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Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden

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XX. CEPHALOZIELLACEAE Douin

David H. Wagner

 

Plants forming mats or mixed among other bryophytes; branches usually intercalary or axillary, rarely terminal, arising laterally or postically, half leaves not produced.  Leaves alternate, transverse or obliquely inserted,  succubous, plane or concave, 2-lobed, entire or toothed; underleaves present or absent, simple, seldom 2-lobed.  Rhizoids scattered, sparse on erect shoots, sometimes numerous on prostrate shoots. Specialized asexual reproduction by 2-celled ovoid or angular, gemmae, sometimes frequent at shoot apices.  Androecia immediately below female bracts, on short lateral or postical branches, or terminal on leading shoots and then often becoming intercalary; bracts in 1--3 pairs when paroicous or on short side branches, up to 10--15 pairs on long shoots. Gynoecium terminal on leading, leafy shoots or on short lateral or postical branches, bracts larger than leaves, 2-lobed, bracteoles free to connate with bracts up to 0.7.  Perianth with or without subfloral branches, perianth exserted over half, weakly to strongly plicate, usually narrowed to mouth, mouth cells usually thick-walled, elongate and often free at tips, perigynium absent. Capsules ovoid, the valves 2-stratose; seta with four large epidermal cells and four minute internal cells.

 

Genera  2 , species ca. 90 (2 genera and 22 species in the flora): Widespread in all continents, North America, South America, Europe, Asia, Africa, Australia, Pacific Islands, Antarctica..

 

Cephaloziellaceae is widespread but poorly known because the species are small and inconspicuous. As often as not, species are first noticed in bulk collections where mixed among other bryophytes. It is characterized by its small size, 2-lobed leaves and well developed perianths. Most taxonomically useful characters are not easily determined in the field. Accurate identification of species requires careful study of microscope preparations.

 

 

1. Leaves mostly inserted transversely, occasionally at angles up to 30--40°; perianth mouth cells differentiated from those of body of perianth, mostly very long, usually more than 3:1, usually thick-walled                1. Cephaloziella, page XXX

1. Leaves inserted at an oblique angle, 45° or more; perianth mouth cells usually not differentiated, occasionally slender with acute apex  2. Cylindrocolea, p. XXX

 

 

SELECTED REFERENCES  Damsholt, K. 2013. The Liverworts of Greenland. Nordic Bryological Society, Lund, Sweden. Schuster, R. M.  1980. The Hepaticae and Anthocerotae of North America. Vol. IV. Columbia University Press, New York, U.S.A. Stotler, R. E. and B. Crandall-Stotler. 2017. A synopsis of the liverwort flora of North America, North of Mexico. Annals of the Missouri Botanical Garden 102: in press1. CEPHALOZIELLA (Spruce) Schiffner, Hepat. in Engler & Prantl, Nat. Pflanzenfam. 1(3): 98. 1893 * [Greek kephale, head, ozot, branch, and Latin -ella, small, alluding to small Cephalozia, a genus with an enlarged terminal infloresence]

 

Plants 2--12 x 0.2--0.65 mm; in thin mats or open turf, in patches, intertwined with other bryophytes, or ascending to suberect shoots overtopping other bryophytes, light yellowish green to dark green, often pigmented red, purple to brown or black. Stems 50--180 \um in diameter, slender, flexuous, terete, wiry to rigid, fleshy or brittle; cortical cells quadrate to short retangular, walls thin to strongly thickened, smooth to verrucose or mammillose, 12--18 x 8--11 \um to 27--52 x 16--24 \um; sparingly branched, axillary or intercalary from terminal, lateral or postical positions; subgynoecial innovations often frequent. Rhizoids sparse on erect shoots, often numerous on recumbent shoots. Leaves mostly transverse, three species with weakly to strongly oblique insertion, from erect to widely spreading, distant to contiguous or tightly imbricate, ovate to quadrate or obtrapezoidal, deeply 2-lobed, typical leaf length 150--250 \um, divided 0.4--0.9. Leaf lobes narrowly lanceolate, long-triangular to short, rounded triangular, 3--10(--14) cells wide at base, spreading to concave, incurved or conduplicate; apex usually acute but some species with most lobe tips obtuse to rounded; sinus acute, rectangulate, obtuse or rounded, plane or reflexed; margins entire, serrate, dentate to spinose; cells at base of lobes (7--)9--14(--20) \um wide; cell walls thin to strongly thick-walled; cuticle smooth, verrucose, papillose, mammillose or spinose. Oil bodies ovoid to ellipsoidal, small, faintly papillose to clear, equal to or smaller than chloroplasts, 3--4 x 1.3--2.4 \um. Underleaves absent to large, subulate, ovate, lanceolate, or 2-lobed, from only 1--3 cells long to 0.5 length of the leaves. Gemmae gemmae 2-celled, ovoid, ellipsoidal to fusiform, usually smooth or in two regional species angulate, green, often tinged with dark red, purple or brown. Sexual condition dioicous, autoicous, heteroicous or paroicous. Androecia immediately below female bracts, on short lateral or postical branches, or terminal on leading shoots and then often becoming intercalary; bracts in 1--3 pairs when paroicous or on short side branches, up to 10--15 pairs on long shoots, larger than leaves, strongly concave and imbricate, margins entire to dentate or spinose. Gynoecia terminal on long leafy shoots, on short lateral or postical branches; bracts larger than leaves, ovate, 2-lobed 0.2--0.4, red to purplish with decolorate margins; lobes acute and triangular, sometimes divided; margins usually denticulate to spinulose, rarely entire; cells thick-walled, smooth to verrucose or papillose; bracteoles free to connate with bracts up to 0.7; lobe triangular to 3--lobed. Perianth fusiform, ellipsoid, or ovoid, moderately to strongly plicate, usually emergent over 0.5 its length and narrowed to mouth, green, often tinged with red, brown, or purple, sometimes nearly black, 0.5--2.2 mm, mouth usually crenulate, mouth cells sometimes with slender, free tips, thick-walled, becoming thicker at distal end, 1.5--6 times longer than wide.

 

Species 75--85 (20 in the flora); widely distributed on all continents including Antarctica.

 

Cephaloziella is characterized by its small size, 2-lobed leaves, frequent red, purple, or brown pigmentation, oil bodies present in all cells, regular presence of underleaves and bracteoles on fertile shoots, underleaves sometimes also on sterile shoots, and small, opaque stem cells. The genus most likely to be confused with Cephaloziella is Cephalozia, which is uniformly green, lacks oil bodies, lacks underleaves on all shoots except immediately below gynoecia, and has a hyalodermis on the stem. Eremonotus lacks oil bodies except sporadically, lacks underleaves, has vestigial bracteoles, and is usually black or olive brown but not reddish. Sphenolobopsis differs in preponderance of terminal branching and being largely epiphytic.

 

Cephaloziella is notorious for being among the most challenging genera of liverworts. The plants are so small that microscope examination is usually necessary for positive identification. Although some species are distinctive and easily recognized, some groups of similar species have numerous environmental variants that are the source of overlapping character states which blur species distinctions. Almost every commonly used character in keys is subject to exceptions, including sexuality and presence of underleaves. Sterile specimens are frequently impossible to determine confidently. The consequence is a nomenclatural history that is complex, with inconsistent and often contradictory definitions of taxa.

 

The present treatment is based largely on the work of R. M. Schuster (1980, 1988), K. Damsholt (2013) and J. Paton (1999). The arrangement and choice of taxa treated here relies primarily on L. Söderström et al. (2016). This taxonomy is unquestionably provisional. For this reason, no attempt is made to delineate infrageneric taxa. Molecular data are essentially nonexistent; what little has been done merely emphasizes the dire necessity of a massive, global study. Progress will depend on assiduous field work to gather fresh material for both molecular study and creation of a bank of high quality photomicrographs.

 

Distribution records provided here are based in large part on literature records. Because so many herbarium specimens have been misidentified in the past without recent review, these records are likely to include a larger proportion of innacuracies than most of the other liverwort genera.

 

SELECTED REFERENCES Damsholt, K. 2002. Illustrated Flora of Nordic Liverworts and Hornworts. Nordic Bryological Society, Lund, Sweden. Damsholt, K. 2013. The Liverworts of Greenland. Nordic Bryological Society, Lund, Sweden. Hong, W. S. 1986. The family Cephaloziellaceae in North America west of the Hundredth Meridian. Bryologist 89: 155--162. Paton, J. 1999. The Liverwort Flora of the British Isles. Harley Books, Colchester, England. Schuster, R. M.  1980. The Hepaticae and Anthocerotae of North America. Vol. 4. Columbia University Press, New York, U.S.A. Schuster, R. M.  1988. The Hepaticae of South Greenland. Nova Hedw. Beih. 92: 1--225. Söderström, L., A. Hagborg, M. von Konrat, S. Bartholomew-Began, D. Bell, L. Briscoe, Elizabeth Brown, D. C. Cargill, D. P. da Costa, B. J. Crandall-Stotler, E. D. Cooper, G. Dauphin, J. Engel, K. Feldberg, D. Glenny, S. R. Gradstein, X. Hu, J. Heinrichs, J. Hentschel, A. L. Ilkiu-Borges, T. Katagiri, N. A. Konstantinova, J. Larraín, D. Long, M. Nebel, T. Pócs, F. Puche, E. Reiner-Drehwald, M. Renner, A. Sass-Gyarmati, A. Schäfer-Verwimp, J. G. Segarra-Moragues, R. E. Stotler, P. Sukkharak, B. Thiers, J. Uribe, J. Váňa, J. Villarreal, M. Wigginton, L. Zhang, and R. Zhu. 2016. World checklist of hornworts and liverworts. PhytoKeys 59: 1--828 (29 Jan 2016). Stotler, R.E. and B. Crandall-Stotler. 2017. A synopsis of the liverwort flora of North America, North of Mexico. Annals of the Missouri Botanical Garden 102: in press.

