BFNA Title: Frullaniaceae
Author: John Atwood
Date: Dec. 6, 2017
Edit Level: S
Version: 1

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. FRULLANIACEAE Lorch

John J. Atwood

 

Plants dull to lustrous, green, yellowish brown, copper-red, or black, with red-purple coloration, forming patches or mats; intercalary branches sporadic and abbreviated; terminal branches at ventral base of lobe, replacing the lobule, the resulting half-leaf mostly inserted on stem, occasionally on branch; the first branch appendages comprised of variously oriented ventral leaves, 2-fid or 3-fid, mostly flat, with one lobe often modified into a galeate, saccate, or cylindric lobule. Lateral leaves asymmetrically 3-fid, with an incubous to sub-transverse dorsal lobe and often two modified ventral lobes (lobule and stylus, though stylus occasionally absent). Ventral leaves emarginate or 2-fid. Rhizoids few, brownish, branched, clustered near middle or base of ventral leaves. Specialized asexual reproduction as multicellular discoid gemmae on margins of dorsal lobe, plants with caducous leaves on vegetative shoots or specialized branches, or absent. Gynoecia with or without subfloral branches. Perianth well-developed, sometimes dorsiventrally compressed, often weakly to distinctly keeled with lateral and postical keels, sometimes with supplementary keels, mostly contracted into a beak, perigynium absent. Spores with rosettes of tubercles.

 

Genus 1 (species 375), cosmopolitan. Greenland; Canada; United States; Mexico; West Indies; Bermuda; Central America; South America; Arctic; Europe; Asia (including Indonesia); Africa (including Madagascar); Atlantic Islands; Indian Ocean Islands; Pacific Islands (including New Zealand); Australia; Antarctic.

 

The monogeneric Frullaniaceae, containing only Frullania (B. Crandall-Stotler et al. 2009; J. Hentschel et al. 2009), differs morphologically from the Jubulaceae in plant color, the insertion of the dorsal half-leaf, the shape and lobing of the primary branch appendages, and in the ornamentation of the spores. Frullania plants frequently have reddish purple coloration, whereas plants in the Jubulaceae regularly lack these pigments. Furthermore, at terminal branch origins, most Frullania species have the dorsal half-leaf inserted on the stem, although occasionally the insertion is also on the branch (R. E. Stotler 1969). In the Jubulaceae, the dorsal half-leaf is partially inserted on both the stem and branch. Next, the primary branch appendages in Frullania are ovate-lanceolate and 2-fid or trifid with one lobe often modified into a lobule. By contrast, the primary branch appendages in the Jubulaceae are lanceolate or sub-triangular, consistently unmodified, and mostly entire, although they can rarely be shallowly 2-fid (B. Crandall-Stotler and W. Guerke 1980). Lastly, Frullania spores have distinct rosettes of tubercles, whereas the spores in the Jubulaceae have a finely papillose ornamentation.

 

SELECTED REFERENCES  Asakawa, Y., N. Tokunaga, M. Toyota, T. Takemoto, S. Hattori, M. Mizutani, C. Suire. 1979. Chemosystematics of bryophytes II. The distribution of terpenoids in Hepaticae and Anthocerotae. J. Hattori Bot. Lab. 46: 67--76.  Crandall-Stotler, B., R. E. Stotler, and D. G. Long. 2009. Phylogeny and classification of the Marchantiophyta. Edinburgh J. Bot. 66: 155--198.  Hentschel, J., M. J. von Konrat, T. Pócs, A. Schäfer-Verwimp, A. J. Shaw, H. Schneider, and J. Heinrichs. 2009. Molecular insights into the phylogeny and subgeneric classification of Frullania Raddi (Frullaniaceae, Porellales). Molec. Phylogen. Evol. 52: 142--156.  Schuster, R. M. 1992. The Hepaticae and Anthocerotae of North America East of the Hundredth Meridian, Vol. 5. Field Museum of Natural History, Chicago. 1--854.

 

1. FRULLANIA Raddi, Jungermanniogr. Etrusca 9. 1818. [For Leonardo Frullani, 1756--1824, Florence statesman and director of the Tuscan treasury]

 

Plants mostly prostrate, loosely or firmly attached to substrate, sometimes spreading, rarely pendent, regularly or irregularly branched, vegetative branches terminal with an associated ovate or orbicular dorsal half-leaf. Stem 5--20 cells wide in cross section, with 1--2 layers of thick-walled epidermal cells, and several layers of firm-walled cortical cells, 18--40 \um. Lateral leaves obliquely to sub-transversely inserted; dorsal lobe margin entire [toothed]; lobule folded under the dorsal lobe; stylus located between lobule mouth and stem, sometimes terminated by a slime papilla; marginal and median cells irregularly polygonal, subquadrate, or isodiametric, basal cells more elongate. Androecia at apex of short or elongate lateral branch, sometimes on short capitate branch, globose and consisting of 2--4 pairs of imbricate, subequally 2-lobed bracts, or spicate and consisting of 8--20 pairs of imbricate subequally 2-lobed bracts, bracteole at base of inflorescence, containing 1--2 antheridia. Gynoecia consisting of 2--6 pairs of keeled, subequally 2-lobed bracts and bracteoles that are longer and wider than the vegetative leaves, stylar tooth often present, bracteole 2-lobed; perianth emergent to exserted; gynoecia with 2--12 archegonia. Foot small, lenticular. Seta non-articulate, in transverse section with numerous, irregularly arranged cell rows, 8--10 cells wide. Capsule barely to shortly exserted, 2-stratose, the inner and outer layers with variously thickened walls, splitting into four, strongly reflexed, hyaline valves. Elaters reddish brown, parallel to the sporophyte, monospiral. Spores reddish brown, orbicular, oblong or bluntly angular.

 

Subgenera 11 (5 in the flora), 375 species (37 in the flora), cosmopolitan. Greenland; Canada; United States; Mexico; West Indies; Bermuda; Central America; South America; Arctic; Europe; Asia (including Indonesia); Africa (including Madagascar); Atlantic Islands; Indian Ocean Islands; Pacific Islands (including New Zealand); Australia; Antarctic.

 

Frullania, like Jubula, has asymmetrically trifid lateral leaves that are composed of an obliquely to sub-transversely inserted dorsal lobe, a completely or partially inflated, sometimes flattened ventral lobule, and a ventrally positioned (although sometimes reduced or absent) stylus. Whereas the lobule in Jubula shows relatively little variation between taxa, the lobule in Frullania is exceedingly variable in form, shape, size, and orientation. This variation is the basis for much of the historic and present infrageneric classification. Eleven Frullania subgenera are currently recognized worldwide (J. Hentschel et al. 2015). Five of these occur in North America, including: subg. Chonanthelia, subg. Diastaloba, subg. Frullania, subg. Meteoriopsis, and subg. Thyopsiella. Taxa in the following treatment are grouped by their subgenera (L. Söderström et al. 2016) to facilitate comparisons made between morphologically similar species. Aside from characters of the lobule, subgenera can be regionally delimited by a combination of differences in their branching pattern; lobing of their primary branch appendages; shape of the antical margin of the dorsal lobe; shape, size, and form of the styli; shape, size and form of the ventral leaves; sexuality; location of the gynoecia; and ornamentation of the perianth. Utilization of these characters has resulted in slightly different classifications for taxa with lobules that appear to be intermediate between subgenera (A. W. Evans 1897; J. Hentschel et al. 2015; R. M. Schuster 1992).

Most North America Frullania grow on the bark of trees, shrubs, and woody vines. The plants are typically prostrate with their shoots loosely or firmly attached to the substrate. In F. ericoides and F. sabaliana, the plants spread from the substrate, particularly at the tips of mature shoots. In F. cucullata and F. riojaneirensis, the shoots may become pendent; a growth form more frequently encountered in tropical members of the genus. Some species such as F. cobrensis and F. selwyniana appear to be ecologically restricted to specialized habitats. Frullania californica, F. caulisequa, F. donnellii, F. ericoides, F. kunzei, F. nisquallensis, and F. stylifera can occur on rotting wood. Frullania asagrayana, F. californica, F. catalinae, F. caucasica, F. cuencensis, F. eboracensis, F. ericoides, F. franciscana, F. inflata, F. kunzei, F. mexicana, F. nisquallensis, F. plana, F. riparia, F. saxicola, F. subarctica, F. tamarisci, and F. wrightii can occur on boulders and rock ledges, sometimes exclusively. There are several reported occurrences of F. asagrayana, F. caulisequa, and F. eboracensis growing as an epiphyll in humid areas receiving heavy rainfall (P. G. Davison 1997; W. Guerke 1973; A. C. Risk et al. 2011; R. M. Schuster 1959). Most Frullania taxa, however, generally inhabit more mesic to xeromorphic environments in the flora area.

Frullania frequently occurs intermixed with other Frullania or other bryophytes. Locating fertile shoots is often necessary for identification, as the shape of the androecia, as well as the location of the gynoecia, are taxonomically important characters. Specimens consisting exclusively of sterile stems or male plants, such as in the F. eboracensis complex, may not be determinable. Mature dorsal lobes, lobules, styli, and ventral leaves from the stem or main branch should be used for identification. Ventral leaves located immediately beneath branches are often atypical in size, shape of the lobe and sinuses, as well as stance and dentation of the margins. These should not be used for comparisons. Some Frullania species, such as F. caulisequa, have branches that become progressively smaller as they spread from the stem. These branches are also often microphyllose. Oil body characteristics should be obtained from the median cells. Oil body characters in the following treatment were taken from R. M. Schuster (1992) when fresh material could not be obtained. The removal of the ventral leaves is sometimes necessary to examine the lobule, stylus, and/or characters of the primary branch appendages. Measurement and characters of the gynoecial bract and bracteoles are taken from the innermost pair. Dissection or sectioning of the gynoecial bract and bracteoles are often necessary to determine the extent of connation between these structures. Sectioning of the perianth may be necessary to determine the location, extent, and number of keels, as well as the presence or absence of carinae or tubercles. Sectioning of the perianth beak, or mounting the perianth in Hoyer’s solution (L. E. Anderson 1954), is often necessary to establish the presence or absence of papillae on the inner beak surface.

 

SELECTED REFERENCES  Evans, A. 1897. A revision of the North American species of Frullania, a genus of Hepaticae. Trans. Connecticut Acad. Arts Sci. 10: 1--39, pl. 1--15.  Frye, T. C. & L. Clark. 1947. Frullania. Hepaticae of North America, part V. p. 735--782.  Hentschel, J., M. J. von Konrat, T. Pócs, A. Schäfer-Verwimp, A. J. Shaw, H. Schneider, and J. Heinrichs. 2009. Molecular insights into the phylogeny and subgeneric classification of Frullania Raddi (Frullaniaceae, Porellales). Molec. Phylogen. Evol. 52: 142--156.  Hentschel, J., M. J. von Konrat, L. Söderström, A. Hagborg, J. Larraín, P. Sukkharak, J. Uribe, and L. Zhang. 2015. Notes on Early Land Plants Today. 72. Infrageneric classification and new combinations, new names, new synonyms in Frullania. Phytotaxa 220: 127--142.

 

Key to Frullania subgenera in North America

1. Inflated lobule wide-spreading, separated from the stem by a distance greater than to two times its width, the lobule mouth oriented towards stem................. 1b. Frullania subg. Diastaloba

1. Inflated lobule erect or obliquely spreading, contiguous or approximate with stem, separated by a distance that is less than or nearly equal to its width, the lobule mouth oriented towards shoot base or slightly toward or away from stem; or lobule frequently explanate

2. Lobule with inflated distal portion and a ligulate to triangular proximal portion

..................................................................................... 1a. Frullania subg. Chonanthelia

2. Proximal lobule portion not noticeably expanded, or lobule frequently explanate

3. Inflated lobule cylindric or sac-shaped, about two or more times longer than wide; or explanate lobule with strongly deflexed margins

4. First branch appendage 2-fid, consisting of a flattened, lanceolate-ovate segment, and a mostly inflated, though occasionally flattened lobule; dorsal lobe sometimes with ocelli restricted to the basal cells; stylus uniseriate and filiform                 1d. Frullania subg. Meteoriopsis

4. First branch appendage 2-fid and consisting of an unequally divided, flattened, lanceolate-ovate segment, or 3-fid and consisting of an unequally 2-fid, flattened, lanceolate-ovate segment, and a flattened or inflated lobule; dorsal lobe frequently with ocelli arranged in a median line and/or scattered throughout the lobe; stylus subulate to lamellate, often appendiculate................................................................................... 1e. Frullania subg Thyopsiella

3. Inflated lobule cap or helmet-shaped, about as long as wide; or explanate lobule with plane or slightly decurved margins............................................. 1c. Frullania subg. Frullania

 

Key to Frullania species in North America

1. Ventral leaves emarginate........................................................... 14. Frullania davurica

1. Ventral leaves 2-fid.

2. Plants with caducous leaves.

3. Caducous leaves on rigidly erect, flagelliform shoots. Flagelliform shoots having short internodes and persistent, squarrose ventral leaves; dorsal lobe base truncate

        ........................................................................................... 8. Frullania bolanderi

3. Caducous leaves on vegetative stems and branches, internodes evenly spaced, ventral leaves flat; dorsal lobe base cordate.

4. Ventral leaf margins mostly entire or occasionally bluntly angulate near leaf middle; perianths trigonous and smooth or mostly so............... 15. Frullania eboracensis

4. Ventral leaf margins sharply angulate or toothed near the base of the sinus; perianths with accessory keels and carinae or perianths unknown in flora area.

5. Dorsal lobe median cells strongly sinuous; perianths usually present, with accessory keels and carinae.............................................................. 7. Frullania appalachiana

5. Dorsal lobe median cells with firm walls or occasional intermediate thickenings; androecia, gynoecia and perianths absent in flora area 11. Frullania caucasica

2. Plants without caducous leaves.

6. Dorsal lobes with ocelli in a conspicuous median line, scattered, or confined to basal cells.

7. Ventral leaf margins mostly plane at or above the sinuses, margins sometimes slightly recurved near leaf base.

8. Dorsal lobes with ocelli forming a single or double median line (scattered ocells occasionally present).

9. Plants autoicous, gynoecia terminal on stem or main branch; styli lacking appendages and teeth............................................................. 35. Frullania selwyniana

9. Plants dioicous, gynoecia terminal on short lateral branches; styli with appendages or teeth.

10. Ventral leaf margins entire; stylus appendiculate.                                31. Frullania asagrayana

10. Ventral leaf margins with a blunt angulation on one or both sides; stylus with a few single-celled teeth................................... 33. Frullania franciscana

8. Dorsal lobes with scattered ocells or ocelli confined to basal cells, not forming a median line.

11. Plants dioicous; ocelli scattered; lobules longer than wide; stylus filiform or subulate............................................................................. 32. Frullania californica

11. Plants autoicous; ocelli confined to basal cells; lobules wider than long; stylus lanceolate................................................................ 13. Frullania cobrensis

7. Ventral leaf margins reflexed, sometimes only at or above the sinuses.

12. Stylus appendage absent or weakly formed, sometimes only a few teeth present.

13. Plants greenish to yellowish brown, frequently tinged red, shoots 0.9--1.2 mm wide…............................................................. 34. Frullania nisquallensis 

13. Plants dark reddish brown to black; shoots 1.1--1.6 mm wide                           36. Frullania subarctica

12. Stylus appendage well developed as a semi-circular disk with lateral teeth.

14. First branch appendage divided into three, unequally sized lobes with acute to apiculate apices, and entire margins; first branch appendage insertion almost sheath-like and extending from the ventral side of the branch to the lateral surface of the stem; dorsal lobe apices mostly deflexed, rounded to bluntly acute, sometimes apiculate; ocelli forming a long median line, sometimes broken, scattered ocells occasionally present; ventral leaf margins reflexed at or above the sinus  31. Frullania asagrayana

14.  First branch appendage divided into two, unequally sized acute lobes, with the acroscopic lobe margin toothed, and sometimes appendiculate; first branch appendage insertion appearing asymmetrical and extending obliquely from the ventral side of the branch to the basioscopic lateral surface of the branch; dorsal lobe apices mostly decurved, acute to apiculate; ocelli forming a short median line, scattered ocells frequently present; ventral leaf margins reflexed throughout 37. Frullania tamarisci

6. Dorsal lobes lacking ocelli or ocelli inconspicuous.

15. Inflated lobule wide-spreading, separated from the stem by a distance greater than its width, the lobule mouth oriented towards stem.

16. Plants yellowish green to light brown, shoots 0.8--1.7 mm wide, autoicous; margins of dorsal lobe and lobule without differentiated marginal cells; lobule margin smooth near mouth.......................................................................... 5. Frullania caulisequa

16. Plants copper-red to blackish, shoots 0.6--0.7 mm wide, dioicous; margins of dorsal lobe and lobule with differentiated marginal cells that are sometimes pigmented and bulging; lobule margin crenulate, incised near mouth              

...................................................................................  6. Frullania taxodiocola

15. Inflated lobule erect or obliquely spreading, contiguous or approximate with stem, separated by a distance that is less than or nearly equal to its width, the lobule mouth oriented towards shoot base or slightly toward or away from stem; or lobule frequently explanate.

17. Lobule with inflated distal portion and a ligulate proximal portion that extends to or beyond the basal lobe margin and is connected to the dorsal lobe by a keel that is parallel or sub-parallel to the stem; free lobule margin sinuous

.................................................................................... 4. Frullania riojaneirensis

17. Proximal lobule portion triangular or not noticeably expanded, forming a short keel with dorsal lobe that is perpendicular or obliquely spreading from stem; free lobule margin plane; or lobule frequently explanate.

18. Inflated lobule cylindric or sac-shaped, about two or more times longer than wide, not beaked on the external margin; or explanate lobule with strongly deflexed margins; ornamentation of perianth smooth.

19. Ventral leaf margins reflexed, sometimes only at or above the sinuses.

20. Dorsal lobes acuminate; lobules rarely explanate; stylus subulate, sometimes with appendages or teeth.

20. Dorsal lobes rounded, bluntly acute, or apiculate; lobules in part explanate; stylus uniseriate and filiform............................................................

..................................................................... 28. Frullania cucullata

21. Plants greenish to yellowish brown, frequently tinged red, 0.9--1.2 mm wide…......................................... 34. Frullania nisquallensis 

21. Plants dark reddish brown to black; shoots 1.1--1.6 mm wide       36. Frullania subartica

19. Ventral leaf margins plane, slightly recurved or contorted.

22. Dorsal lobe bases cordate; plants diocious, gynoecia terminal on short lateral branches..................................... 32. Frullania californica

22. Dorsal lobe bases truncate; plants autoicous, gynoecia terminal on stem or main branch.

23. Lobules separated from the stem by a distance that is less than or nearly equal to its width; gynoecial bracts and bracteole dentate         29. Frullania donnellii

23. Lobules mostly superimposed or contiguous with stem; gynoecial bracts and bracteole entire.................... 30. Frullania kunzei

18. Inflated lobule cap or helmet-shaped, about as long as wide, sometimes beaked on the external margin; or explanate lobule with plane or slightly decurved margins; ornamentation of perianth often tuberculate, sometimes smooth.

24. Plants autoicous or paroicous, androecia mostly capitate, occasionally short spicate.

25. Ventral leaf bases cordate to auriculate, sometimes only on one side.

26. Dorsal lobes squarrose when moist.... 2. Frullania gibbosa

26. Dorsal lobes flattened when moist...... 20. Frullania plana

25. Ventral leaf bases cuneate to slightly decurrent.

27. Dorsal face of perianth flattened or concave, beak lacking; gynoecial bracts often single, replaced by innovations 23. Frullania sabaliana

27. Dorsal face of perianth weakly to distinctly keeled, acuminating in a beak; gynoecial bracts multiple, not replaced by innovations.

28. Innermost gynoecial bracteole free from bract.............

............................................................ 18. Frullania inflata

28. Innermost gynoecial bracteole connate with bract on one or sometimes both sides.

29. Perianth with 8--12 keels, innermost gynoecial bracteole sometimes connate on both sides with bract (often low connate on one side and high connate on the other) 3. Frullania mexicana

29. Perianth 4--5-keeled, innermost gynoecial bracteole connate on one side with bract.

30. Ventral leaves narrow, not or slightly wider than stem, entire or with a blunt angulation on one or both sides; stylus filiform to subulate, mostly 3--5 cells long and 1--2 cells wide at base.

