BFNA Title: Cephaloziaceae
Date: Aug. 25, 2020
Edit Level: R
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Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden
BFNA Web site: http://www.mobot.org/plantscience/BFNA/bfnamenu.htm

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XX. CEPHALOZIACEAE Migula

Richard H. Zander

Terry T. McIntosh

Plants forming thin mats or turfs. Branches either intercalary from underside (seldom laterally) of stem, or replacing ventral half of a leaf; often with ventral stolons.  Leaves alternate, succubous, plane or partly involute to form a sac, unlobed or 2-lobed, entire. Underleaves large or absent, unlobed.  Rhizoids scattered over ventral stem. Specialized asexual reproduction by gemmae when present at apices of stems and branches.  Gynoecium on a short or long ventral branch, with or without subfloral branches, Perianth well developed, cylindrical, weakly 3-angled, mouth only slightly narrowed, perigynium absent.

Genera 11 (4 in flora area): widely distributed across North Temperate Zone.

Pleurocladula Grolle and Schofieldia Godfrey are recognized at the genus level in FNA although the species are treated as Fuscocephaloziopsis in the North American synopsis of liverwort species by R. E. Stotler and B. Crandall-Stotler (2017).

1. Stem lacking hyalodermis.

2. Cells with (1--)2--8 oil bodies, leaves not lobed or lobes blunt  2. Odontoschisma, p. xxx

2. Cells lacking oil bodies, leaves lobed, lobes acute ...................  5. Schofieldia, p. xxx

1. Stem with hyalodermis.

2. Underleaves large, ovate-lanceolate; perianth mostly 3--6-stratose proximally, found at apex of shoot; gemmae rare .................................................  3. Pleurocladula, p. xxx

2. Underleaves absent or very small, often as slime-hairs; perianth 1--3-stratose proximally, at apex of often short ventral intercalary branch; gemmae often present.

3. Leaves obliquely to almost horizontally and long-inserted, not approaching the midline of the stem dorsally, flat to somewhat concave, ventral margin not inflated and folded inward ....................................................................  4. Cephalozia, p. xxx

3. Leaves transversely and shortly inserted, approaching the midline of the stem dorsally, concave, with ventral margin inflexed, forming a water sac  1. Nowellia, p. xxx

SELECTED REFERENCES

1. NOWELLIA  Mitt., Nat. Hist. Azores, 321. 1870 * [For John Nowell, 1802--1867, of Todmorden, a Lancashire cotton operative, contributor to Flora of Yorkshire]

Richard H. Zander

Plants in thin mats, green with red, purple or reddish brown tints.  Stem hyalodermis present; ventral-intercalary branching common but Frullania-type (lateral-terminal) branching rare; stolons absent. Rhizoids long, scattered, transparent. Leaves transversely very narrowly inserted or slightly succubous, not decurrent, concave, usually distant, 2-lobed with lobes ending in a fine acumen, leaf below lobes saclike; oil bodies absent.  Underleaves absent.  Specialized asexual reproduction by gemmae, rare, 1-celled, spherical to ellipsoidal, at shoot apex.  Sexual condition dioicous or monoicous, often with autoicous or paroicous intermixed plants.  Androecia apical on a short ventral branch or intercalary, 3--10(--20) pairs of imbricate bracts, each bract saccate, 2-lobed, dorsal lobe margin sometimes toothed; antheridium 1 per bract, stalk 2-seriate.  Gynoecia terminal on short ventral branch. bracteole similar to bracts; perianth narrowly ovate to elliptic, trigonous, unistratose, mouth laciniate or spinose-dentate. Sporophyte capsule ellipsoidal, capsule wall 2-stratose; elaters 2-spiral.

Species 8--10 (1 species in the flora): North America, Central America, South America, West Indies, Eurasia, Atlantic Islands, Pacific Islands, Australia.

The genus Nowellia is nearly cosmopolitan but, unlike most of the other Cephaloziaceae, is mainly tropical and subtropical. Only one species ranges into the North Temperate Zone.

SELECTED REFERENCES Grolle , R. 1968. Monographie der Gattung Nowellia. J. Hattori Bot. Lab. 31: 20--49.  Robinson, H. A. 1970. Notes on the genus Nowellia. Bryologist 73: 150--152.

1. Nowellia curvifolia (Dicks.) Mitt., Nat. Hist. Azores, 321. 1870

Jungermannia curvifolia Dicks., Fasc. Pl. Crypt. Brit. 2: 15. 1790

Plants prostrate, 1--2 cm.  Stems 90--100 \um in diameter, 6--8 cells in width.  Leaves oblong, slightly overlapping, lobes asymmetrical, sinus broadly rounded, deeply concave, 500--700   x 300--400 \um; lobed ca. 0.5 leaf length, lobe apices abruptly and narrowly acuminate, cuticle mostly smooth.  Leaf cells medially 20--25(--30) x 13--17 \um, rounded-hexagonal; trigones absent but cell corner walls often somewhat thickened.  Underleaves absent.  Specialized asexual reproduction rare, by one-celled gemmae borne at plant apex on leaf lobes.  Androecial bracts in several pairs, saccate, 2-lobed, often marginally denticulate.  Gynoecial bracts 2-lobed, ca. 1 x 0.5 mm, marginally strongly dentate, apically sharply acute.  Perianth elongate, 2--2.5 x 0.5--0.6 mm, mouth not or weakly narrowed, spinose-ciliate. Sporophyte capsule longitudinal walls with nodular thickenings. Elaters 9--11 \um in diameter.  Spores 8--10 \um.

Xylicolous, a pioneer on decorticated logs, occasionally on sandstone or moist soil; N.B., Nfld. and Labrador (Nfld. & Labr.), N.S.,  Ont., P.E.I., Que.,; Ala., Ark., Conn., Del., D.C., Ga., Ill., Ind., Iowa, Ky., Maine, Md., Mass., Mich., Minn., Mo., N.H., N.J., N.Y., N.C., Ohio, Pa., S.C., Tenn., Vt., Va., W.Va., Wis.; Mexico; West Indies; Central America (Guatemala, Honduras, Costa Rica); South America; Europe; Asia; Atlantic Islands (Azores, Madeira).

Nowellia curvifolia is easily recognized in the field by its distant leaves with strongly inflated bases, the leaves often vineous in color, with a narrow base, distal lamina bilobed with each lobe ending in a long acumen of uniseriate cells, the basal leaf margins closely incurved to form an open sac, and the perianths often present, distinctly three-angled. The habitat is distinctive, generally restricted to decorticated fallen logs, where it may be monocultural or mixed with other liverworts. It is absent in the western portion of the flora area, but commonly encountered in forests of the East. The androecial bracts are often 1--2 dentate laterally on the lobes.

2.  ODONTOSCHISMA (Dumortier) Dumortier, Recueil Observ. Jungermann., 19.  1835 *  [Greek ondonto, tooth, and schisma, a split; possibly alluding to sinused bracts with one or more marginal teeth]

David M. Krayesky

Lorinda Leonardi

Jerry G. Chmielewski

Pleuroschisma sect. Odontoschisma Dumortier, Syll. Jungerm. Europ., 68. 1831

Plants pale to yellow green to reddish or yellowish or brownish or purplish in color.  Stem cortical and medullary cells similar, without differentiated superficial cells, except in O. sphagni; ventral branching and flagelliform branches common; rhizoids postical on primary shoots and flagelliform branches, hyaline.  Leaves succubous, not decurrent, wide to erect-spreading, distant to imbricate, rotuntate to oblong, entire or rarely with a few teeth; unlobed to 2-lobed, lobes slightly asymmetrical and sinus acute to obtuse, flat or slightly to distinctly concave; cuticle papillose to slightly papillose or smooth. Leaf cells round or quadrate to elongate; thin- to thick-walled; trigones absent or small to large. Oil bodies papillose to granular-botryoidal, ellipsoidal to ovoid or round, (1--)2--8 per cell.  Underleaves 0--1 per pairs of lateral leaves, 2--7 cells in width at base, mostly lanceolate to quadrate, entire to 2-lobed, lobes ciliate or lanceolate, margins and apices smooth with or without slime papillae; cells round or quadrate to elongate, medium to thick-walled, trigones absent to small or large.  Rhizoids hyaline, scattered ventrally or absent.  Specialized asexual reproduction by gemmae in some species, green to red, 1--2 celled, ellipsoidal to oval to triangular and occasionally stellate, forming at shoot apex on leaf margins of erect growing stem tips.  Sexual condition dioicous.  Androecia terminal or intercalary on stems or branches; bracts imbricate, 3--18 pairs, mostly 2-lobed, erect, concave, toothed or entire at base, with or without slime papillae at apices or along margins; bracteoles similar to underleaves but smaller; antheridia orbicular, hyaline, yellow-orange, bronze or red-purple, 60--200 \um in width, 1 per bract; jacket cells irregularly orientated; stalks 2 cells in width, 3--8 cells in length.  Gynoecia terminal on postical or lateral intercalary branches, bracts 2--4 pairs, 2-lobed, concave, longer than leaves, with or without slime papillae along margins, bracteoles mostly similar in size and shape to bracts; connate, adnate, or free; stem-derived protective structure absent.  Perianth 0.5--0.9 \um in diameter, exerted, oblong to cylindrical; mouth tapered to slightly tapered, sinuate to lobed, crenate to ciliate to entire; (1--)2(--3)-stratose near base, 1-stratose near mouth; without surface ornamentation.  Sporophyte seta cross section (99--)104--221 \um in diameter; 8 or 13--18 exterior cell rows and 4 or 5--16 interior cell rows depending on species; capsule 0.7--1.2 x 0.3--0.5 mm; ovoid, 4-valved, outer cells with nodular rust-colored thickenings, inner cells with semiannular rust-colored thickenings.  Elaters with tapered or truncate ends, 2(--3)spiraled, spirals 2.5--4 \um in width, rust-red.  Spores papillose, rust-red to purple.

