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BFNA
Title: Pilotrichaceae |
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PILOTRICHACEAE
Kindberg Patricia M. Eckel Plants pleurocarpous, small or
slender to robust, rather soft or delicate, shriveled when dry, in loose or moderately
dense, mostly flat dark green to yellow-green, dull or glossy mats. Primary stems creeping and spreading,
leaves reduced, regularly or irregularly pinnately branched, moderately
radiculose, sometimes in tufts; secondary stems spreading, suberect and
frondose, not or sparingly branched. Stems
mostly freely branched, green, more or less prostrate, not ascending, more
or less radiculose, irregularly to pinnately branched; paraphyllia lacking; pseudoparaphyllia
absent [filamentous]; hyalodermis present or absent, central strand absent;
axillary hairs of two to several cells, commonly with an elongate hyaline
distal cell, basal cell small, brown. Leaves
mostly more or less complanate, often asymmetric
and dorsiventrally oriented, somewhat dimorphic, vertical (dorsal and
ventral) leaves erect, broadly oblong, oblong-ovate, to ovate-lanceolate,
usually abruptly or gradually acuminate, symmetrical, lateral leaves tending
narrower, erect-spreading to spreading, less symmetric; frequently bordered
with elongate cells; margins often dentate; laminal cells smooth to 1-papillose,
often lax to firm-walled, isodiametric or linear-rhomboidal, essentially
undifferentiated in alar regions; costa double and well-developed. Specialized
asexual reproduction infrequent, by cylindric gemmae, variously in leaf
axils, specialized branches, and ventrally on stems. Sexual condition polygamous, autoicous, synoicous, dioicous. Seta elongate, red to reddish-yellow;
smooth or papillose-scabrous to spinose distally. Capsule inclined to
pendulous, less frequently erect, ovoid to oblong-cylindric, symmetric, smooth,
annulus none or weakly differentiated; peristome double, dark red or
red-brown, of 16 teeth lanceolate to lance-subulate, furrowed,
cross-striolate basally, papillose distally, trabeculate, sometimes bordered;
endostome yellow to yellow-brown, finely papillose to nearly smooth, basal
membrane generally high, keeled, segments lanceolate, about as long as the
teeth; cilia absent; operculum long-rostrate to subulate from a conic base. Calyptra conic-mitrate to
mitrate-campanulate, shortly lobed or deeply fringed-laciniate at the base,
smooth or papillose-scabrous distally, naked [in our species]. Spores smooth to very finely
papillose. Genera
21, species ca. 400 (3 genera, 3 species in the flora): pan- and subtropical.
The
Pilotrichaceae are diverse and abundant in the humid tropic. The family is
mainly Neotropical (W. Frey 2009). Three
species, representing different genera, have reached the flora area, where
they occur mainly in the South. The
family as currently understood is characterized by a double costa, generally
with a stem hyalodermis present, undifferentiated laminal alar region, and hairy
calyptra. Leaves are generally complanate, with generally a differentiation
between lateral and dorsal or ventral leaves, seta variously papillose to
spinose. The recent, rapid and complex changes in the circumscription of the
Pilotrichaceae are outlined by T. F. Vaz-Imbassahy and D. P. da Costa (2008).
These authors also present a chart comparing this and related families,
indicating how fluid and complex the taxonomy of the family and its
relationship to the traditional Hookeriaceae has been. Such reconsiderations of the family
boundaries began rather recently with a revision by H. A. Miller (1971). W. R. Buck (1998) suggested two central
lines of evolution in the family by which genera may be organized based on
the presence or absence of a differentiated outer layer of inflated stem
cortical cells (a hyalodermis), compared to an outer cortical layer of relatively
thick-walled cells. A pilose calyptra is characteristic of the
Pilotrichaceae, but the calyptra is naked in the three species, one in each
of the three genera, represented in the flora area. There is extensive
floristic literature on the family, but little of relevance to the flora area.
Critical literature associated with the pre-1971 concept of the Hookeriaceae
in the flora area is that of Welch (1962, 1966, 1972). All representatives of
the family are rare in the flora area. SELECTED
REFERENCES Allen, B. 2010. Moss Flora of 1.
Laminal cells with a central papilla; leaf apex rounded-truncate and short-apiculate;
seta papillose; stem hyalodermis absent, leaf border absent . . . 3. Callicostella,
p. xxx. 1.