31. Plants greenish to yellowish brown; lobules frequently explanate, or when inflated, galeate and obscuring less than half of the lobe’s underlying surface; perianth globose or obovate...... 24. Frullania saxicola

31. Plants reddish-brown; lobules consistently inflated, sub-quadrate and often obscuring 3/4ths or more of the lobe’s underlying surface; perianth oblong, sometimes stipitate.

32. Dorsal lobe median cells with trigones small or indistinct; ventral leaf margins conspicuously toothed on one or both sides 19. Frullania oakesiana

32. Dorsal lobe median cells with trigones conspicuous and bulging; ventral leaf margins entire................................................................

................................ 21. Frullania rappii

30. Ventral leaves broad, usually two times wider than stem, with blunt angulations on one or both sides; stylus conspicuous, lamelliform, 6--14 cells long and 2--10 cells wide at base.

33. Dorsal lobe base truncate; innermost gynoecial bracteole with deeply notched, acute sinuses 10. Frullania catalinae

33. Dorsal lobe base cordate; innermost gynoecial bracteole with shallowly notched, obtuse sinuses.

34. Stylus lamelliform, 10--15 cells long and 4--10 cells wide; perianth beak narrow and short 25. Frullania stylifera

34. Stylus lanceolate, 4--10 cells long and 2--4 cells wide; perianth beak wide, mouth flaring.

35. Shoots 0.3--0.6 mm wide

.................... 17. Frullania hattoriana

35. Shoots 0.9--1.2 mm wide..............

.......................... 27. Frullania wrightii

24. Plants dioicous, androecia spicate, though sometimes abbreviated, or unknown.

36. Leaves deflexed and somewhat rolled around the stem when dry; squarrose when moist; gynoecia terminal on short lateral branches with no subfloral innovations................................................... 16. Frullania ericoides

36. Leaves imbricate when dry; flattened, spreading, or wide-spreading when moist; gynoecia terminal on stem or main branch.

37. Lobules mostly explanate or cap-shaped when inflated; stylus filiform................................................................... 22. Frullania riparia

37. Lobules mostly inflated and galeate, sometimes explanate; stylus subulate or lanceolate.

38. Proximal portion of lobule triangular, appearing to have a widened, somewhat decurrent mouth...................................................

.......................................................... 1. Frullania cuencensis

38. Proximal portion of lobule not noticeably expanded

39. Perianths smooth or obscurely tuberculate.

40. Shoots 0.4--0.5 mm wide; dorsal lobe antical base auriculate-appendiculate; stylus lanceolate, 6--8 cells long and 2--4 cells wide; perianth with multiple keels        12. Frullania chilcootiensis

40. Shoots 0.5--1 mm wide; dorsal lobe antical base cordate; stylus subulate, 3--6 cells long and 2--3 cells wide; perianth trigonous............................................... 15. Frullania eboracensis

39. Perianths with conspicuous tuberculae and/or carinae.

41. Shoots 1.0--1.2 mm wide; stylus lanceolate, 5--8 cells long and 2--3 cells wide; gynoecial bracts spreading; innermost bracteole free from bract; perianth beak elongate 9. Frullania brittoniae

41. Shoots 0.7--0.9 mm wide; stylus filiform or subulate, 4--5 cells long and 1--2 cells wide; gynoecial bracts squarrose; innermost bracteole connate on one side with bract; perianth beak short

................................................. 26. Frullania virginica

 

1a. Frullania subg. Chonanthelia Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 8. 1884

 

Plants pinnately or bi-pinnately branched. First branch appendage, 3-fid, consisting of a flattened, lanceolate-ovate, unequally 2-fid segment, and a flattened or inflated lobule. Second branch appendage lateral and consisting of two flattened segments or inflated lobules, or a mixture of both (and associated stylus). Lateral leaves loosely to strongly imbricate when dry, imbricate, erect, wide-spreading or squarrose when moist, antical margin base expanded and extending beyond distal edge of stem, apex rounded; lobules approximate or contiguous with stem, parallel or sub-parallel, mouths oriented towards shoot base or slightly towards stem, the distal portion inflated, slightly to strongly compressed at the mouth, the external margin rostrate or uncinate, the proximal portion flattened, shortly expanded or as an elongated appendage; styli filiform to subulate [lanceolate or lamellate], sometimes with an appendage; ocelli lacking. Ventral leaves distant, contiguous, or imbricate, margins entire, bases narrowed, sometimes short decurrent [auriculate to appendiculate]. Specialized asexual reproduction none. Sexual condition autoicous, paroicous or dioicous. Androecia capitate [spicate], sessile on stem or terminal on short lateral branches. Gynoecia terminal on stem, main branch, short lateral branches, or androecial branch; bract lobule margins with prominent stylar tooth. Perianth trapezoidal in section, with at least two lateral and two ventral keels, sometimes with supplementary keels, ornamentation smooth, beak with papillae.

 

Species approximately 40 (4 in flora): North America; Mexico; West Indies; Central America; South America; Asia (including Indonesia); Africa (including Madagascar); Pacific Islands.

 

Subgenus Chonanthelia has variably modified lobules that are distally inflated and proximally flattened. In F. riojaneirensis, the proximal portion of the lobule is a ligulate appendage that is attached to the inflated portion along an elongated keel. By comparison, the proximal portion of the lobule in F. cuencensis, F. gibbosa, and F. mexicana are shortly expanded with triangular or narrowly triangular appendages and less distinct keels. The lobules in these taxa appear similar to those of F. catalinae, F. ericoides and F. saxicola (subg. Frullania) in that they seem to have widened, somewhat decurrent mouths. Similarities between subg. Chonanthelia and subg. Frullania also extend to the shape of the dorsal lobe, expanded dorsal lobe antical base, connation of innermost gynoecial bracteoles with the bract, and strongly keeled perianths with sometimes supplementary keels. North America species of subg. Chonanthelia can mostly be separated from subg. Frullania by their tendency to have fewer explanate lobules and the ventral leaves to be more shallowly 2-fid with entire margins.

 

SELECTED REFERENCES  Schuster, R. M. 1992. The Hepaticae and Anthocerotae of North America East of the Hundredth Meridian, Vol. 5. Field Museum of Natural History, Chicago. 1--854.  Yuzawa, Y. 1991. A monograph of subgen. Chonanthelia of gen. Frullania (Hepaticae) of the world. J. Hattori Bot. Lab. 70: 181--291.

 

1. Frullania cuencensis Taylor, London J. Bot. 5: 406. 1846

 

Plants reddish brown, 1.0--1.5 mm wide. Stems 120--140 \um wide. Lateral leaves loosely imbricate when dry, erect spreading when moist, dorsal lobe broadly ovate, 0.7--0.9 x 0.6--0.8 mm, convex, apex decurved, base cordate; lobule distal portion 0.2--0.3 mm, slightly compressed at mouth, external margin weakly rostrate, proximal portion triangular, keel 60--100 \um long, sub-parallel with stem, free margin plane; stylus subulate, consisting of 4--6 cells long and 3--4 cells wide; marginal cells 14--20 \um, median cells 18--26 \um, basal cells 24--32 \um, with occasional intermediate thickenings, trigones large, [oil bodies ellipsoidal, 3--4 per cell]. Ventral leaves distant, obovate, 0.3--0.4 x 0.2--0.4 mm, 2-fid in upper 80--120 \um, lobes and sinus acute to broadly acute, margins slightly recurved. Sexual condition dioicous. Gynoecia terminal on stem or main branch, [bract lobe elliptical, 0.9--1 x 0.5--0.7 mm, apex rounded, margins entire; lobule narrowly elliptical, 0.9 x 0.4 mm, apex acute, margins entire; bracteole oblong, 0.8 x 0.6 mm, 2-fid to 100 \um, lobes and sinus acute, margins entire, unevenly connate on both sides]. Perianth [oblong, 1.4 x 1 mm, abruptly narrowed, lacking supplementary keels, beak short].

 

On shaded rhyolite cliff face; high elevations; Tex.; Mexico; West Indies; South America.

 

Frullania cuencensis is known in North America from a single specimen (Worthington 61, colo) that was collected in the Davis Mountains in southwestern Texas. The species is similar to F. riparia in its habit; dioicous sexuality; broadly ovate dorsal lobes; decurved dorsal lobe apices; cordate dorsal lobe bases; and distant, ventral leaves that are about three times the width of the stem, and have narrow to short decurrent bases. Frullania cuencensis differs from F. riparia in having consistently inflated lobules with slightly longer distal portions and narrowly triangular proximal portions. The lobules of F. riparia are frequently explanate, and when inflated are cap-shaped, broad and short with the external margin weakly rostrate, and the mouth somewhat compressed. The proximal portion of the lobule is never expanded along a short keel, although occasionally the mouth will appear broad and enlarged as a result of the lobule spreading from the stem, as well as the asymmetrically formed ventral and dorsal sides. Furthermore, the styli of F. cuencensis differ from F. riparia in their subulate shape, with bases up to six cells wide. Frullania riparia, by comparison, has styli that are mostly uniseriate and filiform. Lastly, F. cuencensis has entire, recurved ventral leaf margins. The ventral leaf margins of F. riparia frequently have blunt angulations on one or both sides, and are plane. The only other Frullania species in western North America that may be confused with F. cuencensis is F. caucasica. It too is dioicous, occurs on rocks, and has dorsal lobes and styli of similar size and shape. However, the dorsal lobes and lobules of that species are caducous, resulting in numerous bare sections along the stem. Furthermore, F. caucasica has angulate or toothed ventral leaf margins.

The Texas material lacks androecia, gynoecia, and perianth. The description of these structures, along with characters of the oil bodies are based on Mexico specimens and the description of F. cuencensis by Y. Yuzawa (1991).

 

2. Frullania gibbosa Nees, in Montagne, Ann. Sci. Bot. sér. 2, 14: 333. 1840

 

Plants reddish brown, 1.0--1.2 mm wide. Stems 180--200 \um wide. Lateral leaves densely imbricate when dry, squarrose when moist, dorsal lobe broadly ovate to orbicular, 0.5--0.6 x 0.6--0.8 mm, base cordate or auriculate; lobule distal portion 0.3--0.4 x 0.2--0.3 mm, slightly compressed at mouth, external margin weakly rostrate, proximal portion triangular, keel 0.1 mm long, sub-parallel to obliquely [widely] spreading from stem, free margin plane [sinuose]; stylus subulate, consisting of 1--4 cells long and 1--2 cell wide, usually with an ovate [reniform] appendage 6--8 [--20] cells wide, margins plane or crispate; marginal cells 14--18 \um, median cells 18--28 \um, basal cells 22--38 \um, occasional intermediate thickenings, trigones large, nodulose, oil bodies fusiform to ellipsoidal, 3--6 per cell. Ventral leaves imbricate, obovate, 0.3--0.5 x 0.3--0.6 mm, contorted, 2-fid in upper 140--180 \um, lobes broadly acute, sinus rounded, margins sinuous, sometimes recurved. Sexual condition autoicous [paroicous]. Gynoecia terminal on stem or short lateral branch [androecial branch], bract lobe ovate, 0.8--1 x 0.2--0.3 mm, apex rounded, margins entire to sinuate; lobule lanceolate, 0.8--0.9 x 0.1, apex acute, margins entire, recurved; bracteole oblong, 0.8--1 x 0.4--0.5 mm, 2-fid to 0.3 mm, lobes and sinus narrowly acute, margins entire or with an occasional tooth on each side, unevenly connate on both sides. Perianth oblong, 0.9--1 x 1.2--1.4 mm, abruptly narrowed, lacking supplementary keels, beak long.

 

Tree bark [or rocks]; low elevations; Fla.; Mexico; West Indies; Central America (Guatemala, Honduras, Costa Rica, Panama); South America.

 

Frullania gibbosa is rare in North America, but common elsewhere in its range. Excluding a historical specimen that was collected by C. T. Mohr dubiously from Alabama (R. M. Schuster 1992), the species is known in the United States only from southern Florida. R. M. Schuster (1992) noted that North America specimens are smaller and less developed in comparison to neotropical ones described by A. W. Evans (1914) and L. Clark (1956). The Mohr specimen, illustrated by R. M. Schuster (1992), therefore suspiciously stands out as a morphological anomaly compared to other specimens from the flora area. Since the specimen is more similar to neotropical specimens, it seems likely that it may have been collected in Mexico as pointed out by W. C. Steere (1946) in his citation of Mohr’s letters to Lesquereux. In fact, Florida specimens of F. gibbosa are perhaps more similar morphologically to F. ericoides than the related F. riojaneirensis. Frullania gibbosa and F. ericoides both have a dark reddish brown color, similar shoot widths, densely imbricate leaves when dry, and squarrose leaves when moist. Furthermore, the two species have similar broadly ovate dorsal lobes, slightly compressed lobules with weakly rostrate external margins, subulate styli, and ventral leaves that are about two to three times the width of the stem. Frullania gibbosa differs from F. ericoides in having lobules that are rarely explanate, and attached to the lobe by a short keel that is sub-parallel or obliquely spreading from the stem. Furthermore, the ventral leaves are imbricate, shallowly 2-fid with rounded sinuses, and have entire to sinuous margins. The ventral leaves of F. ericoides vary from being contiguous to imbricate, and have more deeply divided and narrower sinuses. Additionally, the ventral leaf margins of F. ericoides, although variable, generally have slight angulations on one or both sides. Finally, F. gibbosa has subulate styli with an appendage that is comparable to styli of the F. tamarisci complex (subg. Thyopsiella). In Neotropical specimens, the appendage can sometimes approach the size of the lobule (A. W. Evans 1914) although in North American material, excluding the dubious Mohr specimen, it tends to be much smaller and even may occasionally be absent. Fertile specimens of F. gibbosa and F. ericoides are more easily separated due to their differences in sexuality and perianth ornamentation.

 

3. Frullania mexicana Lindenberg, in C. M. Gottsche, J. B. W. Lindenberg & C. G. D. Nees, Syn. Hep.: 425. 1845

 

Plants green, purplish or reddish brown, 0.8--1.2 mm wide. Stems 100--150 \um wide. Lateral leaves imbricate when dry, widely spreading when moist, dorsal lobe broadly ovate, 0.5--0.8 x 0.5--0.7 mm, apex slightly decurved, base cordate; lobule distal portion 260--400 x 220--340 \um, slightly compressed at mouth, external margin weakly rostrate, proximal portion narrowly triangular, keel 20--84 \um, sub-parallel to obliquely spreading from stem, free margin plane; styli subulate, composed of 3--6 cells long and 2--4 cells wide; marginal cells 12--20 \um, median cells 14--24 \um, basal cells 20--36 \um, sinuous with intermediate thickenings, trigones large, oil bodies spherical, 3--5 per cell. Ventral leaves distant, obovate, 0.2--0.5 x 0.2--0.3 \mm, flat, 2-fid for about 80--100 \um, lobes and sinus bluntly acute. Sexual condition autoicous. Gynoecia terminal on stem or main branch, bract lobe ovate to oblong, 0.8--1 x 0.6--0.8 mm; apex rounded, margins entire; lobule lanceolate, 0.7--1 x 0.3--0.5 mm apex acute to acuminate, margins entire; bracteole obovate, 0.6--0.9 x 0.3--0.4 mm, 2-fid to 60--100 \um, lobes and sinus rounded to bluntly acute, margins entire, unevenly connate on both sides. Perianth obovoid, 1.1--1.5 x 0.8--1 mm, abruptly narrowed, with numerous supplementary keels, beak short, broad.

 

On the branches and trunks of trees, occasionally on rocks; high elevations; Ariz., N.Mex., Tex.; Mexico; Central America (Guatemala); South America.

 

Frullania mexicana has been confused and overlooked in southwestern North America. A. W. Evans (1915) first reported the species from the flora area based on specimens from the Catalina Mountains of Arizona (May 1912, Johnson s.n., fh) and the Sacramento Mountains of New Mexico (Aug. 1899, Wooton s.n., ny) that closely matched the type. Later reports by W. S. Hong (1989) of unusual F. inflata specimens from the Chiricahua Mountains of Arizona (Weber & Shushan 10635, colo, mo) and White Mountains of New Mexico (Weber 4365, colo) can also be attributed to F. mexicana. An Arizoa report of F. brittoniae from the Catalina Mountains (Hermann 19747, us) also belongs to F. mexicana. Similarly, the report of F. pluricarinata by J. Hentschel et al. (2009), based on another specimen collected in the Sacramento Mountains of New Mexico (Worthington 32194, ny), belongs to this species.

Frullania mexicana has been treated as a synonym of the North American F. inflata (A. W. Evans 1923; T. C. Frye & L. Clark 1947; R. M. Schuster 1992) as well as a synonym of the South American F. tetraptera (F. Stephani 1910; V. Schiffner & S. Arnell 1964; Y. Yuzawa 1991). The species, however, is morphologically distinct from both of these taxa. Frullania mexicana differs from F. inflata in having lobules with narrowly triangular proximal portions; subulate styli that are 2–4 cells wide at the base; differentiated marginal and median cells; shallowly 2-fid ventral leaves with entire margins; innermost gynoecial bracteoles that are highly connate with the bract on one side, while partially connate on the other; and obovoid perianths with eight or more keels. Frullania inflata, by comparison, has lobules that when inflated, lack an expanded proximal portion; mostly filiform styli that are sometimes 2 cells wide at the base; similar sized marginal and median cells; ventral leaves that are 2-fid to 1/2 or more their length and often have angulations on one or both margins; innermost gynoecial bracteoles that are consistently free from the bract; and obovate, 4–5 keeled perianths. Frullania tetraptera, although similar to F. mexicana in its lobule morphology, shape and width of the styli, and having shallowly 2-fid ventral leaves with entire margins, differs in having marginal and median cells of similar size; innermost gynoecial bracteoles that are 2/5ths connate with the bract on both sides; and clavate, 4-keeled perianths (Y. Yuzawa 1991). Frullania mexicana specimens from the southwestern United States and Mexico key in Y. Yuzawa (1991) to F. pluricarinata Gottsche, a widespread Neotropical species ranging from Mexico to Chile (Y. Yuzawa 1991). Both species have the innermost gynoecial bracteoles highly connate with the bract, as well as obovoid perianths with eight of more keels. However, a preliminary examination of the type specimen of F. pluricarinata found that it differs somewhat from F. mexicana in having less ramified, slightly wider shoots (1.2–1.4 mm wide); leaves touching and less imbricate when dry; leaves with somewhat wavy margins when moist; and partly squarrose innermost gynoecial bract lobes. A morphological or phylogenetic study that utilizes specimens from a wider distribution is needed to determine if the taxa are conspecific. If so, F. mexicana has priority as the older name by nearly 20 years.

 

4. Frullania riojaneirensis (Raddi) Ångström, Öfvers. Kongl. Vetensk.-Akad. Förh. 33 (7): 88. 1876

Frullanoides riojaneirensis Raddi, Critt. Bras.13. 1822; Frullania arietina Taylor ex C. M. Gottsche, J. B. W. Lindenberg & C. G. D. Nees

 

Plants reddish to light-brown, 1.5--2 mm wide. Stems 200--240 \um wide. Lateral leaves loosely imbricate when dry, erect spreading when moist, dorsal lobe oblong-ovate, 0.7--1 x 0.8--1.1 mm, apex sometimes decurved, base cordate; lobule distal portion 380--540 x 320--460 \um, strongly compressed at mouth, external margin uncinate, proximal portion elongated, ligulate, appendage extending beyond the basal lobe margin, keel 180--220 \um, sub-parallel to stem, free margin sinuous; stylus filiform, uniseriate, 1--3 cells long; marginal cells 12--20 \um, median cells 20--30 \um, basal cells 20--38 \um, sinuous with frequent intermediate thickenings and large, nodular trigones; oil bodies ellipsoidal, 3--6 per cell.  Ventral leaves contiguous to imbricate, orbicular, 0.3--0.6 x 0.4--0.6 mm, 2-fid in upper 80--100 \um, lobes and sinus sub-acute to obtuse. Sexual condition autoicous and paroicous. Gynoecia terminal on stem, short lateral branch, or androecial branch; bract lobe oblong-elliptical 1.4--1.6 x 0.8--1 mm, apex acute, margins irregularly dentate, lobule ovate-lanceolate, 1.2 x 0.5--0.7 mm, apex acuminate, margins sparsely dentate; bracteole 1.0--1.3 x 0.8--1 mm, 2-fid 0.3 mm, lobes sub-acute, sinus narrow, margins entire, unevenly connate on both sides. Perianth oblong, 1.6--2 x 0.8--1.1 mm, abruptly narrowed, lacking supplementary keels, beak short.