Species  21 (6 in the flora): North America; West Indies; Bermuda; Central America; South America; Europe; Asia; Africa; Atlantic Islands; Indian Ocean Islands; Pacific Islands.  J. A. Patton (1999) suggested approximately 30 species worldwide; however, S. R. Gradstein and A. L. Ilkiu-Borges (2015) and S. C. Aranda et al. (2014) provide ample evidence to support the delimitation of only 21 Odontoschisma species, based on molecular evidence, and anatomical characters (i.e. a stem lacking a hyalodermis, large trigones, large oil bodies, and non-tiered antheridial jacket cells); therefore, we follow their assessment of the genus.

SELECTED REFERENCES  Aranda, S. C., S. R. Gradstein, J. Patiňo, B. Laenen, A. Désamoré, and A. Vanderpoorten.  2014.  Phylogeny, classification, and species delimitation in the liverwort genus Odontoschisma (Cephaloziaceae).  Taxon 63: 1008--1025.  Bednarek-Ochyra, H., J. Váňa, R. Ochyra, and R. I. Lewis-Smith.  2000.  The liverwort flora of Antarctica.  Polish Academy of Sciences, Institute of Botany, Cracow.  Chen, F.-D., X.-D. Li, and H. Kanada. 1993.  An investigation of the bryophytes in the Fildes Peninsula, King George Island, Antarctica.  Antarctic Reseach (Chinese Edition) 5: 46--50. [In Chinese with English summary].  Chen, F.-D., X.-D. Li, and J.-X. Ziu.  1995.  Handbook of the Fildes Peninsula Antarctica mosses. Ocean Press, Beijing, China. [In Chinese].   Evans, A.W.  1903.  Odontoschisma macounii and its North American allies.  Botanical Gazette 36: 321-348.  Gradstein, S. R. and A .L. Ilkiu-Borges.  2015.  A taxonomic monograph of the genus Odontoschisma.  Nova Hedwigia 100: 16-100.  Jensen, C.  1897.  Mosser fra Øst-Grønland.  Meddelelser om Grønland 15: 365--443.  Li, X.-D.. J.-X. Liu and F.-D. Chen.  1998.  A preliminary study of the bryoflora of the Fildes Peninsula, King George Island, Antarctica.  J. Hattori Bot. Lab. 84: 199--202. Paton, J. A. 1999.  The Liverwort Flora of the British Isles.  Harley Books, Colchester, England.  Schuster, R. M.  1974.  Odontoschisma.  In: R. M. Schuster.  The Hepaticae and Anthocerotae of North America east of the hundredth meridian. New York, Vol. 3. Pp. 678--695 and 829--870. 

1.      Leaves 2-lobed; seta with 13--18 exterior cell rows and 5--16 interior cell rows in cross section.

2. Leaves typically wide-spreading, mostly flat, 0.5--1.1 mm; gemmae never present…..… …………………………………………………………..….. 1. Odontoschisma fluitans

2. Leaves typically erect-spreading, mostly concave, 0.3--0.5 mm; gemmae often present...  ……………………………………………………………… 2. Odontoschisma francisci

1.      Leaves not lobed; seta with 8 exterior cell rows and 4 interior cell rows in cross section

3. Leaves with 1--4 rows of thick-walled marginal cells; median leaf cells not stellate (cell walls evenly thickened throughout) ………………………... 3. Odontoschisma sphagni

3. Leaves without marginal cells; median leaf cells stellate.

4.      Middle lamella of the leaves distinct, trigones deeply pigmented golden to brownish and well defined ……………….…………..... 4. Odontoschisma elongatum

4.   Middle lamella of the leaves not visible, trigones not deeply pigmented or well defined.

5.   Leaves generally emarginate at apex; large underleaves large, mostly the length of the vegetative leaves; cuticle of leaves smooth …………………………………………………. 5. Odontoschisma macounii

5.   Leaves not distinctly emarginate at apex; underleaves minute; cuticle of leaf papillose ……………………………….…. 6. Odontoschisma denudatum

1. Odontoschisma fluitans (Nees) L. Söderström. & Váňa, Phytotaxa 112: 12. 2013

Jungermannia fluitans Nees, Flora 6: 30. 1823;  Cladopodiella fluitans (Nees) Jörgensen

Plants yellow-brown, green or purple-black, ca. 20--45(--100) mm, prostrate, growing among other bryophytes or in mats. Stems 150--200 \um in diameter, without cortical differentiation, cortex and medulla 6--8 cells in width, cortical cells in transverse section 30--90 x 20--27 \um; flagelliform branches common.  Leaves oblong, 2-lobed, lobes slightly asymmetrical, sinus acute to obtuse, mostly distant but contiguous to slightly imbricate, flat to slightly concave, 500--1100 x 400--800 \um; lobed 0.2--0.4 leaf length, largest leaf lobe 7--15 cells in width at base, lobe apices obtuse to occasionally acute; cuticle smooth.  Leaf cells at margins 15--38 \um, at midleaf in lobes 20--40 \um, at base 15--60 \um; trigones absent or small; oil bodies 3--10 per cell, 4--10 \um.  Underleaves distant, 200--500 x 50--150 \um, cells 12--75 \um.  Specialized asexual reproduction absent.  Androecial bracts 0.4--0.7 x 0.4--0.6 mm.  Gynoecial bracts 0.8--1.2 mm.  Perianth 2.5--4 x 0.7--1 mm; cylindrical; mouth sinuate, entire to crenulate. Sporophyte capsule wall 3 cell layers, 30--35 \um in thickness, outer cells 10--18 \um in width, inner cells 10--13 \um in width. Elaters 125--250 x 10--13 \um, 2-spiraled.  Spores 15--20 \um.

Acidophile, often growing intermingled with sphagnum or peat in pools, less commonly on moist soil or rock; bogs, lake and pond edges, moist open areas; mostly low to moderate elevations; Greenland; Alta., B.C., Man., N.B., Nfld. & Labr., N.W.T., N.S., Ont., P.E.I, Que.; Alaska, Conn., Maine, Mass., Mich., Minn., N.H., N.J., N.Y., Ohio, R.I., Vt., Wash., W.Va., Wis.; Europe; Asia; Africa.

Sometimes confused with the more common Gymnocolea inflata. Odontoschisma fluitans may be easily identified if perianths are present.  The perianths of O. fluitans, which are more elongate than the more globose perianths of G. inflata, do not become detached from the stem.  Also O. fluitans does not develop terminal or lateral branches as does Gymnocolea.

2. Odontoschisma francisci (Hooker) L. Söderström & Váňa, Phytotaxa 112: 12. 2013

Jungermannia francisci Hooker, Brit. Jungermann., pl. 49. 1813;  Cladopodiella francisci (Hooker) Jörgensen

Plants green or purple-red, ca. 3--15 mm, prostrate with erect branches, growing in mats or among other bryophytes. Stems 120--150 \um in diameter, without cortical differentiation, cortex and medulla 5--7 cells in width, cortical cells in transverse section 20--65 x 10--30 \um; flagelliform branches common.  Leaves ovate, 2-lobed, lobes slightly asymmetrical, sinus acute to obtuse, mostly imbricate but contiguous to distant, concave to slightly concave, 300--500 x 200--400 \um, lobed 0.1--0.3 leaf length, largest leaf lobe 5--8 cells in width at base, lobe apices acute to obtuse; cuticle smooth.  Leaf cells at margins 10--28 \um, at midleaf in lobes 10--25 \um, at base 18--45 \um; trigones absent or small; oil bodies 1--5 per cell, 4--8(--14) \um.  Underleaves distant to imbricate, 110--180 x 60--110 \um, cells 7--38 \um.  Specialized asexual reproduction by gemmae often present, forming on leaf margins of erect stem tips, stellate, 3--6-angled, 1---2 cells, 17--33 \um.  Androecial bracts 0.1--0.5 x 0.1--0.3 mm.  Gynoecial bracts 0.6--1 mm.  Perianth 1.8--2.5 x 0.7--0.9 mm; cylindrical; mouth sinuate to lobed, entire to crenulate.  Sporophyte capsule wall 2--3 cell layers, 20--30 \um in thickness, outer cells 10--15 \um in width, inner cells 5--8 \um in width.  Elaters 65--170 x 8--10 \um, 2-spiraled.  Spores 12.5--15 \um.

Acidophile, often forming uniform patches on moist sandy to gravelly soil and less common on peat; wetlands, heathlands, lake and pond edges, roadsides, and cliff ledges; low elevations to alpine summits; Greenland; Nfld. and Labr. (Nfld.), N.S., Que.; Conn., Maine, Mass., N.H., N.Y., R.I.; Europe.

Because of its size and form, Odontoschisma francisci may be confused with species of Cephalozia, but absence of a hyalodermis and the non-decurrent leaves will readily distinguish this species.   In ideal conditions, O. francisci can be form abundant uniform patches.