 

1. Leaves of sterile shoots regularly dentate, serrate or spinose to the base.

                2. Leaves divided 0.5--0.6(--0.7); dioicous.

                                3. Stems and leaves evenly and densely covered with sharp papillae

                                    …..14. Cephaloziella spinicaulis

                                3. Leaves with abaxial, round papillae, mammillae or with conical projections,

                                    papillae seriate on stems below leaf insertion     …..2. Cephaloziella aspericaulis

                2. Leaves divided 0.6--0.8; autoicous or paroicous.

                                4. Underleaves absent     …..20. Cephaloziella turneri

                                4. Underleaves prominent, to half size of leaf.

                                                5. Leaf lobes narrowly lanceolate, 2--4(--5) cells wide at base

                                                    …..15. Cephaloziella spinigera (in part)

                                                5. Leaf lobes acute triangular to ovate triangular; 4--7 cells wide at base.

                                6. Autoicous; leaves shallowly dentate with few basal

                                      projections; cuticle verrucose       …..10. Cephaloziella massalongi

                                6. Paroicous; leaves with long marginal teeth; abaxial leaf

                                    surface with numerous projections; cuticle smooth or papillose..12. Cephaloziella phyllacantha

1. Leaves of sterile shoots entire, rarely weakly or sporadically dentate, sometimes with 1or few

     basal teeth.

                7. Leaves divided 0.75--0.8; lobes 2--4(--5) cells wide at base; basal teeth frequent.

                                8. Leaf cells 12--16 /um wide, walls thin to moderately thickened; leaves erect to

                                    squarrose; growing almost exclusively on Sphagnum     …..4. Cephaloziella elachista

                                8. Leaf cells (7--)9--12(--16) /um wide, walls usually strongly thickened; leaves

                                    erect to erecto-patent; growing in a variety of wetland habitats .15. Cephaloziella spinigera (in part)

                7. Leaves divided 0.5--0.7; lobes (5--)6--11(--14) cells wide at base; basal teeth rare or

                    absent.

                                9. Underleaves absent on sterile, non-gemmiferous shoots.

                                                10. Paroicous; lobes 3--6 cells wide at base       …..13. Cephaloziella rubella

                                                10. Autoicous; lobes 6--9 cells wide at base.

                                                                11. Leaves contiguous to remote on sterile shoots; bracts and

                                                                      bracteole united to 0.65, tightly sheathing at base; gemmae

                                                                      ellipsoidal; widespread, common in temperate regions

                                                                      …..7. Cephaloziella hampeana

11. Leaves closely imbricate on all shoots; bracts and bracteole

      connate to 0.8 and forming a wide mouthed, tubular involucre;

      gemmae angular; rare, subarctic only      …..19. Cephaloziella integerrima

                                9. Underleaves present on sterile, non-gemmiferous shoots, small to 0.5 leaf

                                    length.

                                                12. Paroicous.

                                                                13. Female bracts dentate; arctic and subarctic       .….1. Cephaloziella arctogena

                                                                13. Female bracts edentate; mostly from temperate latitudes.

                                                                                14. Lobe base cells 14--20 /um wide       …..16. Cephaloziella stellulifera

                                                                                14. Lobe base cells 7--10 /um wide       …..5. Cephaloziella elegans

                                                12. Autoicous or dioicous.

                                                                15. Leaf lobe apex obtuse to rounded on mature shoots,

                                                                      occasionally acute.

                                                                                16. Leaves patent, concave; lobe apex obtuse to

                                                                                      occasionally acute; autoicous; widespread in arctic to

                                                                                      subarctic and alpine zones       ..…18. Cephaloziella varians

                                                                                16. Leaves spreading, plane; lobe apices consistently

                                                                                      rounded; dioicous; endemic to North Carolina

                                                                                      …..11. Cephaloziella obtusilobula

                                                                15. Leaf lobe apex consistently acute to acuminate.

                                                                                17. Dioicous; leaf lobes ascending and more or less

                                                                                      conduplicate from a spreading base; abaxial leaf surface

                                                                                      often with multicellular projections, mammillae or

                                                                                      spines; common in all regions      …..3. Cephaloziella divaricata

                                                                                17. Autoicous; leaf lobes divergent; leaves plane to evenly

                                                                                      concave; abaxial leaf surface smooth (except

                                                                                      C. mammillifera); rare or with restricted distribution.

                                                                                                18. Leaves widely spreading; lobes plane; endemic

                                                                                                      to the Southeast       …..8. Cephaloziella hyalina

                                                                                                18. Leaves erect to patent; lobes concave to

                                                                                                      incurved; boreal to subarctic.

                                                                                                                19. Abaxial leaf surface mammillose below

                                                                                                                      sinus       …..9. Cephaloziella mammillifera

                                                                                                                19. Abaxial leaf surface smooth.

                                                                                                                                20. Leaf apex incurved to hooked at

                                                                                                                                      apex; apical cells elongate and

                                                                                                                                      sharp pointed . 17. Cephaloziella uncinata

                                                                                                                                20. Leaf apex acute but neither

                                                                                                                                      incurved nor hooked

                                                                                                                                      …..6. Cephaloziella grimsulana

 

 

1. Cephaloziella arctogena (R. M.  Schuster) Konstantinova, Arctoa 3: 126. 1994

 

 Cephaloziella rubella (Nees) Warnstorf var. arctogena R. M. Schuster, Meddel. Grønland 199: 314. 1974; C. rubella (Nees) Warnstorf subsp. arctogena (R. M.  Schuster) R. M. Schuster & Damsholt

 

Plants 2--6 x 0.2--0.5 mm; in patches or intertwined with other bryophytes, pale green but leaf lobes often claret-red. Stems 75--80 \um in diameter, flexuous; cortical cells quadrate, thick-walled, smooth, 16--21 x (10--)12--14(--15) \um; branches all intercalary. Leaves transverse, suberect, obliquely spreading to more or less squarrose, contiguous to remote, narrowly quadrate to obtrapezoidal, 175--200 x 155--200 \um, divided 0.5--0.75. Leaf lobes ovate lanceolate to lanceolate, (5--)6--8(--9) cells wide at base, erect, moderately concave; apex acute, terminating in 1 or 2 superposed cells; sinus acute; margins entire; cells at base of lobes (7--)8--12(--13) \um wide; cell walls thick, smooth; cuticle smooth. Oil bodies to 10--16 per cell. Underleaves distinct throughout, usually well developed, lanceolate or bidentate to 2-lobed, to 0.35--0.5 the length of leaves. Gemmae absent. Sexual condition paroicous, rarely heteroicous. Androecia below female bracts, rarely spicate and intercalary on long shoots; bracts in 2--3 pairs, lobes (10--)12--15 cells broad, entire or subentire. Gynoecia terminal on leafy shoots; bracts 0.3 2-lobed, connate with bracteole and forming a tight sheath around lower 0.3--0.5 of perianth; lobes acute and triangular, sharply denticulate with 1-celled teeth set at nearly right angles to margins; bracteoles united with bracts. Perianth elongate and exserted for more than half its length, reddish tinged medially, decolorate near mouth, 0.8--0.9 mm, mouth crenulate, mouth cells 1.6--3.5 times longer than wide.

 

On damp, peaty soil; low to moderate elevations; circumboreal at subarctic latitudes; Greenland; Alaska; Eurasia.

 

Cephaloziella arctogena has been recognized at the species rank by both L. Söderström et al. (2016) and R. E. Stotler and B. Crandall-Stotler (2017). It is not a strong species. R. M. Schuster (1988) refrained from recognizing this as a species and K. Damsholt (2013) likewise maintained it as a subspecies of Cephaloziella rubella. Cephaloziella arctogena is distinguished by presence of underleaves, wider leaf lobes (8--12 cells wide versus 3--6 cells wide for C. rubella), and a more northern distribution.

                                       

2. Cephaloziella aspericaulis Jørgensen, Bergens Mus. Skr.[Norg. Leverm.] 16: 197. 1934

 

Cephaloziella byssacea var. aspericaulis (Jørgensen) R. M.  Schuster

 

Plants 10 x 0.25--0.36 mm; in patches or as solitary shoots among other bryophytes, green with reddish brown, orange or purplish tinge. Stems 80--90 \um in diameter, filiform, terete; cortical cells quadrate, thick-walled,  23 x 10--12 \um; with papillae or mammillae below abaxial leaf base, otherwise smooth; branches intercalary or sometimes terminal. Leaves transverse, erect to erecto-patent, distant, broadly quadrate to obtrapezoidal, 160--180 \um, divided 0.6--0.7. Leaf lobes ovate triangular, 6--8 cells wide at base, patent, moderately concave; apex acute, often terminated by a unicellular spine; sinus narrow, acute; margins spinose to dentate, teeth 1-celled; cells at base of lobes 7--10 \um wide; cell walls thick; cuticle papillose; leaf cells abaxially mammillose to spinose and usually mammillose down the stem immediately below the leaves. Oil bodies unknown. Underleaves distinct, frequent, oblong ovate, 2-lobed or lanceolate, 0.5 of the size of the leaf. Gemmae frequent, ellipsoidal, purplish. Sexual condition presumed dioicous. Androecia unknown. Gynoecia terminal; bracts larger than leaves, 2-lobed 0.3--0.4 bract length; lobes triangular, denticulate; bracteoles united to one or both bracts. Perianth known only in an immature state; mouth crenulate, mouth cells 3--5 times longer than wide.