 

On twigs, branches, and trunks of trees and shrubs, sometimes on vines; low elevations; Fla.; Mexico; West Indies; Central America (Belize, Honduras, Nicaragua, Costa Rica, Panama); South America; Africa; Asia (including Indonesia).

 

Frullania riojaneirensis is known in North America from southern Florida. The species was first reported from the flora area by A. W. Evans (1915), who distinguished it from F. arietina based on differences in its sexuality. Autoicous expressions have been referred to as F. riojaneirensis (A. W. Evans 1915), whereas paroecious expressions have been named F. arietina (A. W. Evans 1897). T. C. Frye and L. Clark (1947) and R. M. Schuster (1992) note however, that the location of the gametangia is variable within a single population. A large specimen of F. arietina (Grout 25, duke) has gynoecia that are terminal on the main branches, as well as arising from short, lateral, androecial branches.

The light-brown color of F. riojaneirensis, stout shoot width, and frequency of perianths are similar to F. caulisequa. That species, however, has more strongly flattened shoots that adhere closely to the substrate, as well as trigonous perianths. The lobules of F. riojaneirensis are somewhat similar to neotropical specimens of F. gibbosa in having compressed inflated portions, and expanded proximal portions. However, in F. riojaneirensis, the inflated portion of the lobule is uncinate on the external margins, while the proximal portion extends beyond the basal margin of the lobe. By comparison, the inflated portion in F. gibbosa is slightly rostrate on the external margin, and the proximal portion is narrow, sometimes indistinguishable, terminating before or at the basal lobe margin. Frullania riojaneirensis further differs from F. gibbosa in having irregularly toothed gynoecial bract lobe and lobule margins. In F. gibbosa, the gynoecial bract lobe and lobule margins are entire or somewhat sinuate.

 

1b. Frullania subg. Diastaloba Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 55. 1884

 

Plants pinnately, bi-pinnately or tri-pinnately branched, branches becoming progressively shorter and narrower, with leaves and lobules more densely spaced. First branch appendage, 3-fid, consisting of an unequally 2-fid, flattened, lanceolate-ovate segment, and a mostly inflated, though occasionally flattened lobule. Second branch appendage lateral and consisting of two inflated lobules (and associated stylus). Lateral leaves contiguous to imbricate, flattened to deflexed when dry, erect spreading to widely spreading when moist, antical margin base truncate, rarely cordate [auriculate], not or slightly expanded and extending across stem, apex rounded to bluntly narrowed, rarely acute [apiculate], occasionally deflexed, lobes slightly convex; lobules remote from stem, erect [curved], obliquely to wide-spreading with mouth oriented towards stem, inflated throughout; styli triangular to subulate [lanceolate or lamellate]; ocelli lacking [or present]. Ventral leaves distant to contiguous, margins plane, bases cuneate. Specialized asexual reproduction none. Sexual condition autoicous or dioicous. Androecia capitate, terminal on short lateral branches. Gynoecia terminal on stem or main branch; bract lobe apex acute, sometimes apiculate, lobule apex acute, sometimes sharply so, margins with prominent stylar tooth. Perianth trigonous in section with one ventral and two lateral keels, ornamentation smooth, beak with papilloid cells.

 

Species approximately 125 (2 in flora): North America; Mexico; West Indies; Central America; South America; Asia (including Indonesia); Africa (including Madagascar); Indian Ocean Islands; Pacific Islands (including New Zealand); Australia.

 

Subgenus Diastaloba is polyphyletic as presently circumscribed, and contains at least four groups of taxa (J. Hentschel et al. 2009). The taxa tend to have lobules separated from the stem by a distance that is greater than or equal to the lobule width. Numerous morphological differences separate the two North American representatives, illustrating some of the diversity within this contrived subgenus.

 

SELECTED REFERENCES  Hentschel, J., M. J. von Konrat, T. Pócs, A. Schäfer-Verwimp, A. J. Shaw, H. Schneider, and J. Heinrichs. 2009. Molecular insights into the phylogeny and subgeneric classification of Frullania Raddi (Frullaniaceae, Porellales). Molec. Phylogen. Evol. 52: 142--156.  Schuster, R. M. 1992. The Hepaticae and Anthocerotae of North America East of the Hundredth Meridian, Vol. 5. Field Museum of Natural History, Chicago. 1--854.  Yuzawa, Y. 1988. Some little-known species of Frullania subgen. Diastaloba described from Latin America. J. Hattori Bot. Lab. 64: 437--449.

 

5. Frullania caulisequa (Nees) Montagne, Ann. Sci. Nat. Bot. sér. 2, 12: 51. 1839 Jungermannia caulisequa Nees, Fl. Brasil 1: 373. 1833; Frullania brunnea Austin; F. caroliniana Sullivant; F. gymnotis Nees & Montagne; F. obcordata Lehmann & Lindenberg

 

Plants yellow-green to light-brown, 0.8--1.7 mm wide. Stems 70--100 \um wide. Lateral leaves imbricate and flattened when dry, erect spreading to wide spreading when moist, dorsal lobe broadly ovate, 0.5--0.7 x 0.6--0.8 mm; lobule saccate or sometimes sub-cylindric, 140--200 x 80--120 \um; stylus subulate, 4--5 cells long and 1--3 cells wide; marginal cells 14--22 \um, median cells 16--22 \um, basal cells 22--40, with occasional intermediate thickenings and small trigones; oil bodies ellipsoidal, 3--5 per cell. Ventral leaves ovate to orbicular, 0.3--0.5 x 0.2--0.3 mm, 2-fid to about 0.2 mm, lobes sub-acute to acute, sinuses narrowly to broadly acute, margins entire or with an obtuse tooth on one or both sides. Sexual condition autoicous. Gynoecia bract lobe elliptical, 1.0--1.3 x 0.6--0.8 mm, margins entire, lobule ovate-lanceolate, 0.6--0.8 x 0.4--0.5 mm, margins mostly entire to minutely dentate [with 1--2 blunt teeth]; bracteole oblong, 0.6--0.8 x 0.3--0.5 mm, 2-fid to about 0.2 mm, lobes and sinuses acute, margins entire, free or connate on one side with bract. Perianth obovate, 1.1--1.5 x 0.8--1.2 mm, dorsally flattened, gradually broadened to the apex, beak short.

 

On the trunks, branches, and twigs of trees and shrubs, occasionally on logs, rarely epiphyllous; low to moderate elevations; Ala., Fla., Ga., La., Miss., N.C., S.C., Va; Mexico; West Indies; Central America (Belize, Honduras, Nicaragua, Costa Rica, Panama); South America.

 

Frullania caulisequa is an autoicous species with distinctly flattened shoots, broadly ovate dorsal lobes, ventral leaves mostly 1--3 times wider than the stem, and mostly saccate lobules that are distant from the stem by about two times their width. In North America, F. caulisequa frequently occurs with F. ericoides and F. kunzei, although neither of these species are likely to be confused with it. The leafy shoots of F. kunzei are mostly narrower (0.4--0.8 mm wide), dark-red in color, and have lobules that are nearly contiguous with the stem. The leafy shoots of F. ericoides (0.8--1.3 mm wide) overlap more in size with F. caulisequa, but the stems typically spread from the substrate instead of closely adhere to it, and the lobules are galeate, slightly compressed, and tend to be rostrate on the external margin. Although both F. caulisequa and some expressions of F. ericoides share a trigonous perianth shape, F. ericoides is a dioicous species with the gynoecia produced on short lateral branches. Furthermore, the perianths of that species often have short, supplementary keels and a consistently tuberculate ornamentation.

L. Clark (1958) reported the morphologically similar F. gymnotis from Florida, Louisiana, and South Carolina. Outside of North America, F. gymnotis occurs in the West Indies, Central America, northern South America, and Brazil. It differs most notably from F. caulisequa in its minutely dentate versus entire gynoecial bract margins. Y. Yuzawa (1988) regards F. caulisequa to be a polymorphic species that includes synonyms with entire or sub-entire gynoecial bract margins (F. obcordata) as well as those with margins having 1--2 blunt teeth (F. miradorensis Lindenberg & Gottsche). Following this concept, the gynoecial bract margins appear to express clinal variation, to which F. gymnotis has been included as another conspecific expression (S. R. Gradstein & D. Costa 2003). All of the North America specimens examined for this treatment had entire gynoecial bract margins. The Florida (Koch 7215) and Louisiana (Svihla 344) specimens reported by L. Clark (1958) could not be located for re-examination, but the South Carolina specimen (Batson s.n., f) was re-determined by R. M. Schuster as F. obcordata (= F. caulisequa). L. Clark (1958) gave the date of the Batson specimen as 21 Dec. 1956, whereas the f specimen examined by Schuster has the date 12 Dec. 1956. Despite the different dates, the specimens are believed to be the same, due to the fact that the f specimen’s label is re-typed, and not original. The synonymy of F. gymnotis with F. caulisequa is followed here. Although reevaluation of the Florida and Louisiana specimens is needed, the finding of a close relationship between F. obcordata (= F. caulisequa) and other morphologically similar species (J. Hentschel et al. 2009) suggests that F. caulisequa is in fact polymorphic.

 

6. Frullania taxodiocola R.M. Schuster, Phytologia 53(5): 364. 1983

 

Plants copper-red to blackish, 0.6--0.7 mm wide. Stems 80 \um wide. Lateral leaves contiguous to imbricate, erect spreading to widely spreading when moist, dorsal lobe ovate, 0.2--0.3 x 0.3 mm; lobule galeate, 150--160 x 80--90 \um, external margin crenulate-sinuate, with protuberant cell, incised near mouth; stylus triangular, 2--3 cells long and 2--3 cells wide; marginal cells 8--13 \um, occasionally with differentiated cells that are sometimes pigmented and bulging, median cells 10--18, basal cells 16--32 \um, with occasional intermediate thickenings and small to large trigones; oil bodies ellipsoidal to fusiform, 2--3 per cell. Ventral leaves oblong, 0.2 x 0.1--0.2 mm, 2-fid to 50--70 \um, lobes acute to subacute, sinuses narrowly to broadly acute, margins with a blunt angulation or pointed tooth on one or both sides. Specialized asexual reproduction by caducous leaves. Sexual condition dioicous. Gynoecia bract lobe ovate-lanceolate, 0.6--0.7 x 0.3--0.4 mm, margins entire to coarsely serrate; lobule ovate-lanceolate, 0.5--0.6 x 0.3 mm, margins dentate to coarsely serrate; bracteole ovate, 0.5 x 0.4 mm, 2-fid for 0.2 mm, margins coarsely serrate, free from bracts on both sides. Perianth [oblong-clavate, 1.3 x 0.6 mm dorsally flattened, gradually broadened to the apex, with occasional differentiated cells that are pigmented and bulging, beak short.]

On the buttressed bases of Taxodium; low elevations; Fla.

 

Frullania taxodiocola was described from two specimens that were collected in 1958 at the base of a cypress in a Florida swamp (R. M. Schuster 1983). The species has not been reported outside of its type locality, although a specimen from Belize, with mature perianth, was recently discovered in mo (Allen 19044). The Belize specimen has mature perianths, not previously described for F. taxodiocola. However, unlike the perianth in other Frullania subgenera, those in subg. Diastaloba s. lat., appear to be stenotypic and have few taxonomically significant characters. The perianth are terminal on the stem or main shoot, slightly exserted, smooth, oblong-clavate, 1.3 x 0.6 mm and trigonous in section with one ventral and two lateral keels. Furthermore, the perianth has a few scattered, pigmented cells, similar to the marginal cells of the dorsal lobe and lobule. Lastly, the perianth beak is short-cylindric, 0.1 x 0.08 mm and has numerous single celled protuberances densely covering the inner beak surface. It seems likely that the Florida station is at the northern edge of the distribution for F. taxodiocola; an otherwise tropical or sub-tropical species. However, resolution of the nomenclature requires a better understanding of the Frullania species in the West Indies. Regionally, F. taxodiocola is most similar to F. caulisequa in its lobules positioned remotely from the stem. The two species are otherwise unlikely to be confused. Frullania taxodiocola differs from F. caulisequa in its copper-red color; narrower leafy shoots; dioicous sexuality; and coarsely serrate margins of the ventral leaves, gynoecial bract, and bracteoles. Superficially, F. taxodiocola is more similar to F. donnellii (subg. Meteoriopsis), which is also a small, dark red species distributed in the southeastern United States. Although F. donnellii is an autoicous species, the androecia and gynoecia are sometimes distantly separated. Furthermore, F. donnellii has rounded, deflexed dorsal lobe apices; similarly narrow, galeate lobules; and toothed gynoecial bract margins like that of F. taxodiocola. The lobules, however, differ from those of F. taxodiocola in their orientation and position to the stem. In F. donnellii, the lobules are mostly parallel with the stem, and separated by a distance that is mostly less than or equal to the lobule width. By comparison, the lobules of F. taxodiocola are obliquely to widely spreading, and separated from the stem by a distance greater than the width of the lobule, often about two times wider. Lastly, the external margins of the lobules of F. taxodiocola are crenulate-sinuate and incised near the mouth. In F. donnellii, the external margins of the lobules are smooth, and while the lobule mouth can be somewhat crenulate, it is never incised.

 

1c. Frullania subg. Frullania

 

Plants un-branched or pinnately branched, rarely bi-pinnately branched. First branch appendage, 3-fid, consisting of an unequally 2-fid, flattened, lanceolate-ovate segment, and a flattened or inflated lobule. Second branch appendage lateral and consisting of two flattened segments or inflated lobules, or a mixture of both (and associated stylus). Lateral leaves loosely to strongly imbricate when dry, erect spreading, widely spreading, or squarrose when moist, with antical margin not, slightly, or strongly expanded and extending across stem; lobules approximate or contiguous with stem, parallel or subparallel, inflated throughout, galeate, cap-shaped, or flattened and lanceolate; styli filiform to lamellate; ocellii occasionally present and scattered. Ventral leaves approximate, contiguous, evenly-spaced, distant or imbricate, margins entire or toothed, bases narrowed and cuneate, sometimes short decurrent or cordate. Specialized asexual reproduction none, gemmae, or by caducous leaves. Sexual condition autoicous, paroicous or dioicous. Androecia capitate or spicate, terminal on short lateral branch, ocassionally sessile on stem or main branch. Gynoecia terminal on stem, main branch, or lateral branch; with or without innovations; bract lobule margins often with prominent stylar tooth. Perianth trigonous in section, sometimes with supplementary keels, ornamentation smooth or tuberculate, beak sometimes with papilloid cells, [or perianth unknown].

 

Species approximately 300 (21 in flora): North America; Mexico; West Indies; Central America; South America; Asia (including Indonesia); Africa (including Madagascar); Indian Ocean Islands; Pacific Islands (including New Zealand); Australia.

 

Subgenus Frullania is characterized by its mostly inflated, helmet-shaped lobules that are about as long as wide. The lobules can sometimes be cap-shaped as in F. riparia (somewhat wider than long), compressed at the mouth, or slightly to strongly rostrate at the external margin. Unlike subg. Chonanthelia, the lobules are connected to the dorsal lobe by a keel that is perpendicular or nearly so with the stem. Furthermore, unlike North America members of subg. Diastaloba and subg. Thyopsiella, the lobules have a tendency to sometimes be explanate. Flat lobules in F. ericoides, F. inflata, F. plana, F. riparia, F. sabaliana, and F. saxicola are lanceolate in shape, have rounded to acute apices, and plane to slightly decurved margins. Flat lobules are also found in F. cucullata (subg. Meteoriopsis), but in that species the lobules have strongly deflexed margins.

Autoicous, paroicous and dioicous species are included in subg. Frullania. A molecular phylogenetic study by J. Hentschel et al. (2009) mostly supports this grouping, with the exception that F. inflata, an autoicous species, is aligned with F. bolanderi, a dioicous species, in a clade that is independent from the rest of subgenus. Since no defining morphological characters currently delineate these two species from subg. Frullania, they are retained within that subgenus (J. Hentschel et al. 2009).

The autonym, subg. Frullania, has been used as a replacement name for subg. Trachycolea and subg. Thyopsiella, depending on which of these is interpreted to include the type species. When G. Raddi (1818) described Frullania, he simultaneously described two superfluous species, F. major and F. minor, both containing older names in their synonymy. Under F. major, G. Raddi (1818) included F. dilatata, whereas under F. minor was F. tamarisci. A. W. Evans (1918) selected F. dilatata as the type of Frullania. Most typifications made by Evans are now considered to be mechanical since he often designated the type as the first species listed. Mechanical typifications can be superseded (J. McNeil et al. 2012). As argued by J. Hentschel et al. (2015), however, Evans’s selection was not mechanical, because he selected the older name, F. dilatata, instead of F. major, the first name listed. Since F. dilatata is also the type of subg. Trachycolea, that subgenus is replaced by the autonym, subg. Frullania.

 

SELECTED REFERENCES  Bombosch, A., A. Wieneke, A. Busch, R. Jonas, J. Hentschel, H.-P. Kreier, B. Shaw, A. J. Shaw, and J. Heinrichs. 2010 Narrow species concepts in the Frullania dilatataappalachianaeboracensis complex (Porellales, Jungermanniopsida): evidence from nuclear and chloroplast DNA markers. Plant Syst. Evol. 290:151--158.  Evans, A. W. 1897. A revision of the North American species of Frullania, a genus of Hepaticae. Trans. Connecticut Acad. Arts Sci. 10: 1--39, pl. 1--15. Frye, T. C. and L. Clark. 1947. Frullania. Hepaticae of North America, part V. p. 735--782. Schuster, R. M. 1992. The Hepaticae and Anthocerotae of North America East of the Hundredth Meridian, Vol. 5. Field Museum of Natural History, Chicago. 1--854.

 

7. Frullania appalachiana R.M. Schuster, Phytologia 53 (5): 366. 1983  E

 

Plants green to red-brown, 0.6--1 mm wide. Stems 100--140 \um wide. Lateral leaves distant to contiguous, antical base arching over stem, dorsal lobe broadly ovate 0.3--0.5 x 0.4--0.7 mm, slightly convex, apex rounded, sometimes decurrved, base cordate; lobules helmet shaped, 160--240 x 180--260 \um, truncate at the mouth, weakly to strongly compressed, slighty rostrate on the external margin; styli subulate, 4--7 cells long, 2--4 cells wide; marginal cells 16--20 \um, median cells 16--24 \um, basal cells 16--36 \um, with intermediate thickenings and small trigones; oil bodies oval to ellipsoidal, 2--5 per cell. Ventral leaves obovate, 0.1--0.3 x 0.1--0.4 mm, 2-fid to 80--120 \um, lobes and sinus narrowly acute, margins bluntly angulated, base narrowed. Specialized asexual reproduction by caducous leaves. Sexual condition dioicous. Androecia spicate on short lateral branch. Gynoecia terminal on stem or abbreviated lateral branch; bract lobe ovate to elliptical, 0.3--0.4 x 0.2--0.3 mm, apex rounded, margins entire, lobule lanceolate to ovate-lanceolate, 0.3 x 0.1--0.2 mm, apex acute, margins entire; bracteole oblong, 0.2--0.3 x 0.4--0.5 mm, 2-fid to 140 \um, lobes and sinus narrowly acute, margins with slender tooth on each side, free. Perianth oblong-obovate, 1.1--1.3 x 0.8--1 mm, distinctly compressed, postical keel broad with several accessory keels, tuberculate, beak short, broad.

 

On trunks and branches of trees; moderate to high elevations; Ala., Ark., Ga., Mo., N.C., Okla., S.C., Tenn., Va. 