3.      Odontoschisma sphagni (Dickson) Dumortier, Recueil Observ. Jungermann., 19.  1835

Jungermannia sphagni Dickson, Fasc. Pl. Crypt. Brit. 1: 6. 1785; Odontoschisma prostratum (Swartz) Trevisan

Plants pale tinged with brown to reddish brown (all pale green in shade forms), ca. 0.7--29 x 0.7--2.0 mm, prostrate, growing in mats or singly among other bryophytes.  Stems 145--174 \um in diameter, with cortical differentiation, cortex and medulla 13--14 cells in width, cortical cells in transverse section 5--8 x 7--11 \um; flagelliform branches common.  Leaves suborbicular to rounded-oblong to subrotund with apex rounded, distant to imbricate, flat to moderately concave, 418--498 x 456--535 \um; cuticle slightly papillose.  Leaf cells at margins 10--15 x 10--15 \um, at midleaf 12--20 x 12--19 \um, at base 14--23 x 24--31 \um; trigones small; oil bodies 2--4(--5) per cell, 3--12 \um.  Underleaves distant, 125--148 x 56--60 \um, cells 14--20 \um.  Specialized asexual reproduction gemmae absent.    Androecia bracts 0.3--0.4 x 0.3--0.4 mm.  Gynoecia bracts 1--1.2 mm.  Perianth 3.0--4.5 x 0.8--0.9 mm; cylindrical; mouth lobate, ciliate with teeth.  Sporophyte capsule wall of 2 cell layers, 20--25 \um in thickness, outer cells 11--13 \um in width, inner cells 9--11 \um in width.  Elaters 122--236 x 10--11 \um, 2-spiraled.  Spores (7--)9--11 \um. 

Temperate and tropical from near sea level to upland forests, on peat moss (rarely on other mosses), peaty soils, sandy soils, gravelly soils, decaying logs, rarely on tree bases, bogs, swamps, and upland forests in noncalcareous habitats, from hydric to xeric.; low to high elevations; Greenland; St. Pierre and Miquelon; Alta., N.B., Nfld. and Labr., N.S., Ont., Que., Yukon; Ala., Alaska, Ark., Conn., Del., D.C., Fla., Ga., Ill., Ky., La., Maine, Md., Mass., Miss., Mo., Mont., N.H., N.J., N.Y., N.C., Ohio, Pa., R.I., S.C., Tenn., Tex., Vt., Va., W.Va., Wyo.; Mexico; West Indies; Bermuda; Europe; Atlantic Islands. 

Gradstein and Ilkiu-Borges (2015) suggest O. sphagni does not occur in South America or Asia as collections cited from these localities belong to O. variabile and O. denudatum, respectively.  Schuster (1974) indicated that the Spruce 1882 report of O. sphagni from Africa is erroneous.  The Spruce collection is unavailable; however, since we have been unable to find other reports of O. sphagni from that region we support Schuster’s hypothesis. 

4.      Odontoschisma elongatum (Lindb.) A. Evans, Rhodora 14: 13.  1912

Odontoschisma denudatum fo. elongatum  Lindberg, Helsingfors Dagblad 1874(45): [2]. 1874

Plants yellow green tinged with brown or plants blackish brown (rarely all green), ca. 5.0--15 x  1--1.9 mm, gregarious, prostrate with ascending apices, growing in mats or singly among other bryophytes.  Stems 163--216 \um in diameter, without cortical differentiation, cortex and medulla 8--9 cells in width, cortical cells in transverse section 9--27 x 14--21 \um; flagelliform branches uncommon.  Leaves subrotund with apex rounded, distant to imbricate, slightly to moderately concave, 427--525 x 446--590(--739) \um; cuticle smooth.  Leaf cells at margins 11--17 x 10--15 \um, at midleaf 10--19 x 15--22 \um, at base 14--25 x 15--23 \um; trigones small to moderate to large; oil bodies (1--)2--3(--4) per cell, 5--13 \um.  Underleaves distant, 123--199 x 86--89 \um, cells 19--25 \um.  Specialized asexual reproduction gemmae uncommon, forming at the tips of ascending shoots, suborbicular to ovate to elliptic, 1--2 cells, 50--70 \um.  Androecia bracts rare, 0.3 mm long.  Gynoecia bracts rare, 0.7--0.9 mm.  Perianth rare, 2.0 x 0.5 mm; cylindric; mouth crenulate.  Sporophyte capsule wall not observed.  Elaters 163--229 x 8--9 \um, 2-spiraled.  Spores 8--10 \um. 

Circumpolar in acidic areas, muddy, sandy, gravelly, or rocky substrates, occasionally peaty humus, rotten wood, moorlands, heaths, and in moors around lakes, ponds, bogs, and marshes with Sphagnum;  near sea level to alpine; Greenland; B.C., Nfld. and Labr. (Nfld.), N.W.T., N.S., Ont., Que.; Alaska, Maine, Mass., Mich., N.H.; Europe; Asia. 

Sporophytes are very rare; however, a prepared slide of capsule material from RMS 11730 in (F) had some elater and spore material.  Due to the deteriorating condition of the slide only spore and elater anatomy could be determined.  Additionally, mature perianths are rare; however, sample RMS 12239 in (F) was reported to have few perianths (by Schuster), but presently the sample appears barren.  As sexual structures of the gametophyte were not observed in the material examined measurements for androecial bracts, gynoecial bracts, and perianths are derived from Patton (1999).  

5.      Odontoschisma macounii (Austin) Underwood, Bull. Illinois State Lab. Nat. Hist. 2: 92.  1884

Sphagnoecetis macounii Austin, Bull. Torrey Bot. Club 3: 13. 1872

Plants yellowish to pale green, ca. 6.0--13.0 x 0.7--1.2 mm, prostrate, growing in small mats or singly among other bryophytes.  Stems (141--)163--183 \um in diameter, without cortical differentiation, cortex and medulla 9--10 cells in width, cortical cells in transverse section 10--16 x 7--12 \um; flagelliform branches common.  Leaves subrotund, apex emarginate, loosely to highly imbricate, strongly concave, 341--579 x 252--771 \um; cuticle smooth.  Leaf cells at margins 15--21 x 13--14 \um, at midleaf 15--18 x 14--24 \um, at base 22--34 x 15--19 \um; trigones small to large; oil bodies 1--3--(4) per cell, 6--14 \um.  Underleaves distant, 62--176(--407) x 73--121(--274) \um, cells 20--28 \um.  Specialized asexual reproduction by gemmae common, forming at the tips of erect shoots, subelliptic to elliptic, 1--2 cells, 20--35 \um.    Androecia bracts 0.2--0.3 x 0.2--0.2 mm.  Gynoecia bracts 1.2--1.4 mm.  Perianth 3.5--4.0 x 0.8--0.9 mm; cylindrical; mouth irregularly lobate, entire to slightly crenate.  Sporophyte capsule wall of 2 cell layers, 25--55 \um in thickness, outer cells 22--34 \um in width, inner cells 10--21 \um in width.  Elaters (78--)117--224 x 10--13 \um, 2(--3)-spiraled.  Spores 12--16 \um. 

Circumpolar, arctic-alpine, and with meta-populations along Lake superior, restricted to calcareous and exposed sites, on organic matter (generally humus or peat) atop calcium-rich rock or associated with calcium rich seep; elevation low to high; Greenland; B.C., Man., Nfld. & Labr., N.W.T., N.S., Nunavut, Ont., Que., Yukon; Alaska, Mich., Minn., Wis.; Europe; Asia. 

Schuster (1974) reported populations of Odontoschisma macounii occasionally occur at locations further south in boreal forest zones which represent relic sites for the present distribution of this species.  The reports by Chen et al. (1993; 1995) and Li et al. (1998) of O. macounii from Antarctica are erroneous as these reports have been demonstrated by Bednarek-Ochyra et al. (2000) to be Herzogobryum teres (Carrington & Pearson) Grolle. 

6.      Odontoschisma denudatum (Nees) Dumortier, Recueil Observ. Jungerm., 19. 1835

Jungermannia denudata Nees, Fl. Crypt. Erlang., 14.  1817

Plants pale green or yellow green tinged brown, red, or purplish black or blackish green to blackish brown or violet purple with a metallic like sheen, 8.0--12.0 x 0.5--1.1 mm, gregarious or prostrate, growing in mats or singly among other bryophytes.  Stems 130--186 \um in diameter, with slight cortical differentiation, cortex and medulla 8--9 cells in width, cortical cells in transverse section 17--22 x 16--24 \um; flagelliform branches common.  Leaves suborbicular to broadly ovate with apex rounded (occasionally moderately emarginate), moderately imbricate, strongly concave, 282--690 x 389--516 \um; cuticle slightly papillose.  Leaf cells at margins 13--24 x 8--18 \um; at midleaf 12--26 x 13--29 \um, at base 18--31 x 27--34 \um; trigones small to large; oil bodies (1--)2--6(--8) per cell, 5--16(--19) \um.  Underleaves distant, 75--209 x  65--125 \um, cells 14--23 \um.  Specialized asexual reproduction gemmae common, forming at the tips of ascending shoots, orbicular to ovate to elliptic, 1--2 cells, 25--35 \um.   Androecia bracts 0.3--0.3 x 0.1--0.2 mm.  Gynoecia bracts 0.9--1.1 mm.  Perianth 2.5--4.0 x 0.5--1.0 mm; cylindrical; mouth crenulate.  Sporophyte capsule wall of 2 cell layers; 30--65 \um in thickness outer cells 17--41 \um in width; inner cells 11--22 \um in width.  Elaters 105--256 x 10--11 \um, 2-spiraled.  Spores 8--11(--15) \um. 

Subspecies 3 (1 in the flora)

1a. Odontoschisma denudatum (Nees) Dumort. ssp. denudatum

Odontoschisma denudatum var. laevissima R. M. Schuster; O. gibbsiae A. Evans

Leaves ovate to suborbicular, concave, and straight in stance, never curved or subfalcate.

