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BFNA
Title: Pterobryaceae |
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PTEROBRYACEAE Plants 0.2--3.0(--6.0) cm, creeping stolons becoming
erect, inclined or horizontal to form simple or pinnately branched leafy
stems, often with a basal stipe region; flagelliform branches frequent; dark
or yellowish green. Stem cortex of orange-brown, thick-walled cells,
central strand not differentiated; paraphyllia absent; pseudoparaphyllia
filamentous, uniseriate, hyaline; rhizoids mostly restricted to stolons;
axillary hairs 2--3 per axil, of 1--2 short brownish basal cells and 2--4
longer hyaline cells. Leaves on stolons scale-like, 0.2--1.25 mm,
short-acute or long-acuminate, usually ecostate; in stipe region appressed,
erect or recurved from a clasping base, triangular to ovate-acuminate, costa
double or single, shorter in distal stem leaves; in stem region leaves erect
to spreading or squarrose, lanceolate, ovate-acuminate or orbicular, 1.3--2.6
mm; margins plane, rarely revolute to apex, entire to serrulate distally;
costa single to mid-leaf or percurrent, or short and double; cells at insertion
dark yellow, thick-walled, porose and irregular; medial laminal cells
sinuose, smooth or prorate, rarely with one or two papillae over the cell
lumen, walls often porose; distal laminal cells shorter; alar cells little
differentiated or rhombic to quadrate in several rows up margin, rarely
oblate in many rows extending up margin in lower third of leaf; branch leaves
concave, narrowly ovate-acuminate, smooth or weakly plicate, smaller or
similar to distal stem leaves. Specialized asexual propagation frequent,
clavate gemmae in clusters in leaf axils, reddish brown, uniseriate, of 4--12
quadrate or rectangular cells, thick-walled and verrucose. Sexual
condition dioicous. Perigonia on stem and branches, bud-like, with
a few scale-like leaves, uniseriate paraphyses and 5--10 antheridia. Perichaetia
on stem and branches, urceolate; perichaetial leaves 8--12, the interior
ovate-acuminate, abruptly narrowed to long erect awns; multiseriate
paraphyses often present. Seta red-brown, 3.0--12.4 mm. Capsule
erect, exserted, red-brown, ovate-cylindrical, narrowed to mouth, smooth;
stomates absent; operculum rostrate from low, rounded or conic base, rostrum
inclined. Peristome with prostome; exostome reduced, teeth 16, often
in 8 irregular pairs, incurved when dry, erect when moist, lanceolate, often
irregular, inserted below mouth, mostly smooth; endostome consisting of
remnants adhering to back of exostome. Calyptra cucullate, fully or
partially covering the capsule, with scattered archegonia and paraphyses or
smooth, not plicate. Spores spherical, 20--35µm diameter, granulose,
green. Genera ca. 16, species ca. 200 in
Pterobryaceae s.str. (3 genera, 4 species in the flora): tropical and
subtropical regions worldwide. The family is highly diverse, and has
frequently been the depository of odd tropical taxa excluded from other
families. The description above applies only to taxa in the flora area. Few
features unite the family as currently circumscribed, and even these are not
present in all taxa: tropical, primarily epiphytic habitats; a frondose habit
frequently with a differentiated stipe, although this may only be apparent in
juvenile plants; uniseriate filamentous pseudoparaphyllia (B. H. Allen 1987);
no stomates; reduced peristomes, often with a prostome, and a highly reduced
endostome. The inclusion of Henicodium in the
family is questionable on morphological grounds, since the presence of
papillae centrally located over the lumen and a revolute leaf margin is
otherwise not known in the family. However molecular studies indicate some
affinity between Henicodium and Orthostichopsis (W. R. Buck et
al. 2000), although Pterobryon, the type genus, is usually placed
distantly from both these exemplars (A. E. Newton et al., in prep.). Henicodium
is accepted here pending resolution of the family circumscription. SELECTED REFERENCES Allen, B. H. 1987. On distinguishing
Pterobryaceae and Meteoriaceae by means of pseudoparaphyllia. Bryol. Times
42: 1-3. Arzeni, C. B. 1954. The Pterobryaceae of the southern United
States, Mexico, Central America and the West Indies. Amer. Midl. Naturalist.
52: 1--67. Buck, W. R. 1989. Henicodium replaces Leucodontopsis
(Pterobryaceae). Bryologist 92: 534. Buck, W. R. 1998. Pleurocarpous
mosses of the West Indies. Mem. New York Bot. Gard. 82. Buck, W. R., B.
Goffinet and A. J. Shaw. 2000. Testing morphological concepts of orders
of pleurocarpous mosses (Bryophyta) using phylogenetic reconstructions based
on trnL-trnF and rps4 sequences. Mol. Phylog. Evol. 16: 180-198. Newton,
A. E. 1993. Phylogenetic systematics of the tropical moss genus Pireella
(Pterobryaceae: Musci). Ph.D. Dissertation, Duke University, Durham, North
Carolina.
