurrent for 1/4 of stem width, ventrally subtransversely inserted, insertion line loosely arcuate to straight, 775--1185 x 775--1400 \um (0.95--1.3:1), widest at the middle or slightly below, flattened to loosely concave and concave-channeled, obliquely lingulate to obliquely ovate, oval, transversely oval, reniform and suborbicular; cells at mid leaf 30--45(--50) x 20--38(-45) \um, subisodiametric, mostly 5--6-gonal, lumen mostly rounded, walls colorless to yellowish, slightly thickened, trigones moderate in size, mostly concave, more rarely triangular and large in size and then convex; cells along margin (15--)28--45(--50) \um, very thick-walled to thin-walled (in shady phases), but external wall almost invariantly thickened, walls colorless, yellowish to brownish and yellowish brownish with more deeply colored external wall, cuticle smooth to verrucose near ventral leaf base or rarely everywhere. Oil-bodies (1--)2--3(--5) per cell, ellipsoidal to shortly fusiform, 6--9 x 3--6 \um. Sexual condition dioicous. Perianth terminal on main axis, conical, in distal part pluriplicate, but with 3--5 main plicae, gradually narrowed to the mouth, ca. 700--1050 x 700 \um, immersed to exerted up 1/4, mouth crenulate, composed of more or less short (ca. 1.5 as long as wide) cells, cells at mid leaf 25--43 x 15--20 \um, rectangular to oblong rectangular, strongly thick-walled, walls pink-rose to purplish, cuticle smooth. Perigynium 1/2--3/4 of perianth length; bracts just below perianth, in 1--2 pair, similar to sterile leaves, but longer, ca. 1050--1225 x 1070--1190 \um, oval to oblong-lingulate, sometimes (pairs developed on the perigynium) shortly retuse at apex, widest at middle or in the proximal 1/3 of the bract, adherent to perianth and perigynium at the base and channeled and deflexed away from the perianth in distal half. Androecia intercalary in 3--4 pairs of bracts (frequently androecial branch dying distally), with 2 antheridia per bract, stalk 2-seriate, ca. 50 x 22 \um; bracts with slightly, but distinctly developed border rim. Sporophytes unknown.

On circum-neutral to acidic rocks, sandstone, along shaded rocky streams, or in disturbed areas in forested lowlands; 0--550 m; Ont., Que.; Ark., Conn., Ga., Ill., Ind., Kans., Ky., Mich., N.J., N.Y., N.C., Ohio, Pa., S.C., Tenn., W.Va., Wis. Endemic.

2. Plectocolea fossombronioides (Austin) Mitten, Trans. Linn. Soc. London, Bot., ser 2. 3: 198 1891  F

Jungermannia fossombronioides Austin, Proc. Acad. Nat. Sci. Philadelphia 1869: 220. 1869;  Solenostoma fossombronioides (Austin) R. M. Schuster

Plants very soft, 3--6 x 0.7--1 mm, prostrate to ascending (near apex of fertile shoot), pale brownish, whitish, yellowish brown; fertile plants slightly larger than sterile. Stem 105--175 \um in width and 100--165 \um in height, dorsal surface cells ca. 70--87 x 25--30 \um, thin-walled or slightly thickened, trigones indistinct; stem cross section composed of mostly thin-walled cells, mostly penta- to hexagonal in cross section, ca. 20--38 \um in diameter, smaller near ventral side, walls colorless to brownish, trigones absent or barely visible. Rhizoids dense, brown-red to (more rare) purplish and pale brownish, rigid, separated one from other, or sometimes in indistinct fascicles, originated at 90º with the stem. Leaves distant to contiguous, inserted at angle 15--20º with stem, dorsally shortly decurrent (not more than 1/5 of stem width), ventrally inserted at angle 50--60º with axis, not decurrent; 600--700 x 490--600 \um (0.8--0.9:1), lingulate to ovate, widest between 1/2 and 1/3 of leaf length, flattened to obscurely concave, the biggest leaves frequently undulate at margin; cells at mid leaf thin-walled, 38--60 x 35--43 \um, walls brownish to colorless, trigones small, concave; along margin 25--35 \um, slightly elongated along margin, walls thin, colorless; cuticle smooth everywhere. Oil-bodies 2--5(--8) per cell, 6--8 x 8--22 \um. Sexual condition paroicous, but occasionally with some branches androecious only (heteroicous). Perianth terminal on main axis, no innovations, emergent for 1/3--2/3 of its length, tubular, pentaplicatae when mature (1-dorsal, 2-lateral, 2-ventral) or pluriplicate, obscurely tightened to the mouth, perianth composed by uniformly elongated cells, ca. 70--85 x 25--33 \um, walls thin, trigones small to indistinct, cuticle smooth. Perigynium low, to 1/3 of the perianth length, with 2 pairs of bracts; bracts just below perianth, in 2 pair, the same shape as male bracts. Androecia just below female bracts (or, occasionally only in separate branches, in heteroicous plants), in 2--3(--4) pairs of bracts (if in separate branches then 3--4 pairs of bracts), with 1--2 antheridia per bract, antheridial body shortly elliptical to spherical, ca. 120--170 x 120--170 \um, stalk 2-seriate, ca. 38 x 25 \um; bracts strongly inflated near the base with margin (especially the pair nearest to the perichaetium) deflexed away of the perianth. Seta ca. 150 \um in diameter. Capsule shortly ellipsoidal, ca. 700 x 630 \um, 2-stratose, inner cells, long-rectangular 50--100 x 7.5--13 \um, with 5--8 annular thickenings, outer cells subquadrate to rectangular, ca. 33--55 x 40--43 \um with 2--3 nodular thickenings in vertical wall, and 0--2 on the horizontal one. Elaters 2-spiral, ca. 165 x 7.5 \um, with homogenous endings, ca. 30 \um length, spores brown to brown reddish, spherical, finely papillose, 15--16 \um in diameter.

Soil-covered rocks along streams, occasionally pioneer on sedimentary or slightly metamorphosed rocks, temperate species distributed in forested areas; 0--900; Conn., D.C., Ga., Ill., Ind., Kans., Minn., Miss., N.J., N.C., Ohio, S.C., W.Va.