Temperate and boreal, mesic sites, also sites of low pH, moist decaying logs and peaty soils in moorlands, heaths, rocky slopes, and bogs; low to high elevations; St. Pierre and Miquelon; B.C., N.B., Nfld. and Labr. (Nfld.), N.S., Ont., Que.; Ala., Ark., Conn., Del., D.C., Fla., Ga., Ky., La., Maine, Md., Mass., Mich., Minn., Miss., N.H., N.J., N.Y., N.C., Ohio, Pa., S.C., Tenn., Vt., Va., W.Va., Wis.; Mexico; West Indies; Central America; South America; Europe;  Asia; Pacific Islands. 

Schuster (1974) indicated that the reports of O. denudatum ssp. denudatum to Greenland are erroneous due to confusion with O. elongatum.  We also treat those records as erroneous.  Gradstein and Ilkiu-Borges (2015) provide little information as to why O. denudatum var. laevissima was placed in synonymy with O. denudatum ssp. denudatum other than stating  that, based on the original description of O. denudatum var. laevissima and topotype material, it is conspecific with O. denudatum ssp. denudatum.  Additionally, Gradstein and Ilkiu-Borges (2015) provide no information for synonymizing O. gibbsiae with O. denudatum ssp. denudatum; however, Godfrey (1977) indicated that the difference in leaf stance, degree of leaf imbrication, underleaf stance and gemmae anatomy fell within the circumscription of O. denudatum and hence indicated the synonomy.  After examining the type material of O. gibbsiae (YPM) we support hypothesis that O. gibbsiae is conspecific with O. denudatum ssp. denudatum, as the leaves are not emarginate or lobed, the leaf margins lack thick-walled marginal cells, and the cells of the leaves lack a deeply pigmented middle lamella. 

3.  PLEUROCLADULA Grolle, J. Bryol. 10: 269.  1979 * [Greek pleuro side, Greek clado branch; alluding to lateral branching]

David M. Krayesky

Jerry G. Chmielewski

Pleurocladula º Pleuroclada Spruce 1882 (nom. illeg.), non Pleurocladia A. Braun 1855

Plants light pellucid green, in full sun or shade, nearly white when dry (occasionally reddish brown).  Stem possessing a hyalodermis; Frullania type (lateral-terminal) branching common and lateral-intercalary branching rare; flagelliform branches occurring in some populations.  Leaves transverse-succubous, not decurrent, distant to imbricate, entire; 2-lobed, lobes triangular, and concave; oil bodies absent.  Underleaves large, nearly as long as leaves but only half as wide, often possessing 1--2 sub-basal teeth.  Specialized asexual reproduction by gemmae uncommon, 1-celled, spherical to ellipsoidal, at shoot apex.  Sexual condition dioicous.  Androecia intercalary, each bract with basal antical tooth; antheridia occurring singly per bract, ovoid, with non-tiered jacket cells and a 2-seriate stalk.  Gynoecia terminal, bracteole always narrower and sometimes shorter than bracts; perianth clavate to oblong or fusiform, fleshy and multistratose at base, mouth lobate with a crenulate margin.  Sporophyte seta yielding 8--12 epidermal and 5--8 inner cell rows; capsule ellipsoidal, four valved, capsule wall 2-stratose. 

Species 1 (1 in the flora): North America, Europe and Asia.

Pleurocladula albescens (Hooker) Grolle and P. islandica (Nees) Grolle have been treated as separate species (e.g. M. Krzakowa and J. Sweykowski 1981; R. Grolle 1983).  We concur with J. A. Paton’s (1999) assessment that the genus is monotypic as Paton (1976) provided ample evidence for the inclusion of P. islandica into synonymy with P. albescens. 

The status of Pleurocladula is a point of debate.  A. A. Vilnet et al. (2012) proposed that Cephalozia is polyphyletic forming two clades.  One Cephalozia clade included the generitype, as well as other Cephalozia species.  The second clade included species of Cephalozia, Pleurocladula, and Schofieldia.  J. Váňa et al. (2013) have transferred those species in the second clade to Fuscocepaloziopsis.  K. Feldberg et al. (2016) concur with Váňa et al. (2013).  Although the Feldberg et al. (2016) analysis of the Cephaloziaceae is the most comprehensive molecular assessment of the group to date, the maximum likelihood value of the Fuscocepaloziopsis clade does not provide the same level of support as evidenced in the recent treatment of Odontoschisma (S. C. Aranda et al. 2014).  For this reason we continue to recognize Pleurocladula, Schofieldia, and Cephalozia sensu lato as distinct genera.  In order to support the reorganization of these specific Cephaloziaceae taxa into two genera, namely, Cephalozia and Fuscocephaloziopsis, both an extensive anatomical study and a molecular evaluation with a very strong level of support will need to be demonstrated, as is the case with the recent treatments of Odontoschisma (S. C. Aranda et al. 2014; S. R. Gradstein and A. L. Ilkiu-Borges 2015). 

SELECTED REFERENCES  Aranda, S.C., S.R. Gradstein, J. Patiňo, B. Laenen, A. Désamoré, and A. Vanderpoorten.  2014.  Crandall-Stotler, B., R.E. Stotler, and D.G. Long.  2009.  Phylogeny and classification of the Marchantiophyta.  Edinburgh J. Bot. 66: 155--198.  Feldberg, K., J. Váňa, J. Krusche, J. Kretschmann, S. D. F. Patzak, O. A. Pérez-Escobar, N. R. Rudolf, N. Seefelder, A. Schäfer-Verwimp, D. G. Long, H. Schneider, and J. Heinrichs.  2016.  A phylogeny of Cephaloziaceae (Jungermanniopsida) based on nuclear and chloroplast DNA markers.  Organisms Diversity and Evolution 16 [doi: 10.1007/s13127-016-0284-4].  Frye, T. C. and L. Clark.  1943.  Hepaticae of North America.  Univ. Washington Publ. Biol. 6(2): 187--190.  Geissler, P. and H. Bischler.  1989.  Index Hepaticarum: Vol. 11 Naiadea to Pycnoscenus,  J. Cramer, Berlin, Germany.  Gradstein, S. R. and A. L. Ilkiu-Borges.  2015.  A taxonomic monograph of the genus Odontoschisma.  Nova Hedwigia 100: 16--100.  Grolle, R.  1979.  Miscellanea hepaticologica 171--180.  J. Bryol. 10: 263--272.  Grolle, R.  1983.  Hepatics of Europe including the Azores: An annotated list of species, with synonyms from the recent literature.  J. Bryol. 12: 403--459.  Hooker, W. J.  1816.  British Jungermanniae.  84 plates and 4 supplementary plates.  London.  Krzakowa, M. and J. Szweykowski.  1981.  A species concept in the genus Pleurocladula (Hepaticae).  Acta Soc. Botanic. Poloniae 39: 465--479.  Paton, J. A.  1976.  Pleuroclada albescens (Hook.) Spruce with gemmae.  J. Bryol. 9: 1--5.  Paton, J. A.  1999.  The Liverwort Flora of the British Isles.  Harley Books, Colchester, England.  Schuster, R. M.  1974.  Pleurocladula.  In:  R. M. Schuster.  The Hepaticae and Anthocerotae of North America east of the hundredth meridian. New York, Vol. 3, Pp. 66-6-671.  Smith, A. J. E.  1990.  The Liverworts of Britain and Ireland.  Cambridge University Press.  Cambridge, UK.  Spruce, R.  1882.  Pleuroclada.  In: R. Spruce.  On Cephalozia, its subgenera and some allied genera. Malton, pp. 77--80, Malton.  Váňa, J., L. Söderström, A. Hagborg,  and M. von Konrat.  2013.  Notes on early plants today. 41. New combinations and synonyms in Cephaloziaceae (Marchantiophyta).  Phytotaxa 112: 7--15.  Vilnet, A. A., N. A. Konstantinova, and A. V. Troitsky.  2012.  Molecular phylogeny and systematics of the suborder Cephaloziineae with special attention to the family Cephaloziaceae s.l. (Jungermanniales, Marchantiophyta).  Arctoa 21: 113--132. Wolf, J. and E. Hall.  1878.  A list of mosses, liverworts, and lichens of Illinois.  Bull. Illinois State Lab. Nat. Hist. 1: 18--35.