 

R. M. Schuster (1992) regarded F. appalachiana to be restricted to the southern ranges of the southern Appalachian Mountains, particularly at elevations at or above 1,500 meters. Recent fieldwork and herbarium studies by Paul Davison, however, have revealed this species to be more broadly distributed in the southeastern and interior United States. Frullania appalachiana is similar to F. eboracensis in its color, size, and frequency of caducous leaves. Furthermore, both species have ovate dorsal lobes with cordate, sometimes auriculate bases; large inflated lobules; sinuous median cells; differentiated marginal cells that are sometimes pigmented and bulging; sublate styli; ventral leaves that are about one to one and a half times the stem width; and mostly erect gynoecial bracteoles when moist. Fertile stems of the two species readily differ in their perianth ornamentation: strongly tuberculate in F. appalachiana, while smooth or mostly smooth in F. eboracensis. Frullania appalachiana also has toothed gynoecial bracteole margins, whereas the bracteoles margins of F. eboracensis are entire. Sterile shoots are unfortunately more difficult to distinguish from one another, particularly in fragmentary or mixed specimens. On well-developed shoots, the lobules of F. appalachiana tend to have strongly compressed mouths, as well as ventral leaves with 1--2 teeth on each margin. By comparison, the lobules of F. eboracensis tend not to be compressed at the mouth, and the ventral leaf margins are mostly entire or have blunt angulations. These characters are subject to much variability as noted by the numerous caveats listed in R. M. Schuster’s (1992) discussion of the complex. To further complicate matters, F. virginica is sometimes treated as conspecific with F. eboracensis. Frullania virginica has tuberculate perianth similar to those of F. appalachiana. The species is further subject to confusion due to its similar shoot width, sinuous leaf cells, angulate to toothed ventral leaf margins, toothed gynoecial bracteoles, and perianth with accessory keels. Frullania virginica, however, does not produce caducous leaves, nor does it have scattered differentiated marginal cells that are deeply pigmented and bulging as in F. appalachiana. Lastly, the gynoecial bracts of F. virginica tend to be squarrose when moist, while the innermost bracteole is low connate on one side with the bract.

 

8. Frullania bolanderi Austin, Proc. Acad. Nat. Sci. Philadelphia 1869: 226. 1869

Frullania petalumensis Gottsche, in Bolander; F. hallii Austin

 

Plants dark green to reddish, 0.6--0.9 mm wide. Stems 100--130 \um wide. Lateral leaves loosely imbricate, weakly squarrose when moist, antical margin arching over stem; doral lobe ovate, 0.3--0.7 x 0.3--0.5 mm, slightly convex, apex rounded, base truncate; lobule galeate, 250--320 x 250--400 \um, compressed at the mouth; styli lamellate, 6--10 cells long and 3--6 cells wide; marginal cells 12--16 \um, median cells 16--22 \um, basal cells 18--32 \um, mostly lacking intermediate thickenings, trigones large; oil bodies spherical, 6--9 per cell. Ventral leaves distant, obovate, 0.1--0.3 x 0.1--0.2 mm, 2-fid to 100 \um, lobes and sinus bluntly acute, margins entire or with one or two teeth on each side, base narrowed. Specialized asexual reproduction by caducous leaves on short, flagelliform branches. Sexual condition dioicous. Androecia spicate on short lateral branch. Gynoecia terminal on the stem or main branch; bract lobe lanceolate, ovate, or elliptical, 0.7--0.8 x 0.5--0.6 mm, apex acute to rounded, margins entire, lobule lanceolate to ovate-lanceolate, 0.5--0.6 x 0.3--0.4 mm, apex bluntly acute to rounded, margins entire but with a stylar tooth; bracteole lingulate to oblong, 0.4--0.7 x 0.2--0.4 mm, 2-fid to 100--300 \um, lobes and sinus acute to narrowly acuminate, margins entire or with blunt angulations, unevenly connate on one or both sides. Perianth obovate, 1.25 x 0.8 mm, two-angled postical keel and 1 or more antical keels, plus additional supplementary ridges, narrowed at apex, beak short, broad.

 

On trunks and branches of trees, on logs, as well as sometimes on rocks and soil; low to high elevations; B.C., Nfld., N.S., Ont., Que., Alaska, Calif., Iowa, Maine, Mich., Minn., N.Y., Ore., Vt., Wash., Wis; Europe; Asia.

 

Frullania bolanderi can be readily recognized by its leafless, erect or curving, flagelliform shoots, with short internodes and persistent and squarrose ventral leaves. Aside from these specialized shoots, F. bolanderi might be confused with F. caucasica as both species are dioicous; have ovate dorsal lobes; large, sub-quadrate lobules; and obovate ventral leaves with blunt angulations or teeth on the margins. Frullania bolanderi differs, however, in that the antical margin of the dorsal lobe base is truncate, whereas in F. caucasica it is cordate. Furthermore, F. bolanderi has larger, lanceolate styli, that are 5--11 cells long and 3--6 cells wide at the base, whereas the styli of F. caucasica are short-triangular and are only 3--5 cells long and 2--3 cells wide at the base. In North America, F. catalinae, F. chilcootiensis, F. cobrensis, F. hattoriana, and F. stylifera all have lanceolate or lamellate styli that are similar in size to that of F. bolanderi. The styli of F. catalinae are 6--8 cells long and 2--4 cells wide, but that species differs in its autoicous sexuality, having squarrose leaves when moist, and frequency of explanate lobules. The styli of F. chilcootiensis are also 6--8 cells long and 2--4 cells wide, but that species has an auriculate-appendiculate dorsal lobe base. The styli of F. cobrensis are 12--16 cells long and 4--6 cells wide at the base, however it is also an autoicous species, has strongly cordate dorsal lobe bases, ocelli in the basal cells of the dorsal lobe, and lobules that narrow towards the mouth. The styli of F. hattoriana are 7--12 cells long and 4--6 cells wide. However, F. hattoriana is somewhat narrower than F. bolanderi (leafy shoots 0.36--0.56 versus 0.9--1.2 mm wide), autoicous, has strongly cordate dorsal lobe bases, and more prominently toothed ventral leaves. Lastly, the styli of F. stylifera are 10--15 cells long and 4--10 cells wide, typically larger than the styli of F. bolanderi. Frullania stylifera further differs in its autoicous sexuality and cordate dorsal lobe bases.

Except for outlying populations near the Great Lakes, the distribution of F. bolanderi is largely restricted to the eastern and western coasts (R. M. Schuster 1992). Eastern North America material often lacks perianths, whereas western specimens readily have perianths. Moreover, the innermost gynoecial bracteole and bract in eastern North America populations differ somewhat from western populations in that they have narrow bracts with acute apices, and margins that are free from the bracteole or low connate with it on one side (R. M. Schuster 1992). In western specimens, such as the California plants illustrated by A. W. Evans (1897), the bracts are broad, rounded at the apices, and highly connate on one or both sides with the bracteole. Additional study is needed to determine if the eastern populations belong to the same taxon as the western populations.

 

9. Frullania brittoniae A. Evans, Trans. Connecticut Acad. Arts Sci. 10 (1): 15. 1897

Frullania dilatata Underwood   E

 

Plants green to reddish brown, 0.8--1.2 mm wide. Stems 100--140 \um wide. Lateral leaves imbricate, flat or slightly convex, antical margin arching over stem, dorsal lobe reniform-orbicular, 0.4--0.6 x 0.5--0.7 mm, apex rounded, slightly decurved, base cordate; lobule galeate, 0.2--0.3 x 0.2 mm, slightly to strongly compressed at mouth, slightly rostrate in the external margin; stylus lanceolate, 5--8 cells long and 2--3 cells wide at base; marginal cells 14--16 \um, median cells 20--24 \um, basal cells 22--32 \um, with occasional intermediate thickenings and distinct trigones; oil bodies oval to ellipsoidal, 4--7 per cell. Ventral leaves distant, broadly orbicular to elliptical, 0.2--0.3 x 0.2--0.4 mm, 2-fid to about 80--100 \um, lobes and sinus acute, margins irregularly dentate above the middle with 2--3 sharp to blunt angulations on one or both sides, base narrowed. Sexual condition dioicous. Androecia capitate, terminal on short lateral branches. Gynoecia terminal on the stem or main branch, lobes ovate, 1.3 x 0.8 mm, apex rounded or obtuse, sometimes apiculate, margins entire to slightly crenulate, lobule ovate to lanceolate, 1 x 0.5 mm, apex sub-acute to apiculate, with stylar tooth, bracteole ovate, 0.9 x 0.5 mm, 2-fid to 0.2--0.3, lobes and sinus acute, margins entire or unidentate on one or both sides, free or slightly connate on one side with bract. Perianth obovate, 1.9 x 1.2 mm, truncate above and abruptly narrowed, compressed at the sides and with a postical keel and 1 or more short, supplementary antical and postical ridges, the whole surface with scattered tubercles, especially numerous on keels, beak long and slender, tapering.

 

On the trunks and branches of trees, as well as sometimes on rocks; low to high elevations; Ala., Ark., Conn., Fla., Ga., Ill., Ky., La., Mass., Miss., Mo., N.H., N.Y., N.C., Okal., Pa., S.C., Tenn., Va., W.Va.

 

Frullania britoniae has been subject to much confusion in North America with many older reports requiring verification. Due to its ventral leaves with numerous teeth or angulations on both margins, the species has been confused in eastern North America with F. appalachiana and F. virginica, whereas it has been confused with F. caucasica and F. wrightii in the west. Compared to the eastern species, the shoots of F. brittoniae are mostly wider (0.8--1.2 mm wide); the lobules are lightly to strongly compressed at the mouth and have slightly rostrate external margins; and the perianths have long, slender, tapering beaks. Furthermore, the gynoecial bract lobes are erect, free from the bracteole, and have entire margins. Frullania ericoides has lobules that are similarly rostrate on the external margin. However, the perianths are terminal on short lateral branches instead of terminal on the stem or main branch. Furthermore, the ventral leaves of F. brittoniae are distant, while those of F. ericoides are more or less imbricate.

Several literature reports of F. brittoniae in western North America, such as from Colorado and New Mexico, belong to F. caucasica. The two species are similar in having decurved dorsal lobe apices and cordate bases; styli that are 2--3 cells wide at base; and ventral leaf margins that are irregularly dentate above the middle. The ventral leaves of F. caucasica, however, have fewer teeth than those of F. brittoniae. Furthermore, F. brittoniae never has caducous leaves, and the cells are more distinctly trigones with intermediate thickenings. When compared to F. wrightii, sexuality and connation of the innermost gynoecial bracteole and bract differentiate the two species. Frullania brittoniae is dioicous and has the innermost gynoecial bracteole free from the bract, whereas F. wrightii is autoicous and has the innermost gynoecial bracteole connate on one side with the bract. The ornamentation of the perianth, as well as the length of the perianth beak provides additional differences.

 

10. Frullania catalinae A. Evans, Trans. Connecticut Acad. Arts Sci. 10(1): 11. 1897  E

 

Plants green to reddish-brown, 0.8--1.2 mm wide. Stems 100 \um wide. Lateral leaves imbricate, squarrose when moist, antical margin arching over stem; dorsal lobe ovate, 0.6 x 0.5 mm, apex rounded, slightly decurved, base truncate; lobule broadly galeate, 160--180 x 200--230 \um, often explanate, 300 x 230 \um; styli subulate, 6--8 cells long and 2--4 cells wide; marginal cells 16 \um, median cells 16--20 \um; basal cells 16--36 \um, slightly thick-walled, no intermediate thickenings, trigones inconspicuous. Ventral leaves broadly rhombic, 0.3 x 0.2--0.3 mm, 2-fid to 0.5 their length, lobes and sinus obtuse to sub-acute, margins with blunt to sharp angulations on each lateral margin. Sexual condition autoicous. Androecia capitate, sessile or on short lateral branch. Gynoecia terminal on stem or main branch, bract lobe ovate, 1.2 x 0.7 mm, apex rounded, bract lobule narrowly ovate, 0.8 x 0.4 mm, apex rounded to sub-acute, margin with stylar tooth, bracteoles narrowly ovate, 0.7 x 0.3 mm, 2-fid to 1/3, lobes lanceolate, sinus narrow, margins entire, connate on one side with bract. Perianth obovate, 1.5 x 0.7 mm, gradually narrowed from middle to a short broad beak, 1 postical keel, 1 or more less pronounced antical keels and postical ridges, beak with papillae.  

 

On rocks as well as the trunks and branches of trees and shrubs; low to moderate elevations; Calif.

 

Frullania catalinae is geographically restricted to California’s Catalina Islands and the adjacent neighboring mainland counties. The species is similar to F. wrightii in shoot width; autoicous sexuality; dorsal lobe size and shape; subulate styli; ventral leaf size, shape and dentation; and cell dimensions. However, the leaves of F. catalinae are squarrose instead of spreading when moist, the dorsal lobe base is truncate instead of cordate, and the innermost gynoecial bracteole is deeply versus shallowly 2-fid. The lobules of F. catalinae also tend to be flattened, whereas those of F. wrightii are more often inflated. Although F. catalinae has been treated as a synonym of F. inflata (A. W. Evans 1923; L. Clark & R. D. Svihla 1944; T. C. Frye & L. Clark 1947; R. M. Schuster 1992), it differs significantly from that species in having bracteoles connate on one side with the gynoecial bract, gradually narrowed perianths, and shorter perianth beaks. By comparison, F. inflata has the innermost gynoecial bracteoles free from the bract, abruptly narrowed perianths, and tubular perianth beaks. Frullania catalinae is more comparable to F. saxicola in that it too has frequently explanate lobules, bracteoles connate on one side with the inner gynoecial bract, gradually narrowed perianths, and short perianth beaks. Frullania saxicola, however, differs in its narrower styli (1--2 cells wide at base versus 2--4 cells wide at base) and narrower ventral leaves with less pronounced dentation on the margins.

 

11. Frullania caucasica Stephani, Sp. Hepat. 4: 440. 1910

Frullania eboracensis subsp. caucasica (Stephani) R.M. Schuster, Hepat. Anthocerotae N. Amer. 5: 143. 1992.

 

Plants green to yellowish brown, 0.7--1 mm wide. Stems 110--150 \um wide. Lateral leaves contiguous to imbricate, antical margin arching over stem; dorsal lobe broadly ovate, 0.3--0.8 x 0.2--0.7 mm, convex, apex rounded, decurved, base auriculate; lobule galeate, 150--270 x 130--300 \um, truncate compressed at the mouth, sometimes rostrate at the external margin, lanceolate when explanate; styli lanceolate, 5--8 cells long and 2--3 cells wide at base; marginal cells 10--14 \um, median cells 14--22 \um, basal cells 20--30 \um, thin-walled, flexuous, some intermediate thickenings, tigones small, triangular to nodular, oil bodies ellipsoidal, 4--9 per cell. Ventral leaves distant to approximate, ovate to rhombic-ovate, 0.2--0.4 x 0.1--0.4 mm, 2-fid to 1/3 the length, lobes and sinus sub-acute, margins sharply angulate to toothed on both sides near base of sinus, base cuneate. Specialized asexual reproduction by caducous leaves [or gemmae]. Sexual condition dioicous. Androecia unknown. Gynoecia terminal on stem or main branch, immature. Perianth unknown.

 

On shaded boulders and rock outcrops; moderate to high elevations; Ariz., Colo., N.Mex., Mexico; Europe; Asia.

 

Frullania caucasica is morphologically similar to F. parvistipula in having caducous leaves. S. Hattori (1978) considered the two taxa to be synonymous, whereas R. M. Schuster (1992) recognized them as subspecies of F. eboracensis. The two differ in that F. caucasica has slightly larger lobules than F. parvistipula, in addition to more distinctly toothed or angulate ventral leaf margins. Examination of the Arizona material briefly discussed by R. M. Schuster (1992) found the specimens to be identical with the New Mexico material cited as F. parvistipula by A. Bombosch et al. (2010). A. Bombosch et al. (2010) advocated for species level recognition within the F. eboracensis complex and showed that New Mexico specimens named F. parvistipula phylogenetically differed from eastern North America specimens of F. eboracensis. The Arizona and New Mexico specimens further differ from typical F. caucasica and F. parvistipula in that the styli are lanceolate, 5—8 cells long and 2--3 cells wide at the base. According to S. Hattori (1978), the styli in both of these species are uniseriate and filiform. Nevertheless, the name F. caucasica is applied to the western North America specimens due to the overall similarities in plant color, dorsal lobe size and shape, decurved apical margins, and ventral leaf margin dentation. A preliminary examination of F. caucasica specimens from the Caucaucus region found the styli to occasionally be 2--3 cells wide at the base. More study is needed to determine the extent of this variability.

Frullania caucasica differs from F. eboracensis in its green to yellowish brown color, slightly wider stems, and lack of mature gynoecia and androecia in North America populations. Furthermore, the shoot width is often noticeably wider, but this difference is obscured by the tendency for the lobe apices to be decurved. Of the numerous F. caucasica specimens examined, none had the black or deep-green color frequently associated with F. eboracensis. Occasionally though, F. caucasica plants are tinted with a similar red-brown color. Further differences between the two species include the shape of the median cells and the extent and location of teeth on the ventral leaf margins. Frullania caucasica has leaf cells with small trigones and a few intermediate thickenings. The ventral leaves have a tooth or sharp angulation on both lateral margins at about the base of the sinus. By comparison, the cell walls in F. eboracensis freely develop intermediate thickenings and often have nodular trigones. The ventral leaf margins of F. eboracensis are mostly entire or sometimes have a blunt angulation on one or each side, typically at about the middle of the leaf.

 

12. Frullania chilcootiensis Stephani, Bot. Jahrb. Syst. 8 (2): 98. 1886   E

 

Plants brownish, 0.4--0.5 mm wide. Stems 60--70 \um wide. Lateral leaves imbricate, distant to contiguous, antical base auriculate-appendiculate, arching over the stem; dorsal lobe broadly ovate, 230--280 x 230--280 \um, convex, apices rounded, decurved; lobules saccate, 170 x 110 \um, truncate at mouth; styli triangular-lanceolate, 2--3 cells wide; marginal cells 10 \um, median cells 18 \um, basal cells 18--24 \um, thick-walled. Ventral leaves obovate, 160--200 x 80--120 \um, 2-fid to about ½ the length, lobes and sinus obtuse, base cuneate, margins with blunt angulation or obtuse tooth. Sexual condition unknown. Androecia unknown. Gynoecia bract lobe oblong-ovate, apex obtuse, lobule ovate lanceolate, acute, with stylar tooth; bracteole oblong, 2-fid to about 1/3 the length, lobes sub-acute, sinus obtuse. Perianth immature, 500 x 360 \um, with several keels and a broad, flaring beak.

 

On the bark or trees; moderate elevations; Alaska.

 

Frullania chilcootiensis, like F. sabalina, is known only from a single North American locality. The species was collected in the late 1800s by brothers Arthur and Aurel Krause, who botanized in the Alaskan Panhandle (E. Hultén 1968). A. W. Evans (1897) provided a description and short discussion of the species based on drawings that were sent to him by F. Stephani. He considered F. chilcootiensis to be incompletely known, but did not speculate on the taxon’s validity. Stephani’s drawings were later published by T. C. Frye and L. Clark (1947), who speculated that the species might be a depauperate expression of F. bolanderi. The type specimen of F. chilcootiensis was examined by J. D. Godfrey and G. A. Godfrey (1980), who further compared and contrasted the species with F. hattoriana and F. eboracensis. A fragment of the type specimen in fh has narrow shoots, dorsal lobes with auriculate bases, and large triangular-lanceolate styli. Based on these characters, F. chilcootiensis would seem to be a good match for F. hattoriana. Frullania hattoriana is, however, autoicous whereas the sexuality of F. chilcootiensis is unknown. More specimens are needed to determine the affinities and variability of F. chilcootiensis.