 

On rock, in cliff crevices, or on gravel; low to moderate elevations; Greenland, Alaska; Eurasia.

 

The distinction between Cephaloziella aspericaulis and the scabra phenotype of Cephaloziella divaricata is subtle. The leaves of C. aspericaulis are distinctly dentate and the protuberances on the abaxial surface of the leaves continue down the stem, sometimes quite dramatically.

 

3. Cephaloziella divaricata (Smith) Schiffner, Hepat. in Engl. & Prantl, Nat. Pflanzenfam. 1(3): 99. Sep. 1893

 

Jungermannia divaricata Smith, Engl. Bot. 10: tab. 719. 1800.; Cephaloziella divaricata (Smith) Schiffner var. scabra (M. Howe) Haynes; C. divaricata (Smith) Schiffner var. asperifolia (Taylor) Damsholt; C. polystratosa (R. M. Schuster & Damsholt) Konstantinova

 

Plants 3--8(--12) x (0.12--)0.16--0.45 mm; scattered, lax shoots among other bryophytes or in dense, tight cushions, dark green, reddish brown to black. Stems (65--)70--90 \um in diameter, relatively stout, terete; cortical cells quadrate to short-rectangular, moderately thick-walled, smooth, 16--24(--30) x 12--13(--14) \um; branches intercalary, lateral or postical; often with subgynoecial innovations. Leaves transverse, base usually more or less spreading but lobes loosely folded upward and erect, remote, contiguous to imbricate, broadly quadrate to obtrapezoidal, 150--200 \um, divided 0.5--0.8. Leaf lobes ovate-triangular to ovate lanceolate, (5--)6--9(--10) cells wide at base, moderately folded inwards to erect; apex acute, terminated by 1 or 2 cells; sinus acute to rectangulate, often somewhat reflexed; margins entire or rarely with occasional tooth; cells at base of lobes (8--)9--12(--13) \um wide; cell walls thin to strongly thickened; cuticle smooth, rarely faintly papillose. Oil bodies 2--7(--8) per cell. Underleaves distinct throughout even though small, variable, minute and subulate to ovate and 2-lobed, 2--3 cells wide to 4--5(--6--7) cells wide. Gemmae frequent, green to purplish black. Sexual condition dioicous. Androecia prominent, spicate, becoming intercalary on long, leafy shoots; bracts in 5--15 pairs, strongly concave, divided 0.5, margins usually entire, often purplish red. Gynoecia terminal on long leafy shoots; bracts much larger than leaves, 0.3--0.4 2-lobed, purplish red, cell walls thick; lobes subacute to acuminate, denticulate or serrulate to spinulose dentate; bracteoles united with one or both bracts up to 0.5 their length. Perianth fusiform-ovoid, 0.5--0.65 emergent, obtusely 4-plicate distally, green to reddish purple, 0.7--1.4 x 0.3--0.5 mm, mouth crenulate, mouth cells often less than 2 times longer than wide.

 

On diverse substrates from among Sphagnum in bogs, on rocks, sand, inorganic soil, wood to embedded in cryptobiotic soil crusts of dry grasslands; sea level to high elevations; Greenland; Alta., B.C., N.B., Nfld. and Labrador, N.S., Nunavut, Ont., Que.,Yukon; Alaska, Ark., Calif., Colo., Conn., Idaho, Ill., Iowa, Kans., Ky., Maine, Md., Mass., Mich., Minn., Mo., Mont., Nebr., Nev., N.H., N.J., N.Mex., N.Y., N.C., Ohio, Oreg., Pa., R.I., S. Dak., Tenn., Tex., Utah,  Vt., Va., Wash., West Va., Wis., Wyo.; on all continents.

 

Cephaloziella divaricata is the most variable of all the species of Cephaloziella yet most specimens of this species are placed here with reasonable confidence. Most remarkable is the difference between forms having leaves which are quite smooth on their abaxial surface to forms with leaves bearing branched, spinose-lamellate projections below the sinus. This range of cellular ornamentation may be found in a single colony, even on different parts of the same shoot, a well known phenomenon that has probably prevented an even greater proliferation of names in the past. It is to be expected that future work, particularly using data from molecular analysis, will uncover quit distinct genetic entities from among the various forms.

 

The species with closest resemblance is C. aspericaulis but the restricted, boreal-arctic distribution of that presents few problems. In western North America C. stellulifera is equally abundant in the southern part of its range, especially Oregon and California. It is critical to recognize the paroicous inflorescences of C. stellulifera as distinct from C. divaricata. Cephaloziella divaricata is always described as dioicous but occasional autoicous plants have been observed. In either case, the presence of androecia in a collection will rule out the paroicous species. In western North America Cephaloziella stellulifera is distinguished by edentate female bracts and long perianth mouth cells which may be free at the tip. Its leaves are consistently smooth abaxially in contrast to the forms of C. divaricata with armed leaves. Female bracts of C. divaricata are always dentate and the perianth mouth cells are mostly short and form a crenulate margin without free tips.

 

4. Cephaloziella elachista (J. B. Jack) Schiffner, Sitzungsber. Deutsch. Naturwiss.-Med. Vereins Böhmen "Lotos" Prag 48[n.s. vol. 20]: 338. 1900

 

Jungermannia elachista J. B. Jack ex Gottsche & Rabenhorst, Hepat. Eur., 574. 1873; Cephaloziella grimsulana (J. B. Jack ex Gottsche & Rabenhorst) Lacouture var. angustiloba (Douin) Jørgensen

 

Plants 4--12 x 0.25--0.33 mm; creeping among Sphagnum, rarely forming dense patches, pale to yellow-green. Stems (50--)60--80(--90) \um in diameter, flexuous; cortical cells rectangular, thin-walled, smooth, 27--52(--74) x 16--24 \um; branches lateral intercalary. Leaves almost transversely inserted, erect spreading to squarrose, remote, obtriangular to obtrapezoidal, 135--215 \um, divided 0.6--0.8. Leaf lobes narrow, divaricate, lanceolate, 3--4(--5) cells broad at base, erect, spreading; apex acuminate, usually terminating in 2--3 superposed cells; sinus acute to obtuse; margins entire or with one or two sharp, 1--2 celled teeth near lobe bases; cells at base of lobes (10--)12--16(--20) \um wide; cell walls thin or rarely somewhat thickened; cuticle smooth or weakly papillose. Oil bodies 5--6 per cell. Underleaves frequent, distinct, filiform to lanceolate or 2-lobed with linear divisions, (40--)70--110 \um long. Gemmae occasional, pale green, clustered on ascending, flagelliform shoots. Sexual condition autoicous. Androecia on short postical shoots, spicate, occasionally intercalary; bracts somewhat larger than leaves, contiguous to imbricate, margins spinose-dentate. Gynoecia on short postical branches or short lateral branches; bracts much larger than leaves, imbricate, 2-fid to 0.5--0.6 of bract length, often reddish purple; lobes acuminate, sharply spinose-dentate, teeth often curved or reflexed, 1--2-celled; bracteoles divided 0.4--0.45, free when young, often united 0.2--0.3 with bracts when mature. Perianth fusiform-ovoid, long exserted when mature, green to reddish purple, 2 x 0.6 mm, mouth crenulate, mouth cells strongly narrowed cells, 3--6 times longer than wide.

 

Closely associated with Sphagnum in boreal peatlands, typically found as thin strands embedded in Sphagnum; low to moderate elevations; Alta., B.C., N.B., Nfld. and Labrador, N.S., Nunavut, Ont., Que.; Alaska, Conn., Ill., Ind., Maine, Mass., Mich., Minn., N.H., N.J., N.Y., N.C., R.I., Vt., West Va., Wash., Wis.; Eurasia.

 

Cephaloziella elachista is largely restricted to Sphagnum bogs, where it is nearly ubiquitous in tundra ecosystems from Alaska across Canada. Cephaloziella spinigera occurs regularly in the same habitats but its distribution extends further southwards into wetlands which are not dominated by Sphagnum. They may be difficult to differentiate; C. elachista has larger leaf cells with thinner cell walls and usually has basal spinose teeth well developed.