1. Pleurocladula albescens (Hooker) Grolle, J. Bryol. 10: 269.  1979

Jungermannia albescens Hooker, Brit. Jungermann., pl. 72 and suppl. pl. 4.  1815; Pleurocladula islandica (Nees) Grolle

Plants ca. 4.7--9.4 x 0.2--0.5 mm; prostrate to gregarious, growing in mats, turfs, or singly among other bryophytes.  Stems fleshy 120--185 \um, cortical cells thin walled and 23--39 x 24--39 \um; medullary cells thick walled and 18--24 x 18--22 \um, medulla 112--123 \um in diameter and 5--8 stratose; flagelliform branches occurring in some populations.  Leaves distant to imbricate; erecto-patent; concave; suborbicular 240--397 \um long x 200--400 \um broad; margins entire; 2-lobed, lobes triangular, incurved; sinus 1/3--1/2 the length of the leaf, base of sinus rounded to acute; marginal cells 17--31 x 17--31 \um; median cells in lobes 17--26 x 17--35 \um; median leaf cells 17--29 x 23--32 \um; basal cells 23--39 x 23--36 \um. Oil-bodies absent.  Rhizoids scarce to abundant, and abundant on flagelliform branches, colorless.  Underleaves large 273--307 x 99--177 \um; erecto-patent; ovate to sublanceolate; apices acute to obtuse; with or without basal teeth, often present on one side, occasionally on both.  Specialized asexual reproduction by gemmae 23--26 \um in diameter.  Androecia bracts 0.5--0.6 x 0.4--0.5 mm; in 15--18 pairs; 2-lobed, and with basal antical tooth.  Gynoecia bracts 1--1.3 x 0.9--1.2 mm and bracteoles 0.5--0.6 x 0.4--0.5 mm; in series of 2; bracts 2-lobed, and much larger than the leaves; bracteoles free or connate to the base of the bract, unlobed or shallowly 2-lobed with 1--3 teeth.  Perianth 1.9--2.9 x 0.8--0.9 mm; clavate to oblong or fusiform; fleshy and multistratose at base; mouth lobate with a crenulate margin.  Seta 250--322 \um in diameter.  Capsule 1.1--1.5 x 0.4--0.5 mm, capsule wall 2 cell layers thick; epidermal cells 15--38 x 22--40 \um; interior cells 15--20 x 19--76 \um in diameter.  Elaters 7--10 \um in diameter, 127--318 \um in length, and (1--)2-spiraled.  Spores brown, vermiculate-verrucose 14--16 \um . 

Northern Hemisphere Arctic and temperate (alpine to montane) areas, on fine to gravelly soils, peaty soils, and rock, and on the banks of small streams and ephemeral ponds associated with peatland areas; low to high elevations; Greenland; Alta., B.C., Nfld. and Labr., N.W.T., Nunavut, Que., Yukon; Alaska, Calif., Mont., Wash.; Europe; Asia. 

R. M. Schuster (1995) indicated Pleurocladula albescens has a tendency toward being acidophilic. There are some doubtful records in the United States and these published records will be treated here as erroneous.  These aforementioned records include: the Wolf and Hall (1878) report from Illinois and the Watson s.n. 1972 collection in TROPICOS at MO from New Hampshire.  The Wolf and Hall (1878) report was treated by Schuster (1974) as an erroneous, as we have been unable to locate this collection we continue to follow the Schuster (1974) assessment of this population.  The Watson s.n. 1972 collection from New Hampshire is not Pleurocladula albescens, but is Orthocaulis kunzeanus (Huebener) H. Buch. 

4. CEPHALOZIA (Dumortier) Dumortier, Recueil Observ. Jungerm., 18. 1835 * [Greek kephale, head, and ozos, bud, alluding to enlarged bracts and bracteoles]

Richard H. Zander

Terry T. McIntosh

Jungermannia sect. Cephalozia Dumortier, Syll. Jungerm. Europ., 60. 1831; Fuscocephaloziopsis Fulford

Plants in patchy, thin mats, yellow-green or green with brown tints.  Stem hyalodermis often present or occasionally absent; simple or commonly with ventral-intercalary branching common but Frullania-type (lateral-terminal) branching rare; stolons often present. Rhizoids long, scattered, transparent. Leaves transversely very narrowly inserted or slightly succubous, often decurrent, flat to concave, distant to contiguous, 2-lobed with lobes broadly acute to acuminate, sac-like lobes absent; oil bodies absent.  Underleaves rarely present, often present in gynoecious as bracteoles.  Specialized asexual reproduction by gemmae, rare, spherical to ellipsoidal, 1(--2)-celled, at shoot apex.  Sexual condition dioicous or autoicous.  Androecia apical on a short ventral branch or intercalary on longer branches, 2--20 pairs of imbricate bracts, each bract 2-lobed, dorsal lobe margin often toothed; antheridium 1 per bract.  Gynoecia terminal on short ventral branch. bracteole similar to or narrower than bracts; perianth narrowly ovate to elliptic or fusiform, cells trigonous distally, 1--3-stratose proximally, mouth crenulate to laciniate or spinose-dentate. Sporophyte capsule ellipsoidal, capsule wall 2-stratose; elaters 2-spiral.

Species 45 (11 in the flora): North America, Central America, South America, Eurasia, Africa, Atlantic Islands.

Fuscocephaloziopsis is not considered distinct from Cephalozia as M. H. Fulford’s (1968) criteria are weak, and the molecular study of A. A. Vilnet et al. (2012) does not distinguish the two genera on the basis of evolutionary fact. The careful study of Cephalozia in Russia by A. D. Potemkin and E. V. Sofronova (2013) includes Nowellia, Pleurocladula, Schofieldia in Cephalozia, which sinks good genera unnecessarily simply because they are molecularly monophyletic in the cladistic sense.

SELECTED REFERENCES  Fulford, M. H. (1968). Manual of the leafy Hepaticae---part III. Memoirs of the New York Botanical Garden, 11, 277--392.  Potemkin, A. D. and E. V. Sofronova. 2013. Taxonomic study on the genus Cephalozia in Russia and circumscription of the genus. Arctoa 22: 173--206.  Schuster, R.M. 1974. The Hepaticae and Anthocerotae of North America East of the Hundredth Meridian. Columbia University Press, New York. Volume 3.  Söderström, L., A. Hagborg, M. von Konrat, et al. 2016. World checklist of hornworts and liverworts. Phytokeys 59: 1--828. Stotler, R. E. and B. Crandall-Stotler. 2012. A synopsis of the liverwort flora of North America north of Mexico. Ann. Missouri Bot. Gard. 102: 574--709. Vilnet, A. A., N. A. Konstantinova and A. V. Troitsky. 2012. Molecular phylogeny and systematics of the suborder Cephaloziineae with special attention to the family Cephaoziaceae S.L. (Jungermanniales, Marchantiophyta). Arctoa 21: 118--132.

1. Leaves quadrate to ovate; gemmae rare; plants usually with brownish secondary pigmentation on exposed shoots.

2. Plants brownish; leaves inserted nearly transversely, rectangular to ovate, not decurrent; stolons common ................................................ 1. Cephalozia bicuspidata

2. Plants green to yellowish; leaves inserted obliquely or horizontally, rarely subtransversely, commonly decurrent; stolons rare.

3. Female bracts 3--5-lobed; medial leaf cells 30--46 x 40--46 \um, thin-walled; perianth mouth laciniate .................................................. 2. Cephalozia lacinulata

3. Female bracts 2-lobed; medial leaf cells 12--20 x 16--20 \um, thick-walled; perianth mouth crenulate to dentate ................................. 3. Cephalozia macounii

1. Leaves rounded to ovate; plants with exposed shoots yellow-green to olive or brown;

4. Leaves ovate or ovate-rectangular, somewhat longer than wide; sinus deep, 0.5--0.7 leaf length; lobes not connivent; leaves not decurrent ......... 6. Cephalozia leucantha

4. Leaves round to rounded-quadrate, 1:1; sinus less deep, 0.3--0.5 leaf length; lobes often connivent; leaves often decurrent.

5. Leaf cells large, to 45--60 x 45--50 \um medially.

6. Leaves usually 8--12 cells wide, strongly decurrent; lobes sharp, strongly connivent,  3--5(--8) cells wide; leaf cells usually 45--55 x 49--48 \um; ......... ............................................................................................................................ ..................................................................................... 5. Cephalozia connivens

6. Leaves usually 12--25 cells wide, little decurrent; lobes short-acute, little connivent,  somewhat wider,  often 6--10 cells wide; leaf cells 20--60 x 24--50 \um .....................................................................................................................  ...................................................................................  11. Cephalozia pleniceps

5. Leaf cells small, to 18--40 x 15--45 \um; stolons absent.

7. Leaves long-decurrent; lobes not connivent to touching; plants pale green, transparent.

8. Leaf lobes narrowly acute, ending in 1--2(--3) uniseriate cells, each 2:1; leaf cells usually thick-walled; perianth mouth with cilia 2--6 cells long; autoicous ........................................................... 7. Cephalozia loitlesbergeri

8. Leaf lobes broadly acute, ending in 1--2 uniseriate cells, each 1:1; leaf cells usually thin-walled; perianth mouth with teeth 1--2 cells long; dioicous  8. Cephalozia lunulifolia

7. Leaves short-decurrent or decurrencies lacking, lobes mostly little connivent; plants deep green.

9. Leaves about 20 cells at widest, not decurrent; lobe tips very narrowly acute; laminal cells thin-walled; stem lacking a hyalodermis ................................ ....................................................................................................................... ........................................................................... 10. Cephalozia pachycaulis

9. Leaves about 9--10 cells at widest, short-decurrent; lobe tips broadly acute; laminal cell walls sometimes thickened; stem with hyalodermis.

10. Cells very thick-walled, 15--18 x 20--25 \um at lobe base 4. Cephalozia catenulata

10. Cells thin- or thick-walled, 20--30 x 30--35 \um at lobe base ...........  9. Cephalozia macrostachya

1. Cephalozia bicuspidata (Linnaeus) Dumortier, Recueil Observ. Jungerm., 18. 1835

Jungermannia bicuspidata Linnaeus, Sp. Pl., 1132. 1753; Cephalozia bicuspidata var. lammersiana (Hübener) Breidler; Cephalozia bicuspidata subsp. lammersiana (Hübener) R. M. Schuster; Jungermannia lammersiana Hübener

Plants brownish green, secondary pigments brown. Terminal branching common, stolons common. Stem with hyalodermis. Leaves rectangular to ovate, 1:1, insertion subtransverse, not decurrent, leaf width at widest 10--12 cells; lobe tips acute, lobe tip ending in (1--)2 uniseriate cells, lobe width 5--7 cells, lobes not connivent, sinus descending 0.5--0.6 leaf. Leaf cells thin or equally thick, 18--50 x 20--50 \um at lobe base. Specialized asexual reproduction by gemmae rarely present, round to ovoid. Sexual condition autoicous or dioicous. Female bracts 2-lobed  Perianth mouth long-denticulate to short-laciniate.