1. Leaf margins revolute nearly to apex, cells in proximal third of leaf oblate in many rows up the margin, lamina cells with one or two papillae over the lumen 3. Henicodium 1. Leaf margins plane, cells in basal angles undifferentiated or rhombic-quadrate in several rows, lamina cells smooth or prorate. 2. Plants simple or sparsely branched, leaves broadly ovate to orbicular, spreading to squarrose, costa variable on a single plant, double and short, or single, short or reaching to midleaf or above 1. Jaegerina 2. Plants simple or weakly and irregularly pinnate, leaves ovate acuminate, erect to spreading, costa single, percurrent or shortly excurrent 2. Pireella 1. JAEGERINA J. K.
A. Müller, Linnaea 40: 273. 1876 [for August Jaeger, 1842--1877, Swiss
bryologist] Jaegerinopsis V. F. Brotherus, Natürlichen Pflanzenfamilien
1(3):790. Leipzig 1906 Plants 1--5 cm, creeping stolons forming erect stems that
are simple or sparsely branched above a very short stipe-like basal portion,
shiny yellowish green. Stem leaves dense, not seriate, similar wet or
dry, spreading to recurved or squarrose [erect], broad ovate-orbicular
[ovate-lanceolate], 1.5--2.5 X 1.0--1.5 mm, concave, weakly plicate; apex
acute, base not auriculate or decurrent; margins plane, entire to serrulate;
costa short and double or single, reaching to mid-leaf or above, sometimes
forked; medial laminal cells sinuose, 35--65 µm, smooth, walls porose; alar
cells little differentiated. Branch leaves, if present, similar to
stem leaves. [Seta 4--5 mm. Capsule cylindrical; operculum
long-rostrate from a low conical base, rostrum inclined, blunt. Calyptra
partially covering capsule, hairless, rough with projecting cell ends.] Species 12 (1 in the flora); tropical and
subtropical regions, se United States, Mexico, West Indies, Central America,
n S. America, tropical Africa, Asia, Africa. 1. Jaegerina scariosa (Lorentz)
Arzeni, American Midland Naturalist 52: 12. 1954 Meteorium scariosum Lorentz, Moostudien. p. 165. Leipzig. 1865;
Jaegerinopsis scariosa (Lorentz) Brotherus; Jaegerinopsis
squarrosa E. Britton Plants 1--5 cm; not or rarely branched. Stem leaves
spreading to recurved or squarrose, ovate-orbicular, 1--2 X 0.75--1.5 mm,
apex acute; costa variable, from short and double to long and single to above
mid-leaf; medial laminal cells sinuose, 33--60 X 2.5 µm, smooth; alar cells
little differentiated, 4-8 small quadrate cells. Sporophytes not known from flora region.
Humid hardwood forests and hammocks, tree trunks, especially Magnolia
grandiflora; 0-800 m; Fla., La.; Mexico; West Indies; Central America; n
South America, tropical Africa. The extreme variability of the costa in J.
scariosa has resulted in a surplus of names for this species. Individual
plants may have leaves with a weak or moderately strong single costa reaching
to mid-leaf or above, sometimes forked above, or a short double costa, or a mere
trace at the leaf base. The plant is readily recognized in the field,
however, where it forms small tufts of erect stems on tree trunks. The leaves
are glossy yellow and always broadly spreading to squarrose, giving the
plants a distinctive appearance. Leucodon julaceus and species of Cryphaea
occupy the same habitat, but are smaller and have the leaves appressed when
dry, amongst other differences. Neither gametangia nor sporophytes are known
for J. scariosa in the flora area. 2. PIREELLA Cardot,
Rev. Bryol. 40: 17. 1913 [for Louis Piré, 1827--1887, Belgian
bryologist] Plants 1--5 cm, creeping stolons forming erect stems that
are simple or irregularly pinnate from a short or long stipe-like region,
dull dark green to glossy yellowish green. Stem leaves dense,
obscurely seriate; in basal stipe region appressed to erect,
triangular-lanceolate, 0.5--1.7 mm; in distal region erect dry, spreading
wet; broadly ovate-acuminate, 1.3--2.6 mm; concave, not plicate; base
rounded, auriculate or decurrent; margin plane; costa single, short in
proximal leaves, percurrent or shortly excurrent in distal leaves; medial
laminal cells sinuose, prorate, firm-walled, porose; alar cells rhombic or
quadrate, in small groups or extending up margin in a few rows. Branch
leaves weakly or strongly seriate in five rows, spreading wet and dry;
narrow ovate-acuminate, concave with flattened apex, not plicate; costa
percurrent; medial laminal cells sinuose, distinctly prorate. Seta
5.4--12.4 mm. Capsule cylindrical; annulus not differentiated;
operculum rostrate from a low rounded or conic base. Calyptra covering
the capsule, hairy. Species 13 (2 in the flora): se United
States, Mexico, West Indies, Central America, n and c South America. Pireella, with 13 species, is the largest genus of the
Pterobryaceae in the Neotropics, and is composed of three distinct groups,
two of which are not represented in the flora area, and are not covered by
the description above. The two species in North America are rather similar
and have frequently been misidentified in the past, but differ in the shape
of the leaf base, the number, size and distribution of alar cells, the shape
and stance of the leaves, and the branching pattern. The leaf-base characters
are most useful, but can only be distinguished reliably on the distal stipe
or stem leaves. C. B. Arzeni (1954) clarified the distinction between these
species, but misidentifications have continued. This may in part be
attributed to misidentified material in herbaria, in particular to exsiccati
issued by W. Bauer and by A. J. Grout, and material annotated by C. B. Arzeni
and by E. Britton. The distributions of the species differ in the flora area,
but elsewhere their ranges overlap.