The capsule of Plectocolea fossombronioides matures in mid April.

3. Plectocolea hyalina (Lyell) Mitten, Trans. Linn. Soc. London, Bot. Ser. 2, 3: 198. 1891

Jungermannia hyalina Lyell, Brit. Jungermann. Pl. 63. 1814; Solenostoma hyalinum (Lyell) Mitten

Plants 3--8 x 0.7--1.4 mm (--1.75 in perianthous shoots), prostrate, loosely adhering to the substratum, pale green to pale yellowish brownish to brownish green; female plants larger for 1.5 times than male and sterile. Stem 140--210 \um in width and 100--170 \um in height, branching rarely occur ventral; in the stem cross section outer cells ca. 17--25 \um in diameter, mostly 5--6-gonal, walls thin-walled, inner cells ca. 23--30 \um in diameter, mostly 5--6-gonal, thin-walled, trigones indistinct. Rhizoids dense to isolated, brownish to reddish brown, at angle 90º with stem, often in dense fascicles (mostly in weak and male branches) or loosely decurrent down the stem. Leaves distant, inserted at angle of 10--20° with axis, dorsally barely decurrent, ventrally subtransversely inserted, sometimes insertion line arcuate, not decurrent; 400--975 x 400--1175 \um (1:0.85--1.1), transversely oval to widely lingulate, ovate and reniform, the biggest leaves (near the shoot apex) are frequently retuse and undulate, down the stem leaves become ovoid to widely ovate, widest in the middle in larger leaves or at proximal 1/3 in smaller, mostly flattened to (near apex) loosely concave-channeled; cells at mid leaf thin-walled, 33--50 x 28--50 \um, subisodiametric, trigones indistinct or very small and concave to convex, walls colorless; along margin 20--43 \um, thin-walled, trigones small concave to convex and bulging, sometimes with intermediate thickening in tangential wall, external wall thickened, sometimes loosely; cuticle smooth with the exception of the proximal third, where it is obscurely papillose. Oil-bodies 2--5(--10) per cell, 8--15(--22) x (4--)6--8 \um, finely granulate. Sexual condition dioicous. Perianth terminal on main axis, immersed to emergent for 1/2 of its length, mostly fusiform to fusiform-conical, pluriplicate and turbinate to the mouth, ca. 1000--1225 x 500--600 \um. Perigynium ca. 1/2 of perianth length. Androecia with bracts in 3--6 series, reniform to subrotund, rounded at the apex, with 1--3 antheridia per bract. Capsule ovoid. Elaters 2-spiral, 9 \um wide. Spores brownish, 14-17 \um in diameter.

On wet soil or soil covered rocks, sometimes among mosses in wet mossy tundras; 0--2500; Greenland; Alta., B.C., Ont., Que., Ala.; Alaska, Ariz., Calif., Colo., Conn., D.C., Ga., Ill., Ind., Iowa, Kans., Ky., Maine, Mass., Mich., Minn., Miss., Mont., N.H., N.Mex., N.Y., N.C., Ohio, Oreg., Pa., S.C., Va., Wash., Wis., Wyo.; Eurasia, Africa.

4. Plectocolea obovata (Nees) Mitten Fl. Vit., 405 1873

Jungermannia obovata Nees Naturgesch. Eur. Leberm., 1: 332 1833

Solenostoma obovatum (Nees) R.M. Schuster Hep. Anth. N. Amer., 2: 1007 1969

Plants 4--15 mm in length and 2.2--2.5 mm width, ascending, dirty yellowish green, yellowish greenish, dirty yellowish brown. Stem 240--290 \um in width and 120--180 \um in height, branching rarely occur lateral or ventral; dorsal surface cells 63--125 x 20--23 \um, long rectangular, obliquely rectangular to linear, walls slightly thickened, trigones indistinct to concave and small; in the stem cross section outer cells slightly larger than inner ones, ca. 20--25 \um in diameter, walls slightly thickened, inner ca. 12--23 \um in diameter, mostly 5--6-gonal, trigones indistinct. Rhizoids deep purple, sparse, decurrent down the stem. Leaves distant to contiguous, inserted at angle 15--20° with axis, dorsally clearly decurrent up 1/3 of the stem width, ventrally inserted subtransversely, insertion line arched, not decurrent; 1050--1225 x 1050--1120 \um (1: 0.9--1.15), lingulate to trapezoidal, sometimes shortly 2--lobed or retuse at apex, flattened to concave; cells at mid leaf 20--25 x 15--22 \um, mostly 4--5--angular, walls colorless, slightly thickened, trigones distinct, mostly concave, rarely triangular; along margin 15--18 \um, thin-walled, walls yellowish to yellowish brownish, trigones triangle to slightly convex, cuticle smooth. Oil-bodies 2--5(-6) per cell, sphaerical 4--8(-10) \um in diameter to ellipsoidal 6--15(-20)x4--8(-10) \um. Sexual condition paroicous. Perianth terminal, emergent from bracts for 1/4 of its length, conical, pluriplicate, sometimes turbinate, ca. 750 x 750 \um, perigynium 3/2--4/2  equal to perianth length, bracts the same size with antheridial bracts, adherent to perigynium in the base, apex deflexed. Androecia below perianths, commonly divided from the latter by 1--3 pairs of sterile leaves, in 3--4 pairs of bracts, ca. 900--1000 x 600--1080 \um, trapezoidal to ovate, with 1--2 antheridia per bract, body ellipsoidal 170--200 x 140--180 \um, brown, stalk 2-seriate, ca. 100--140 x 15--20 \um. Seta ca 150 \um in diameter and 9000 \um length. Capsule 500--1000 \um in length, ellipsoidal, capsule wall 2-stratose, outer cell subquadrate to rectangular, 20--40 x 28--35 \um with 2 nodular thickenings in each vertical wall, inner cells irregularly linear, 30--90 x 8--20 \um with 6--11 annular thickenings. Elaters ca. 150 x 12, 2-spiral, with narrowed homogenous ends. Spores finely papillose, spherical, 20--22 \um in diameter.