Excluded from the FNA floral area is Cephalozia bicuspidata subsp. otaruensis (Stephani) S. Hattori, following A. D. Potemkin and E. V. Sofronova (2013). These authors treated the taxon as C. hamatiloba Stephani, and cast doubt on the identity of the Southern Appalachian material reported by R. M. Schuster (1974), who was originally also uncertain.  R. E. Stotler and B. Crandall-Stotler (2016), however, did recognize this species as C. hamatiloba but without discussing the flora material. A similar plant from the northwestern coastal area of the flora merits additional study (D. H. Wagner, pers. comm.) The infraspecific taxon C. bicuspidata subsp. lammersiana (Hübener) R. M. Schuster was treated by A. D. Potemkin and E. V. Sofronova (2013) as a synonym of the typical species, which is followed here.

Subspecies 2 (1 in the flora). North America, Central America, South America, Eurasia, n Africa.

1. Leaves widest at midleaf, leaf lobes usually broadly acute; leaf cells 18--24 x (22--)30--35 \um near lobe base; underleaves often present; autoicous or dioicous ....................... .......................................................................................................................................... .......................................................................................................................................... ...................................................................  1a. Cephalozia bicuspidata subsp. ambigua

1. Leaves widest in proximal 1/3 of leaf, leaf lobes narrowly acute; leaf cells 25--50 \um wide near lobe base; underleaves lacking except in gynoecia and androecia; autoicous .......................................................................................................................................... .......................................................................................................................................... ...............................................................  1b. Cephalozia bicuspidata subsp. bicuspidata

1a. Cephalozia bicuspidata subsp. ambigua (A. Massalongo) R. M. Schuster, Hepat. Anthocerotae N. Amer. 3: 723. 1974

Cephalozia ambigua A. Massalongo, Malpighia 21: 310. 1907

Leaves widest at midleaf, leaf lobes usually broadly acute; leaf cells 18--24 x (22--)30--35 \um near lobe base. Underleaves often present. Sexual condition dioicous.

On acidic rock; low to moderate elevations; Greenland; Nunavut, Que.; Alaska, Maine, Mich., Minn., N.H., Pa.; n Europe; Asia.

Cephalozia bicuspidata var. ambigua apparently replaces the var. bicuspidata in the north of the floral area. Unlike var. bicuspidata, it is restricted to acidic rock. The perianth mouth is merely crenulate and gemmae are rare. Collections from Wisconsin and North Carolina are considered intermediates with var. bicuspidata by R. M. Schuster (1974: 721, 725).

1b. Cephalozia bicuspidata (Linnaeus) Dumortier subsp. bicuspidata

Leaves widest in proximal 1/3 of leaf, leaf lobes narrowly acute; leaf cells 25--50 \um wide near lobe base. Underleaves lacking except in gynoecia and androecia. Sexual condition autoicous.

On acidic rock, ledges, peaty and sandy soils, decorticated logs; low to high elevations; Greenland; Miquelon; Alta., B.C., Nfld. & Labrador (Nfld. & Labrador), N.B., N.S., Nunavut, Ont., P.E.I., Que., Yukon; Alaska, Calif., Colo., Conn., Del., Ga., Idaho, Ill., Ind., Iowa, Kans., Ky., Maine,  Mass., Mich., Minn., Mo., Mont., N.H., N.J., N.Y., N.C., Ohio, Oreg., Pa., Tenn., Vt., Va., Wash., W.Va., Wis., Wyo.; Mexico; Central America; South America; n Africa; Atlantic Islands (Canary Is.); Eurasia.

2. Cephalozia lacinulata (J. B. Jack ex Gottsche & Rabenhorst) Spruce, Cephalozia, 45. 1882

Jungermannia lacinulata J. B. Jack ex Gottsche & Rabenhorst, Hepat. Eur., 624. 1877

Plants green or yellow-green, secondary pigments absent. Terminal branching often present, stolons common. Stem with hyalodermis. Leaves ovate to ovate-rectangular, 1.2--1.5:1, insertion subtransverse, decurrent , leaf width at widest 4--5 cells; lobe tips sharp, lobe tip ending in 1--2(--3) cells, lobe width 2 cells, lobes not connivent, sinus descending 0.5--0.6 leaf. Leaf cells thin-walled, 30--46 x 40--46 \um at lobe base. Specialized asexual reproduction is by gemmae, rare, ovoid to ellipsoid. Female bracts 3--5-lobed. Perianth mouth laciniate.

On decaying logs in deep forest;  low to moderate elevations; Ont., N.W.T.; Mich., Minn., S.C., Wis.); Eur.; Asia (China, Japan).

Cephalozia lacinulata is rare and sporadic in occurrence in largely northern areas.

3. Cephalozia macounii (Austin) Austin, Hepat. Bor.-Amer., 14. 1873

Jungermannia macounii Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 222. 1869 [1870]

Plants green or yellow-green, secondary pigments absent. Terminal branching common, stolons often present. Stem with hyalodermis. Leaves ovate to ovate-rectangular, 1.5--1.8:1, insertion subtransverse, decurrent, leaf width at widest 6--8 cells; lobe tips narrowly lanceolate, lobe tip ending in 2 uniseriate cells, lobe width 3--4(--5) cells, lobes not connivent, sinus descending 0.5--0.7 leaf. Leaf cells thick-walled, 12--20 x 16--20 \um at lobe base. Sexual condition dioicous. Specialized asexual reproduction is by gemmae, rarely present. Female bracts 2-lobed. Perianth mouth crenulate to dentate.

On logs; low to moderate elevations; Greenland; B.C., Nfld. & Labrador (Labrador), Ont.; Alaska, Maine, Mich., Minn., N.H., Vt., Wash., Wis.; Eur., Asia (China, Japan, e Russia).

Cephalozia macounii is a rare and apparently circumboreal species.

4. Cephalozia catenulata (Hübener) Lindberg, Contr. Fl. Crypt. As., 262. 1872 [1873]

Jungermannia catenulata Hübener, Hepatocol. Germ., 169. 1834; Fuscocephaloziopsis catenulata (Huebener) Váňa & L. Söderström

Plants deep green, secondary pigments absent or olive or brown. Terminal branching occasional, stolons absent. Stem with hyalodermis. Leaves round to rounded quadrate, 1:1, insertion oblique, short-decurrent, leaf width at widest 9--10 cells; lobe tips broadly acute, lobe tip ending in 1--2 uniseriate cells, lobe width 4--5 cells, lobes little connivent, sinus descending 0.3--0.5 leaf. Leaf cells thick-walled, 15--18 x 20--25 \um at lobe base. Specialized asexual reproduction is by gemmae frequent, ovoid to angular. Sexual condition dioicous. Female bracts 2-lobed. Perianth mouth laciniate.

On decaying logs in shady forested sites; low to moderate elevations; Miquelon; Alta., B.C., Nfld. and Labrador (Nfld.), N.B., N.S., Ont., Pa., P.E.I., Que.; Ala., Calif., Conn., Del., Fla., Idaho, Kans., Ky., La., Maine, Md., Mass., Mich., Minn., Miss., N.H., N.J., N.Y., N.C., Ohio, Pa., R.I., S.C., Tenn., Vt., Va., Wash., W.Va., Wis.; Mexico; Eur., Asia; Atlantic Islands.

5. Cephalozia connivens (Dickson) Lindberg, Contr. Fl. Crypt. As., 238. 1872

Jungermannia connivens Dicks., Fasc. Pl. Crypt. Brit. 4: 19. 1801; Fuscocephaloziopsis connivens (Dicks.) Váňa & L. Söderström

Plants pale green, secondary pigments absent or olive or brown. Terminal branching common, stolons absent. Stem with hyalodermis. Leaves round to rounded quadrate, 1:1, insertion oblique or horizontal, strongly decurrent, leaf width at widest 8--12 cells; lobe tips sharp, lobe tip ending in 1 cell, lobe width 3--5(--8) cells, lobes little connivent, sinus descending 0.2--0.4 leaf. Leaf cells thin-walled, 45--55 x 40--48 \um at lobe base. Sexual condition autoicous. Specialized asexual reproduction is by gemmae rare, round or ellipsoid. Female bracts 2--5 lobed. Perianth mouth laciniate.

Given that Cephalozia connivens var. bifida R. H. Schuster and C. connivens var. compacta (Warnstorf) Nichols are not mentioned in the world checklist of hornworts and liverworts (Söderström et al. 2016), these probable synonyms are not treated in the FNA.

On moist soil, decaying logs, ditches, mires; low to moderate elevations; Greenland;  Miquelon; Alta., B.C., Nfld. and Labrador (Labrador), N.S., Ont., P.E.I., Que.; Alaska, Calif., Conn., Del., Fla., Ga., Ill., Ind., Iowa,  Kans., Ky., La., Maine, Md., Mass., Mich., Minn., Miss., N.H., N.J., N.Y., N.C., Ohio, Pa., R.I., S.C., Tenn., Tex., W.Va., Vt., Va., Wis.; Eurasia; Africa; Atlantic Islands.