 

5. Cephaloziella elegans (Heeg) Schiffner, Sitzungsber. Deutsch. Naturwiss.-Med. Vereins Böhmen "Lotos" Prag 48[n.s. vol. 20]: 336. 1900

 

Cephaloziella elegans Heeg, Rev. Bryol. 20 (5):82. 1893; Cephaloziella rubella (Nees) Warnstorf var. elegans (Heeg) R. M. Schuster

 

Plants 2--6 x 0.2--0.3 mm; small, dense mats, decumbent to ascending, deep green to brownish green, only rarely reddish brown. Stems 70--80 \um in diameter, flexuous; cortical cells quadrate, thick-walled, smooth, 17--25 x 15--18 \um; branches originating from lateral leaf axils or as innovations from below perianth. Leaves transverse, relatively erect, little spreading, distant to imbricate towards stem apices, quadrate to obtrapezoidal, relatively broad, circular-quadrate when flattened, 120--150(--210) \um, divided 0.5--0.75. Leaf lobes broadly triangular or ovate triangular, 6--9(--11) cells wide at base, moderately concave; apex acute, terminating in 1 or 2 superposed cells; sinus acute; margins entire; cells at base of lobes (7--)9--10 \um wide; cell walls thin to moderately thickened; smooth. Oil bodies 3--5 per cell. Underleaves distinct throughout on sterile shoots, lanceolate to 2-lobed, when lanceolate of 3--4 cells to 2-lobed and of 6--8(--16) cells. Gemmae occasional, greenish, thin-walled. Sexual condition paroicous. Androecia always located immediately below female bracts; bracts in 3--4 pairs, 2 x sterile leaves, saccate, the lobes with margins entire to crenulate denticulate. Gynoecia terminal on leafy shoots; bracts often carmine red or vinaceous, somewhat larger than male bracts; lobes 0.5 2-fid into triangular lobes, finely but sharply to strongly denticulate; bracteoles united for about half their length with the bracts, forming a 5--6 lobed sheath around perianth base. Perianth cylindrical, exserted beyond bracts, deeply plicate, green to reddish, 0.7 mm, mouth crenulate, mouth cells 1.5--4 times longer than wide.

 

On thin soil, often over rocks; low to moderate elevations; Ont., Que.; Alaska, Iowa, Maine, Minn., N.Y., Vt., Wis.; Eurasia.

 

Cephaloziella elegans is a segregate from Cephaloziella rubella that is based on vegetative characters: wide leaf lobes with small cells and large underleaves. K. Damsholt (2002, 2013) maintains it as a subspecies of C. rubella.

 

 

6. Cephaloziella grimsulana (J. B. Jack ex Gottsche & Rabenhorst) Lacouture, Hépat. France: 52. 1905

 

Jungermannia grimsulana J. B. Jack ex Gottsche & Rabenh., Hepat. Eur., 526. 1872

 

Plants 5--18(--20) x 0.25--0.3 mm; slender and attenuate, often crowded in dense tufts, green with ruby-red to purplish-red tinge, often blackish, shiny. Stems 85--130 \um in diameter, filiform and slender; cortical cells rectangular, thin to moderately thick-walled smooth, 30--50 x 15--20 \um; branches mostly lateral intercalary. Leaves transverse, erect to suberect, distant, oblong to rotund-quadrate, 200--235 \um, divided 0.5--0.6. Leaf lobes triangular to ovate-triangular, (5--)6--8(--10) cells wide at base, often incurved; apex acute to obtuse; sinus acute; margins entire; cells at base of lobes (13--)14--18 \um wide; cell walls thin to slightly thick-walled; smooth. Oil bodies (4--)5--12 per cell. Underleaves distinct throughout on sterile shoots, ovate to lanceolate, sometimes 2-lobed, less than 0.5 the length of the leaves. Gemmae often abundant, ruby red to deep purplish red. Sexual condition autoicous. Androecia terminal, becoming intercalary on elongate shoots; bracts up to 10--15 pairs, imbricate, erect, saccate, crenulate to serrate. Gynoecia on elongated leafy shoots; bracts larger than leaves; lobes 2--3 lobed, acute, coarsely crenulate-dentate, decolorate at margin; bracteoles divided 0.25--0.35, often united with bracts and forming an urn like sheath around base of perianth. Perianth ovoid to ellipsoid, 3--4 plicate, narrowed to mouth, red to blackish-purple, apex decolorate, average 0.8 mm, mouth crenulate, mouth cells 3--6 times longer than wide.

 

On alpine tundra soil, vertical rock faces, or in crevices, often with other bryophytes; low elevations; Greenland; N.B., Nunavut, Ont., Que.; Alaska; Northern Europe, Russia.

 

7. Cephaloziella hampeana (Nees) Schiffner ex Loeske, Moosfl. Harz.: 92. 1903

 

Jungermannia hampeana Nees, Naturgesch. Eur. Leberm. 3: 560. 1838

 

Plants 2--8 x 0.3 mm; prostrate in dense patches or creeping sparsely, erect only when growing among other bryophytes, green to brownish green. Stems 80--90 \um in diameter, terete, flexuous; cortical cells short to long rectangular, walls thin to moderately thick, smooth, 35--56 x 15--17 \um, irregularly and distantly branched. Leaves transverse, spreading to squarrose, distant to contiguous, subquadrate to obtrapezoidal, 190--210(--225) \um, divided 0.5--0.75. Leaf lobes narrowly ovate-triangular to lanceolate-triangular, (5--)6--9(--10) cells wide at base, more or less divaricate; apex acute; sinus acute to rectangulate; margins entire, rarely with a small, sharp basal tooth; cells at base of lobes (10--)11--15(--19) \um wide; cell walls thin; cuticle smooth. Oil bodies 4--6 in median leaf cells. Underleaves absent on sterile shoots. Gemmae frequent, greenish or brownish, ellipsoidal. Sexual condition autoicous. Androecia on postical branches below female shoot, occasionally on leading shoot; bracts imbricate, erect, reddish brown to purplish, erect, strongly concave, 0.35--0.5 2-lobed, entire to sparingly denticulate. Gynoecia on long, leading leafy shoots, infrequently on short postical branches; bracts larger than leaves, with spreading lobes; lobes acute to subacuminate, denticulate with small but sharp, 1-celled teeth; bracteoles smaller and connate with bracts for 0.45--0.65 their length, forming a distinct sheath at base of perianth. Perianth oblong-ovate to cylindrical, exserted for 0.35--0.6 its length, 4--5 plicate, green or red to purple or black, distal third decolorate in pigmented forms, to 2.2 x 0.7 mm, mouth crenulate, mouth cells narrow, fingerlike, 3--6 times longer than wide.

 

Most common on organic substrates or on soil, in pure patches or mixed in with mosses, occasionally on damp, shady rocks; low to moderate elevations; Greenland; Alta., B.C., N.B., Nfld. and Labrador, N.S., Nunavut, Ont., Que.; Ala., Alaska, Ariz., Calif., Colo., Conn., Idaho, Ill., Ind., Iowa, Kans., Ky., Maine, Md., Mass., Mich., Minn., Mo., Mont., N.H., N.J., N.Mex., N.Y., N.C., Ohio, Oreg., Pa., R.I., Tenn., Tex. ,Vt., Va., Wash., West Va., Wis., Wyo.; Eurasia.

 

The complete absence of underleaves on all shoots of Cephaloziella hampeana is characteristic. Autoicous shoots are usually readily observed as the androecia and gynoecia are typically developed in proximity.

 

8. Cephaloziella hyalina Douin, Mém. Soc. Bot. France 29: 77. 1920 E

 

Cephaloziella hyalina Douin var. rappii (Douin) R. M. Schuster

 

Plants 2--12 x 0.3--0.5 mm; creeping or suberect, light green to (rarely) light brown. Stems 50--60(--75) \um in diameter, terete, slenderly flexuous; cortical cells quadrate, slightly to strongly thick-walled, smooth, 20--35 x 18--20 \um; branches postical-intercalary, often flagelliform. Leaves slightly to strongly obliquely inserted, usually widely spreading but not squarrose, plane to weakly concave, remote to subcontiguous, obovate to obdeltoid, 180--210 \um, divided 0.6--0.7. Leaf lobes slender and narrowly lanceolate-triangular, (3--)4--6(--7) cells wide at base, spreading to subsquarrose, divergent; apex acute, terminating in 1(--2) superposed cells; sinus acute; margins entire; cells at base of lobes (12--)13--17(--19) \um wide; cell walls moderately thickened; cuticle verrucose or papillate. Oil bodies 6--12 per cell. Underleaves vestigial to small but distinct, rarely absent, tiny projections to 2--3-fid , 1--2(--3--4) cells wide, to 5--7 cells long. Gemmae rare, greenish to reddish, ellipsoidal-fusiform. Sexual condition autoicous (rarely paroicous). Androecia on short postical shoots, occasionally intercalary on long shoots; bracts in 4--6 pairs, 2-lobed, forming a distinctly spicate androecium, erect to secund, entire or weakly crenulate. Gynoecia on short postical branches; bracts twice the size of leaves, ovate, 2-fid 0.35--0.5; lobes 8--12 cells broad, acute, margins subentire to somewhat dentate serrate; bracteoles free (rarely up to 0.35 fused with bracts). Perianth shortly cylindrical, narrowed above, 3--4-plicate distally, 0.5--0.6 emergent, hyaline, green or pale brownish, mouth not decolorate, 0.9--1 x 0.4--0.5 mm, mouth cells sharply projecting , often curved to incurved at their tips, 2.5--4.5(--5) longer than wide.

 

On soil or among mosses; low elevations (up to 765 m); Ala., Ark., Fla., Ga., Miss., N.Y., N.C., Ohio, endemic to eastern U.S.A.

 

Cephaloziella hyalina is an interesting local species. It resembles C. hampeana in its spreading leaves with plane, divergent leaf lobes but is distinguished by the distinctly oblique insertion of leaves.