On shaded wet rocks and soil near streams, rarely submerged, mostly in tundra zone or belt; 0--2000 m; Greenland; Alta., B.C., Nfld. and Labr. (Nfld.), Que.; Alaska, Calif., Maine, Mont., N.H., Oreg., Vt., Wash.; Eurasia.

5. Plectocolea obscura (A. Evans) A. Evans in Buch, A. Evans & Verdoorn, Ann. Bryol. 10: 42. 1938

Nardia obscura A. Evans, Rhodora 21: 159. 1919; Jungermannia evansii Váňa

Plants 4--8 x 1.4--2.1 mm, prostrate to ascending, green brownish, with light purplish traces near apices; female plants larger. Stem 200--400 \um in width and 120--300 \um in height, dorsal surface cells long-rectangular to obliquely rectangular, ca. 75--150 x 15--23 \um, mostly thin-walled with indistinct trigones; in the stem cross section there are 3 strata: outer (hyaloderm) with thin external wall and moderately thickened other walls, ca. 12--25 \um in diameter, medium (scleroderm) composed by thick-walled cells, walls yellowish in color, ca. 12--20 \um in diameter, inner with cell wall thin to moderately thickened, cells mostly polygonal, ca. 12--22 \um in diameter, cell walls colorless. Rhizoids purple, rigid, in a bundle or separated one from other, angled at 90º with stem. Leaves distant to contiguous, inserted at angle of 15--30° with axis, dorsally shortly, but distinctly decurrent; 1050--1225 x 980--1190 \um (0.85--0.95:1), oval to ovate, widest at midleaf or slightly below, with rounded to retuse apex, adherent to the stem in the base and flattened and erect above; cells at mid leaf thin-walled, subisodiametric 28--38 x 25--33 \um, trigones convex; along margin 15--23 \um, thin-walled with convex trigones; cuticle distinctly papillose in the proximal third of the leaf, in distal parts papillae very obscure or absent. Oil-bodies 2--5(--10) per cell, ovoid to ellipsoidal, 5--9 x 6--14 \um. Sexual condition dioicous. Perianth terminal on main axis, immersed, conical loosely pluriplicate with 3--4 main plicae those are undulate to the mouth, ca. 700 x 600 \um, perianth mouth crenulate, perianth in distal part 1-stratose, cells mostly rectangular, 25--43 x 17--18 \um, walls thin, trigones small, cuticle smooth or finely papillose, perianth in proximal part 2-stratose, cells mostly oblong rectangular, 38--50 x 10--25 \um, walls thin to thickened, trigones small and concave, cuticle distinctly striolate papillose. Perigynium ca. 1/1--3/2 of the perianth length, with 1--1.5 pairs of bracts; bracts just below perianth, in 1--2 pairs, lingulate, adherent to perianth and perigynium at the base and deflexed away the perianth in distal half, 1470--1750 x 1015--1470 (0.7--0.9:1). Androecia intercalary in 2--3 pairs of bracts, with 1--3 antheridia per bract, stalk 2-seriate, ca. 50--63 x 15--18 \um; bracts strongly inflated in base, but deflexed away the stem in distal part. Sporophytes not seen.

On wet acidic rocks near streams, largely a montane in subalpine habitats; 0--2000 m; Greenland; B.C.; Alaska, Ga., Maine, Mass., N.H., N.Y., N.C., Oreg., Tenn., Vt., Wash.; e Asia.

The sporophyte of Plectocolea obscura is rare, and was described by R. H. Schuster (1969: 1015).

6. Plectocolea schusteriana (J. D. Godfrey & G. Godfrey) Bakalin, Bot. Pacif. 3: 92. 2014 E

Jungermannia schusterana J. D. Godfrey & G. Godfrey, J. Hattori Bot. Lab. 46: 109, fig. 1--3. 1979; Solenostoma schusterianum (J. D. Godfrey & G. Godfrey) Váňa, Hentschel & J. Heinrichs

Plants 20--150 x 2.1--5 mm, ascending to erect, deep green, brownish deep green and green brown and green-purple (due to purple colored ventral bases of leaves); female plants slightly larger than male and sterile and have mainly retuse leaves (even far down of perianth). Stem 280--525 \um in width and 220--400 \um in height, dorsal surface cells thin-walled, mostly elongate-rectangular to obliquely rectangular and linear, 70--125 x 15--20 \um, trigones small and concave, cuticle distinctly striolate; in the stem cross section there are 3 strata: outer (hyaloderm) moderately thick-walled to almost thin-walled, ca. 12--25 \um in diameter, walls brown (external wall more deeply) to purplish and purple colored, medium (scleroderm) very thick-walled with commonly visible median lamina of cell wall, ca. 17--25 \um in diameter, inner thin-walled to moderately thick-walled, ca. 17--38 \um in diameter, with indistinct trigones and walls mostly colorless. Rhizoids uncommon in male and sterile shoots, but more or more dense on female, purple to brown-purple, in the bundle or separated one from other or in indistinct fascicles. Leaves distant to contiguous, inserted at angle of 50° with axis, dorsally decurrent for 1/2--2/2 of the stem width, ventrally subtransversely inserted, line insertion arcuate, decurrent for 1/2--2/2 of stem width; (875--)1100--2400 x (830--)1000--2000 \um (0.6--0.9(--15):1), oval to ovate-oval, obliquely ovate, widest slightly between 1--2/1/3 of proximal part of leaf, with rounded to acuminate or retuse apex, adherent to the stem in the base, but channeled and deflexed or erect in distal 2/3 of leaf; cells at mid leaf thin-walled, subisodiametric to oblong 35--50(-58) x 33--43 \um, trigones large and bulging; along margin 20--38 \um, with slightly to obviously thickened external wall; cuticle finely striolate everywhere but more markedly in proximal part or sometimes loosely developed. Oil-bodies of the grape-cluster type, consisting of distinctly glistening, homogenous globules, (1--)2--6(--8) per cell., spheric (5--)6--15 \um in diameter to ovoid 6--14 x 7--19 \um. Sexual condition dioicous. Perianth terminal on main axis, immersed to emergent for 1/3 of its length, conical pluriplicate and turbinate to the mouth, ca. 490 x 490 \um, perianth in distal part 1-stratose, cells rectangular to obliquely rectangular, 38--75 x 10--13 \um, walls thickened, yellowish colored, perianth in proximal part 2-stratose, cells linear to oblong rectangular, 125--200 x 20--25 \um, walls moderately thickened. Perigynium rhizogenous ventrally, 2 times longer than perianth; bracts just below perianth, in 2 pairs (due to high and robust perigynium), ovate shortly 2-lobed, 1250--1400 x 1250--1300 (0.95--1.1:1). Androecia intercalary in 8--13 pairs of bracts, with 1--2 antheridia per bract, stalk 2-seriate, ca. 63 x 18 \um; bracts slightly inflated in base, but flattened and deflexed away the stem above, purplish to deep-green purple in inflated area, more or less similar in shape to sterile leaves, but complicated on a micro-slide due to inflation in the base. Capsule with outer cells 30--58 x 22--34 \um, each cell with 2--4 nodular thickenings on each vertical wall and with 0--2 on horizontal wall; inner cells 40--110 x 10--22 \um. Elaters 8--12 \um in width, 2-spiral. Spores reddish to golden-brown, 14--18 \um in diameter.