6. Cephalozia leucantha Spruce, Cephalozia, 88. 1882

Fuscocephaloziopsis leucantha (Spruce) Váňa & L. Söderström

Plants pale green, secondary pigments absent or olive or brown. Terminal branching common, stolons absent. Stem with hyalodermis. Leaves ovate to ovate-rectangular, 1,2--1.5:1, insertion subtransverse, not decurrent, leaf width at widest 8--9 cells; lobe tips narrowly acute, lobe tip ending in 2 uniseriate cells, lobe width 3--4 cells, lobes not connivent, sinus descending 0.5--0.7 leaf. Leaf cells thick-walled, 10--20 x 16--30 \um at lobe base. Specialized asexual reproduction is by gemmae not uncommon, spheric to ovoid. Sexual condition dioicous. Female bracts 2--3-lobed. Perianth mouth crenulate to laciniate.

On decaying logs, cliffs and ledges, humus, peaty soil; low and moderate elevations; Greenland; B.C., Nfld. and Labrador (Nfld., Labrador), Que., Yukon; Alaska, Minn., N.Y.,  Vt., Wash.; Eur.; Asia.

7. Cephalozia loitlesbergeri Schiffner, Oesterr. Bot. Z. 62: 10. 1912

Fuscocephaloziopsis loitlesbergeri (Schiffner) Váňa & L.Söderström

Plants pale green, secondary pigments absent or olive or brown. Terminal branching common, stolons absent or rare. Stem with hyalodermis. Leaves round to rounded quadrate, 1:1, insertion oblique or horizontal, long-decurrent, leaf width at widest 8--12 cells; lobe tips narrowly acute, lobe tip ending in 2--3 uniseriate cells, lobe width 2--4 cells, lobes connivent or touching, sinus descending 0.3--0.5 leaf. Leaf cells thin-walled, 24--40 x 20--40 \um at lobe base. Specialized asexual reproduction not seen. Sexual condition autoicous. Female bracts 3--5 lobed. Perianth mouth laciniate and lobed.

On rotten wood, humic or peaty soil; low to moderate elevations; Greenland; B.C., Nfld. and Labrador (Nfld.), Que., Yukon; Alaska, Conn., Mich., Minn., Wash.; Eur.; Asia (Siberia).

8. Cephalozia lunulifolia (Dumortier) Dumortier, Recueil Observ. Jungerm., 18. 1835  F

Jungermannia lunulifolia Dumortier, Syll. Jungerm. Europe., 61. 1831; Cephalozia affinis Lindberg ex Stephani; Cephalozia media Lindberg; Fuscocephaloziopsis affinis (Lindb. ex Steph.) Váňa & L.Söderstrom; F. lunulifolia (Dumortier) Váňa & L. Söderström; Pleurocladula lunulifolia (Dumortier) Konstantinova, Vilnet & A.V. Troitsky

Plants pale green, secondary pigments absent or olive or brown. Terminal branching common, stolons absent. Stem with hyalodermis. Leaves round to rounded quadrate, 1:1, insertion almost longitudinally, long-decurrent, leaf width at widest 8--12 cells; lobe tips broadly acute, lobe tip ending in 1(--2) uniseriate cells, lobe width 2--4 cells, lobes not or little connivent, sinus descending 0.3--0.4(--0.5) leaf. Leaf cells thin-walled, 20--30 \um at lobe base. Specialized asexual reproduction is by gemmae common, angular to pyriform. Sexual condition dioicous. Female bracts 2-lobed . Perianth mouth crenulate to dentate.

Cephalozia affinis Lindberg ex Stephani is a synonym of C. lunulifolia following the argument of K. Damsholt (2002: 510), and also as reflected in the world checklist of hornworts and liverworts (L. Söderström et al. 2016). R. E. Stotler and R. Crandall-Stotler (2016) accept this species citing the molecular study of Vilnet et al. (2012), although A. D. Potemkin and E. V. Sofronova (2013) distinguish C. affinis by lobulate-dentate perianth mouth with teeth often 2--3 cells in length, while C. lunulifolia has a crenulate-dentate perianth mouth with teeth 0.5--1.5(--2) cells in length. It is, however, recognized here as simply a monoicous phase of C. lunulifolia.

In bogs among Sphagna, decaying wood, humus, soil, slopes, banks; low to moderate elevations; Greenland; Miquelon; Alta., B.C., N.B., N.S., Nunavut., Ont., Que., Yukon; Alaska, Calif., Conn., Del., Fla., Ga., Idaho, Ill, Ind., Iowa, Kans., Ky., La., Maine, Md., Mass., Mich., Minn., Mo., Mont., Nev., Ohio, Oreg., Nev., N.H., N.Y., N.C., Ohio, Oreg., Pa., Tenn., Vt., Va., Wash., W.Va., Wis., Wyo.; Eurasia; Atlantic Islands (Azores, Iceland, Madeira).

9. Cephalozia macrostachya Kaalaas, Rev. Bryol. 29: 8. 1902

Fuscocephaloziopsis macrostachya (Kaalaas) Váňa & L.Söderström; Pleurocladula macrostachya (Kaalaas) Konstantinova

Plants deep green, secondary pigments absent or olive or brown. Terminal branching occasional, stolons absent. Stem with hyalodermis. Leaves round to rounded quadrate, 1:1, insertion oblique or horizontal, short-decurrent, leaf width at widest 9--10 cells; lobe tips broadly acute, lobe tip ending in 1--2 uniseriate cells, lobe width 5--7 cells, lobes straight or connivent, sinus descending 0.3--0.5 leaf. Leaf cells thin- or thick-walled, 20--30 x 30--35 \um at lobe base. Specialized asexual reproduction is by gemmae rare, spheric to rounded ellipsoidal or pyriform. Sexual condition dioicous. Female bracts 2-lobed. Perianth mouth irregularly dentate.

Subspecies 2 (2 in the flora): Greenland, North America, Europe, Atlantic Islands.

1. Perianth strongly emergent, perianth mouth crenulate. Bracts 2- or 4-lobed, lobes acute. ..................................................................  10a. Cephalozia macrostachya subsp. australis

1. Perianth weakly emergent, perianth mouth ciliate. Bracts 2-lobed, lobes acuminate...... .......................................................... 10b. Cephalozia macrostachya subsp. macrostachya

9a. Cephalozia macrostachya Kaalaas subsp. australis R. M. Schuster, Hepat. Anthocerotae N. Amer. 3: 754, fig. 450: 7--13; 453: 11--17. 1974   E

Fuscocephaloziopsis macrostachya subsp. australis (R.M.Schuster) Váňa & L.Söderström

Plants deep green to yellow-brown. Bracts 2- or 4-lobed, lobes acute. Perianth strongly emergent, perianth mouth crenulate.

Soil in crevices, soil near stream; moderate elevations; Miss., N.C.

9b. Fuscocephaloziopsis macrostachya subsp. macrostachya

Plants yellow to olive. Bracts 2-lobed, lobes acuminate. Perianth weakly emergent, perianth mouth ciliate.

Peaty soil, bogs, occasionally sandy soil on slopes; low to moderate elevations; Greenland; Conn., Fld., Ga., La., Maine, Mass., Miss., N.H., N.Y., N.C., R.I., Tex., Vt., Va.; Europe; Atlantic Islands (Iceland).

10. Cephalozia pachycaulis R. M. Schuster, Bryologist 96: 632, fig. 1--2. 1993

Fuscocephaloziopsis pachycaulis (R. M. Schuster) Váňa & L.Söderström

Plants deep green, secondary pigments absent or olive or brown. Terminal branching occasional, stolons absent. Stem without hyalodermis. Leaves round to rounded quadrate, 1:1, insertion oblique, not decurrent, leaf width at widest about 20 cells; lobe tips narrowly acute, lobe tip ending in (1--)3 uniseriate cells, lobe width 8--12 cells, lobes little connivent, sinus descending 0.4--0.6 leaf. Leaf cells thin-walled, 30--40 x 40--50 \um at lobe base. Specialized asexual reproduction is by gemmae occasional, subspheric to ellipsoidal. Sexual condition autoicous. Female bracts 2-lobed. Perianth mouth crenulate-denticulate, occasionally lacinulate.

Soil under rock overhang. tundra slope; moderate elevations; Alaska; Asia (Kuril Islands, Russian Far  East, Siberia).

11. Cephalozia pleniceps (Austin) Lindberg, Morgonbladet (Helsinki) 1882(388): 3. 1882

Jungermannia pleniceps Austin, Proc. Acad. Nat. Sci. Phyiladelphia 21: 222. 1869; Fuscocephaloziopsis pleniceps (Austin) Váňa & L. Söderström

Plants pale green, secondary pigments absent or rarely olive or brown. Terminal branching occasional, stolons often present. Stem with hyalodermis. Leaves round to rounded quadrate, 1:1, insertion oblique or horizontal, little-decurrent, leaf width at widest 12--25 cells; lobe tips short-acute, lobe tip ending in 1(--2) uniseriate cells, lobe width 6--10 cells, lobes little connivent, sinus descending. Leaf cells thin-walled, 20--60 x 24--50 \um at lobe base. Specialized asexual reproduction is by gemmae not uncommon, ellipsoidal to fusiform. Sexual condition autoicous. Female bracts 2-lobed. Perianth mouth acute-crenulate.

