BFNA Title: Liochlaena
Author: V. Bakalin
Date:April 4, 2016
Edit Level: R
Version: 1

Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden
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XX. LIOCHLAENA Nees, Syn. Hepat., 150 1844 [Greek, leios, smooth, and chlaina, cover, alluding to perianth not folded distally]

Vadim Bakalin


Plants 10--40 mm x 2--3(--5) mm, mostly prostrate, rarely ascending (perianthous plants), pale green, yellowish green to deep green, blackish in very wet places, rarely tinted orange, loosely adhering to the substrate by tawny rhizoids. Branching intercalary. Rhizoids often dense, colorless to yellowish or brownish, never red or purplish. Leaves contiguous, inserted at 30--40° with stem, but near apex of perianthous plants leaves subtransversely inserted, dorsally shortly but distinctly decurrent, ventrally inserted subtransversely, insertion line arched, not decurrent; leaves obliquely lingulate to oval, frequently retuse in larger leaves, sheathing the stem proximally, erect-flattened distally; midleaf cells with walls thin to slightly thickened, walls yellowish to colorless, trigones moderate in size, mostly concave, rarely slightly convex; cuticle verrucose, sometimes indistinctly so. Specialized asexual reproduction by gemmae, very rare, ellipsoidal to shortly fusiform, 2-celled.  Sexual condition paroicous or dioicous. Androecia below perianth or in separate branches, of 1--13 pairs of bracts, oblique-lingulate, slightly larger than sterile leaves, strongly inflated near the base, antheridia 1--3 per bract. Perianth terminal on main axis, cylindric to clavate, abruptly constricted to the beaked mouth, apex often concave with beak projecting from the concavity, surface smooth, emergent from bracts for 1/2--3/4 times the perianth length. Capsule oblong-ellipsoidal, wall 2-layered. Elaters 2-spiral, ca. 8--10 \um in width. Spores 12--14 \um, verrucose.


Species 2; North America, Eurasia, Pacific Islands.


Liochlaena includes two closely related species sometimes regarded as subspecies of one species. The main distinctions are in inflorescence type, gemmae production and (to a lesser degree) in mid leaf cell size. Liochlaena subulata is distributed mostly in Asia, but area of L. lanceolata mostly confined to Europe and North America.


SELECTED REFERENCES Grolle, R. 1966. Die Typisierung von Jungermannia L. und Jungermannia lanceolata L. Taxon 15: 186--191.  Váňa, J. 1973. Studien über die Jungermannioideae (Hepaticae). 3. Jungermannia subg.Liochlaena. Folia Geobot. Phytotax.. 8: 397--416. Váňa J. and W. S. Hong. 1999. The genus Jungermannia in western North America. Lindbergia 24: 133--144.


1. Plants paroicous, freely fertile, gemmae virtually absent or very rare, midleaf cells 35--65 x 30--45 \um  … 1. Liochlaena lanceolata

1. Plants dioicous, rarely fertile, gemmae common, midleaf cells 25--40 x 25--35\um … 2. Liochlaena subulata



1. Liochlaena lanceolata Nees, Nat. Pflanzenfam., 150. 1845


Jungermannia leiantha Grolle


Midleaf cells 35--65 x 30--45 \um . Oil bodies (4--)6--11(--14) per cell, spheric or oblong, 5--13 x 5--7 \um. Gemmae very rare, ellipsoidal. Sexual condition paroicous; frequently fertile. Androecia below perianth, with 1--5 pairs of bracts.


Shaded soil, decaying wood, forests, mostly coniferous; 0--3500 m; Alta., B.C., Nfld. and Labr., Ont., Que., Ala.; Alaska, Ariz., Calif., Colo., Conn., Idaho, Ill., Ind., Iowa, Kans., Ky., Maine, Mass., Mich., Minn., Mont., Nev., N.H., N.J., N.Y., N.C., Oreg., R.I., Tenn., Utah, Vt., Va., Wash., W.Va., Wis., Wyo.; Eurasia.



Liochlaena lanceolata has long been classified under the name Jungermannia lanceolata L.  According to Grolle (1966), the type of this name is a different species (i.e. Jungermannia atrovirens Dumort.), and it is not correct to apply it to Liochlaena lanceolata. The chromosome number is 16--18.



2. Liochlaena subulata (A. Evans) Schljakov, Pečen. Mkhi severa SSSR, 71. 1981


Jungermannia subulata A. Evans, Trans. Connect. Acad. Arts Sci. 8: 258. 1892

Jungermannia lanceolata ssp. stephanii Amakawa Journ. Hattori Bot. Lab. 22: 71. 1960


Midleaf cells 25--40 x 25--35\um. Oil-bodies 6--8 per cell, spheric or oblong, 5--17 x 5--7 \um. Gemmae ellipsoidal to shortly fusiform colorless to brownish, commonly present on specialized attenuate shoots with scale-like, eroded leaves. Sexual condition dioicous. Rarely fertilized. Androecia in separate branches, semispicate, intercalary, with 3--12 pairs of bracts.



Moist decaying wood, wet soil; ~500-1500; Minn., Miss.; Eurasia, Pacific Islands.


The chromosome number of Liochlaena subulate is 9.