On wet rocks in streams, mostly in coniferous forest and crooked forests in oceanic regions; 0--1200 m; B.C.; Alaska, Wash. Endemic.

7. Plectocolea subelliptica (Lindberg ex Heeg) A. Evans in H. Buch, A. Evans & Verdoorn, Ann. Bryol., 10: 42. 1938

Nardia subelliptica Lindberg ex Heeg, Verh

. K.K. Zool.-Bot. Ges. Wien

 43(Abh. 1): 69. 1893; Jungermannia subelliptica (Lindberg ex Heeg) Levier; Plectocolea obovata var. minor (Carrington) Schljakov; Solenostoma subellipticum (Lindberg ex Heeg) R.M. Schuster

Plants 3--5 x 0.6--1 mm, erect to (rarely) ascending, brownish green, yellowish brown, sepia-brown; fertile plants larger than the sterile. Stem 210--280 \um in width and 200--270 \um in height; dorsal surface cells 43--75 x 10--13 \um, long-rectangular to linear, cuticle distinctly striolate-papillose; in the stem cross section thick-walled, with the exception of external wall of outer layer(!), walls brownish to colorless in inner portion, trigones small, concave, outer cells ca. 30--40 \um in diameter. Rhizoids reddish to purplish and brown to nearly colorless. Leaves inserted at 40--80° with axis, dorsally shortly decurrent ca. 1/5 of the stem width, ventrally inserted subtransversely, not decurrent; 525--735 x 525--875 \um (1: 0.85--1), suborbicular to widely ovate and ovate-lingulate, widest near the middle or at proximal 1/3, distinctly, but not strongly concave-channeled; cells at mid leaf thin-walled, subisodiametric, 22--30 x 22--28 \um, walls colorless, trigones distinct, convex; along margin 20--25 \um, thin-walled, walls colorless to brownish, trigones triangular to convex, cuticle striolate in proximal 1/3 of the leaf. Oil-bodies (2--)3--5(--6) per cell, 4--5(--5.5) x 6--10 \um, granulate. Sexual condition paroicous, some innovations with 2 male generations divided by some pairs of sterile leaves---heteroicous plants). Perianth terminal on main axis, conical, with ca. 8 archegonia per gynoecium. Perigynium 1--1.3 of the perianth length; bracts just below perianth, in 1--2 pairs, ca. 1000 x 950 \um, wider in proximal 1/3, frequently with undulate margin. Androecia below perianth, commonly divided from the latter by 1--3 pairs of sterile leaves, spicate, in 2--3 pairs of bracts, ca. 630--805 x 810--1050 \um, suborbicular to orbicular triangle, wider in proximal 1/3 of the leaf, strongly inflated near the base, purple near the inflated area, bract margin incurved, with 2 antheridia per bract, body grayish to almost colorless, stalk 2-seriate, ca. 87--100 x 15--20 \um. Capsule 500--1000 \um, ellipsoidal, capsule wall 2-stratose, outer cell subquadrate to rectangular, 20--45 x 26--35 \um with 2--3 nodular thickenings in each vertical wall; inner cells irregularly linear, 30--90 x 8--20 \um with 6--18 annular thickenings. Seta 100-- 150 \um in diameter and 1500--9000 \um in length. Spores finely papillose, spheric, 18--24 \um. Elaters 120--150 x 9--12, 2-spiral, with narrowed homogenous ends.

On moist soil and rocks near stream, almost exclusively in mountains tundra or slightly descending to forest belt along streams; 0--600; Greenland; B.C., N.S., Que.; Alaska, Wash.; Eurasia.

3. SOLENOSTOMA Mitten, emend. Zerov, J. Linn. Soc. Bot., 8: 51 1865 * [From Greek solen, canal and stoma, mouth, alluding to perianth suddenly narrowed to mouth]

Vadim Bakalin

Jungermannia subg. Solenostoma (Mitten) Amakawa

Plants 3--20 mm x 0.5--3.4 mm, ascending to erect, rarely creeping, greenish, pale green yellowish, yellowish brownish to dirty green, in the most species with characteristically red or purple coloration (at least as a tint at the tips of perianths and leaf rim). Stem orbicular to transversely oval in cross section 100--520 \um in width and 80--500 \um in height, dorsal surface cells mostly subquadrate, more rarely shortly elongate with walls colorless to slightly brownish, trigones mostly small and concave, cuticle smooth to striolate; in the stem cross section outer cells smaller and with thicker cells. Rhizoids mostly dense, rarely scattered, in bundles, colorless to brownish, rarely reddish or purplish. Leaves contiguous to imbricate, subtransversely to (more rarely) obliquely inserted, dorsally decurrent or not, ventrally insertion line arcuate, not or long-decurrent; cells at midleaf mostly thin-walled, walls colorless to yellowish, never deep brownish or orange-brown, trigones mostly distinct, concave to convex. Sexual condition paroicous or dioicous. Specialized asexual reproduction absent. Perianth terminal on main axis, innovations rare, emergent from bracts for 1/4--3/4 of its length, tubular obtrapezoidal, 1-stratose to base or sometimes 2-stratose in proximal 1/5(--1/3) of the length, mostly contracted to the beaked mouth and distinctly 3--5-keeled (1--2- dorsal, 0--2- lateral, 1--2- ventral), perianth mouth crenulate, composed by elongate cells, cells in mid perianth mostly isodiametric, hexagonal to rectangular, never linear, trigones mostly distinct and convex; perigynium absent, or, if present, short, less than 1/3 of perianth length. Androecia intercalary, in (1--)2--5 pairs of bracts, with 1--3(--4) antheridia per bract, stalk 2-seriate. Capsule spheric to shortly ellipsoidal, 2-stratose. Elaters 2-spiral, ca. 100 x 7.5 \um, 2-spiral, with or without homogenous ends. Spores brown to brown reddish, spheric, finely papillose, 12--19 \um.