Laminal cells smooth; leaf apex short to long-acuminate; seta smooth; stem
hyalodermis present or absent; leaf border present or absent. 2. Leaf apex shortly acuminate, entire:
laminal margins bordered by one row of linear cells, median cells large,
nearly isodiametric, laxly hexagonal or oblong-hexagonal; stem hyalodermis
present . . . 1. Cyclodictyon, p.
xxx 2. Leaf apex long and slenderly pointed,
serrate; laminal margins not bordered, median cells narrow, linear, not noticeably lax; stem hyalodermis absent
. . . 2. Trachyxiphium, p. xxx 1.
CYCLODICTYON Mitten, J. Proc. Linn. Soc., Bot. 7: 163. 1863 * [Greek cyclos-, circular, and dictyon, network, alluding to large, nearly
circular laminal cells] Plants small and slender to medium-sized,
soft, delicate, in white to pale green or bluish glaucous, thin and flattened
mats. Stems and branches prostrate and rooting along the
stem, irregularly to pinnately branched; with a hyalodermis 1-stratose over
firm-walled median cells surrounding large thin-walled central cylinder; sparsely
radiculose. Leaves asymmetric, complanate, shriveled or more or
less crisped when dry, broadly oblong or oblong-ovate, abruptly narrowing to
a short or fairly long acumen, rarely ovate-lanceolate and gradually
acuminate, dorsiventral and lateral leaves not much differentiated in shape, margins
plane or occasionally narrowly recurved,
bordered by 1 to occasionally several rows of elongate and narrower cells,
entire or serrulate distally, otherwise similar to the laminal cells; costa
double, divergent, extending about 2/3--3/4 the leaf length, ending beyond
the leaf middle, smooth or somewhat serrulate distally; laminal cells hexagonal
or oblong-hexagonal, lax, thin-walled, transparent, smooth, becoming linear in 1 to
few rows at the margins and forming a concolorous border, oblong or
rhomboidal at leaf base. Specialized
asexual reproduction not known. Sexual
condition synoicous, dioicous or autoicous; perichaetial leaves small,
ovate to lanceolate, short- to long-acuminate. Seta elongate, dark red, smooth. Capsule inclined to horizontal, oblong-cylindric, annulus narrow,
operculum straight and long-rostrate from a conic base; exothecial cells not
collenchymatous; peristome trabeculate. Calyptra
mitrate, rostrate, short-lobed at the base, scarcely longer than the
operculum, smooth. Species
90 (1 in the flora): largely tropical and occasionally subtropical. Cyclodictyon species occur largely
in the Neotropics, though they are widespread in tropical SELECTED
REFERENCE: Gradstein, S. R., S. P. Churchill, and N. Salazar-Allen. 2001. Guide
to the Bryophytes of Tropical 1. Cyclodictyon
varians (Sullivant)
Kuntze, Revis. Hookeria varians Sullivant, Proc. Amer.
Acad. Arts 5: 285. 1861 Plants in delicate, intricate flat mats, with metallic, luster when dry. Stems much branched. Leaves 1.3--2 mm, ovate to broadly ovate, with
short acumination, marginal border of a
single row of narrow, thin-walled and elongate cells; median cells 29-40 \um
wide, 1.5--2:1, lax, oblong-hexagonal; costa divergent, slender, ending only in the base of acumen,
essentially smooth, without dorsal projections. Specialized asexual reproduction unknown. Sexual condition polygamous: synoicous and autoicous. Seta 11--15.5
mm, somewhat sinuose. Capsule inclined,
horizontal to pendulous, more or less asymmetric, contracted below the mouth
when dry; operculum long and slenderly rostrate. Calyptra naked, lobed at base. Wet
places, occasionally aquatic, soil, humus, leaf litter, occasionally soil
over limestone and limy sandstone rocks, springs, hammocks, stream banks, limestone
sinks, deep ravine bottoms in deeply shaded forests; near sea level; Fld.,
La.; West Indies; Central America; n South America; Atlantic Islands
(Bermuda). Cyclodictyon varians is rare in the flora,
found in southeastern SELECTED
REFERENCES: Crum, H. A. and L. E. Anderson. 1981. Mosses of Guide
to the Bryophytes of Tropical 2.