 

13. Frullania cobrensis Gottsche ex Stephani, Hedwigia 33 (3): 142. 1894

 

Plants blackish green, 0.5--0.6 mm wide. Stems 80--100 \um wide. Lateral leaves imbricate, antical margin arching over stem; dorsal lobe broadly ovate, 275--500 x 300--440 \um, convex, apex rounded, slightly decurved, base auriculate; lobule saccate, oval, 200--240 x 160--230 \um; styli triangular-lanceolate; marginal cells 24 \um, median cells 26 \um, basal cells 14--28 \um, firm-walled, intermediate thickenings sparse, trigones present, ocelli 24--28 \um in basal group, oil bodies ellipsoidal 2--4 per cell. Ventral leaves distant to contiguous, obovate to oblong 110--210 x 60--150 \um, 2-fid to about 60 \um the length, lobes triangular, bluntly acute, sinuses acute, base cuneate, margins entire or with a blunt tooth on one or both sides. Sexual condition autoicous. Androecia capitate or short spicate, sessile or on a short lateral branch. Gynoecia terminal on stem or main branch, bract lobe obovate to oblong, 350--580 x 270--330 \um, 0.6--0.7 2-fid, apex rounded, margin entire; bract lobule obovate to lanceolate, 275--550 x 160--200 \um, apex rounded, margins entire, antical margin with a large sublinear stylar tooth; bracteole ovate to oval 340--500 x 170--250 \um, 2-fid 0.25--0.4 its length, lobes acute, sinus narrowly acute, margins entire, connate on one side with bract. Perianth obovate, 0.7--1 x 0.5--0.7 \um, dorsiventrally flattened, rounded into truncate apex, 1 dorsal keel, 1--2 lateral keels, and 2 ventral keels, sometimes with additional supplementary keels, beak with papillae.

 

On the trunks, branches, and twigs of trees and shrubs; low elevations; Ala., Fla.; West Indies.

 

Frullania cobrensis is the only North America species in subg. Frullania with ocellate dorsal lobes. The ocelli are 24--28 \um wide and confined to the basal cells. Apart from this character, the species is likely to be confused with F. inflata and F. rappii, due to similarities in their autoicous sexuality, presence of inflated lobules, and obovate perianth with several undulate keels. Frullania cobrensis differs from these two taxa in having auriculate versus rounded dorsal lobe bases and lanceolate versus filiform to subulate styli. Frullania stylifera is also morphologically similar to F. cobrensis in that it too has auriculate dorsal lobe bases and large styli. The styli of F. stylifera, however, are lamellate, and the perianth is globose to obovoid, with fewer keels. D. Griffin and D. A. Breil (1982) made a systematic search for F. cobrensis in Florida and found it to be ecologically restricted, but otherwise more common than previous literature reports suggested.

 

14. Frullania davurica Hampe ex Gottsche, Lindenb. & Nees, Syn. Hepat. 3: 422. 1845

 

Plants yellowish green to red-brown, or black, 1.2--1.5 mm wide. Stems 160--180 \um wide. Lateral leaves imbricate, antical margin arching over stem; dorsal lobe broadly ovate to orbicular, 0.6--0.8 x 0.9--1 mm, apex rounded, slightly decurved, base cordate; lobules galeate, 240--270 x 180--210 \um, slightly to strongly compressed at mouth, frequently rostrate and incurved in the external margin, sometimes oriented towards the stem; styli filiform to subulate, 2--4 cells long and 1--2 cells wide; marginal cells 16--20 \um, median cells 20--28 \um, basal cells 20--34 \um, thick-walled, intermediate thickenings frequent, trigones large, oil bodies spherical to ellipsoidal, 5--9 per cell. Ventral leaves somewhat distant to contiguous, sub-orbicular to reniform, 0.4--0.6 x 0.6--0.7 mm, emarginate, narrowed to base or slightly decurrent. Specialized asexual reproduction lacking. Sexual condition dioicous. Androecia and Gynoecia unknown in the flora area.

 

In moist depression, on sedge tussock of tundra; low elevations; Alaska; Europe; Asia.

 

Frullania davurica is unlike any other North America Frullania in having sub-orbicular to reniform ventral leaves that are emarginate and about four times the width of the stem. Furthermore, the galeate lobules have a rostrate external margin that, along with the lobule mouth, is sometimes oriented towards the stem. The single known North America specimen was collected by W.C. Steere in 1951 in the Alaskan Brooks Range, and had originally been determined as F. jackii (W. C. Steere & H. Inoue 1978). Frullania jackii has been considered a subspecies of the closely related F. davurica (Hattori 1976), as well as a synonym of that species (R. N. Schljakov 1982). R. Mues et al. (1984) regarded the two taxa as distinct, with the distribution of F. jackii restricted to central and southern Europe, while that of F. davurica is more widespread throughout eastern, far-eastern, and central Asia. Recent molecular studies (J. Hentschel et al. 2009) reiterate the close relationship of these taxa. The Alaska specimen (Steere 16851, ny) agrees morphologically with Asiatic specimens of F. davurica in the mo herbarium, as well as R. N. Schljakov’s (1982) description and illustration of the species.

 

15. Frullania eboracensis Gottsche, in Lehmann, Nov. Stirp. Pug. 8: 14. 1844  E

Frullania sullivantii Austin, Proc. Acad. Nat. Sci. Philadelphia 1869: 226. 1869

 

Plants black or deep-green, often tinged with brown or red, dull, 0.5--1 mm wide. Stems 80--110 \um wide. Lateral leaves contiguous to imbricate, antical margin arching over stem, dorsal lobe oval to sub-orbicular, 0.4--0.6 x 0.5--0.6 mm, convex, apex rounded, slightly decurved, base cordate; lobule galeate, 260--300 x 200--240 \um, truncate at base; stylus subulate, 3--6 cells long and 2--3 cells wide at base; marginal cells 10--14 \um, median cells 16--20 \um, basal cells 24--30 x 16--20 \um, sinuous, thick-walled with trigones and intermediate thickenings, oil bodies oval 2--5 per cell. Ventral leaves distant to approximate, ovate to rhombic-ovate, 200--250 x 150--230 \um, 2-fid 1/3 the length, lobes and sinus sub-acute, margins entire, unidentate on one or both sides, or bluntly angulate, base cuneate. Specialized asexual reproduction by gemmae or caducous leaves. Sexual condition dioicous. Gynoecia terminal on the stem and main branch, lobe ovate, 800--850 x 450--550 \um, apex rounded, margins entire or slightly crenulate, lobule narrower, ovate, 600--750 x 300--350 \um, apex acute or obtuse, bearing a small stylar tooth at middle; bracteole ovate, 800--850 x 550, 2-fid 0.2--0.3, lobes and sinus acute, margins entire to irregularly dentate, free or connate on one side. Perianth obovate or obcuneate, 1.1--1.3 x 0.8--1 mm, more or less compressed, abruptly narrowed into a short, broad keel, but without distinct supplementary ridges, mostly smooth, sometimes weakly tuberculate, keels sometimes undulate.

 

On the trunks and branches of trees and shrubs, as well as on boulders and rock outcrops, occasionally as an epiphyll; low to high elevations; N.B., Nfld., N.S., Ont., Que., Ala., Ark., Conn., D.C., Del., Ga., Ill., Ind., Iowa, Kans., Ky., Maine, Md., Mass., Mich., Minn., Mo., N.H., N.J., N.Y., N.C., Ohio, Okla., Pa., S.C., Tenn., Tex., Vt., Va., W.Va., Wis.

 

Frullania eboracensis is an abundantly common species in northeastern North America, where it occurs on the bark of trees in both natural and disturbed habitats. A. W. Evans (1897) remarked that except for sterile specimens, F. eboracensis and F. virginica are recognizably distinct based on differences in their perianth ornamentation and number of keels. He further noted that the two species are somewhat geographically correlated, with F. eboracensis distributed predominantly in northern regions, whereas F. virginica occurs in the south (A. W. Evans 1897). Later, A. W. Evans (1906) re-considered the distinctiveness of F. virginica and treated it as a synonym of F. eboracensis due to apparent morphological and geographic overlap. Although R. M. Schuster (1992) similarly recognized F. eboracensis and F. virginica to have fairly discreet distribution patterns, he considered the taxa, along with F. caucasica and F. parvistipula, to be subspecies within a complex. Using molecular data, A. Bombosch et al. (2010) has argued for recognition of species, noting that F. eboracenis and F. virginica remain morphologically distinct despite their partially sympatric distributions. Fertile shoots of F. eboracensis can be separated from F. virginica by their dark green to blackish brown, rounded or trigonous perianth that are mostly smooth, but occasionally have a few tubercles on the keels. By comparison, the perianths of F. virginica are green to yellowish, and differ in having more extensive tuberculae, as well as carinae. Furthermore, the perianths of F. virginica have numerous supplementary keels. Additional differences are found in the perianth beaks, which are short and broad in F. eboracensis, while more elongated, slender, and flaring in F. virginica. Lastly, the gynoecial bracts of F. eboracensis are mostly erect, with the bracteole margins free and entire on both sides. In F. virginica, the innermost gynoecial bracts are mostly squarrose, and the innermost bracteole margins, by comparison, have several distinct teeth and are connate on one-side with the bract. Caducous leaves of F. eboracensis, like those of F. appalachiana and F. caucasica, have scattered differentiated marginal or sub-marginal cells on the dorsal lobe that tend to be bulging and pigmented in contrast to the surrounding leaf cells. The cells elongate and multiply, developing eventually into thick-walled, short, hyaline rhizoids, shoots, or cells masses. The development of these adventive branches and propagula has been detailed in F. eboracensis by A. Lorenz (1912).

 

16. Frullania ericoides (Nees) Montagne, Ann. Sci. Nat., Bot., sér. 2, 12: 51. 1839

            Jungermannia ericoides Nees, in Martius, Fl. Bras. Enum. Pl. 1: 346. 1833;

            Frullania squarrosa Dumortier

 

Plants yellowish green to reddish brown, 0.8--1.3 mm wide. Stems 150--200 \um wide. Lateral leaves densely imbricate and rolled around the stem when dry, squarrose when moist, antical base extending beyond stem; dorsal lobe ovate, 0.6--0.8 x 0.5--0.7 \um, apex rounded, base auriculate; inflated lobule saccate, 220--400 x 250--350 \um, truncate at mouth and usually rostrate, lanceolate when explanate, 200--400 x 150--250 \um, acute; stylus subulate, composed of 4--5 cells long and 2--3 cells long at base; marginal cells 18--20 \um, median cells 20--24 \um, basal cells 30--40 x 20--24 \um, firm-walled, sinuous, with intermediate thickenings, and nodulose to triangular trigones, oil bodies ellipsoidal, 2--4 per cell. Ventral leaves distant to imbricate, orbicular, 0.4--0.6 x 0.4--0.7 mm, 2-fid 0.5--0.7 \um, lobes and sinus acute, base narrowed, margins plane, entire, or with an obtuse tooth or angulation on one or both sides above middle. Sexual condition dioicous. Androecia capitate, sessile on short lateral branches. Gynoecia terminal on short lateral branch, bract lobes ovate 1.0--1.3 x 0.8--1.2 mm, 2-fid to ¾ the length, apex rounded, margins entire, lobules ovate to lanceolate, acute, with 2--3 small teeth on free margins, bracteole ovate, 0.8--0.9 x 0.5--0.6 mm, 2-fid to 1/3 the length, lobes narrowly elliptic, acute, sinus acute, margins with a tooth on both lateral margins, connate with one or both bracts. Perianth obovoid, 1.2--1.6 x 0.9--1.2 mm, dorsally compressed, keels 3 or more, with numerous scattered tubercles, abruptly narrowed into a long beak.

 

On the trunks and branches of trees and shrubs, logs, boulders and rocks outcrops, as well as on soil; low to moderate elevations; Ala., Ariz., Ark., Conn., Fla., Ga., Ill., Ind., Kans., Ky., La., Miss., Mo., N.J., N.C., Ohio, Okla., Pa., S.C., Tenn., Tex., Va., W.Va.; Mexico; West Indies; South America; Africa; Asia; Australia.

 

The morphological plasticity of F. ericoides in North America has been thoroughly outlined and illustrated by R. M. Schuster (1992). Most specimens, despite their variability, can be distinguished from other Frullania species in the flora area by their sub-convolute leaves that are often rolled around the stem when dry, and become squarrose when moist. Frullania cucullata has similarly sub-convolute leaves when dry, but when moist become erect-spreading. Conversely, F. catalinae and F. gibbosa have squarrose leaves when moist, but the leaves in those taxa are imbricate when dry. Sterile, lax and weakly squarrose leaved expressions of F. ericoides occur at the western and northern edge of its distribution in eastern North America, and are morphologically comparable to F. riparia (R. L. McGregor 1955; R. M. Schuster 1953, 1992). Both species are similar in their green to reddish brown color, shoot widths (0.7--1.3 mm wide), broadly ovate dorsal lobes with expanded bases, and dorsal lobe median cells with similar bulging trigones and intermediate wall thickenings. Both species also have a tendency to not form lobules. Lastly, both species have filiform styli as well as large ventral leaves that are 2--4 times the stem width, often with blunt angulations on one or both margins. In mixtures of the two species from Missouri and Arkansas, the shoots of F. ericoides are generally less ramified than the shoots of F. riparia, whose numerous secondary branches are frequently branched again. Furthermore, the first branch appendage in F. ericoides usually consists of a 2-fid ventral segment and a weakly formed lobule. In F. riparia, the first branch appendage is trifid. Next, the lateral leaves of F. ericoides are often more or less densely imbricate, while those of F. riparia tend to be more remote, contiguous, or loosely imbricate. Also in F. ericoides, at least a few inflated lobules can usually be found. When inflated, the galeate lobules of F. ericoides are unlike the cap-shaped lobules of F. riparia. Lastly, the ventral leaves are often more closely spaced: contiguous to imbricate in F. ericoides, while remote to contiguous in F. riparia.

 

17. Frullania hattoriana J.D. Godfrey & G. Godfrey, J. Hattori Bot. Lab. 48: 321. f. 1--2. 1980   E

 

Plants golden or reddish-brown, 0.3--0.5 mm wide. Stems 62--96 \um wide. Lateral leaves loosely imbricate, antical margin arching over stem, dorsal lobe 0.2--0.3 x 0.3--0.4 mm, apex rounded, decurved, base auriculate on both the antical and postical margin; lobules obovate, 184--256 x 128--160 \um, somewhat narrowed at mouth; styli lanceolate-subulate, 7--12 cells long and 4--6 cells wide; marginal cells 10--20 \um, median cells 10--26 \um, basal cells 20--40 x 10--20 \um, thin-walled, occasional intermediate thickenings, bulging trigones. Ventral leaves distant, rectangular to ovate-spatulate, 184--280 x 120--240 \um, 2-fid to 0.3--0.5 the length, lobes triangular, acute to obtuse, sinus narrow, acute to obtuse, margins with 1--2 angular projections, rarely entire. Sexual condition autoicous. Androecia capitate, on short lateral branches. Gynoecia terminal on stem or main branch, with innovations, bract lobe 0.3--1 x 0.2--0.8 mm, apex rounded, bracteole ovate, 0.4--0.9 x 0.2--0.5 \um, 2-fid 0.3--0.45, connate on one side with bract. Perianth obovate, with 6--9 folds, narrowed to a broad, stout beak with flaring mouth, occluded with clavate projecting cells.

 

On bark of Tsuga and Alnus; moderate elevations; B.C.

 

Frullania hattoriana is known only from a few stations in western British Columbia, Canada. The species is similar to F. wrightii in its autoicous sexuality, broad styli, toothed ventral leaf margins, innermost gynoecial bracteole that are connate on one side with the bract, pluriplicate perianth, and broad, stout perianth beak. Frullania hattoriana differs from that species in its narrower shoots (0.3--0.5 mm wide versus 0.9--1.2 mm wide), narrower stem width, and more frequent branches. Furthermore, the dorsal lobe bases of F. hattoriana are auriculate-appendiculate, while the lobules are slightly longer than wide. By comparison, F. wrightii has cordate dorsal lobe bases and lobules that are about as long as wide. Both species have been overlooked in the flora area and require more study.

 

18. Frullania inflata Gottsche, in Gottsche, Lindenberg, & Nees, Syn. Hepat. 424. 1844

Frullania inflata var. communis R.M. Schuster   E

 

Plants green, brown, sometimes reddish, dull, 0.7--1.3 mm wide. Stems 80--120 \um wide. Lateral leaves imbricate, flat or slightly decurved when dry, spreading when moist, antical base arching over stem; dorsal lobe orbicular to broadly ovate, 550--640 x 360--600 \um, convex, apex rounded, decurved, base truncate; lobules galeate, 250 x 250 \um, truncate at the base,  when explanate lanceolate-lingulate; stylus filiform to subulate, 4--8 cells long and (1)2--3 cells wide at base; marginal cells 16--20 \um, median cells 24--28 \um, basal cells 28--32 \um, firm-walled, lacking intermediate thickenings, trigones small or inconspicuous, oil bodies ellipsoidal, 4--5 per cell. Ventral leaves evenly spaced, plane, orbicular to obovate, 300--320 x 240--300 \um, 2-lobed to 1/3 the length, lobes and sinus acute, bases cuneate, margins entire or with a blunt angulation or tooth on one or both sides. Sexual condition autoicous. Androecia capitate, sessile or on short branches. Gynoecia terminal on stem or main branch, bract lobes ovate to obovate, 850 x 500 \um, apex rounded, margins entire, lobule narrowly ovate, 650 x 400 \um, apex rounded to sub-acute, margins with a small stylar tooth, bracteole ovate, 520--650 x 380--430 \um, deeply 2-fid with acute lobes and acute to obtuse sinus, margins entire or with minute teeth at base, free from bracts. Perianth obovate, 0.6--0.9 mm, abruptly narrowed to a short, broad beak, with 1 postical keel and one or more supplementary keels.

 

On the trunks and branches of trees, as well as on rocks; low to moderate elevations; Ala., Ark., Conn, D.C., Del., Fla., Ga., Ill., Iowa, Kans., Ky., La., Md., Miss., Mo., Nebr., N.J., N.C., Okla., Pa., S.C., Tenn., Tex., Va., W.Va.

 

Frullania inflata is one of the most common Frullania species in the southeastern United States. The species was described from a specimen collected by H. Beyrich near ‘Whastite’ and the ‘Red River’. The locality ‘Whastite’, has been attributed to Louisiana (T. C. Frye and L. Clark 1947) and Arkansas (R. M. Schuster 1992), but is seemingly best placed in Oklahoma as it appears to be a misspelling of ‘Washita’, a river that joins the Red River at the Oklahoma and Texas border.

Frullania inflata has been treated in North America as a complex of morphologically similar taxa sharing autoicous, sometimes paroicous sexuality; capitate androecia that are sessile or borne on abbreviated lateral branches; gynoecia that terminate the stem or main branch; perianth with a smooth ornamentation; perianth beaks bearing papilloid cells on the inner beak surface; and lack of gemmae (A. W. Evans 1923; R. M. Schuster 1992). Differences in the shape and size of the innermost gynoecial bracteoles, as well as the extent (or lack of) connation between the bracteoles and innermost bract, however, provides taxonomically useful characters for resolving the complex. Frullania inflata is separated from F. catalinae, F. rappii, and F. saxicola by its innermost gynoecial bracteole margins that are free from the bract. By comparison, F. catalinae, F. rappii, and F. saxicola have innermost gynoecial bracteole margins that are connate on one-side with the bract.

R. M. Schuster (1985) described F. inflata var. communis for plants with smooth inner perianth beak surfaces, after R. Grolle (1970) noted that the type of F. inflata has papillae on the inner perianth beak surface, similar to that of F. saxicola. Specimens with this morphology correspond to what had been previously regarded by A. W. Evans (1897) as F. inflata. A. W. Evans (1923) later noted that the inner beak surface of F. inflata can also have papillae, best seen in cross section of the beak, to which he credits C. C. Haynes for the discovery. In cross section of the beak, the type specimen of F. inflata var. communis (Blomquist 29231, duke) also has papillae on the inner beak surface. It is regarded as a synonym of F. inflata.