Species ca. 70 (6 in the flora): North America, Central America, South America, Eurasia, Africa, Australia, Pacific Islands, Antarctica.

SELECTED REFERENCES

Crum, H. A. 1991. Liverworts and Hornworts of southern Michigan. Ann Arbor, Michigan. Damsholt K. and J. Váňa 1977. The genus Jungermannia L. emend. Dumort. (Hepaticae) in Greenland.  Lindbergia. 4: 1--26. Doyle W.T., Stotler R.E. 2006 Contribution toward a bryoflora of California III. Keys and Annotated species Catalogue for Liverworts and Hornworts. Madrono. 53: 89--197. Hicks, M. L. 1992. Guide to the Liverworts of North Carolina. Durham, North Carolina. Schuster R.M. 1969. The Hepaticae and Anthocerotae of North America. Vol. 2. Columbia University Press., New York. Schuster R. M. 1988. The Hepaticae of South Greenland. Beih. Nova Hedwigia. Heft 92. Váňa J. and W. S. Hong 1999. The genus Jungermannia in western North America. Lindbergia 24: 133--144.

1. Plants paroicous, leaves without distinct rim of swollen cells along margin, midleaf cells mostly with trigones moderate in size.

2. Plants mostly erect, more rarely ascending, green in shady places, to mostly sepia-brown in color, never purplish, rhizoids originated from ventral side of stem only …6. Solenostoma sphaerocarpum

2. Plants ascending to creeping, green in shady places, to brown-red to purplish in color, rhizoids originated from both ventral side of stem and from some cells in lower half of leaves … 1. Solenostoma confertissimum

1. Plants dioicous, leaves with or without distinct rim of swollen cells along margin, midleaf cells commonly with trigones moderate to large in size and bulging, or trigones indistinct.

3. Plants lax, large (mostly more than 2 mm wide), with undulate leaf margins, leaves turned to dorsal side of the branch, sheathing the stem, blackish to reddish black, but always with the stem whitish, plants mostly erect in dense patches … 2. Solenostoma fusiforme

3. Plants not lax, small (mostly less than 2 mm wide), leaf margin not undulate, leaves horizontally spreading, not sheathing the stem or sheathing only near the leaf base, not blackish, commonly reddish to red brown, the stem not whitish, plants creeping to loosely ascending.

4. Leaves bordered with swollen cells with equally thickened cell walls, size of marginal cells at least two times larger than midleaf cells; rarely leaves not bordered, but in the latter case trigones in the leaf are not developed … 3. Solenostoma gracillimum

4. Leaves not bordered with a rim of swollen subequally thickened cell walls, size of marginal cells nearly the same or only slightly larger than midleaf cells.

5. Leaves bordered with rim of strongly unequally thick-walled cells, with large, sometimes semi-confluent trigones, size of marginal and midleaf cells nearly the same …5. Solenostoma rubrum

5. Leaves not bordered, cell walls evenly thin, with moderate to large convex trigones … 4. Solenostoma appalachianum

1. Solenostoma confertissimum (Nees) Schljakov, Pechen. Mkhi Severa SSSR 4: 5.1 1981

Jungermannia confertissima Nees, Naturgesch. Eur. Leberm. 1: 277, 291. 1833; J. pyriflora subsp. purpurea (R.M. Schuster & Damsholt) Stotler & Crandall-Stotler; J. pyriflora subsp. purpurea var. innovata (R.M. Schuster & Damsholt) Stotler & Crandal-Stotler; Solenostoma levieri (Stephani) Stephani; S. pyriflorum var. innovatum R. M. Schuster & Damsholt; S. pyriflorum subsp. purpureum R. M. Schuster & Damsholt

Plants 3--8 mm x 1--1.7 (up to 2.4 near perianth) mm, ascending, greenish, pale green yellowish, yellowish brownish to dirty green almost constantly with purplish tint or deeply purple colored apex of shoots and marginal part of leaves; fertile plants slightly larger than sterile. Stem 245--520 \um in width and 240--500 \um in height, branching lateral, dorsal surface cells ca. 30--50 x 25--30 \um thin-walled, walls colorless, trigones small and concave, cuticle smooth; in stem cross section outer cells smaller than inner ones, with thin to slightly thickened walls, ca. 22--25 \um in diameter, walls colorless to brownish and brown (especially ventrally), inner thin-walled, ca. 20--43 \um in diameter, walls colorless, trigones indistinct to small and concave. Rhizoids very dense, mostly in bundles, colorless to brownish. Leaves contiguous to subimbricate, subtransversely inserted, dorsally not or barely decurrent, ventrally not decurrent; 525--735 x 805--1050 \um (1:0.7--0.8), transversely oval to reniform, widest at middle, concave to (more frequently) channeled; cells in the midleaf thin-walled, 28--40 x 22--28 \um, walls colorless to yellowish, trigones distinct, concave to slightly convex; cells along margin 12--20 \um, with thickened external walls and other walls thin, colorless to yellowish, trigones large. Sexual condition paroicous. Perianth terminal on main axis, rarely with subfloral innovations, emergent for 1/2--1/3 of its length, tubular obtrapezoidal, suddenly tightened to the mouth, distinctly 5-keeled (1- dorsal, 2- lateral, 2- ventral), ca. 1470--1575 x 665--700 \um, perianth mouth crenulate, composed of elongate cells, perianth distally 1-stratose, cells isodiametric, mostly hexagonal, 20--30 \um in diameter, trigones distinct, convex, perianth basally 2-stratose (for 1/3 of perianth length), cells obliquely rectangular to hexagonal, 25--45 x 17--21 \um, walls thin, trigones small and concave to indistinct; perigynium indistinct; bracts just proximal to perianth in 1 pair, the same size or slightly larger as male bracts. Androecia just proximal to female bracts, in (1--)2 pairs of bracts, with 2(--3) antheridia per bract, stalk 2-seriate, ca. 38--40 x 15--18 \um; bracts similar with sterile leaves, transversely oval, widest in the middle, obscurely inflated near the base. Seta ca. 200 \um in width. Capsule spheric to shortly ellipsoidal, ca. 700 x 700 \um, 2-stratose, inner cells, obliquely rectangular to irregular in shape 38--63 x 12--18 \um, with 5--8 annular thickenings, outer cells quadrate to shortly rectangular, ca. 30--43 x 22--35 \um with 2--3 nodular thickenings in vertical wall, and 0--2 on horizontal wall. Elaters 2-spiral, ca. 100 x 7.5 \um, 2-spiral, with homogenous endings, ca. 15--25 \um in length. Spores brown to brown reddish, spheric, finely papillose, 15--17 \um.