 

9. Cephaloziella mammillifera R. M.  Schuster & Damsholt, Phytologia 63: 327. 1987

 

Plants 5--12.5 x 0.18--0.25 mm; shoots long and stringy, usually mixed with other bryophytes, greenish brown, always with purplish or purplish black secondary pigmentation. Stems 80--110 \um in diameter, long and stringy, rigid; cortical cells short retangular, thick-walled, distal cortical cells protuberant, often serially, 14---20 x 10--14 \um; branches remote, lateral and ventral intercalary, rarely terminal. Leaves transverse, erect or suberect, slightly concave, distant, rounded-quadrate to short oblong, 160--170 \um, divided 0.6--07. Leaf lobes narrow, ovate or ovate triangular, acute, (6--)7--9(--10) cells wide at base, slightly divergent; apex acute; sinus mostly acute; margins entire; cells at base of lobes 8--10(--13) \um wide; cell walls thin to equally thick-walled; cuticle smooth, frequently with mammillate, balloon-shaped or tumid abaxial leaf cells. Oil bodies unknown. Underleaves small, distinct below gynoecia, smaller or vestigial on vegetative shoots, deeply 2-lobed when maximal, to 8 cells long x 10 cells wide. Gemmae frequent, green to reddish at maturity. Sexual condition autoicous, possibly dioicous. Androecia terminal, becoming intercalary; bracts spicate, somewhat saccate, of 8--10 pairs, 2-lobed 0.3--0.6. Gynoecia terminal on longer main shoots; bracts larger than leaves, ovate, 2-lobed 0.2--0.4, red to purplish with decolorate margins; lobes triangular, acute or subacute, denticulate to obscurely serrulate; bracteoles 2--3-lobed, connate less than 0.35. Perianth oblong ovoid, narrowed and plicate above, exserted 0.5--0.75 its length, reddish to purplish, decolorate near mouth, mouth crenulate, mouth cells with alternating elongate and shorter rectangular cells, 3.5--6 times longer than wide.

 

On wet ledges; low elevations; s Greenland.

 

Cephaloziella mammillifera is known from so few collections that a clear distinction from either Cephaloziella varians or C. divaricata is questionable. R. M. Schuster (1988) mentioned that he was often unable to affirm that autoicous material was at hand and therefore practically indeterminable except when the distinctive mammillae are well developed.

 

10. Cephaloziella massalongoi (Spruce) Müller Frib., Lebermoose 2(17): 191. 1913

 

Cephalozia massalongoi Spruce, Cephalozia: 71. 1882 (as massalongi)

 

Plants 3--6(--8) x 0.15--0.3 mm; in patches or intertwined with other bryophytes, pure green to brownish green, rarely reddish brown or blackish. Stems 50--70(--75) \um in diameter, flexuous, smooth; cortical cells short rectangular, very thick-walled, lumina elliptical and guttulate, smooth,14--20(--23) x 9--16 \um; freely but irregularly branched, postical and lateral. Leaves transverse, stiffly spreading but not squarrose, patent, lobes erect, often conduplicate, remote to subcontiguous, subquadrate, 120--155 \um, divided 0.6--0.9. Leaf lobes lanceolate to narrowly triangular, acute, (5--)6--7(--8--9) cells wide at base, incurved or sometimes adaxially concave; apex acute, terminating in 2 superposed cells; sinus acute, plane to slightly reflexed; margins dentate-serrate with spinose teeth, rarely edentate; cells at base of lobes (8--)10--14(--15) \um wide; cell walls usually thick; cuticle typically coarse and often with contiguous papillae, never totally smooth; leaves sometimes with abaxial protuberances. Oil bodies 2--5(--6) per cell. Underleaves regularly present, prominent, usually 2-lobed, with 1-several teeth, their apices often incurved, large and conspicuous, apical cell 15--42 \um. Gemmae brown to brownish or purplish red. Sexual condition autoicous, often freely fertile. Androecia usually loosely spicate, often close to gynoecia; bracts similar to leaves in size and dentition but more concave and with suberect lobes. Gynoecia terminal on a leading leafy stem or on short postical branches; bracts larger than leaves and similar in shape or somewhat longer than wide, similarly 2-fid or rarely 3-fid; lobes more spinose dentate than leaves, many teeth 2-celled, cells thick-walled, cuticle prominently verrucose; bracteoles as wide as bracts, rarely narrower; slightly connate. Perianth ellipsoidal, plicate above, green to brownish green, to 0.8 mm, mouth cells dentate to spinous dentate, 4--6 times longer than wide.

 

On moist soil or rocky detritus; low to moderate elevations; Greenland, B.C., Nfld. and Labrador; N.C., Tenn., Vt.; Eurasia.

 

Cephaloziella massalongoi is largely restricted to copper-bearing substrates in Europe. In North America, its occurrence on substrates not clearly associated with copper indicates a fundamental genetic difference might be at hand.

 

11. Cephaloziella obtusilobula R. M.  Schuster, Hepat. Anthocerotae N. Amer. 4: 108. 1980 E

 

Plants 5--12 x 0.3 mm; shoots mostly as loose, decumbent strands, grass green. Stems 90--100 \um in diameter, somewhat fleshy; cortical cells short rectangular, walls thin to slightly thickened, smooth, 17--25 x 11--14 \um; branches very rare, lateral or postical. Leaves obliquely inserted on mature shoots, spreading to squarrose, remote, broad, rounded oblong, 200--280 \um, divided 0.6--0.7. Leaf lobes very broad, ovate-triangular to ellipsoidal, 8--11 cells wide at base, nearly plane, moderately spreading; apex obtuse-rounded at apex; sinus obtuse, occasionally rectangular; margins entire; cells at base of lobes 12--14(--15) \um wide; cell walls thin to moderately thickened; very delicately striolate. Oil bodies (2--)4--7(--8) per cell. Underleaves frequent, variable, ovate or ovate lanceolate to shallowly 2-lobed, 2 cells wide by 3--4 cells long to 6--7 cells wide by 10--11 cells long. Gemmae frequent, pale green. Sexual condition dioicous. Androecia unknown. Gynoecia on short, postical branches; bracts connate 0.5--0.6 and similarly united to bracteole, forming a tubular sheath; lobes irregularly 3-lobed, margins subentire, serrate, cells thick-walled; bracteoles obscurely 2-lobed. Perianth known only from immature gynoecia, size unknown, green, mouth cells slightly narrowed and free at tips, 2--3 times longer than wide.

 

On exposed cliff faces or moist soil along streams; rarely at low elevation, otherwise at moderate elevations (1400--1500 m); Ga., N.C., endemic to southeast U.S.A.

 

12. Cephaloziella phyllacantha (C. Massalongo & Carestia) Müller Frib., Lebermoose 2(17):194. 1913

 

Anthelia phyllacantha C. Massalongo & Carestia, Nuovo Giorn. Bot. Ital. 12: 340. 1880

 

Plants 10 x 0.2--0.3 mm; in thin patches or intertwined with other bryophytes, green to whitish green, usually with pale brown or purplish tinges, distal parts decolorate. Stems 50--70(--75) \um in diameter, brittle, smooth, rigid, stout; cortical cells quadrate to short rectangular, thick-walled, smooth, 12--18 x 8--11 \um; simple or with lateral or ventral intercalary banches. Leaves transverse, erect to erecto-patent, almost sheathing the stem, distant to contiguous, roounded-quadrate, 150 \um, divided 0.5--0.8. Leaf lobes ovate triangular, 4--6 cells wide at base, incurved; apex acute to acuminate; sinus acute, often reflexed or convoluted; margins dentate with 1--2-celled, hyaline teeth; cells at base of lobes 10--12 \um wide; cell walls very thick; smooth or somewhat verrucose on marginal teeth, often with abaxial, spiny projections. Oil bodies (1--)2--4(--5) per cell. Underleaves regularly present, distinct, oblong-ovate, 1--2-lobed, dentate, almost half the size of the leaves. Gemmae ellipsoidal, 2-celled, green to brown, red or purple. Sexual condition autoicous or paroicous. Androecia terminal or intercalary; bracts like leaves, with both marginal and abaxial teeth. Gynoecia terminal or on short lateral branches; bracts larger than leaves, divided 0.4--0.5 into 2--3 lobes, with abaxial spines; lobes dentate serrate with oblique, thick-walled teeth; bracteoles basally connate to one or both bracts, 1--2-lobed, dentate. Perianth ovoid, exserted 0.6--0.7 of its length, plicate, contracted at mouth, pale green, occasionally tinged purplish or brown and decolorate at mouth, to 0.7 mmm, mouth crenulate to setose, mouth cells elongate, thick-walled, 4--6 times as long as wide.

 

On rock or damp soil in rock crevices, rarely on wood; low to high elevations, Greenland; B.C.; Oreg., Wash.; Europe.

 

Cephaloziella phyllacantha appears to be much more abundant in western than eastern North America. The western forms are characterized by large, prominent papillae on leaf cells. This species is reported to be associated with C. massalongi on copper bearing substrates in central Europe but no such association is evident in North America.

 

13. Cephaloziella rubella (Nees) Warnstorf, Krypt.-Fl. Brandenburg, Leber- Torfm. 1(2): 231. 1902

 

Jungermannia rubella Nees, Naturgesch. Eur. Leberm. 2: 236. 1836; Cephaloziella rubella (Nees) Warnstorf var. bifida (Schreber ex Schmidel) Douin; C. rubella (Nees) Warnstorf var. pulchella (C. E. O. Jensen) R. M. Schuster; C. rubella (Nees) Warnstorf var. sullivantii (Austin) Müller Frib.