 

19. Frullania oakesiana Austin, Proc. Acad. Nat. Sci. Philadelphia 1869: 226. 1869

 

Plants reddish or purplish brown, sometimes green, dull, 0.6--0.9 mm wide. Stems 60--1 \um wide. Lateral leaves contiguous to imbricate when dry, slightly squarrose when moist, antical side not or rarely extending across stem; dorsal lobe ovate to orbicular, 0.3--0.4 x 0.2--0.3 mm, slightly convex, apex rounded, base rotundate to truncate, margins somewhat incurved; lobule consistently galeate, subquadrate, 220--240 x 190--220 \um, mouth truncate; stylus subulate, 4--5 cells long and 1--3 cells wide at base; marginal cells 12--20 \um, median cells 18--32 \um, basal cells 20--26 x 14--20 \um, firm walled, trigones very small and concave-sided, no intermediate thickenings; oil bodies spherical to oval, 3--5 per cell. Ventral leaves distant to approximate, obovate, 160--180 x 130--160 \um, 2-lobed 0.30--0.50 \um of the length, lobes narrowly triangular, acute, sinus acute, base cuneate, margins entire or unidentate on one or both sides. Sexual condition autoicous. Androecia capitate, terminal on short lateral branches. Gynoecia terminal on stem or leading branch, bract lobe ovate, 520--550 x 280--310 \um, apex rounded, margins entire, bract lobule narrowly ovate, 500--550 x 230--250 \um, apex acute, margins entire except for prominent stylus tooth near middle, bracteole ovate, 650--750 x 350--500 \um, lobes and sinuses acute, margins entire, connate on one side with lobule. Perianth emergent, obovate, 1--1.5 x 0.8--1 mm, gradually broadened to the apex, dorsiventrally flattened, trigonous, with one ventral and two lateral keels of equal size, smooth or slightly crenate, beak short, the mouth fringed with projecting, papillae.

 

On tree trunks and branches; moderate to high elevations; Nfld., N.S., Ont., Que., Maine, Mass., Mich., Minn., N.H., N.Y., N.C., Tenn., Vt., Wis.; Europe; Asia.

 

Frullania oakesiana is a small, reddish brown, autoicous species similar to F. kunzei. The dorsal lobes of F. oakesiana are, however, ovate to orbicular, rounded or truncate at the base, and rarely arch across the stem. Those of F. kunzei, by comparison, are more reniform, with bases slightly cordate and extending across the stem on the antical margin. Frullania oakesiana also has lobules that differ from F. kunzei in their subquadrate versus cylindric shape, and ability to obscure most of the dorsal lobe surface when viewed from the ventral side. Small expressions of F. eboracensis can be confused with F. oakesiana due to their similarly narrow shoot width as well as presence of large, inflated lobules. However, F. eboracensis differs in having cordate dorsal lobe bases, strongly sinuous median cell walls of the dorsal lobe, spicate androecia, as well as a dioicous sexuality. Frullania rappii might also be confused with F. oakesiana due to its similarly rounded or truncate dorsal lobe base, inflated lobules, gynoecial bracts that are connate on one side with the bracteole, and autoicous sexuality. The two species differ in that F. rappii lacks teeth on the ventral leaf margins and has an oblong versus obovate perianth. Unlike F. eboracensis, F. kunzei, and F. rappii, the perianths of F. oakesiana become somewhat stipitate at maturity and are partially exserted above the bracts.

 

20. Frullania plana Sullivant, Mem. Amer. Acad. Arts (n.ser.) 4: 175. 1849 [1850]  E

 

Plants green, sometimes tinged brown, 1.2--1.4 mm wide. Stems 160--180 \um wide. Lateral leaves imbricate, antical margin arching over the stem; dorsal lobe orbicular to reniform, 0.6--0.7 x 0.7--0.8 mm, moderately convex, apex rounded, decurved, antical base auriculate, postical base strongly cordate; lobule galeate, 180--240 x 180--200 \um, approximate to stem, compressed to weakly beaked at mouth, sometimes flattened and lanceolate with recurved margins; stylus mostly uniseriate and filiform, 3--5 cells long and 1(--2) cells wide; marginal cells 14--16 \um, median cells 18--26 \um, basal cells 28--44 x 18--24 \um, cell firm-walled, with intermediate thickenings and conspicuous nodular trigones, oil bodies ellipsoidal, 3--5 per cell. Ventral leaves distant, orbicular, 320--400 x 440--520 \um, 2-fid to about 140 \um the length, lobes obtuse, sinus acute, margins entire, base cordate. Sexual condition autoicous. Androeica capitate, terminal on a short lateral branch. Gynoecia terminal on a short lateral branch, bract lobe ovate, 0.8--1 x 0.5--0.7 mm, apex rounded, margins irregularly crenulate, lobule narrowly-ovate, 0.6--0.8 x 0.3--0.4 mm, apex rounded, margins irregularly crenulate, bearing a stylar tooth, bracteole ovate, 0.5--0.6 x 0.4 mm, 2-fid to 0.2--0.3 mm, lobes and sinus acute, margins variably lacinate, dentate or crenulate, free from bracts. Perianth oblong or obovate, 1.5 x 0.7--0.9 mm, narrowed into a short, broad beak, trigonous, compressed on the sides with a broad postical keel and a shallow antical sulcus, smooth.

 

On rock ledges as well as sides of boulders; sometimes on tree bark; moderate to high elevations; Ala., Ark., Conn., Ga., Ky., Mass., Mo., N.J., N.Y., N.C., Ohio, S.C., Pa., Tenn., Va., W.Va.

 

Frullania plana is endemic to eastern and interior North America. The species is distinct from other similarly sized, saxicolous Frullania in having dilated antical and postical dorsal lobe bases; small (180--240 x 180--200 \um) galeate lobules when inflated; minute styli; and orbicular ventral leaves with entire margins and cordate bases. Frullania riparia is perhaps most likely to be confused with F. plana due to its small lobules (180--300 x 180--400 \um), vestigial styli, and generally saxicolous habit. Both species also have lax shoots, with loosely imbricate leaves and often explanate lobules. Frullania riparia, however, has mostly angulate ventral leaf margins as well as cuneate to short decurrent ventral leaf bases. Frullania riparia is also a dioicous species with gynoecia that terminate the stem or main branch. By comparison, F. plana is an autoicous species with both the androecia and gynoecia borne on abbreviated branches. Although F. ericoides has its androecia and gynoecia borne on similarly short, lateral branches, as well as having variable androecia that range from capitate to short spicate, that species also differs in its dioicous sexuality. Furthermore, F. ericoides has more closely imbricate leaves that roll around the stem when dry and become squarrose when moist.

 

21. Frullania rappii A. Evans, Bryologist 15: 22. 1912  E

 

Plants dull green, brown or dark red, lustrous, 0.6--0.75 mm wide. Stems 80--100 \um wide. Lateral leaves imbricate, flat when dry, but becoming squarrose on older stems, wide-spreading when moist, antical margin arching over stem; dorsal lobe ovate, 0.75 x 0.6 mm, slightly convex, apex rounded, decurved, margins entire and somewhat sinuate; lobules galeate, 0.27 x 0.24 mm, truncate and compressed at mouth; styli subulate; marginal cells 14 \um, median cells 14--23 \um, basal cells 24--, firm-walled, intermediate thickenings few, trigones distinct, triangular with straight or bulging sides; oil bodies ellipsoidal, 4--5 per cell; Ventral leaves distant to imbricate, broadly ovate, 0.3 mm x 0.3 mm, 2-fid ¼ to ½, margins plane, bases cuneate, with a narrow sinus and blunt, broad lobes, rounded to acute at the apex, entire or slightly crenulate on both sides. bract lobe oblong to obovate, about 0.9 x 0.6 mm., apex rounded to bluntly acute, margins entire, lobule ovate, 0.7 x 0.4--0.5 mm, apex rounded to obtuse, with a slender lanceolate stylar tooth near base, bracteole oblong-ovate, 0.7 x 0.4 mm., 2-fid to 2/3 the length, lobes acute, sinus narrow, margins connate on one side with bract. Perianth oblong, 1 x 0.75 mm, compressed with sharp lateral keels, rounded to truncate at the apex with a short beak, entire or very minutely crenulate at the mouth, ventral keel broad and sharply two-angled, dorsal keel sharp, sometimes with 1--2 supplementary keels.

 

On the trunks and branches of trees; low to moderate elevations; Fla., N.C., Tex.

 

Frullania rappii, like some expressions of F. inflata, have stems with similarly large, galeate lobules. Furthermore, both species have a few short, rounded, papillae on the internal surfaces of the perianth mouth. Frullania rappii differs from F. inflata in having more pronounced cell-wall thickenings in the median cells of the lobe, as well as slightly smaller ventral leaves with entire margins. However, the best character for distinguishing these species is the differences in their connation between the gynoecial bracteole and bract. In F. rappii, the bracteoles are connate on one side with the gynoecial bracts, whereas in F. inflata, the bracteoles are free from the bract. The presence of partially connate gynoecial bracteoles align F. rappii with F. catalinae, F. wright, F. saxicola, and F. stylifera.

 

22. Frullania riparia Hampe ex Lehmann, Nov. Stirp. Pug. 7: 14. 1838

Frullania aeolotis Montagne & Nees

 

Plants green to brownish, dull, 1.1--1.7 mm wide. Stems 110--130 \um wide. Lateral leaves distant to imbricate, spreading when moist, antical margin arching over stem; dorsal lobe broadly ovate, 0.6--0.8 x 0.5--0.7 mm, apex rounded, plane to slightly decurved, base auriculate; lobules cap-shaped, 180--300 x 180--400 \um, truncate at base, lanceolate when explanate, 300 x 120 \um; styli minute; marginal cells 14 \um, median cells 18 \um, basal cells 30 \um, slightly thickened walls, occasional intermediate thickenings, distinct trigones. Ventral leaves distant, obovate to orbicular, 0.3--0.6 x 0.3--0.7 mm, 2-fid to 1/3, lobes and sinus sub-acute, margins with blunt angulations on one or both sides or sometimes entire, base narrow, sometimes short decurrent. Sexual condition dioicous. Androecia unknown. Gynoecia terminal on stem or main branch, bract lobe ovate, apex rounded, margins entire, bract lobule lanceolate, apex acute, margin with stylar tooth; bracteole narrowly ovate, 2-fid to 1/3, lobes and sinus sub-acute, margins irregularly dentate, sub-entire, free from bracts. Perianth immature.

 

On rock outcrops, ledges and boulders, also on trees; moderate elevations; Ariz., Ark., Colo., D.C., Del., Ill., Iowa, Kans., Ky., Mass., Minn., Mo., N. H., N.Mex., N.Y., N.C., Ohio, Okla., Pa., Tenn., Tex., Vt., Va., W.Va., Wis.; Mexico

 

Frullania riparia frequently has lax, approximate to loosely imbricate leaves. Due to the frequency of explanate lobules, the species is often confused with F. inflata. The two differ in that F. riparia has auriculate dorsal lobe bases, whereas the lobe bases of F. inflata are cordate. Furthermore, the lobules of F. riparia are cap-shaped when inflated, the styli are uniseriate and filiform, and the ventral leaves are about 3--4 times the width of the stem. In F. inflata, the inflated lobules are larger and helmet-shaped, the styli are often subulate, and the ventral leaves are narrower, approximately 1--1.5 times the width of the stem. Mature perianths and androecia of F. riparia are unknown (A. W. Evans 1897; R. M. Schuster 1992).

 

23. Frullania sabaliana R.M. Schuster, Phytologia 53: 365. 1983  E

 

Plants green to fulvous brown, sometimes copper-red, dull, 1.5--1.8 mm wide. Stems 100--220 \um wide. Lateral leaves imbricate to contiguous when dry, slightly squarrose when moist, antical side not or rarely extending across stem,; dorsal lobe oblong-ovate to ovate-obcordate, 0.7--0.9 x 0.6--1 mm, flat or slightly convex, apex rounded, decurved, base truncate; lobule explanate, ovate-lanceolate, 250--400 x 120--160 \um, apex acute; stylus small-triangular, 1--2 cells long and 2--4 cells wide; marginal cells 12--14 \um, median cells 20--26 \um, basal cells 25--30 x 30--35 \um, thick-walled, some intermediate thickenings, trigones large, oil bodies spherical, 3--5 per cell. Ventral leaves distant, oblong, 300--400 x 190--270 \um, 2-lobed for 300--500 \um the length, lobes lanceolate, acute, sinus acute, flat, bases cuneate, margins entire. Sexual condition autoicous. Androecia capitate, sessile or terminal on short lateral branches. Gynoecia terminal on stem or main branch, with subfloral innovations, bract lobe elliptical to obovate, 0.8--0.9 x 0.5--0.6 mm, 2-lobed for 500--600 \um the length, apex rounded to bluntly acute, sometimes apiculate, margins entire, bract lobule ovate-lanceolate, 0.7--0.8 x 0.5 mm, apex sharply acute to apiculate, margins entire except for prominent styli; bracteole oblong, 0.4 x 0.3 mm, 2-lobed for 1/3 the length, lobes and sinus acute, margins entire, free or short-connate on one side with bract. Perianth obovate, 1.1--1.3 x 0.8--0.9 mm, obtusely narrowed to apex, beak short or lacking, 3--4-carinate in cross-section, with 2 lateral and 1--2 ventral keels, supplementary ridges lacking, smooth or with low papillae, mouth smooth.

 

On the trunks, bases, knees, and exposed roots of Taxodium, as well as on Sabal palm; Fla.

 

Frullania sabalina is known only from the type locality in Florida, where it was first collected in 1958, and then again in 1983, by R. M. Schuster. Compared to other species in the F. inflata complex with which it might be confused, F. sabalina has wider shoots (1.5--1.8 mm wide) that spread from the substrate. In addition, the species has uniformly explanate lobules, and ventral leaves that are not or only slightly larger than the stem. By comparison, the lobules in most expressions of F. inflata are usually at least partially inflated. Based on the size of the plants and the autoicous sexuality, an older name for the species is likely to be found among neotropical Frullania species.

 

24. Frullania saxicola Austin, Proc. Acad. Nat. Sci. Philadelphia 1869: 225. 1869  E

 

Plants yellowish-green to brown, 0.7--0.9 mm wide. Stems 110--120 \um wide. Lateral leaves imbricate, flat or convex when dry, spreading when moist, antical margin extending over stem; dorsal lobe broadly ovate, 320--440 x 380--450 \um, apex rounded, decurved, base truncate; lobule galeate when inflated, 180--200 x 180--200 \um, lanceolate when explanate; stylus uniseriate and filiform, 2--3 cells long; marginal cells 16--22 \um, median cells 18--26 \um, basal cells 24--38 \um, thin-walled, intermediate thickenings absent, trigones small. Ventral leaves distant, ovate, 155--175 x 145­-170 \um, 2-lobed 0.3--0.4, lobes and sinus acute, base cuneate, margins entire. Sexual condition autoicous. Gynoecia terminal on stem or main branch; bract lobe 0.5--0.7 x 0.3--0.5, apex rounded to bluntly acute; bract lobule 0.5--0.6 x 0.1--0.3 mm, apex acute; bracteole 0.4 mm x 120--180 \um, 2-fid to 160 \um, lobes and sinus acute, margins entire, connate on one side with bract. Perianth bulbous to obovate, 0.8--1.1 x 0.8--0.9 mm, beak short.

 

On trunks and branches of trees, as well as on rocks; moderate elevations; Ark., Iowa, Kans., Minn., Mo., Nebr., N.Mex., Tex.

 

Frullania saxicola was described from syntype New Jersey and Texas specimens that differ from F. virginica Lehm. in their shorter, repeatedly branched stems, frequently explanate lobules, perianth that lack tuberculae on the keels, and shorter, broader perianth beaks (C. F. Austin 1869). The New Jersey specimens were later distributed by Austin (1873) as number 104 in his Hepat. Bor. Amer. Exsic. Specimens in fh and mo contain a mixture of F. inflata and F. virginica. However, the Texas material, deposited in Sullivant’s herbarium, now fh, contains only F. saxicola. A. W. Evans (1897) initially considered F. saxicola to be a synonym of F. virginica Lehm., based on an examination of an exsiccatae duplicate. However, after studying additional duplicates, he later re-instated the species (A. W. Evans 1910). Frullania saxicola and F. inflata Gottsche are morphologically similar in their broadly ovate dorsal lobes with rounded bases, median leaf-cells with small trigones and few intermediate thickenings, frequently explanate lobules, autoicous sexuality, and 4–5 keeled perianth that lack tuberculae. According to A. W. Evans (1910), F. saxicola differs from F. inflata in its short, often cup-shaped perianth beak as well as in the papillose ornamentation of the inner beak surface. Frullania inflata, by comparison, is described as having a longer perianth beak with a mostly smooth inner surface (A. W. Evans 1910). A. W. Evans (1923) later found that the inner perianth beak surface of F. inflata can also have papillae, best seen in cross section of the beak, to which he credits Caroline Conventry Haynes (1858–1951) for the discovery. Based on this finding, L. Clark and R. D. Svihla (1944) regarded F. saxicola to be a synonym of F. inflata. T. Frye and L. Clark (1947) accepted this synonymy, as did R. Grolle (1970), who further noted that the type of F. inflata (in S) also has papillae on the inner perianth beak surface. R. M. Schuster (1985, 1992) cited Grolle’s findings as evidence that the species treated by A. W. Evans (1897, 1910) as having a smooth inner perianth beak surface, now lacked a name. He subsequently described this taxon as F. inflata var. communis R.M. Schust. (= F. inflata), whereas F. inflata var. inflata (= F. saxicola) was retained for specimens having papillose inner perianth beak surfaces. The recent re-examination of the type of F. inflata var. communis, however, has found that it also has numerous papillae on the inner beak surface, as seen through cross section of the perianth beak. In fact, all examined North America specimens determined as F. inflata and F. saxicola have papillose cells on the inner perianth beak surface. The two species are better resolved by their differences in the shape of their perianth, shape of the innermost gynoecial bracteole sinues and lobes, and in the extent (or lack of) connation between the bracteoles and innermost gynoecial bracts.

 

25. Frullania stylifera (R.M. Schuster) R.M. Schuster, Hepat. Anthocerotae N. Amer. 5: 210. 1992   E

Frullania inflata var. stylifera R.M. Schuster, Phytologia 53: 366. 1983

 

Plants reddish brown or green, dull, 1.0--1.2 mm wide. Stems 110--120 \um wide. Lateral leaves contiguous to imbricate with somewhat reflexed margins when dry, erect to wide-spreading when moist, antical side extending across stem; dorsal lobe reniform-ovate, 0.5--0.6 x 0.7--0.8 mm, convex, apex rounded, decurved, base cordate; lobule galeate, quadrate, 355--420 x 310--400 \um, compressed in the lower half, mouth truncate; stylus lamelliform, oblong, 10--15 cells long and 4--10 cells wide; marginal and median cells 20--24 \um, basal cells 26--32 x 20 \um, thin-walled, no intermediate thickenings, trigones small; oil bodies spherical to ellipsoidal, 8--16 per cell. Ventral leaves distant, flat, obovate, 330--365 x 340--380 \um, 2-lobed to 1/3 their length, lobes broadly triangular, sub-acute, sinus acute, bases cuneate, margins with an obtuse angulation on one or both sides. Sexual condition autoicous. Androecia capitate, terminal on short lateral branches. Gynoecia terminal on stem or main branch, bract lobe elliptical to ovate, 0.7--0.8 x 0.5--0.6 mm, apex rounded, margins entire, bract lobule lanceolate, 0.7--0.8 x 0.4 mm, apex acute, bracteole narrowly ovate, 0.5 x 0.4 mm, 2-lobed to 0.2 the length, lobes and sinus rounded, margin connate on one side with bract. Perianth globose to obovoid, 1 x 1 mm, abruptly contracted with 1 dorsal keel and 2 ventral keels, no accessory keels, beak with papillae.

 

Trunks and branches of shrubs and trees, occasionally on logs; moderate elevations; Ark., Kan, Minn., Mo., Okla., Tenn.

 

Frullania stylifera is a seemingly endemic species that is common in the North America interior, although it has been consistently overlooked or confused with F. inflata and F. eboracensis (J. J. Atwood 2016). The three species can be distinguished by their differences in dorsal lobe shape, styli size and shape, ventral leaf size, number of oil bodies, and sexuality and androecia shape. Frullania stylifera has broadly reniform-ovate dorsal lobes compared to the more orbicular, ovate, to broadly ovate dorsal lobes of F. inflata and F. eboracensis. Also, F. stylifera has a massive stylus consisting of 10--15 cells long and 4--10 cells wide. Both F. inflata and F. eboracensis have substantially smaller, filiform or shortly-triangular styli that are usually 3--6 cells long and 1--3 cells wide. The large ventral leaves of F. stylifera are 2--3 times the width of the stem, whereas the ventral leaves of F. inflata and F. eboracensis are generally only one to one and half times the width of the stem. Furthermore, F. stylifera can have upwards to 16 oil bodies per cell, an exceptional number compared to F. inflata and F. eboracensis that each have approximately 3--6 oil bodies per cell. Lastly, Frullania stylifera and F. inflata are autoicous species that have capitate androecia, compared to the dioicous F. eboracensis that has spicate androecia.