On moist soil and wet rocks near streams frequently associated with mosses and other hepatics; 0--1100 m or higher; Greenland; B.C., Que. ; Alaska, Calif., Mich., Nev., Wis.; Eurasia.

2. Solenostoma fusiforme (Stephani) R. M. Schuster, Hepat. Anth. N. Amer. 2: 944. 1969

Nardia fusiformis Stephani, Bull. Herb. Boiss., 5(2): 99. 1897; Jungermannia fusiformis (Stephani) Stephani; J. koreana (Stephani) Amakawa; Plectocolea koreana (Stephani) S. Hattori; Solenostoma koreanum Stephani

Plants 15--25 x 1--2.8 mm, ascending to erect in dense patches, pale green to whitish, with characteristically brownish to reddish brown colored leaf margins. Stem 315--420 x 280--390 \um, oval in cross section, dorsal surface cells thin- to slightly thick-walled, cells obliquely rectangular to oblong-hexagonal, with small concave or indistinct trigones, 100--125 x 17--33 \um. Rhizoids dense, colorless to brownish, in indistinct fascicles at 90º with stem; in the stem cross section outer layer 2-stratose mostly composed by moderately thick-walled cells, ca. 12--25 \um in diameter, inner cells thin-walled with small to indistinct trigones, in central part nearly the same size with outer layer, but between center and outer layer larger up to 45 \um in diameter. Leaves distant to contiguous, inserted at angle of 10--40° with axis, dorsally decurrent for ca. stem width, ventrally subtransversely inserted, not or barely decurrent, (700--)900--1750 x (665--)1120--2135 \um (1:0.6--0.95(--1.2)), transversely oval to rounded- widely triangular, mostly flattened to slightly concave, moderately and large-sized frequently undulate at margin; mostly with distinct brown to reddish brown and purplish colored rim along margin; cells in the midleaf thin-walled or slightly thickened, colorless, subisodiametric to oblong hexagonal, 30--45 x 25--38 \um, trigones small and concave; near margin 28--33 \um, with mostly thickened, but not strongly, or (rarely) thin-walled but with thickened external wall, brownish to reddish brown or, occasionally, colorless; in basal part cells oblong rectangular to oblong hexagonal, 50--100 x 30--43 \um; cuticle smooth or very loosely striolate. Sexual condition dioicous. Sporophyte unknown for the flora region.

On wet rocks and soil near sluggishly flowing streams, more rarely in bogs; 0--500 m; B.C., Colo., Oreg., Wash.; East Asia.

3. Solenostoma gracillimum (Smith) R. M. Schuster, Hp. Anthocerotae N. Amer. 2: 972. 1969

Jungermannia gracillima Smith, Engl. Bot. 32: tab. 2238. 1811; J. crenulata Smith non J. crenulata Schmidel; Solenostoma crenulatum (Smith) Mitten

Plants 3--10 x 0.4--0.9 (fertile up to 1 mm) mm, prostrate to (fertile) ascending, yellowish brownish to pale green to pale yellowish and brown; female plants yellow red to brownish purplish. Stem (100--)175--245 \um in width and (90--)150--220 \um in height, branching lateral as normal branch or slender innovations; dorsal surface cells 35--75 x 17--25 \um, rectangular to oblong hexagonal and obliquely rectangular, walls thin; cells in the stem cross section nearly the same size in outer and inner layers, but outer with cell walls thickened with yellow to brownish colored walls, inner cells more or less thin-walled, with walls colorless, ca. 15--20 \um in diameter. Rhizoids sparse, colorless, in indistinct fascicles, tawny or decurrent down the stem. Leaves distant to subimbricate (especially in area 5--7 pairs below the perianth), inserted at angle (10--)15--25º, although sometimes to subtransverse in leaves proximal to female bracts; (190--)470--900 x (200--)500--940 \um (1: (0.7--)0.95), orbicular to rounded-triangular, lingulate and reniform, widest at proximal 1/3 of leaf length, more rarely near the middle; cells in the midleaf (22--)30--45 x (17--)25--28 \um, thin-walled, subisodiametric, mostly 5--6-gonal, trigones indistinct to small; along margin (25--)30--70 \um, very thick-walled or not differs from midleaf cells (then leptodermous, without trigones to slightly equally thickened), walls brownish, cuticle striolate at proximal 1/2 of the leaf. Sexual condition dioicous. Perianth terminal on main axis, no innovations, emergent for 1/3--1/2 of its length, conical, tubular-conical, ellipsoidal to short-clavate, with the suddenly tightened mouth 4--5-plicae (1--2- dorsal, 2- lateral, 1- ventral), perianth mouth crenulate, composed elongate thick-walled cells, slightly dilated on the external end, perianth in the middle part rectangular to isodiametric, ca. 30--45 x 18--25 \um, cuticle distinctly striolate in lower half; perigynium ca. 1/4 of the perianth length or smaller; bracts not differentiable from sterile leaves, slightly larger only, in 1 pair, ca. 950 x 950--1070 \um. Androecia intercalary, spicate, in 2--3 pairs of bracts, strongly inflated in the base, with incurved margin of bracts, with 1 antheridium per bract, body brownish, shortly ellipsoidal ca. 125 x 112 \um, stalk 2-seriate, ca. 33 x 17 \um. Seta ca. 3000--5000 x 125 \um. Capsule shortly ellipsoidal, 2-stratose, inner cells long rectangular 38--63 x 7--13 \um, with 8--10 annular thickenings, outer cells subquadrate, ca. 25--30 x 17--35 \um with 1--2 nodular thickenings in both vertical and horizontal walls, elaters 2-spiral, ca. 100 x 7.5 \um, with homogenous endings ca. 10 \um length, spores red-brown, spheric, finely papillose, 10--11 \um in diameter.  Elaters 600--800 x 6-7 \um. Spores 10--12 \um, red brown, papillose. 