 

Plants 1--6 x 0.3 mm; creeping or in dense patches, pale or brownish green, usually tinged bright red, at least in reproductive shoots. Stems 70--85 \um  in diameter, flexuous; cortical cells quadrate to rectangular, thick-walled, smooth, 13--16 x 10--13 \um; branches postical and lateral. Leaves transverse, suberect, obliquely spreading to more or less squarrose, contiguous to slightly spaced, quadrate to obtrapezoidal, 170(--200) \um, divided 0.5--0.75. Leaf lobes ovate triangular to lanceolate, (3--)4--5(--6--10) cells wide at base, erect to squarrose; apex acute, terminating in 2(--3) superposed cells; sinus acute; margins entire or rarely with small basal tooth; cells at base of lobes (8--)11--17(--19) \um wide; cell walls thin to (usually) thick; cuticle smooth. Oil bodies (4--)6--11 per cell. Underleaves absent or present in some forms, subulate, of few cells when present, minute. Gemmae frequent, green. Sexual condition autoicous, paroicous, or heteroicous. Androecia below gynoecia, on separate shoots or intercalary; bracts 4--6 or more on separate shoots, 1--3 pairs below gynoecia, margins denticulate. Gynoecia on elongate, leafy shoots; bracts larger than leaves, usually 2-lobed; lobes ovate-triangular, sharply denticulate to dentate-serrate, erect to spreading, rarely squarrose; bracteoles united to bracts, forming a sheath closely surrounding the perianth base. Perianth fusiform-ovoid, exserted for 0.5--0.75 its length, 4-plicate and sulcate, green, often reddish purple below, 0.6--0.8 mm, mouth crenulate, mouth cells narrow, elongate, 1.5--4 longer than wide.

 

On a variety of terrestrial habitats, rarely on wood or trees, at all elevations; Greenland; Alta., B.C., Man., N.B., Nfld. and Labrador, N.S., Nunavut, Ont., Que., Sask., Yukon; Ala., Alaska, Ariz., Calif., Colo., Conn., Del., Idaho, Ill., Ind., Iowa, Kans., La., Maine, Md., Mass., Mich., Minn., Mo., N.H., N.J., N.Mex., N.Y., N.C., Ohio, Oreg., Pa., R.I., S.C., S. Dak., Tenn., Vt., Va., Wash., West Va., Wis., Wyo.; Eurasia.

 

Of all the species of Cephaloziella, C. rubella has the most confusing nomenclatural history. There is a welter of named forms which may or may not have taxonomic validity. The subspecific taxa listed here as synonyms are probably as distinctive as the two segregates treated as species: C. arctogena and C. elegans. Their sexuality may be labile within a single collection, so whether or not a plant is autoicous, paroicous or heteroicous is not a consistently reliable character. The vegetative characters that supposedly chacterize them are likewise variable and overlap insensibly.

 

 

 

14. Cephaloziella spinicaulis Douin, Mém. Soc. Bot. France 29: 62. 1920  C

 

Plants 1--5 x 0.25--0.33 mm; ascending to suberect shoots over other bryophytes, pale grayish green. Stems 80--180 \um in diameter, rigid and stout, conspicuously papillose; cortical cells quadrate, thin-walled, stongly spinose, 18--20 x 12 x 15 \um; usually freely branched, lateral or postical intercalary, sometimes axillary or abundantly terminal. Leaves transverse, erect or erect-spreading, distant, quadrate to rotund quadrate, 120--160 \um, divided 0.5---0.6. Leaf lobes ovate to ovate triangular, (4--)6--7(--9) cells wide at base, folded upwards; apex acute to subacuminate; sinus acute; margins spinose dentate; cells at base of lobes (9--)11--14(--15) \um wide; cell walls thin; cuticle smooth. Oil bodies unknown in North America. Underleaves present throughout, lanceolate to 2-fid, 50--135 x 50--80(--90) \um. Gemmae rare, green, elliptical to ovate. Sexual condition dioicous. Androecia on leading shoots; bracts concave, ornamented like vegetative leaves. Gynoecia terminating a clavate shoot; bracts similar to leaves, free; lobes triangular; bracteoles free, unlobed. Perianth poorly known, mouth lobulate, greenish, probably to 0.7 mm, mouth cells tapered, acute, free 0.3--0.7 their length, 2--3 times longer than wide.

 

On damp soil in hardwood forests; low to high elevations; Conn., Mass., N.C., Tenn., Vt., Va.; e Asia (Japan).

 

Cephaloziella spinicaulis is a very rare and distinctive species. The disjunction between eastern North America and eastern Asia is well known in flowering plants and appears to be demonstrated here.

 

15. Cephaloziella spinigera (Lindberg) Jørgensen, Hedwigia 53: 224. 1913

 

Cephalozia spinigera Lindb., Musci Scand. 4. 1879 [1880]. Cephaloziella spinosa Douin

 

Plants 2--8 x 0.1--0.2 mm; creeping among mosses, rarely forming dense patches, pale or yellowish green through brownish red to purplish copper. Stems 50--60 \um in diameter, flexuous, filiform; cortical cells long rectangular, thick-walled, smooth, 25--35 x 12--15 \um; branches postical or rarely postical. Leaves transverse, suberect to patent, remote to subcontiguous, obtriangular to obtrapezoidal, 200 \um, divided 0.7--0.8. Leaf lobes narrowly lanceolate, 2--4(--5) cells broad at base, erect, widely spreading; apex acuminate , usually terminated by 2 elongated cells; sinus narrow, acute; margins entire to sinuose-dentate, often basally spinose; cells at base of lobes (8--)9--12(--20) \um wide; cell walls thick; with delicate, hyaline, elliptical papillae on lobes and striolate at leaf base and on stems. Oil bodies (4--)6--11 per cell. Underleaves distinct on sterile shoots, 2-fid or subulate, small, often inconspicuous. Gemmae occasional, greenish, 2-celled. Sexual condition autoicous. Androecia on short postical shoots adjoining gynoecia; bracts of 3--4 pairs, often purplish, concave, erect, imbricate, 0.5--0.6 2-lobed, coarsely dentate on lobes. Gynoecia on postical, ventral intercalary branches; bracts large, not connate, 0.5--0.75 2-lobed, rarely 3--4 lobed; lobes slenderly triangular lanceolate, sharply pointed, sinuose to spinose dentate, curved teeth rare; bracteoles usually 2-lobed, connate at base only. Perianth fusiform-ovoid, long exserted, trigonous or obscurely 4-gonous, narrowing distally, hyaline, usually copper red below mouth, 0.8--1.5 mm, mouth crenulate, mouth cells narrow and elongated, 3--6 times as long as wide.

 

Growing over exposed peat, living Sphagnum or other fen mosses; low elevations; Greenland; Alta., B.C., N.B., Nfld. and Labrador, N.W.T., Nunavut, Que.,Yukon; Alaska, Calif., Fla., Ill., Maine, Maine, Mich., Minn., Mont., N.H., N.Y., Oreg., Wash.; Eurasia.

 

Cephaloziella spinigera is very similar to C. elachista, from which it differs in having smaller leaf cells with thicker walls. Unlike C. elachista it may grow in fens and wetlands in the absence of Sphagnum.

 

16. Cephaloziella stellulifera (Taylor ex Carrington and Pearson) Crozals, Rev. Bryol. 30(2): 31. 1903

 

Jungermannia stellulifera Taylor ex Carrington and Pearson, Hep. Brit. Exsicc., 32. 1878

 

Plants 8(--15) x 0.5(--0.65) mm; in small patches to dense mats, pale yellowish or brownish green, rarely dark reddish brown to purplish or almost black. Stems 60--130(--150) \um in diameter, flexuous, sometimes geniculate; cortical cells rectangular quadrate, moderately thick-walled, smooth, 23--33 x 14--24 \um; branches sparse, nearly always postical. Leaves transverse, erect to patent, sometimes squarrose, distant to contiguous, quadrate to obtrapezoidal, 220(--280) \um, divided 0.6--0.8. Leaf lobes lanceolate to lanceolate triangular, acute, 4--8 cells wide at base, erect to squarrose; apex acute, terminating in 2 superposed cells; sinus subacute to obtuse, plane or slightly recurved; margins entire; cells at base of lobes (12--)14--18(--20) \um wide; cell walls thin or slightly thickened; cuticle smooth or occasionally obscurely verrucose. Oil bodies (4--)5--8(--11) per cell. Underleaves present on all shoots, subulate to lanceolate, apical cell 12--28 \um, few celled and subulate to lanceolate or orbicular, 0.3 the length of the leaves. Gemmae fairly common, green, reddish or brownish purple. Sexual condition paroicous or rarely autoicous. Androecia hypogynous or sometimes on a lateral branch; bracts erect to squarrose, entire or minutely serrate, often only slightly saccate. Gynoecia on elongated, leafy shoots; bracts much larger than leaves, erect to patent or squarrose; lobes acute, ovate-triangular, entire (in North America), apex 2-celled; bracteoles 0.3--0.7 connate, unlobed. Perianth cylindrical, gradually tapered to relatively open mouth, brownish green, to 1.7 x 0.6 mm, perianth mouth crenulate, mouth cells to 0.3 free at tips, 3.5--6 times longer than wide.

 

On shaded soil or stony substrate, rarely tree bases or organic substrates; low to high elevations; Calif., Oreg., Utah, Wash.; South America; Eurasia; Africa.

 

Although Cephaloziella stellulifera has been reported from eastern North America (R. M. Schuster 1980; R. Stotler and B. Crandall-Stotler 2017), none of the records can be verified as valid. Even though he treated this species as occurring in the east, R. M. Schuster (1980) explicitly stated he had not seen any clearly authentic specimens. It appears to be a member of the biographical group whose distribution is disjunct between northern Europe and western North America while absent from eastern North America. See C. divaricata for differentiation from that species, which is very similar.