TRACHYXIPHIUM W. R. Buck, Brittonia
39: 219. 1987 * [Greek trachy-, rough, and xiphion, sword, alluding to sword-shaped
leaves marginally roughened by serrations] Plants small, medium-sized to
fairly large, in glossy, soft, dense, light to brownish or golden green to
bluish-green mats. Stems prostrate
and creeping, closely irregularly and freely branched, hyalodermis lacking, the
cortical layer with small cells somewhat thicker walled than the larger,
thin-walled central cells. Leaves
slightly asymmetric, lateral
leaves generally spreading, vertical leaves erect, somewhat to occasionally clearly
complanate, flaccid, somewhat flexuose with the margins plane, incurved when
dry, infrequently asymmetric, sometimes secund or falcate-secund, lanceolate
to ovate-lanceolate, the short and broad to long and slenderly pointed acumen
serrate marginally; margins broadly or narrowly recurved on one side, sharply
serrate, especially distally, with bifid, tumid teeth, entire below, marginal
cells not differentiated into a border; costa double, slender and well
developed, 1/2--2/3\x length of the leaf, ending at about the base of the
acumen, parallel to slightly divergent,
distally toothed on the dorsal side; laminal cells homogeneous, long-rhomboidal
to linear, narrow, elongate, smooth or
sometimes prorulose by projecting distal cell ends. Specialized
asexual reproduction none. Sexual
condition polygamous: synoicous but may also have male and female
gametoecia; perichaetial leaves subulate from a broader base. Seta elongate, reddish, thick, smooth.
Capsule black, dark brown or
red-brown, ovoid, erect, inclined to
pendent, contracted below the mouth when dry, smooth; annulus none; operculum
obliquely and generally narrowly long-rostrate. Calyptra conic-mitrate, covering the distal portion of the urn, naked,
fringed at the base, often split on one side, smooth or distally slightly
scabrous. Species
15 (1 in the flora): pantropical. Ten
species of Trachyxiphium occur in
the Neotropics, growing on wet soil, rocks and logs in humid lowlands to mid
elevations, occasionally upper montane forests, from near sea level to 3400 m
(S. R. Gradstein et al. 2001). SELECTED
REFERENCE Gradstein, S. R., S. P.
Churchill and N. Salazar-Allen. 2001. Guide to the Bryophytes of Tropical 1. Trachyxiphium hypnaceum
(Müll. Hal.) W. R. Buck, Brittonia 39: 220. 1987 C F Hookeria hypnacea Müll. Hal., Bot. Zeit.
( Plants slender, flaccid, pale
whitish yellow to brownish green to yellow-green, nearly hyaline when wet, not
particularly complanate. Leaves 0.9--1.5 mm, somewhat distant (green or
red stems visible between the leaf bases especially when dry), loosely
erect-spreading, ovate- to narrowly lanceolate, becoming lightly to strongly
falcate-secund near the stem tips, not undulate or plicate, somewhat flexuose
but not crisped when dry, concave in proximal 2/3, distally gradually or sometimes
abruptly narrowing to a long, flat, broad or slender and sharply pointed
acumen that is somewhat flexuose when dry; margins serrulate in distal 1/3, tumid
teeth single or double often tipped with a papilla; costa slender, cristate
distally, dorsally strongly serrulate distally; distal laminal cells 7.5--10:1,
linear-rhomboidal, pellucid with thin walls, smooth, the cell apices mostly
pointed and often sharply prorulose. Seta
orange-brown, smooth, 6.5--12 mm. Capsule
black on a red seta, 0.7--1.5 mm. Spores
spherical, 8--10.5 \um, essentially smooth. Capsules
mature early March. Rocks, wet places on soil, tree bases, wet, rotted wood, near
spring seepage and stream banks, deep shade in wet forests, concrete culvert
around a spring, cut-over woods of Pinus, Fagus, Magnolia grandiflora; moderate
elevations; La., Miss.; Mexico; Central America (Costa Rica in Cocos Island);
South America. Both
the SELECTED
REFERENCES Reese, W. D. and B. E.
Lemmon. 1970. Mosses of Clear Springs Recreation Area, 3.
CALLICOSTELLA (Müller Hal.) Mitten, J. Proc. Linn. Soc., Bot. Suppl. 1: 136.
1859, conserved name * [Latin calli, hardened
or thick, and costa, rib, alluding
to the strong costae] Hookeria sect. Callicostella Müller Hal., Syn. Musc.
Frond. 2: 216. 1851; Hookeria subg.