On fine-grained soil along streams and (more frequently) in human disturbed areas; 0--500; N.B., Nfld. and Labr. (Nfld.), N.S., Ont., P.E.I., Que.; Ala., Ark., Calif., Conn., D.C., Fla., Ga., Ill., Ind., Iowa, Ky., Maine, Mich., Minn., Miss., N.H., N.J., N.Y., N.C., Ohio, Pa., R.I., S.C., Tenn., Vt., Va., W.Va., Tex.; Central America, Eurasia.

There are two forms of Solenostoma gracillimum in the flora area. The fo. crenulatum (Mitten) Bakalin is distinguished from the widespread fo. gracillimum by leaves with the rim of swollen thick-walled cells, 1.5--2 times larger than midleaf cells and is known only from Arkansas, Kentucky, Texas, and distribution in Eurasia.

4. Solenostoma appalachianum R. M. Schuster ex Bakalin, Arctoa, 23: 127. 2014

Plants 3--5 mm x 0.7--1.4 mm, erect to ascending, bright green, green yellow, yellowish green brownish; female plants the same size as the sterile, but males are narrower. Stem 175--245 \um in width and 160--220 \um in height; dorsal surface cells 30--38 x 15--20 \um, walls thickened, walls brownish sometimes with reddish tint; in the stem cross section outer cells ca. 22--30 \um in diameter, walls thickened, yellowish, trigones large, convex, inner cells 20--30 \um in diameter, less thickened, walls colorless, trigones small. Rhizoids very dense, varying from purple, purplish and brown to nearly colorless, at right angles with stem, in dense fascicles, or decurrent down the stem. Leaves contiguous to imbricate, inserted at angle 45--50º with axis, dorsally shortly decurrent (not more than 1/3 of the stem width), ventrally inserted subtransversely, insertion line arcuate, decurrent for 1/5 of stem width; 500--700 x 650--1050 \um (1: 0.7--0.9), orbicular, widest near the middle, concave; cells in the midleaf thin-walled or loosely thickened, subisodiametric, mostly 4--6-gonal, 22--30 x 20--25 \um, walls colorless, trigones distinct, triangle to slightly convex, lumen rounded; along margin 12--18 \um, thin-walled to slightly thickened, walls colorless, cuticle smooth throughout. Sexual condition dioicous. Perianth terminal on main axis, innovations rare, emergent for 1/3--1/2 of its length, 1120--1190 x 560--630 \um, oblong to shortly fusiform, with 4--5 distinct plicae (1--2- dorsal, 2- lateral, 1- ventral), suddenly tightened to the beaked mouth, perianth mouth crenulate to denticulate, perianth in upper part 1-stratose, cells isodiametric, 15--20 \um in diameter, trigones convex, cuticle smooth, in the middle part 2-stratose, oblong to isodiametric, ca. 30--40 x 17--23 \um, walls thin, rarely with intermediate thickenings, cuticle smooth, in proximal 1/4 2-stratose, cells rectangular to oblong-rectangular, 33--45 x 15--20 \um, thin-walled, trigones concave, cuticle papillose; perigynium indistinct; bracts just proximal to perianth in 1 pair, ca. 750 x 1000--1050 \um, orbicular to transversely oval, channeled. Androecia intercalary, but sometimes end of male branch dying, spicate, in 5--8 pairs of bracts, ca. 630--700 x 840--930 \um, orbicular to transversely oval, cupped, but with incurved margin, with 2(--3) antheridia per bract, stalk 2-seriate, ca. 63--68 x 17--23 \um. Sporophyte unknown. 

In moist rock crevices or on rocky surfaces, rarely on bare soil in human disturbed areas; 700--1000 m; Ga., Mich., N.C., S.C., Va.; Asia.

Solenostoma appalachianum has been misidentified in the past under the names Solenostoma pyriflorum or Jungermannia pyriflora.