 

26. Frullania virginica Gottsche, in Lehmann, Nov. Stirp. Pug. 8: 19. 1844  E

            Frullania eboracensis subsp. virginica (Lehmann) R.M. Schuster, Hepat. Anthocerotae N. Amer. 5: 147. 1992

 

Plants green to brownish, 0.7--0.9 mm wide. Stems 100--110 \um wide. Lateral leaves imbricate, antical margin arching over stem, dorsal lobes sub-orbicular, 0.6 x 0.5 mm, apex rounded, decurved, base cordate; lobules galeate, 280 x 180 \um, slightly truncate at base; stylus filiform to subulate, 4--5 cells long and 1--2 cells wide; marginal cells 14 \um, median cells 18 \um, basal cells 24 \um, sinuous, thick-walled with intermediate thickenings and conspicuous trigones, oil bodies oval to ellipsoidal 2--4 per cell. Ventral leaves rhombic-ovate, 0.2--0.3 x 0.2--0.3 mm, 2-fid to 1/3, lobes and sinus sub-acute, entire to rarely unidentate. Sexual condition dioicous. Gynoecia terminal on the stem or main branch, bract lobes squarrose, ovate to orbicular, 0.7--0.8 x 0.5--0.6 mm, apex rounded, lobule ovate to lanceolate, 0.7--0.8 x 0.3--0.4 mm, apex acute to apiculate, with stylar tooth, bracteole ovate, 0.8 x 0.5 mm, 2-fid 0.4--0.5, lobes and sinus acute, margins entire, crenulate or slightly dentate, connate on one or both sides. Perianth obovate, 1.35 x 1 mm, abruptly narrowed to a short, broad beak, somewhat compressed on sides, distinctly angled postical keel and usually with two or more supplementary antical and postical ridges, more or less tuberculate, particularly on keels.

 

On trees, and sometimes on rocks; low to moderate elevations; Ala., Ark., Del., Fl., Ga., Ill., La., Miss., Mo., N.C., Okla., S.C., Tenn., Tex., Va.

 

Frullania virginica can be confused with F. brittoniae and F. appalachiana. However, it has ventral leaves that are 2--2.5 times the stem width, and ventral leaf margins that have mostly one to sometimes two teeth. By comparison, the ventral leaves of F. brittoniae tend to be slightly wider (2--3.5 times the stem width) and have 2--3 teeth along the margins. The ventral leaves of F. appalachiana tend be slightly narrower (1.5--2 the stem width) and often have an angulation or tooth on each side of the leaf. Frullania virginica has gynoecial bract that are distinctly squarrose, with margins with several strong teeth; the innermost gynoecial bracteoles that are deeply 2-fid for about ½ their length and connate on both sides with the bract (R. M. Schuster 1992). The bract and bracteole of F. brittoniae and F. appalachiana are more or less identical in having gynoecial bracts margins that are entire except for a prominent stylar tooth, as well as non-squarrose bracts (R. M. Schuster 1992). Also the bracteoles are only shallowly lobed. Frullania virginica has variable styli that are sometimes uniseriate and filifrom (R. M. Schuster 1992) and sometimes subulate (4--5 cells long and 1--2 cells wide at the base).

 

27. Frullania wrightii Austin, Bull. Torrey Bot. Club 3: 15. 1872  E F

 

Plants light green to yellow-brown, dull, 0.9--1 mm wide. Stems 100--120 \um wide. Lateral leaves distant to contiguous, loosely imbricate when dry, spreading when moist, antical margin extending across stem; dorsal lobes reniform-ovate, 640--660 x 800--820 mm, convex, apex rounded, decurved, base cordate; lobule galeate, orbicular, 200 x 200 \um, lower half compressed, mouth truncate, or lobule explanate, 160--180 x 180--200 \um, styli subulate, 6--8 cells long and 2--4 cells wide; margins cells 14--18, median cells 18--20 \um, basal cells 26--32 x 20 \um, thin-walled, no intermediate thickenings, trigones small. Ventral leaves distant, obovate, 0.26 x 0.28 mm, flat, 2-lobed to one-half their length, lobes broadly triangular, apices sub-acute, sinus acute, bases cuneate, margins with a blunt angulation on both sides. Sexual condition autoicous. Androecia capitate, sessile on stem or main branch, in approximation to the gynoecia, but not below it. Gynoecia terminal on stem or main branch. Bracteole shallowly 2-fid, connate on one side with bract. Perianth pyriform to oblong, rounded to the apex, with sinuous, supplementary keels, beak short, mouth with papillae.

 

On rocks; moderate to high elevations; Ariz., Col., N.Mex., S. Dak., Utah.

 

The study of several western United States specimens previously determined as F. brittoniae were found to consistently have repand-dentate ventral leaf margins with several blunt to sharp angulations beneath the base of the sinus. Frullania riparia has somewhat similar ventral leaf margins, but often has only a single, blunt angulation on each side near the base of the sinus (R. M. Schuster 1992). By comparison, the ventral leaves of F. brittoniae have 1--3 prominent teeth along each margin, extending from the shoulders of the leaf to beneath the base of the sinus (A. W. Evans 1897; R. M. Schuster 1992). Overlaps between the morphology of F. brittoniae and the polymorphic F. riparia were discussed by R. M. Schuster (1992), but pertain mainly to lax leaved expressions with explanate lobules. Rather, the western United States specimens have imbricate leaves and frequently inflated, galeate lobules similar to typical F. brittoniae. Moreover, these specimens, like F. brittoniae, have cordate dorsal lobe bases and lanceolate styli, whereas F. riparia has auriculate dorsal lobe bases and filiform styli (R. M. Schuster 1992). Oddly, the western United States specimens have convex dorsal lobes with strongly decurved apices, characters not found in either F. brittoniae or F. riparia. Those species have mostly flat or only somewhat convex dorsal lobes, with apices scarcely, narrowly, or not decurved. An examination of the synonyms of F. brittoniae and F. riparia revealed F. wrightii Austin, a little known species from the western United States, but a good match for the confusing specimens in question. Frullania wrightii was described from New Mexico based on a specimen collected by Charles Wright (1811--1885). C. F. Austin (1872) compared F. wrightii to F. aeolotis (= F. riparia), but noted several differences in its leaf stance; curvature of the dorsal lobes and apices; cell areolation, lobule size; stylus shape; ventral leaf size, shape and dentation; and shape of the gynoecial bracts apices. A. W. Evans (1897) subsequently accepted F. wrightii in his revision of Frullania for North America, and reiterated the superficial affinities of F. wrightii with F. riparia. He nevertheless regarded the species to be incompletely known due to the lack of specimens outside of the type specimen. T. Frye and L. Clark (1947), however, considered F. wrightii to be a synonym of F. riparia, and incorrectly stated that the shape of the gynoecial bract apices, rounded in F. wrightii, while acute to bluntly right-angular in F. riparia, was the only morphological character that distinguished the species. Later, R. M. Schuster (1992) accepted this synonymy, while W. S. Hong (1989) seemingly overlooked F. wrightii in his work on western North America Frullania. Frullania wrightii is, however, morphologically distinct from F. riparia. Although the species was originally described as dioicous, F. wrightii is in fact diffusely autoicous, with androecia intermittently and occasionally positioned on the stem or main branch beneath the gynoecia. The reinstatement of F. wrightii from synonymy clarifies several reports of F. brittoniae from the western United States. The New Mexico stations of F. brittoniae are based on a specimen (Standley & Bollman 11138) that was tentatively determined by Evans as F. brittoniae (P. C. Standley 1915; G. Arsène 1933; L. M. Shields 1954). As indicated by G. Arsène (1933) and in notes by Caroline C. Haynes (1858--1951) on the label of a US specimen. Evans was unsure of his determination due to the specimen’s atypical ventral leaves. Re-examination of this specimen found the ventral leaf margins to have several blunt angulations on each side, characteristic of F. wrightii. The Colorado stations of F. brittoniae are based on several specimens, but only Weber B-9307 is F. wrightii. The other specimens reported by W. A. Weber (1963) are F. caucasica Steph. Frullania wrightii and F. caucasica are very similar morphologically in their narrow, imbricate shoots, as well as having convex dorsal lobes with decurved apices; inflated, galeate lobules with truncate mouths; lanceolate styli; and angulate ventral leaf margins on both sides near the base of sinus. However, F. wrightii lacks caducous leaves, while shoots of F. caucasica are often denuded. Furthermore, androecia and perianths can regularly be found in large populations of F. wrightii, whereas these structures are absent for F. caucasica. W. S. Hong’s (1989) report of F. brittoniae from Arizona is based on Hermann 19747, but this specimen has been re-determined as F. mexicana Lindenb. (J. J. Atwood 2017). Frullania wrightii and F. mexicana are similar in their autoicous sexuality, lobule size and shape, and in having numerously keeled perianths. However, the lobules of F. wrightii lack the narrowly triangular proximal portions that are characteristic of F. mexicana. Furthermore, the ventral leaves of F. wrightii are considerably wider than the stem and have repand-dentate margins. The ventral leaves of F. mexicana are about the width of the stem and have entire margins. Finally, F. wrightii has gynoecial bracteoles that are connate on one-side with the gynoecial bracts, and perianth with short, broad beaks. By comparison, the gynoecial bracteoles of F. mexicana are connate on both sides with the gynoecial bracts, while the perianth beaks are more or less cup-shaped.

 

1d. Frullania subg. Meteoriopsis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 37. 1884

 

Plants irregularly pinnately or bi-pinnately branched, sometimes branched indeterminately, becoming progressively shorter and narrower, with leaves and lobules more densely spaced. First branch appendage, 2-fid, consisting of a flattened, lanceolate-ovate segment, and a mostly inflated, though occasionally flattened lobule. Second branch appendage lateral and consisting of two inflated or flattened lobules (and associated stylus). Lateral leaves slightly to strongly imbricate when dry, flat, loosely deflexed, or sub-convolute, erect spreading [convolute] when moist, antical margin base extending across stem, [appendiculate at postical margin], lobes flat or slightly to strongly convex, apex rounded, bluntly acute, or sometimes broadly apiculate, plane or slightly to strongly deflexed [cucullate]; lobules mostly erect, parallel or sometimes subparallel, mouth oriented towards shoot base, saccate or sub-cylindrical when inflated, lanceolate and with plane or reflexed margins when flattened; stylus uniseriate and filiform; ocellii lacking. Ventral leaves approximate, contiguous, or distant, margins entire or toothed, bases narrowed and cuneate, sometimes auriculate. Specialized asexual reproduction none. Sexual condition autoicous or dioicous. Androecia capitate or short spicate, terminal on short lateral branches. Gynoecia terminal on stem or main branch [or on short lateral branches], frequently with innovations; bract lobule margins with prominent stylar tooth. Perianth trigonous in section with one ventral and two lateral keels, [or terete], ornamentation smooth, beak with papillae.

 

Species approximately 60 (3 in flora): North America; Mexico; West Indies; Central America; South America; Asia (including Indonesia); Africa (including Madagascar); Indian Ocean Islands; Pacific Islands.

 

The morphological boundaries separating subg. Meteoriopsis and subg. Thyopsiella are vaguely defined (S. Hattori 1972; J. Hentschel et al. 2009; R. M. Schuster 1992, J. Uribe-M. 2011; J. Uribe-M. & S. R. Gradstein 2003). Taxa in both subgenera have lobules that are comparable in their longer than wide dimensions, parallel to sub-parallel orientation, and contiguous or approximate position to the stem. R. Spruce (1884) cited differences in habit, insertion of the lateral leaves, and shape of the dorsal lobe bases as characters that differentiate the two subgenera. Subgenus Meteoriopsis was later further restricted by J. Uribe-M. (2008) to include taxa having leaves that are convolute around the stem when both dry and moist, as well as having expanded dorsal lobe bases on the antical and postical margins. Most members of subg. Meteoriopsis are distributed in Central and South America.

Using four molecular markers, J. Hentschel et al. (2009) found that some pendant habit species, formerly placed in subg. Thyopsiella (R. Spruce 1884; R. E. Stotler 1969 as subg. Frullania; J. Uribe-M. 2008), formed a moderately supported clade with the type of subg. Meteoriopsis (sensu M. Kamimura 1961; sensu J. Uribe-M. & S. R. Gradstein 2003). Furthermore, small, compact species, such as F. kunzei, were found to be related. Subgenus Meteoriopsis was thereafter re-circumscribed to include taxa with pendent as well as prostrate stems; leaves that are convolute or flat when dry; and dorsal lobes with truncate, cordate or auriculate bases on the antical, and sometimes postical margin (J. Hentschel et al. 2015). The removal of species from subg. Thyopsiella restricts it to a clade containing mostly taxa with ocellate lobes. Ocelli also occur in the lobes of some members of subg. Frullania and subg. Diastaloba.

 

SELECTED REFERENCES  Evans, A. W. 1897. A revision of the North American species of Frullania, a genus of Hepaticae. Transactions of the Connecticut Academy of Arts and Sciences 10: 1--39, pl. 1--15.  Hentschel, J., M. J. von Konrat, T. Pócs, A. Schäfer-Verwimp, A. J. Shaw, H. Schneider, and J. Heinrichs. 2009. Molecular insights into the phylogeny and subgeneric classification of Frullania Raddi (Frullaniaceae, Porellales). Molec. Phylogen. Evol. 52: 142--156.  Schuster, R. M. 1992. The Hepaticae and Anthocerotae of North America East of the Hundredth Meridian, Vol. 5. Field Museum of Natural History, Chicago. 1--854.  Stotler, R. E. 1969. The genus Frullania subgenus Frullania in Latin America. Nova Hedwigia 18: 397--555.

 

28. Frullania cucullata Lindenberg & Gottsche, in C. M. Gottsche, J. B. W. Lindenberg, & C. G. D. Nees, Syn. Hepat. 782. 1847

 

Plants greenish brown, reddish brown, or black, 0.8--1 mm wide. Stems 110--130 \um wide. Lateral leaves dorsal lobe oblong-ovate, 0.5--0.7 x 0.4--0.5 mm, base cordate [circinate-auriculate]; lobule saccate when inflated, 160--200 x 80--110 \um, constricted near the flaring mouth, or flattened and lanceolate, 200--240 x 60--80 \um, apex acute, margins strongly deflexed; stylus uniseriate and filiform, 3--5 cells long; marginal cells 10--12 \um, median cells 12--22 \um, basal cells 22--32 x 14--26 \um, walls thin, flexuose, occasional intermediate thickenings, trigones large; oil bodies oval to ellipsoidal, 2--3 per cell. Ventral leaves approximate to contiguous, concave, elliptic, 0.3--0.5 x 0.2--0.5 mm, 2-fid for 0.1 mm, lobes acute to obtuse, sinus acute to rounded, margins plane or slightly to strongly reflexed, entire, base auriculate. Sexual condition dioicous. Gynoeica bract lobe broadly ovate, 1.0--2.2 x 0.4--0.7 mm, apex rounded to bluntly acute, 2-fid to 0.1 mm, margins plane, entire, lobule lanceolate, 0.6--1 x 0.1--0.3 mm, apex acuminate, margins reflexed, entire; bracteole elliptic, 0.7--0.9 x 0.4--0.6 mm, 2-fid to about 0.1--0.2 mm, lobes acute, sinus acute, margins entire, plane to reflexed, free from the bracts. Perianth dorsally flattened, oblong-cylindrical, 1.6--2 x 0.8--1 mm, gradually narrowed to the apex, beak long.

 

On the trunks, branches, and twigs of trees and shrubs; low elevations; Fla.; Mexico; West Indies; Central America (Costa Rica, Panama).

 

Frullania cucullata has been collected several times in North America since A. W. Evans (1914) first reported it from southern Florida. Although R. E. Stotler (1969) placed F. cucullata in synonymy with the neotropical and polymorphic F. intumescens, that species differs in having a large basal appendage attached to the stylus (Y. Yuzawa & N. Koike 1989), as well as toothed gynoecial bract and bracteole margins (R. E. Stotler 1969).

R. M. Schuster (1992) reported the morphologically similar F. compacta from Collier County, Florida, based on two specimens containing only androecia bearing stems (Schuster 82-313, 26009, both duke). Frullania compacta differs from F. cucullata in having consistently plane to weakly deflexed dorsal lobe margins, a tendency to have more numerously inflated lobules, plane to slightly reflexed ventral leaf margins, gynoecial bracts that are connate on one side with the bracteole, and terete perianths (L. Clark & R. D. Svihla 1951; R. M. Schuster 1992). The species is known otherwise from Cuba, where it was collected without locality by C. Wright. An examination of the Florida plants found some dorsal lobes with strongly convex margins, inflated and flattened lobules occurring together on the same stem, and variable ventral leaf margins ranging from slightly to strongly reflexed. Similar variation in the lobules and stance of the ventral leaf margins has been noted for F. cucullata by T. C. Clark and L. Frye (1945). Moreover, the F. compacta plants compare well to prostrate, androecia bearing stems of F. cucullata that were collected from nearby or nearly identical localities in Collier County, Florida (Breil 5410, 5432, both mo; Mar. 1979, Griffin s.n., Bryoph. Florida, mo). These specimens have associated gynoecia and perianths clearly matching that of F. cucullata. The two Florida specimens of F. compacta are therefore better treated as weak expressions of F. cucullata. Until gynoecia or perianth bearing stems of F. compacta can be demonstrated from North America, that species should be excluded from the flora area.

 

29. Frullania donnellii Austin, Bull. Torrey Bot. Club 6: 301. 1879  E

 

Plants copper-red, 0.6--1 mm wide. Stems 60--100 \um wide. Lateral leaves dorsal lobe ovate, 0.5--0.6 x 0.6 mm, base truncate; lobule separated from the stem by a distance that is less than or nearly equal to the lobule width, saccate, 1.8--2 x 1.0--1.2 mm, or sometimes flattened; stylus uniseriate and filiform, 3--5 cells long; marginal cells 10--12 \um, median cells 12--18 \um, basal cells 32--40 x 18--24 \um, walls mostly thickened, trigones small; oil bodies spherical to ellipsoidal, 2--3 per cell. Ventral leaves distant, obovate, 0.3--0.4 x 0.3 mm, 2-fid to about 0.1 mm, lobes acute, sinuses narrowly to broadly acute, flat, margins entire or with an obtuse tooth on one or both sides. Sexual condition autoicous. Gynoecia bract lobe elliptical, 0.8--1 x 0.4--0.5 mm, 2-fid to 0.5--0.7 mm, margins dentate, lobule ova28te-lanceolate, 0.7--0.9 x 0.3--0.5 mm, margins mostly dentate; bracteole oblong, 0.6--0.9 x 0.5--0.6 mm, 2-fid to about 0.3 mm, sinuses and lobes acute, margins dentate, free from the bracts. Perianth strongly dorsally flattened, oblong, 1.2 x 0.7 mm, abruptly narrowed to the apex, beak short.

 

On the trunks, branches, and twigs of trees and shrubs; sometimes on logs; low elevations; Ala., Fla., Ga., La., Miss., N.C.

 

Frullania donnellii is endemic to North America and is currently known from stations in the Atlantic and Gulf Coastal Plains of the southeastern United States. Its distribution appears to be more restrictive than that of the morphologically similar F. kunzei, whose North America range is sympatric in the southeastern Coastal Plains, but occurs also in the Piedmont, and Appalachian, Ozark, and Ouachita Mountains. Outside of the flora area, F. kunzei is distributed throughout the West Indies, as well as in Central and South America. In addition to its narrower distribution, F. donnellii has been collected only on wood, whereas F. kunzei has been found on both wood and rock. Due to the commonness of F. kunzei and the difficulties in reliability separating the two species in the field, F. donnellii may perhaps be overlooked.