1. Leaves usually with sinus 0.4--0.5 leaf length, sinus narrowly U-shaped .......................... .............................................................................................................................................. ......................................................................... 11c.  Cephalozia pleniceps var. sphagnorum

1. Leaves usually with sinus 0.3--0.4 leaf length, sinus broadly U-shaped,

2. Stolons common; leaves obliquely inserted, concave, some leaf lobes connivent, median leaf cells 25--45 \um; Arctic and northern areas  11a. Cephalozia pleniceps var.  pleniceps

2. Stolons absent; leaves sublongitudinally inserted, little concave, lobes not connivent, median leaf cells 36--50 \um; Southern Appalachians .......................................................  .......................................................................................................................................... ....................................................................... 11a. Cephalozia pleniceps var. caroliniana

Varieties 3 (3 in the flora): North America, Eurasia, Atlantic Islands.

11a. Cephalozia pleniceps var. caroliniana R. M. Schuster, Hepat. Anthocerotae N. Amer. 3: 780. 1974  E

Fuscocephaloziopsis pleniceps var. caroliniana (R. M. Schuster) Váňa & L.Söderström

Stolons absent.  Leaves sublongitudially inserted, rarely concave, lobes not connivent, sinus 0.3--0.4 leaf length, broadly U-shaped. Leaf cells medially 36--50 \um.

Mineral soil, stream edge; moderate elevation; N.C.

11b. Cephalozia pleniceps var. pleniceps

Fuscocephaloziopsis pleniceps (Austin) Váňa & L. Söderström var. pleniceps

Stolons common.  Leaves obliquely inserted, concave, some leaf lobes connivent, sinus 0.3--0.4 leaf length, sinus broadly U-shaped. Leaf cells medially 25--45 \um.

Wet acidic or calcareous rock, peaty soil, with Sphagna; moderate to high elevations; Greenland; Alta., B.C., Nfld. and Labarador (Nfld., Labrador), N.S., N.W.T., Nunavut, Que., Yukon; Alaska, Calif., Conn., Ind., Maine, Mass., Mich., Minn., Mont., N.H., N.Mex., N.Y., N.C., Ohio, Oreg., Vt., Wash., Wis., Wyo.; Eurasia; Atlantic Islands (Iceland).

11c. Cephalozia pleniceps var. sphagnorum (C. Massalongo) E. Jörgensen, Bergens Mus. Skr. 16: 266. 1934

Cephalozia symbolica var. sphagnorum C. Massalongo, Malpighia 21: 306. 1907

Stolons usually absent. Leaves sublongitudinally inserted, little concave, lobes rarely connivent, usually with sinus 0.4--0.5 leaf length, sinus narrowly U-shaped. Leaf cells medially 36--50 \um.

Peat bogs; low to moderate elevations; Minn., N.Y.; c Europe.

L. Söderström et al. (2016) recognize the var. sphagnorum as Lophoziopsis polaris var. sphagnorum (R. M. Schuster) Konstantinova & Vilnet; however, R. E. Stotler and B. Crandall-Stotler (2017) recognize the species as Fuscocephaloziopsis pleniceps var.  sphagnorum (C. Massalongo) Stotler & Crandall-Stotler. The distribution is little understood but is apparently transcontinental within the range of the var. pleniceps, and this taxon is generally misidentified as the typical variety.

EXCLUDED to other genera

Cephalozia albescens (Hooker) Dumortier (Pleurocladula albescens (W. J. Hooker) Grolle; Pleuroclada albescens (W. J. Hooker) Spruce, Pleuroclada albescens var. islandica (Nees) Spruce) This species is recognized in FNA as Pleurocladula]

5. SCHOFIELDIA J. D. Godfrey, Bryologist 79: 314. 1976 • [For W. B. Schofield, American bryologist and ecologist, 1927--]

Judith Dean Godfrey

Geoffrey A. Godfrey

Plants robust, translucent, clear green. Stem lacking a hyalodermis; both Frullania-type and lateral-intercalary branching. Leaf cells lacking oil-bodies. Underleaves present. Asexual reproduction by (1--)2-celled fasciculate gemmae originating from elongate, specialized stalks produced on or between reduced, clasping leaves at the apices of gemmiparous shoots. Androecia with antheridial jacket cells not tiered, stalks biseriate. Gynoecia anisomorphic; perianths fleshy and multistratose at the base. Sporophyte seta consisting of 8 epidermal plus 4 inner cells rows; capsule wall 2(--3)-stratose.

Species 1 (1 in the flora): North America, restricted to the Pacific Northwest, and ne Asia (Kamchatka Peninsula, Russia).

SELECTED REFERENCES Bakalin V. A. 2005. Schofieldia and Cryptocoleopsis. The new genera of liverworts (Hepaticae) to the flora of Russia. Botanical Journal 90: 594--603. Godfrey, J. D. 1976. Schofieldia, a new Hepatic from the Pacific Northwest. Bryologist 79: 314--320. Godfrey, J. D. and G. A. Godfrey. 1980. Notes on Hepatics from the Pacific Northwest. Bryologist 83: 224--228. Schuster, R. M. 1993. On Cephalozia pachycaulis sp. nov. and the perimeters of Cephalozia. Bryologist 96: 619--625. Schuster, R. M. 1995. Notes on nearctic Hepaticae. XX. On Schofieldia and evolution in the Cephalozioideae. Fragm. Flor. Geobot. 40: 39--46.

1. Schofieldia monticola J. Godfrey, Bryologist 79: 314. 1976

Plants translucent, clear, bright green, lacking secondary pigmentation, with dead shoots becoming whitish or brownish-tinged, ca. 8.0--15 x 1.5--3 mm (varying to 2.3--3.6 x ca. 1.1 mm in southern extreme of range in Oregon); usually robust, succulent, brittle, growing in crowded, dense turfs, horizontal to slightly ascending at stem apices. Stems stout, fleshy, brittle, whitish-green; in transverse section somewhat walled, hyaline; with both Frullania-type and lateral-intercalary branching. Leaves succubous, symmetrically 2(--3)-lobed, with unlobed half-leaf at base of Frullania-type branch; large, somewhat asymmetrical, broader than long (width 1--1.35 x the length), ovate to ovate quadrate, 1280--1960 x 1300--2400 \um; lobes broadly triangular; margins usually entire, occasionally becoming undulate, infrequently with a large hooked tooth or "thumb" near the antical leaf base; cells of lobe apices and margins 30--62 x 20--48 \um; median cells 40--94(--100) x 30--52 \um; basal cells more elongated, 54--125 x 25--62 \um. Underleaves often obscured by rhizoids; distant; highly variable in form, ranging from small flaps 3--5 cells broad at the base by 3 or more cells high, to larger structures, but not exceeding one-third the size of the leaves; irregularly lobed with variably rounded lobes, or occasionally lanceolate. Asexual reproduction by fasciculate gemmae, few to abundantly produced in masses from elongate, specialized stalks produced on or between reduced, clasping leaves at the apices of gemmiparous shoots; greenish, spheric or more usually elliptic to ovoid; 20--34 x 14--20 \um; (1--)2-celled, thin-walled. Sexual condition dioicous. Androecia with bracts ca. 1040 x 760 \um; antheridia spherical to rounded-elliptical, ca. 160--200 \um. Gynoecial bracts 1160--1696 x (680--)794--1440 \um; bracteole smaller, 640--1480 x 600--1256 \um. Perianth mouth crenulate-denticulate with 1--2-celled teeth, 162--248 x 64--112 \um, mostly formed by projecting ends of mouth cells. Sporophyte seta ca. 220 \um in diameter; epidermal cells 60--70 \um, interior cells 32--54 \um; elaters 112.5--280 \um, 7.5--14 \um in diameter; spiral thickenings 3--4 \um wide. Spores light brown, papillose, 12.5--16 \um.

Sporophytes infrequent, capsules mature late July--late Oct., depending on snow conditions. Seepy slopes or banks of snow-melt rills, streamlets or ponds, sometimes more or less exposed, but usually partially to completely shaded by heathers, subalpine meadows or alpine (tundra) areas; 1200--2000 m, descending lower to 500--1200 m in coastal Alaska and extreme nw B.C.; B.C.; Alaska, Oreg., Wash.; ne Asia (Kamchatka, Russia). 

Hepatic species commonly associcated with Schofieldia monticola include Calypogeia azurea, Lophozia opacifolia, L. wenzelii, Moerckia blyttii, Nardia geoscyphus, N. japonica and Pleurocladula albescens. The Oregon plants examined were growing isolated among mosses, or in patches of reduced-sized plants on soil, rather than in the characteristic tight, luxuriant, pure mats that occur in the core area of its range to the north. Leaf cell sizes of the Oregon plants fit the typical range, but elsewhere plants ranged much smaller. The easternmost locality known for Schofieldia (R. M. Schuster 1995) is in the Interior Wet Belt of British Columbia, on the windward western slopes of the Selkirk Mountain Range (one of four ranges in the Columbia Mountains). Many coastal species reappear far inland in this Interior Wet Belt, which is separated from the Coast Mountains by a dry interior zone approximately 350 km wide, in which Schofieldia would not be expected to occur. R. M. Schuster (1995) described plants from the northernmost known location in the Talkeetna Mountains, Alaska, as being few, poorly developed and reduced in size. While placing these plants into Schofieldia monticola, he noted that they are "of considerable interest in that they seem to occupy an “uncomfortable midground between the very plesiomorphic Cephalozia pachycaulis and Pleurocladula.” In contrast, W.B. Schofield's specimen from Chisik Island, in the same general vicinity but a more coastal location further to the west, is quite typical for Schofieldia in growth habit and appearance. Schofieldia monticola is abundant and widespread in its coastal, Pacific Northwest range, and had been assumed for over 30 years to be endemic to western North America.  However, the recent reports of this species, apparently from a single locality in Kamchatka, Russia, extend the known range into northeastern Asia.