5. Solenostoma rubrum (Gottsche ex Underwood) R. M. Schuster, Hepat. Anthocerotae N. Amer. 2: 975. 1969   E  F

Jungermannia rubra Gottsche ex Underwood, Bot. Gaz. 13: 113. 1888

Plants 3--8 x 0.7--1.8 mm, prostrate to (mainly) ascending and erect, green, green purplish (due to purple-colored ventral base of leaves) to yellowish pale green, yellowish brown, brownish, brownish reddish, with markedly purple colored perianth beak and tops of ascending erect shoots; female plants the same size with sterile ones or slightly larger, male are wider due to incurved apices of the bracts. Stem 210--370 \um in width and 160--300 \um in height; dorsal surface cells rectangular, ca. 74--100 x 25--38 \um thin-walled trigones indistinct, cuticle striolate-papillose; in the stem cross section outer cells smaller than inner ones, ca. 17--25 \um in diameter, walls slightly thickened (external wall evidently thickened), yellowish, trigones distinct, concave, inner cells 5-gonal to polygonal, ca. 33--50 \um in diameter, thin-walled, walls colorless, trigones small to indistinct. Rhizoids sparse to common, mostly confined to area near ventral base of leaves and commonly absent in distal 3--4 pairs of leaves, colorless to pale grayish and brownish, in indistinct fascicles decurrent down the stem. Leaves distant to contiguous and subimbricate, inserted at angle ca. 45--65º with axis, dorsally decurrent for 1/4 of the stem width, ventrally inserted subtransversely, insertion line arcuate, barely decurrent (in some shoots are inserted at angle ca. 80--85° with the axis and not decurrent both dorsally and ventrally); 630--1050 x 700--1225 \um (1:0.6--1), suborbicular, transversely oval to orbicular-lingulate and orbicular-ovate, widest near the middle, concave to channeled, sometimes deflexed away the stem; cells in the midleaf thin-walled, subisodiametric, mostly 5--6-gonal, 30--45 x 30--60 \um, walls colorless to purplish, trigones distinct, concave to triangle; along margin 20--45 \um, with external wall only clearly thickened (rarely walls equally thickened), sometimes, with visible median lamina in marginal cell walls, trigones distinct, triangle, cuticle smooth. Sexual condition dioicous. Perianth terminal on main axis, no innovations, emergent for 2/3 of its length, 1470--1750 x 985--1050 \um, clavate to shortly fusiform, with 4 distinct plicae (1-dorsal, 2-lateral, 1-ventral), suddenly tightened to the beaked mouth, perianth mouth crenulate, perianth in distal part 1-stratose, cells subisodiametric to oblong, 25--33 x 38--50 \um, trigones distinct to convex, in the middle part subisodiametric, ca. 33--50 x 30--38 \um, walls thin, trigones distinct, triangle to convex, cuticle smooth, in basal part 2-stratose, cells rectangular to oblique rectangular, 100--150 x 28--38 \um, trigones small and concave, cuticle striolate-papillose; perigynium small, less than 1/6 of the perianth length; bracts just below perianth, in 1 pair, ca. 720--900 x 950--100 \um, lingulate to obtrapezoidal, frequently shortly 2-lobate, clinging to perianth in its basal part, but deflexed distally. Androecia intercalary, spicate due to deflexed tops of bracts, in 3--5 pairs of bracts, ca. 630 x 875 \um, orbicular to transversely oval, strongly inflated at the base and cupped, but with incurved margin, with 2 antheridia per bract, antheridial body shortly elliptical to spheric, ca. 110--140 x 95--140 \um or smaller (probably immature), stalk 2-seriate. Seta ca. 3000 \um x 200 \um. Capsule brown, shortly ellipsoidal, ca. 500--625 x 490--500 \um; inner cells 45--60 x 10--15 \um, oblong-rectangular to irregular in shape, with 7--8 annular thickenings; outer cells rectangular, 25--50 x 15--23 \um with 3--4 nodular thickenings in each of vertical wall and 0--1 on horizontal wall. Elaters reddish brown, 2-spiral ca. 112 x 7.5 \um, homogenous narrowed endings from each side ca. 13--15 \um in the length. Spores brown to reddish brown, papillose, ca. 15--16 \um.

On moist soil along stream, clayey roadsides; 0--2160 m; Alta., B.C.; Alaska, Calif., Idaho, Mont., Oreg., Wash., Wyo.

6. Solenostoma sphaerocarpum (Hooker) Stephani, Sp. Hepat. 2: 61. 1901

Jungermannia sphaerocarpa Hooker, Brit. Jungermann., pl. 74. 1815

Plants 3--7 x 0.35--2 mm, erect to ascending, sepia to brownish yellow and grayish brown, without traces of red pigmentation. Stem ca. 175--300 \um in diameter, branching not seen; dorsal surface cells 33--45 x 22--30 \um, subrectangular, cuticle obscurely striolate; in the stem cross section cells ca. 20--25 \um in diameter, walls brownish, mostly thickened, trigones small but distinct, concave. Rhizoids numerous, dirty grayish to almost colorless, in indistinct fascicles, mostly erect, more rarely decurrent. Leaves contiguous to distant, subtransversely inserted, 400--700 x 550--900 \um (1:0.6--0.8), widest in middle or just below, concave to almost flat; cells in midleaf 20--25 x 19--25 \um, lumen rounded to hexagonal, walls thin to obscurely thickened, brownish, trigones large and convex; cuticle striolate in lower 1/3. Oil-bodies 5--6 per cell, finely granulate, brownish-grayish, oval in projection, 5--11 x 4--7 \um. Sexual condition paroicous. Perianth terminal on main axis, innovations lateral, 1--2 per gynoecium, rarely present; emergent for 1/4--2/3 of its length, 1540--1750 x 1540--1540 \um, tubular with the suddenly tightened beaked mouth, with 3--4 distinct plicae (1--2-dorsal, 2-ventral), perianth mouth crenulate, perianth mainly 1-stratose, but 2-stratose in basal portion, in the middle part of perianth cells 28--35 x 17--23 \um, subisodiametric to oblong, mostly hexagonal; perigynium up to 1/3 of the perianth length; bracts almost not differentiated from sterile leaves, only slightly larger, in 1 pair, ca. 750--800 x 1000--1250 \um, concave to flattened, rarely obscurely undulate at margin or slightly retuse. Androecia below perichaetium, in 1--3 pair of bracts, bracts similar to sterile leaves, slightly inflated near the base, with 2--3 antheridia per bract, stalk 2-seriate, ca. 43 x 17 \um. Seta ca. 1500--2000 x 120 \um. Capsule oval ca. 600--700 x 400 \um, inner cells 50--70 x 12--15 \um, frequently with flexuous walls, with 6--8 annular thickenings, outer cells nearly subquadrate, ca. 30 x 28 \um with 2 nodular thickenings in vertical wall. Elaters 2-spiral, ca. 150 x 8 \um. Spores finely papillose, ca. 17 \um.

On moist soil and rocks near streams, on wet fine-grained soil in anthropogenic or natively disturbed areas; 0--3100m; Greenland; Alta., B.C., Nfld. and Labr. (Nfld.), N.W.T., Nunavut, Que., Yukon; Alaska, Calif., Colo., Idaho, Maine, Mich., Minn., Mont., N.H., N.Y., Oreg., Utah, Wash., Wis., Wyo.; Central America, South America, Eurasia, Africa.