Callicostella (Müller Hal.) Hampe; Schizomitrium Bruch & Schimp. in
Bruch, Schimper & Gümbel, rejected name. Plants small to medium-sized,
in dull, sordid pale-green, yellowish, brownish, sometimes bluish-green, flattened
mats. Stems prostrate and rooting,
branching freely, regularly or irregularly pinnate; hyalodermis lacking,
epidermis composed of small, thick-walled cells, central cylinder of large,
thin-walled cells, sparsely radiculose in tufts. Leaves somewhat asymmetric, more or less complanate, shriveled,
often crisped when dry; ovate-oblong to lingulate, dimorphic, ventral leaves
often more acuminate, dorsal and lateral leaves mostly broadly oblong to
oblong-ovate, apex bluntly and broadly rounded, usually abruptly apiculate, margins
plane, not or obscurely bordered, crenulate
to serrulate to strongly toothed in the apical quarter, smooth in basal
quarter; costa double, strong, ending in the leaf apex, subpercurrent, divergent
from the base, parallel or somewhat convergent distally, distally toothed
dorsally; laminal cells isodiametric or a little longer than wide,
irregularly hexagonal to quadrate, variously short-rectangular and
thin-walled in the proximal half to nearly isodiametric in the distal half
with thicker walls and rounder corners; each lumen with a single, central sharp papilla on both
surfaces, laminal cell corners angular, not thickened; basal cells
rectangular, 6--2:1; smooth, cell walls thin, even, somewhat vesiculose at mid-base,
small alar group turning inward in a curve at the extreme basal margin. Specialized asexual reproduction
rare, of brownish, 1-seriate, papillose gemmae in the axils of the leaf. Sexual condition polygamous,
infrequently dioicous; perichaetial leaves small, inconspicuous, ovate- or
linear-lanceolate, gradually acuminate to acute, with costa double. Seta red to reddish yellow, smooth or
distally papillose, sometimes strongly so, or scabrous throughout. Capsule ovoid or oblong-cylindric, inclined
to horizontal to pendulous, exothecial cells mostly collenchymatous, annulus
none; operculum abruptly short-rostrate or subulate; peristome with
well-developed trabeculae ventrally, projecting from the sides. Calyptra narrowed and shortly or deeply
lobed at the base, mitrate-campanulate, smooth or sometimes split on one side
from base, somewhat scabrous or with a few hairs distally. Species
95 (1 in the flora): pantropical and southern Callicostella is characterized by
broad, lingulate, shortly and abruptly acuminate leaves with no border. The
long, tough double costa is distinctive, as are the small, isodiametric,
firm-walled leaf-cells, especially those of the apex, each with a single,
central papilla. The distal margins of the leaf are also strongly serrate with
teeth often forking. Other species in the genus may have calyptra fimbriate
at base, but our species the calyptra is only lobed. A technical character
that helps to distinguish this genus from closely related genera is the epidermis
of the stem section composed of small, thick-walled cells (i.e. lacking a
hyalodermis). SELECTED
REFERENCES: Bowers, F. D. Callicostella.
In: A. J. Sharp, H. A. Crum and P.
M. Eckel, eds. The Moss Flora of 1. Callicostella pallida
(Hornschuch)
Ångström, Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 33(4): 27. 1876 F Hookeria pallida Hornschuch in Martius,
Fl. Bras., 1(2): 64. 1840; Schizomitrium
pallidum (Hornschuch) H. A. Crum & L. E. Anderson Leaves with distal laminal cells
homogeneous, ca. 8--11 \um wide, not decurrent, differentiated lateral leaves
more asymmetrical than the erect dorsal/ventral leaves, long cells from the
stem may remain attached at the insertion. Specialized asexual reproduction not seen. Sexual condition polygamous (synoicous and autoicous, appearing
to have some male or female buds that also contain both sex organs); perichaetial
leaves ovate-lanceolate, 0.8--1 mm, gradually acuminate to acute. Seta 6--8(--20) mm, moderately to
strongly scabrous throughout with broad, low papillae. Capsule ca. 1 mm, constricted below the mouth and at the short
neck when dry; operculum 0.7--1 mm. Spores
8--11 \um. Capsules
present winter (late December--March). Moist sites, sometimes subaquatic or
submerged, usually moist rock and soil, occasionally rotted stumps, logs, limestone
rocks in springs, wet limy muck, sandy soil, stream banks, water-filled
ditches, hammocks, deeply shaded ravines, dark swampy subtropical forests
with Magnolia and Illicium; may be associated with Cyclodictyon and Hygroamblystegium; low elevations near sea level, 0--80 m; Ala., Fld.,
La.; Mexico; Central America; West Indies; South America. The
presence of Callicostella pallida in
the flora is apparently not relictual, but recently introduced (W. D. Reese
1984). Like Callicostella pallida, Trachyxyphium
heteroicum has a robust double costa and broad leaves lacking a margin of
differentiated cells, but T. heteroicum
has leaves that are long-acuminate (not truncate-lingulate with an abrupt apiculus),
cells smooth (with no central papilla), leaf cells linear-rhomboidal and
thin-walled (not isodiametric with firm walls). |