1. Plants mostly weakly and irregularly pinnate, branch leaves weakly seriate; base of stem and upper stipe leaves rounded to auriculate, with a small area of shorter, rhombic alar cells not reaching the leaf margin 1. Pireella pohlii 1. Plants mostly simple, sometimes weakly pinnate, branch leaves strongly seriate, base of stem and upper stipe leaves weakly decurrent, with several rows of quadrate alar cells extending up the leaf margin 2. Pireella cymbifolia 1. Pireella pohlii (Schwägrichen)
Cardot, Rev. Bryol. 40: 18. 1913 Leucodon pohlii Schwaegrichen, Spec. Musc. Frond. Suppl. 3(1):
232. 1828; Pireella ludoviciae (J. K. A. Müller) Grout Plants 0.5--10 cm; branching irregularly pinnate. Stem
leaves in short basal stipe region ovate-triangular; distal leaves erect
from clasping base, ovate-acuminate, 0.6--2. 5 X 0.4--1.1 mm; base not
decurrent, weakly to strongly rounded, often auriculate; apex narrowly
acuminate; costa filling awl-like apex in distal leaves; medial laminal cells
sinuose, 18-64 X 2.5 µm; alar cells variable, indistinct or (especially in
auriculate plants) with a few rows of irregular-quadrate cells not reaching
leaf margin. Branch leaves in five obscure spiralling rows;
ovate-acuminate, 0.9--1.7 X 0.3--0.6 mm; strongly concave with long narrow
flat acuminate apex; medial laminal cells 18--62 X 2.5 µm, weakly to strongly
prorate, tips of prorae with low conical papillae. Seta 5--12 mm. Capsule
cylindrical, 1.5--2.5 X 1 mm. Calyptra cucullate, covering capsules at
maturity, with scattered biseriate hairs. Spores spherical or ovoid,
23--37 X 18--35 µm. Capsules mature in late winter
(Feb.--Mar.); sporophytes known from only one site in Florida in the flora
area. Dry, mesic or boggy hardwood forests and hammocks, deep wooded ravines,
mixed woodland and dry sandy scrub, tree buttresses and trunks from base to
20 m, branches, also palms, logs, rocks; 0--1900 m; Ala., Fla., La., Miss., c
and s Mexico, West Indies, Central America, South America (to Paraguay and s
Brazil). Few other mosses in the region are likely
to be confused with P. pohlii, other than P. cymbifolia. Plants
of P. pohlii are usually irregularly pinnate, but may be simple if
depauperate. The bases of stipe and stem leaves are variously rounded to
almost auriculate, the alar cells are sub-quadrate or rhombic and few in
number, and do not reach the margin. The species is found from central and
northern Florida to Louisiana.
2. Pireella cymbifolia (Sullivant)
Cardot, Rev. Bryol. 40: 17. 1913 Pilotrichum cymbifolium Sullivant in A. Gray, Manual (ed. 2.) 681. 1856 Plants 0.2--2.5 cm, unbranched or sparsely and
irregularly pinnate. Stem leaves in basal stipe region broad
ovate-triangular, often very small for a large proportion of the stipe, 0.5 X
0.3 mm; distal leaves erect from clasping base, ovate-lanceolate, 1.8 X 0.9
mm; base decurrent, not rounded or auriculate; apex broadly acuminate; costa
percurrent in distal leaves, medial lamina cells sinuose, 16--46 X 2.5 µm;
alar cells in 7--14 rows of 2--23 quadrate or short rectangular cells 9--16 X
3--9 µm, running into decurrent wing and up leaf margin. Branch leaves
in five distinct spiralling rows; ovate-acuminate, 1.1--1.7 X 0.3--0.6 mm,
strongly concave with short broad flat acuminate apex; medial laminal cells
16--46 X 2.5 µm, strongly prorate, tips of prorae with low conical papillae.