 

30. Frullania kunzei (Lehmann & Lindenberg) Lehmann & Lindenberg, Syn. Hepat. 3: 449. 1845.

Jungermannia kunzei Lehmann & Lindenberg, in Lehmann, Nov. Stirp. Pug. 6: 50. 1834; Frullania drummondii Taylor; Frullania kunzei var. maritima R.M. Schuster ex Hentschel & von Konrat  

 

Plants olive-green to copper-red, 0.4--0.8 mm wide. Stems 80--100 \um wide. Lateral leaves dorsal lobe orbicular to elliptical, 0.3 x 0.4--0.5 mm, base truncate to slightly cordate; lobule mostly superimposed or contiguous with stem, saccate to sub-cylindrical, 130--180 x 60--92 \um, or sometimes flattened, approximate or nearly contiguous with each other; stylus uniseriate and filiform, 3--4 cells long; marginal cells 10--12 \um, median cells 12--16 \um, basal cells 18--22 \um, walls thickened, firm, lacking intermediate thickenings, trigones small to large; oil bodies ellipsoidal, 2 per cell. Ventral leaves distant, obovate, 180--210 x 132--170 \um, 2-fid, lobes acute to obtuse, sinuses narrowly to broadly acute, flat, margins entire or with an obtuse tooth on one or both sides. Sexual condition autoicous [dioicous]. Gynoecia bract lobe elliptical, 1.0--1.2 x 0.4 mm, apex rounded to variably acute, sometimes apiculate, rarely short-acminate, margins entire, lobule ovate-lanceolate, 0.6--0.8 x 0.4--0.5 mm, apex acute, margins entire; bracteole oblong, 0.6--0.8 x 0.3--0.5 mm, lobes and sinuses acute, margins entire, free [connate on one side with bract]. Perianth dorsally flattened, obovate, 0.8--0.9 x 0.6--0.8 mm, gradually broadened to the apex, trigonous, ventral keel short to elongate, narrow or broad, beak short, with papillae.

 

On the trunks, branches, and twigs of trees and shrubs; vines; fallen tree limbs; and boulders, ledges, and rock faces; low to moderate elevations; Ala., Ark., Fla., Ga., Ill., Kans., La., Miss., Mo., N.C., Okla., S.C., Tenn., Tex., Va.; Mexico; West Indies; Bermuda; Central America (Guatemala, Honduras, Costa Rica); South America.

 

Frullania kunzei is commonly found intermixed with other bryophytes on trees and rocks in the southeastern United States. The species is similar to F. donnellii in color, size, branching pattern, lobe and lobule size and shape, ventral leaf size and shape, and the overall size and areolation of the cells. The two differ in the distance of the lobule from the stem, as well the dentation of the gynoecial bract and bracteole margins. The apices of the gynoecial bract lobes in F. kunzei are somewhat variable, ranging from mostly rounded to broadly or narrowly acute, to sometimes apiculate. Rarely are they short-acuminate, but never end in a row of three or more superimposed cells as in F. donnellii. According to R. M. Schuster (1992), the number of oil bodies in the basal cells will also separate the two species when gynoecial bracts and bracteoles are absent. Frullania knuzei is noted to have 2 oil bodies per cell, whereas F. donnellii has 3--4 oil bodies per cell.

Frullania kunzei var. maritima was described from a single specimen that was collected on the Florida gulf coast by R. M. Schuster in 1976. Following Article 37.7 of the International Code of Botanical Nomenclature (J. McNeill et al. 2012), R. M. Schuster’s 1991 publication of the variety is invalid because he did not indicate a herbarium for the type specimen. The variety, however, was later validated by J. Hentschel et al. (2015) who cited Schuster’s specimen of var. maritima in f as the holotype. According to R. M. Schuster (1992), several differences separate var. maritima from var. kunzei, including shoot width, dorsal lobe shape and dimensions, lobule spacing, and the ornamentation of the perianth and perianth beak. These differences, however, seem to be restricted to North American specimens alone. When neotropical specimens of F. kunzei are include in the comparison, the noted differences in plant size, dorsal lobe shape and dimensions, as well as spacing of the lobules erode, since these characters do not appear to be intraspecifically correlated. Furthermore, among North America specimens, the ornamentation of the perianth and perianth beak varies significantly in the length and depth of the ventral keel, as well as in the number and size of the papillae on the inner beak surface. The described differences of var. maritima can be accommodated within the variable F. kunzei, as presently described.

 

1e. Frullania subg. Thyopsiella Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 41. 1884

 

Plants bi-pinnately or tri-pinnately branched. First branch appendage 2-fid and divided into two unequally sized lobes, or 3-fid, consisting of an unequally 2-fid, flattened, lanceolate-ovate segment, and a flattened or inflated lobule. Second branch appendage lateral and consisting of two inflated lobules or one inflated and one flattened lobule (and associated stylus). Lateral leaves flat to somewhat convex, imbricate when dry, spreading when moist, antical margin base sub-auriculate to auriculate, extending across stem, dorsal lobe flat to convex, apex reflexed; lobules erect, contiguous with stem or separated by less than or about the lobule width, parallel or sometimes subparallel, mouth oriented toward shoot base, saccate or sub-cylindrical when inflated; stylus minute and subulate, sometimes appendiculate; ocelli typically forming a median line, sometimes scattered or absent none. Ventral leaves distant or contiguous, margins entire or toothed, bases appendiculate or slightly auriculate. Specialized asexual reproduction none. Sexual condition autoicous or dioicous. Androecia capitate or spicate, terminal on short lateral branch. Gynoecia terminal on stem or main branch, or on short lateral branches, with or without innovations; bract lobule margins with prominent stylar tooth. Perianth trigonous in section, with one ventral and two lateral keels, ornamentation smooth, beak smooth.

 

Species approximately 20 (7 in flora): North America; Europe; Asia.

 

With the exception of F. selwyniana, the North America species in subg. Thyopsiella belong to the F. tamarisci complex. This complex includes plants that vary intraspecifically in their plant size and shoot widths; dorsal lobe lengths and widths; ventral leaf widths; shape of leaf lobe apices, cell dimensions; and distribution of ocelli. Consequently, the complex has been treated as a single, polymorphic species composed of more or less geographically isolated subspecies (M. Kamimura 1961; S. Hattori 1972; R. M. Schuster 1992), as well as discreet species having broadly overlapping character states (A. W. Evans 1897; B. Crandall-Stotler et al. 1987; M. Ramaiya et al. 2010; A. A. Vilnet et al. 2014). Discreet species are recognizable, but determination often requires reliance on a combination of characters.

North America species of subg. Thyopsiella differ from most other Frullania species in the flora by their tendency to have ocelate dorsal lobes with the ocelli mostly arranged in a short median line, and/or scattered throughout the leaf. Some expressions of F. californica, F. franciscana, F. nisquallensis and F. subarctica can, however lack ocelli. In the flora area, aside form subg. Thyopsiella, only F. cobrensis (subg. Frullania) also has ocelate dorsal lobes, but the ocelli in are restricted to the basal cells. Subg. Thyopsiella further differs from subg. Chonanthelia and subg. Frullania in having cylindric, versus galeate inflated lobules that are often two or more times longer than wide. The lobules of subg. Diastaloba are also longer than wide, but species in that subgenus have lobules remotely positioned from the stem. Species in subg. Meteoriopsis have cylindric lobules as well, but the stylus in those species are consistently uniseriate and filiform, while in subg. Thyopsiella are typically subulate or lamellate, often with cilia or laciniae.  

 

SELECTED REFERENCES  Evans, A. W. 1897. A revision of the North American species of Frullania, a genus of Hepaticae. Transactions of the Connecticut Academy of Arts and Sciences 10: 1--39, pl. 1--15.  Crandall-Stotler, B. R. E. Stotler, and P. Geissler. 1987. A biosystematics study of the subspecies of Frullania tamarisci (L.). Bryology 90(4) 287--308.  Hattori, S. 1972. Frullania tamarisci complex and the species concept. J. Hattori Bot. Lab.35: 202—251. Heinrichs, J., J. Hentschel, A. Bombosch, A. Fiebig, J. Reise, M. Edelmann, H.-P. Kreier, A. Schäfer-Verwimp, S. Caspari, A. Schmidt, R.-L. Zhu, M. von Konrat, B. Shaw, and A. J. Shaw. 2010. One species or at least eight? Delimination and distribution of Frullania tamarisci (L.) Dumort. s.l. (Jungermanniopsida, Porellales) inferred from nuclear and chloroplast DNA markers. Molecular Phylogenetics and Evolution 56: 1105--1114.  Schuster, R. M. 1992. The Hepaticae and Anthocerotae of North America East of the Hundredth Meridian, Vol. 5. Field Museum of Natural History, Chicago. 1--854.  Stotler, R. E. 1969. The genus Frullania subgenus Frullania in Latin America. Nova Hedwigia 18: 397--555.

 

31. Frullania asagrayana Montagne, Ann. Sci. Nat. Bot. (sér. 2) 18: 14. 1842  E

            Frullania grayana Sullivant, in A. Gray, Manual (ed. 2) 697. pl. 8. 1856; Frullania tamarisci subsp. asagrayana (Montagne) S. Hattori, in H. Hara, Fl. E. Himalaya 1: 528. 1966.

 

Plants pale-green to reddish-brown. Stems 150 \um wide. Lateral leaves dorsal lobe ovate, 0.7 x 0.5 mm, apex rounded or bluntly acute, decurved, antical base auriculate; lobule obovoid-clavate, 0.3 x 0.2 mm, contracted toward base, stylus filiform to subulate, with suborbicular disk like appendage with cilia and 1--2 marginal teeth; marginal cells 14 \um, median cells 18 \um, basal cells 28 x 18 \um, thick-walled, with intermediate thickenings, trigones present, ocelli forming a median line extending from the lobe base to half the lobe length, rarely scattered, ocells 24 \um. Ventral leaves distant, orbicular-ovate, 0.4 x 0.4 mm, 2-fid about 1/3 the length, lobes obtuse, sinus narrow, margins plane or sometimes reflexed, base appendiculate or slightly auriculate with a few teeth appressed to the stem. Sexual condition dioicous. Androecia terminal on short lateral branches. Gynoecia bract lobe ovate, 1.5 x 0.7 mm, 2-fid to ½ the length, apex acute, margins entire or somewhat dentate, lobule narrowly ovate to lanceolate, 0.9 x 0.3 mm, apex acuminate, margins plane, rarely revolute, entire, bracteole ovate, 1.2 x 0.5 mm, 2-fid to ½ the length, lobes acuminate, sinus acute, margins variously toothed, free or connate on one side with bract. The gynoecial bract lobes are entire to dentate, and the lobules have a few teeth to being lacinate at the base with occasional cilia, and a distinct stylar tooth.  Perianth oval to obovate, 1.9 x 0.9 mm, narrowed to a short beak, somewhat compressed at sides, with deep postical keel

 

On boulders and rock ledges, trunks and branches of trees, twigs, and occasionally as an epiphyll; medium to high elevations; Nfld., N.S., Ont., Que.; Ala., Ark., Conn., Del., Ga., Ill., Ind., Iowa, Ky., Maine, Md., Mass., Mich., Minn., Miss., Mo., N.H., N.J., N.Y., N.C., Ohio, Okla., Pa., S.C., Tenn., Vt., Va., W.Va., Wis.

 

Frullania asagrayana is a variable species both locally and throughout its range. The typical expression has dorsal lobes with decurved, rounded to sometimes bluntly acute apices, cordate antical bases, and ocelli arranged in a long median line that extends from the base of the lobe to nearly the apex. The line is mostly 1 cell wide, although occasionally it can be broken or sporadically become 2 cells wide. Rarely are the ocelli scattered. This expression additionally has narrow ventral leaves that are about 2 times the width of the stem, acute lobe apices, and plane to slightly reflexed margins. The styli and disk-like appendage are generally visible as a result of the narrower ventral leaf width. A second expression has some morphological similarities to F. tamarisci, and consequently has been reported as that taxon (R. M. Schuster 1992). It has dorsal lobes with decurved to deflexed, bluntly acute to apiculate apices, auriculate antical bases, and ocelli arranged mostly in a short median line. The line is frequently broken with at least a few scattered accessory ocells. Furthermore, this expression has wider ventral leaves that are about 3 times the width of the stem, with sub-acute to obtuse apices, and slightly to strongly reflexed margins. The styli and disk-like appendage are generally not visible due to the leaf width. As noted by R. M. Schuster (1992) a series of intermediates with transitional morphology links the extremes of these expressions, and is suggestive of a single polymorphic species. Support for recognition of a single species was found by M. Ramaiya et al. (2010) based on an extensive analysis of nucleotide sequences. Microsatellite data from that same study, however, found F. asagrayana to be composed of two geographically discernible clades, sharing little to no gene flow between them (M. Ramaiya et al. 2010).

 

32. Frullania californica (Austin) A. Evans, Trans. Connecticut Acad. Arts Sci. 10 (1): 25. 1897  E

Frullania grayana var. californica Austin ex Underwood, Bull. Illinois State Lab. Nat. Hist. 2: 67. 1884; F. asagrayana var. californica Howe, Erythea 2: 98. 1894; F. asagrayana var. alsophila Howe, Erythea 2: 99. 1894

 

Plants green to brownish-red, 0.6--0.8 mm wide. Stems 110--150 \um wide. Lateral leaves dorsal lobes sub-orbicular, 0.4--0.5 x 0.4--0.5 mm, apex rounded, occasionally acute or sometimes acuminate, more or less decurved, lobule obovate-clavate, 170--200 x 70--90 \um, stylus subulate, sometimes with disk appendage; marginal cells 10--14 \um, median cells 12--24 \um, basal cells 30 x 22 \um, thick-walled, intermediate thickenings few, trigones inconspicuous, ocelli few or lacking, mostly scattered and rarely in a vitta. Ventral leaves orbicular, 0.2--0.3 x 0.2--0.3 mm, 2-fid 1/3 the length, lobes obtuse or sub-acute, sinuses obtuse to sub-acute, margins plane or slightly reflexed, base slightly auriculate. Sexual condition dioicous. Gynoecial bract lobe ovate to ovate lanceolate, 1.2--1.4 x 0.4--0.5 mm, apex acute, lobule lanceolate or subulate, 0.7--0.8 x 0.2--0.3 mm, acuminate, margins more or less reflexed, bracteole ovate, 0.9--1 x 0.5--0.6 mm, 2-fid to about ½ the length, lobes lanceolate-acuminate, sinus acute, margins ciliate at base, connate on one side with bract. Perianth ovoid, 1.7 x 0.9 mm, compressed on sides, narrowed to a short beak, deeply 1-keeled postically.

 

On the trunks and branches of trees, logs, and rock outcrops; low to high elevations; B.C., Calif., Oreg., Wash.

 

Frullania californica is morphologically similar to F. franciscana and F. nisquallensis, and the differences separating it are discussed under those species. Like F. asagrayana, F. franciscana, F. nisquallensis, F. subarctica, and F. tamariscii, it has dorsal lobe ocelli that are variable in their location. While the ocelli are generally few, they may also be scattered or form a short median line in some expressions. As noted by W. S. Hong (1989) and W. T. Doyle & R. E. Stotler (2006), F. californica may sometimes have dorsal lobe apices that are sporadically acute or even sometimes acuminate. These specimens are otherwise identical to specimens with rounded apices. 

 

33. Frullania franciscana M. Howe, Erythea 2 (6): 99. 1894  E

 

Plants green to reddish-brown, 0.9--1.2 mm wide. Stems 150--170 \um wide. Lateral leaves dorsal lobe ovate, 0.7 x 0.6--0.7 mm, apex rounded, obtuse or apiculate, decurved; lobule short-clavate, 220--250 x 120 \um, stylus subulate, occasionally with cilia; marginal cells 12--14 \um, median cells 12--20 \um, basal cells 36 x 22 \um, thick-walled, with intermediate thickenings, trigones present, ocelli usually in vitta, sometimes absent. Ventral leaves distant, ovate, 0.4 x 0.3--0.4 mm, 2-fid to about 1/3 the length, lobes obtuse, sinus acute, margins plane, bluntly unidentate, base not auriculate or appendiculate. Sexual condition dioicous. Gynoecial bract lobe ovate, 1.1--1.4 x 0.6--0.8 mm, lobule ovate to lanceolate, 0.6--0.8 x 0.2--0.3 mm, apex acute, margin with cilia towards base; bracteole ovate, 1.4 x 0.6 mm, 2-fid for about ½ the length, lobes lanceolate, acuminate, sinus acute, margins ciliate at base, connate on one side with bract. Perianth oblong-obovate, 2.2 x 1 mm, abruptly short-rostrate, compressed on sides and with a deep postical keel.

 

On trees as well as on rock outcrops; low to moderate elevations; B.C., Alaska, Calif., Oreg., Wash.

 

Frullania franciscana has variable dorsal lobe apices that are usually rounded, but may sometimes be bluntly acute. Similarly rounded dorsal lobe apices are found in expressions of F. californica, whereas acute apices are more common in F. nisquallensis and F. subarctica. Frullania franciscana differs from F. californica in having ocelli arranged in a median line that is 1--2, sometimes 3 cells wide, versus ocelli sometimes scattered on the dorsal lobe. In F. nisquallensis and F. subarctica, the ocelli are frequently absent, although they may be present in a few dorsal lobes as a short, broken line. Both of these species also have entire ventral leaf margins. The ventral leaf margins of F. franciscana usually have a blunt angulation on one or both sides at about the middle of the leaf.

 

34. Frullania nisquallensis Sullivant, Mem. Amer. Acad. Arts (n.ser.) 4: 175. 1849 [1850]  E

Frullania tamarisci subsp. nisquallensis (Sullivant) S. Hattori, in H. Hara, Fl. E. Himalaya 1: 528. 1966

 

Plants green to golden-green, brownish or purplish-red, never blackish, 0.6--1.2 mm wide. Stems 180--200 \um wide. Lateral leaves dorsal lobe ovate, 0.8--0.9 x 0.6--0.8 mm, apex acute to acuminate-apiculate, margins strongly deflexed, base cordate to auriculate; lobule oblong-clavate, 160--200 x 80--100 \um; stylus subulate 3--6 cells long and 1--2 cells wide, appendage disc composed of 3--5 cells; marginal cells 14--18 \um, median cells 14--26 \um, basal cells 36 x 24 \um,  thick-walled, with intermediate thickenings, trigones present, ocelli vitta usually absent, sometimes present, but broken in a row approximately 5 cells long. Ventral leaves distant or contiguous, orbicular to obovate, 0.4--0.5 x 0.6--0.8 mm, 2-fid to about ¼ the length, lobes obtuse, sinus obtuse, margins strongly reflexed at apex, occasionally bear a small tooth of one or both sides, base auriculate. Sexual condition dioicous. Gynoecia bract lobe ovate, 2 x 0.7--0.8 mm, apex acuminate, margins entire to dentate, lobule subulate, 0.8--1 x 0.2 mm, acuminate, margins revolute; bracteole ovate, 1.2--1.4 x 0.5--0.7 mm, 2-fid to 1/3 the length, lobes acuminate, sinus acute, margins revolute, entire above, dentate or ciliate at base, connate on both sides with bract. Perianth, ovate to oblong, 2.0--2.4 x 1 mm, gradually narrowed to a short beak, concave antically, compressed on the sides, deeply 1-keeled postically.

 

On trunks and branches of trees and shrubs, as well as on decaying logs and stumps, also on soil and rock outcrops; low to moderate elevations; B.C., Alaska, Calif., Oreg., Wash.

 

Frullania nisquallensis has ovate dorsal lobes with acute to acuminate, deflexed apices, and ocelli absent, scattered, or forming a partial or broken median line. Like F. californica, F. franciscana, and F. subarctica, the styli may have a reduced or absent appendage. The ventral leaves can be somewhat auriculate with weakly developed basal appendages, similar to F. californica and F. subarctica, or have gradually narrowed bases that lack appendages, like F. franciscana. In distinguishing F. nisquallensis from the morphologically similar species of subg. Thyopsiella in western North America, the stance of the ventral leaf margins provides a useful characteristic. In F. nisquallensis, the ventral leaf margins are entire and reflexed at or above the sinuses. By comparison, both F. franciscana and F. californica have the ventral leaf margins plane or nearly so. Although F. subarctica also has reflexed ventral leaf margins, it differs from F. nisquallensis in its color, shoot width, and ecology.

 

35. Frullania selwyniana Pearson, List Canad. Hepat. 1, pl. 1. 1890  E

Frullania sullivantiae Austin

 

Plants green to copp