 

17. Cephaloziella uncinata R. M.  Schuster, Meddel. Grønland 199: 316. 1974

 

Cephaloziella uncinata R. M. Schuster var. brevigyna R. M. Schuster & Damsholt; C. uncinata R. M. Schuster var. sphagnicola R. M. Schuster

 

Plants 6--12 x 0.25--0.50(--0.60) mm; in patches or as solitary shoots among other bryophytes, yellow to olive-green with light brown secondary pigmentation, reddish in reproductive shoots. Stems 100 \um in diameter, rigid, fleshy, flexuous; cortical cells quadrate to rectangular, thick-walled, smooth, 17--20 x 12--15 \um; branches mostly lateral intercalary. Leaves transverse, erect or suberect, distant, rounded-quadrate to short ovate, 150--220 \um, divided 0.6--0.7. Leaf lobes narrow, ovate to lanceolate, (5--)6--9(--10) cells wide at base, erect or incurved; apex acute, often uncinately incurved at apex; sinus acute; margins entire or occasionally with a small basal tooth; cells at base of lobes (10--)11--14(--15 \um wide; cells thick-walled to very thick-walled with concave trigones; cuticle smooth to coarsely verrucose, sometime with prominent abaxial mammillose thickenings. Oil bodies (2--)3--7(--10) per cell. Underleaves indistinct, vestigial except at apex of vigorous shoots, small, lanceolate to 2-lobed, 3--5(--7) cells long. Gemmae rare, green, brownish or purplish. Sexual condition autoicous (rarely paroicous). Androecia terminal or on intercalary branches, spicate, of 6--25 pairs of bracts; bracts 2-lobed 0.4--0.5, saccate at base, lobes erect, incurved, usually serrulate to denticulate. Gynoecia at apex of intercalary branch; bracts 2-lobed 0.3--0.4, greenish to rarely reddish; lobes ovate triangular to ovate, acute, usually serrulate, apex acuminate and incurved, reddish tinge; bracteoles 2-lobed, connate 0.2--0.6, forming an urn like sheath around the perianth. Perianth oblong-ovoid, 4 plicate in distal two thirds, narrowed at the mouth, light brown or reddish, decolorate distally, to 0.9 mm or more, mouth crenulate to serrulate-dentate, mouth cell ends up to 0.3 free, 4--7 times longer than wide.

 

On peaty ground, sometimes with Sphagnum, or on calcareous wet ledges and crevices; low to high elevations; Greenland; Que.; Alaska, Colo.; Eurasia.

 

Cephaloziella uncinata  is very similar to Cephaloziella varians but differs in having hooked leaf lobe apices and in often having male and female branches arising in close proximity.

 

 

 

18. Cephaloziella varians (Gottsche) Stephani, Wiss. Ergebn. Schwed. Südpolar-Exped. 1901--1903 4(1): 4. 1905

 

 Jungermannia varians Gottsche, Int. Polarforsch., Deutsch. Exped. 2: 452. 1890

 

Plants 2--5 x 0.180--0.3 mm; in small patches, brownish green to purplish brown or purplish black. Stems 70--85\um in diameter, rigid; cortical cells quadrate to short rectangular, walls thin to strongly thickened, smooth, 23--36 x 11--15 \um; freely branched, normally intercalary. Leaves transverse, erect, suberect, or obliquely patent, weakly to moderately concave, distant to approximate, subquadrate to short-oblong, 170--225 \um, divided 0.6--0.7. Leaf lobes narrowly ovate to broadly ovate triangular, 6--9(10--14) cells wide at base, little divergent, erect; apex acute, often more or less incurved; sinus acute; margins entire; cells at base of lobes 11--14 \um wide; cell walls thin to strongly thick-walled; cuticle smooth to verruculose in thick-walled forms. Oil bodies 2--7 per cell. Underleaves usually present throughout, lingulate to lanceolate, 3--6 cells long x 2--4 cells wide. Gemmae frequent, usually violet. Sexual condition autoicous, usually fertile. Androecia frequent, subtending unfertilized gynoecia or on postical branches; bracts violet black, narrowly spicate, 5--15 pairs, somewhat saccate, 0.5 2-lobed, entire, apices incurved. Gynoecia on elongate, leafy shoots, subinvolucral innovations frequent, these usually male; bracts broadly ovate to subquadrate, 0.45--0.6 2-lobed; lobes broadly ovate-triangular to ovate, entire to finely denticulate, tinged vinaceous or purplish, margins decolorate; bracteoles 2-lobed, usually considerably smaller than bracts, united for 0.1--0.3 its length with bracts. Perianth oblong to oblong clavate, tinged vinaceous or purplish, margins decolorate in 2--5 cell rows, 0.5--0.7 mm, mouth crenulate, mouth cells weakly projecting, 1.5--2 times longer than wide.

 

Over damp soil, in rock crevices, on ledges, often calcareous; low to moderate elevations in northern regions, only high elevations in western North America south of Canada; Greenland; Alta., B.C., Nfld. and Labrador, Man., N.W.T., Nunavut, Que., Yukon; Alaska, Colo., Maine, Mich., Minn., Vt., Wyo.; Eurasia; Antarctic.

 

19. Cephaloziella integerrima (Lindberg) Warnstorf, Krypt.-Fl. Brandenburg, Leber- & Torfm. 1(2): 232. 1902

 

Cephalozia integerrima Lindberg, Acta Soc. Sci. Fenn. 10: 502. 1875

 

Plants 4 x 0.5 mm; in thin mats or open turfs, pale yellowish green to dark brown or reddish brown. Stems 60--100 \um in diameter, flexuous; cortical cells quadrate to short retangular, thin to moderately thick-walled, smooth, 19--26 x 12--20 \um; branches lateral, seldom postical. Leaves transverse, patent to erecto-patent, less often suberect or squarrose, contiguous to subimbricate, obtriangular, broadest above middle, 100--255 \um, divided 0.4--0.7. Leaf lobes triangular, 4--8(--10) cells wide at base, patent, suberect, to squarrose; apex acute or sometimes rounded; sinus acute to slightly rounded; margins entire; cells at base of lobes (10--)12--18(--20) \um wide; cell walls walls thin or somewhat thickened; cuticle smooth. Oil bodies 4--8 per cell. Underleaves absent on sterile shoots. Gemmae common, with 4--6 angles, green or brown to reddish brown or reddish purple. Sexual condition autoicous. Androecia on lateral branches; bracts edentate or weakly serrate, lobes rounded to acuminate. Gynoecia terminal on leading shoots; uppermost subinvolucral leaves and underleaf free; bracts and bracteole to 650 \um, 0.5--0.8 connate, occasionally only on one side of bracteole; lobes of bracts divided to 0.3; bracteole lobes ovate-triangular, acute to rounded. Perianth ellipsoid, slightly plicate, pale green to dark reddish-brown, 0.5--1.6 mm, mouth crenulate, mouth cells 3.5--6 times longer than wide.

 

On damp, exposed to shaded, sandy to clayey soil, sometimes a pioneer on disturbed sites; low elevations; Greenland; Alaska; Europe.

 

Except for Greenland material (R. M. Schuster 1988), most specimens of Cephaloziella integerrima from North America that have been determined as C. integerrima are both sterile and lack gemmae, so that only a general resemblance to authentic material justifies such a placement. Identity with European material of these records remains to be clearly established.

 

20. Cephaloziella turneri (Hooker) Müller Frib., Lebermoose 2(17): 202. 1913

 

Jungermannia turneri Hooker, Brit. Jungermann. pl. 29. 1816

 

Plants 2--7 x 0.4(--0.5) mm; typically in dense, tight mats, pale yellow-green or dark green to brown, reddish brown or dark red. Stems 40--120 \um in diameter, moderately flexuous; cortical cells quadrate, moderately thick-walled, smooth, 20--30 x 10--20 \um; branches infrequent, lateral, seldom postical. Leaves transverse, lower part patent to erecto-patent, lobes abruptly erect, loosely complicate, contiguous to subimbricate, broadly quadrate, 300 \um, divided 0.5--0.8. Leaf lobes triangular, antical lobe 4--10 cells wide, postical lobe 6--14 cells wide at base, incurved or slightly concave; apex acute, terminating 2 superposed cells; sinus acute, plane to slightly recurved; margins coarsely dentate-serrate, rarely subdentate, teeth 1--2-celled; cells at base of lobes 10--20 \um wide; cell walls thick; cuticle smooth or obscurely verrucose. Oil bodies 4--9 per cell. Underleaves absent on sterile shoots. Gemmae frequent, 3--6 angled, pale green to reddish brown. Sexual condition autoicous, often fertile. Androecia on lateral branches, spicate; bracts scarcely saccate sometimes weakly concave, margins serrate. Gynoecia terminal on leading shoots; bracts divided 0.25--0.4, connate 0.5 or less antically; lobes margins dentate-serrate with 1(2)-celled teeth; apex 2-celled; bracteoles sometimes truncated. Perianth ovate-lanceolate, often strongly plicate, sometimes curved, green, to 1.6 x 0.4 mm, mouth crenulate, mouth cells 3.5--4.5 longer than wide.

 

On soil or sand, often on stabilized road cuts and trail sides, occasionally on wood; low to moderate elevations; B.C.; Calif., Oreg., Wash.; Eurasia; Africa.