[Seta 5--9 mm. Capsule cylindrical, 1.8--2.7 X 0.9 mm. Calyptra
cucullate, covering capsules at maturity, with scattered biseriate hairs. Spores
spherical or ovoid, 30--41 X 28--35 µm.] Sporophytes not known from flora area. Dry
evergreen forest, lowland deciduous forest and hammocks, occasionally flooded
palm woodland, and disturbed woodlands, on trees and tree roots, bushes,
decaying wood, limestone and sandstone; 0--700 m; Fla.; s Mexico, West
Indies, Central America, South America (to offshore islands in s Brazil.) Plants of P. cymbifolia are usually
small and scarcely branched, although well-grown plants (for example in the
Florida Keys) may be larger and irregularly pinnate. The leaf bases are
decurrent, with several rows of quadrate alar cells extending up the margin. P.
cymbifolia is restricted to southern Florida. Superficially, Pireella
cymbifolium may resemble Henicodium, but under the microscope the
more extensive area of oblate cells in the leaf base and the distinctly
plicate leaves in the latter species should prevent confusion. 3. Henicodium (J. K. A. Müller) Kindberg, Enum. Bryin. Exot. 16. 1888 William R. Buck Hypnum sect. Henicodium J. K. A. Müller, Linnaea 39: 470. 1875; Leucodontopsis Renauld & Cardot, "Leucodoniopsis" Plants medium-sized, bright- to yellow-green to bronze; creeping stolons turning away from substrate to form erect stems that are not or little branched. Stem leaves dense, not seriate, appressed when dry, spreading when moist, oblong-lanceolate to oblong-ovate, concave, plicate, broadly and gradually acuminate; base somewhat decurrent; margins revolute from just below apex to mid-alar region; costa single, ending about 2/3 the leaf length, often obscured, especially above, by plicae; laminal cells linear-flexuose, 1-papillose, firm-walled, ± porose; alar cells well differentiated in large, conspicuous areas, oblate, collenchymatous. Seta elongate, ca. 6 mm. Capsule well exserted, short-cylindric; annulus and operculum not seen. Calyptra not known. Species probably 1 (1 in flora); North America, West Indies, Central America, South America, Africa, Asia. Henicodium is distinguished by mostly unbranched, erect stems with revolute-margined leaves. The leaf cells are 1-papillose (individual leaves may sometimes be smooth) and the costa ends well below the leaf apex. Gemmae are regularly produced on older stems in large clusters in distal leaf axils. SELECTED REFERENCES Buck, W. R. 1989. Henicodium replaces Leucodontopsis (Pterobryaceae). Bryologist 92: 534. Buck, W. R. 1998. Pleurocarpous mosses of the West Indies. Mem. New York Bot. Gard. 82: 1--400. 3. Henicodium geniculatum (Mitten) W. R. Buck, Bryologist 92: 534. 1989 Leucodon geniculata Mitten, J. Linn. Soc., Bot. 12: 409. 1869; Leucodontopsis floridana (Austin) E. Britton; Leucodontopsis geniculata (Mitten) H. A. Crum & Steere Plants medium-sized to slender, mostly 1--2 cm; erect stems not or rarely branched. Stem leaves appressed when dry, spreading when moist, oblong-lanceolate to oblong-ovate, 1.2--2.0 × 0.25--0.88 mm, apex gradually broadly acuminate or often appearing ± abruptly so because of subapical recurved margins; median laminal cells linear-flexuose, 42--71 × 4 _¼m, 1-papillose, rarely smooth; alar cells in 7--15 rows and extending up the margins by 25--50 cells, oblate, collenchymatous. Sporophytes not known from the flora region. In ± open, mesic to dry forests, branches or less often tree trunks, frequently in, but rarely collected from, the canopy; near sea level in the floral area (to above 1000 m in South America); Fla.; Mexico; West Indies; Central America; South America; Africa; se Asia. Henicodium geniculatum has mostly simple, erect secondary stems with 1-costate leaves. The margins are recurved nearly throughout and the cells are 1-papillose. The species may be distinguished from various Pireella species by leaves appressed when dry and spreading when moist (rather than little altered when dry), and leaf cells papillose over the lumina (rather than smooth or prorulose). Henicodium seems most often confused with poorly developed specimens of Pseudocryphaea domingensis. However, that species has a percurrent costa, leaf cells much wider and usually smooth, and no axillary gemmae but, usually, conspicuous flagellate branches. Version History for Pterobryaceae: Version 2: June 2001 Version 1: May, 2001 |
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