 

Cephaloziella turneri is the most easily recognized species of Cephaloziella in North America by its robustness and prominently dentate leaves. It is undoubtedly more abundant now than in the previous century beause of its affinity for the disturbed soil of trail and road cuts.

 

DUBIOUS AND EXCLUDED SPECIES

 

The following species names are based on single records or specimens whose validity has not been verifiable. Long literature trails are the only reason for mention in this work. Their occurrence in North America awaits rediscovery and verification of their taxonomic reality.

 

Cephaloziella brinkmanii Douin, Mém. Soc. Botan. France 29: 75. 1920. Known only from a collection from Yoho National Park, British Columbia (W. S.Hong 1986), presumably the type. Material in the cited collection is clearly inadequate for placement.

 

Cephaloziella dentata (Raddi) Stephani Bull. Herb. Boissier 5: 78. 1897. Basionym: Jungermannia dentata Raddi, Jungermanniogr. Etrusca, 21. 1818.  Known only from a presumed collection made in British Columbia and not found in North America since the original report in 1934 (W. S. Hong 1986). It possibly represents a misidentified specimen of C. phyllacantha.

 

Cephaloziella patulifolia (Stephani) Douin, Mém. Soc. Botan. France 29: 70. 1920. Basionym: Cephalozia patulifolia Stephani,  Bull. Herb. Boissier, sér. 2, 8: 509. 1908.  Known only from the type collection from California. Not studied since original publication, no recent gatherings fit the description sufficiently to justify association with that name.

 

 

 

2. CYLINDROCOLEA R. M. Schuster, Bull. Natl. Sci. Mus. Tokyo (n.ser.) 12 (3): 666. 1969 * [Greek kýlindros, roll, and koleos, female sheath, alluding to the cylindric perianth]

 

Plants 1--10 x 0.35--0.8 mm; in thin mats, tufts or as solitary, prostrate shoots; green, yellow green, brown or purplish brown. Stems 60--135 \um in diameter, soft textured to somewhat rigid; cortical cells rectangular, thin-walled to moderately thick-walled, 20 x 15 \um to 45 x 24 \um; branches relatively frequent, mostly lateral or postical intercalary, occasionally terminal. Rhizoids few, scattered. Leaves 2-lobed, succubous, oblique to nearly horizontal, with a dorsal leaf free strip 2--3 cells wide, spreading, flat or only slightly curved upwards, remote to contiguous, suborbicular, ovate, obovate to rectangular, 200--480 \um, divided 0.3--0.6. Leaf lobes ovate-triangular to narrowly triangular, 6--14 cells wide at base, flat, spreading to suberect; apex sharply acute to occasionally rounded; sinus acute to slightly rounded; margins entire to crenulate; cells at base of lobes 13--20(--22) x 12--15(--17) \um to (20--)23--35 x (15--)17--20(--21) \um; cell walls thin to slightly thickened; cuticle smooth. Oil bodies average 5--10 per cell, clear. Underleaves usually present, sometimes vestigial or absent, subulate or lanceolate, up to 7 cells long, 5 cells wide. Gemmae consistently absent. Sexual condition autoicous. Androecia on short postical branches, rarely terminal on long shoots and becoming intercalary; bracts 2--3 lobed, lobes entire, acute. Gynoecia terminal on main leafy shoots or on short postical branches; bracts subquadrate, as wide as long, similar to leaves, squarrose to sheathing; lobes triangular to oblong ovate, margin entire, crenulate to denticulate; bracteoles mostly free, rarely united to bracts up to 0.5, unlobed to 2-lobed. Perianth long emergent, cylindrical below, deeply 4-plicate distally, slightly narrowed at mouth and denticulate or broadly truncate and entire, green to brownish, 0.7 x 0.3 mm to 1.2 x 0.4 mm, mouth cells isodiametric, thin-walled, entire to crenulate or slender, elongated and denticulate.

 

Species 18--24 (2 in the flora): southeastern coastal plain of U.S.A., pantropical.

 

Cylindrocolea differs from Cephaloziella by its strongly oblique to nearly horizontal leaf insertion, by the presence of a leaf free strip of at least two rows of cells on the dorsal midline of the stem separating the dorsal leaf base insertion, a complete absence of gemmae, and having a wide or weakly contracted perianth mouth. The leaf bases in Cylindrocolea are more abrubtly narrowed in a tight curve to the insertion as compared to most species of Cephaloziella, where the leaf bases are typically gradually and moderately narrowed to the leaf insertion.

 

SELECTED REFERENCE Schuster, R. M.  1980. The Hepaticae and Anthocerotae of North America. Vol. IV. Columbia University Press, New York.

 

1. Perianth not constricted, broadly truncate, entire, fringed with thin-walled, isodiametric cells       little differentiated from cells of the main body of the perianth; leaves obliquely inserted at      45--60º to the stem ………………..1 Cylindrocolea rhizantha

1. Perianth gradually narrowed to mouth, crenulate to denticulate from projecting ends of slender, elongate, sharp pointed cells; leaves nearly horizontally inserted ………….2  Cylindrocolea planifolia

 

1. Cylindrocolea rhizantha (Montagne) R. M. Schuster, Nova Hedwigia 22: 175. 1971[1972]

 

Jungermannia rhizantha Montagne in Sagra, Hist. Phys. Pol. et Nat. Cuba, Bot., Pl. Cell.: 454. 1842; Cylindrocolea andersonii R. M. Schuster

 

Plants 1--2.5 x 0.35--0.5 mm; in tufts or as solitary, prostrate shoots, green to brown or purplish brown. Stems 60--70 \um in diameter, soft textured, flexuous; cortical cells rectangular, thin-walled, 20--25 x 15--16(--18) \um; branches most often postical, sometimes lateral intercalary or terminal. Leaves obliquely inserted 45--60º on primary shoots, spreading, nearly flat, suborbicular to rectangular, to 200 \um, divided 0.3--0.4(--0.5). Leaf lobes ovate-triangular, 6--9(--12) cells wide at base, nearly flat, spreading; apex acute, rarely obtuse; sinus acute; margins often crenulate from bulging cell walls;  cells at base of lobes 13--20(--22) x 12--15(--17) \um to (20--)23--35 x (15--)17--20(--21) \um wide. Oil bodies 3--7 per cell. Underleaves usually present, subulate or lanceolate, from 3 cells long x 1--2 cells wide to 7 cells long x 5 cells wide. Sexual condition autoicous. Androecia on short postical branches, rarely terminal on long shoots and becoming intercalary; bracts equally 2-lobed, lobes entire, acute. Gynoecia mostly terminal on long leafy shoots, less often on short lateral or postical branches; bracts about the same size as leaves, often squarrose, 0.5 2-lobed; lobes triangular, margin crenulate; bracteoles usually free, shorter than bracts, shallowly 2-lobed, rarely with a tooth on margin. Perianth 0.3--0.4 emergent, truncate, not contracted at mouth, green to brownish, 650--750 x 250--360 \um, perianth mouth entire, mouth cells isodiametric and thin-walled.

 

Usually on tree trunks, less often on soil; Fla., La., N.C., S.C., Tex.; West Indies; Central America; South America.

 

Cylindrocolea andersonii was described from a single, fragmentary specimen not adequate for confident placement. Since R. M. Schuster himself suggested it might well be simply a form of C. rhizantha, it is placed in synonymy here until additional material is discovered to determine its distinctness and placement.

 

2. Cylindrocolea planifolia (Stephani) R. M. Schuster, Nova Hedwigia 22: 164, 1971[1972]

 

Cephaloziella planifolia Stephani, Hedwigia 32 (5): 317, 1893; Cylindrocolea obliqua (Douin) R. M. Schuster

 

Plants 3--10 x 0.6--0.8(--1.2) mm; in thin, loose patches, green to brown, never red. Stems 85--135(--150) \um in diameter, flexuous; cortical cells rectangular, moderately thick-walled, 35--45 x (13--)20--24 \um; branches frequent, mostly postical intercalary. Leaves almost horizontally inserted, flat, laterally patent, oblong-ovate, obovate to subquadrate, 310 x 260 \um to 480 x 440 \um, divided 0.4--0.6. Leaf lobes narrowly triangular, ventral lobe usually slightly larger, (6--)8--12(--14) cells wide at base, flat, weakly divergent or suberect; apex acute, terminated by 1(--2) cells, occasionally rounded; sinus acute, rarely slightly rounded; margins entire; cells at base of lobes (20--)23--35 x (15--)17--20(--21) \um. Oil bodies (4--)7--13(--15) per cell. Underleaves vestigial or absent, rarely subulate, to 4 cells long x 2 cells wide. Sexual condition autoicous. Androecia on short postical branches; bracts smaller than leaves, subequally 2-lobed, lobes entire, acute. Gynoecia mostly on short lateral or postical branches, less often terminal on long leafy shoots; bracts slightly larger than leaves, sheathing, divided 0.35--0.5, 2--3 lobed; lobes oblong-ovate, margin entire or weakly denticulate; bracteoles mostly free, rarely united to bracts up to 0.5, equal to bracts, often unlobed. Perianth 0.4--0.5 emergent, slightly narrowed to mouth, green to brownish, to 1.2 x 0.4 mm, perianth mouth crenulate to denticulate, mouth cells slender and elongated.

 

On sandy or peaty soil, rarely on logs; Fla., La.; West Indies; Central America; South America.

 

 

 

CephaloziellaceaeCephaloziellaTurneri_Art_12-1

 

CephaloziellaceaeCylindrothecaPlanifolia_Art_12-1