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BFNA Title: Thuidiaceae |
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XX.
THUIDIACEAE Schimper William R. Buck Plants
small to robust, in typically stiff, dark-green to golden, loose mats. Stems creeping or ascending, mostly
regularly 1--3-pinnate, not at all stipitate; without a hyalodermis, central
strand small; paraphyllia present, filamentous to foliose, cells papillose;
pseudoparaphyllia foliose; axillary hairs usually a single short brown basal
cell and elongate hyaline distal cells, rarely (Abietinella) all cells brown. Stem and branch leaves strongly differentiated. Stem leaves mostly spreading when
moist, usually broadly ovate, often abruptly acuminate, concave; margins
serrulate to crenulate ± throughout, plane or recurved; costa single, strong,
ending in the acumen to ± excurrent; cells quadrate to short-rectangular, 1-
or pluripapillose on both surfaces or only abaxially, firm-walled; alar cells
not or poorly differentiated. Branch
leaves smaller than stem leaves, usually spreading, lanceolate to ovate,
usually gradually acuminate to obtuse; margins serrulate to crenulate ±
throughout, usually plane; costa single, weaker than in stem leaves; cells ±
quadrate, 1- or pluripapillose on both surfaces or just at back, the apical
cell typically truncate and pluripapillose, firm-walled, not porose; alar
cells not or poorly differentiated. Specialized
asexual reproduction none. Sexual
condition autoicous or dioicous. Perichaetia
conspicuous; leaves mostly erect, oblong-lanceolate, long-acuminate,
sometimes plicate; margins subentire to long-ciliate, plane; costa single,
usually strong; cells rectangular, more elongate than in vegetative leaves,
usually smooth. Seta elongate,
smooth to roughened, either throughout or rarely only distally, reddish. Capsule mostly inclined to
horizontal, rarely erect, mostly asymmetric and arcuate, cylindric;
exothecial cells thin- to thick-walled, not collenchymatous; annulus
differentiated; operculum usually rostrate; peristome double, attached at the
mouth, exostome teeth shouldered, bordered, externally cross-striolate
basally, sometimes with overlying papillae, papillose, trabeculate
internally; endostome mostly with a high basal membrane, segments keeled, not
to narrowly perforate, cilia in groups of 1--3, nodulose. Calyptra cucullate, rarely mitrate,
mostly naked, smooth or roughened. Spores
spheric, usually papillose. Genera 9, species ca. 130 (4 genera, species in the
flora): cosmopolitan (but rare in Arctic), greatest occurrence in North
Temperate region and southern The Thuidiaceae family is characterized by pinnately
branched plants with strongly differentiated branch and stem leaves. The
stems are usually amply clothed with paraphyllia, but in smaller plants
paraphyllia are often sparse. The leaf cells of both stem and branch leaves
are short and papillose. Only Rigodium
(Rigodiaceae) approaches the aspect presented by the Thuidiaceae, but it is
stipitate, there are no paraphyllia, and leaf cells are smooth. Haplocladium
(referred here to the Leskeaceae) might be confused with various Thuidiaceae.
However, in that genus the paraphyllia are not papillose (as they are throughout
the Thuidiaceae) and branch leaves terminate in a smooth, sharp cell, rather
than a truncate, pluripapillose cells (except in Rauiella praelonga). The Thuidiaceae were distilled to a more
natural family by W. R. Buck and H. A. Crum (1990), emphasizing sexuality,
placement of leaf cell papillae, papillosity of the cells of the paraphyllia,
morphology of axillary hairs, and seta ornamentation. More recently, A. Touw
(2001) provided a different generic scheme, which for the most part is not
followed here because it seems to have been based primarily on knowledge of
the Asian flora. A final resolution will require molecular study and a survey
of all the taxa formerly associated with the Thuidiaceae, Leskeaceae and
Anomodontaceae. selected references Buck,
W. R. and H. A. Crum. 1990. An evaluation of familial limits among the genera
traditionally aligned with the Thuidiaceae and Leskeaceae. Contr. Univ. Mich.
Herb. 17: 55--69. Touw, A. 2001. A review of the Thuidiaceae (Musci) and a
realignment of taxa traditionally accommodated in Thuidium sensu amplo (Thuidium
Schimp., Thuidiopsis (Broth.) M.
Fleisch., and Pelekium Mitt.),
including Aequatoriella gen. nov.
and Indothuidium gen. nov. J.
Hattori Bot. Lab. 90: 167--209. 1.
Plants 1-pinnate. 2.
Plants robust, erect-ascending; laminal cells stoutly 1-papillose; apex of
costa of branch
leaves not covered with quadrate cells ……………………….. 1. Abietinella 2.
Plants slender, creeping; laminal cells bulging, pluripapillose; apex of
costa of branch
leaves covered with quadrate cells ……………………………… 3. Rauiella 1.
Plants (1--)2--3-pinnate. 3. Plants robust, dioicous;
paraphyllia branched; leaf cells papillose only abaxially, usually
1-papillose; seta smooth................................................................ 4.
Thuidium 3.
Plants small, autoicous; paraphyllia usually unbranched; leaf cells papillose
on both
free surfaces, usually pluripapillose; seta often roughened........... 2. Cyrto-hypnum XX. Abietinella
Müller Hal., Nuovo Giorn. Bot. Ital. n.ser. 3: 115. 1896 * [Latin abietis, of the conifer genus Abies, and -ella, diminutive, alluding to the habit aspect] William
R. Buck Thuidium
subgen. Abietinella
(Müller Hal.) Brotherus, T. sect.
Abietina Schimper Plants
relatively robust, rigid, 1-pinnate; paraphyllia filamentous to narrowly foliose,
simple or branched; axillary hairs 2-celled, all cells brown. Stem leaves ovate, acuminate,
plicate; margins crenulate-papillose, often recurved at least proximally;
costa strong, extending ca. 3/4\x leaf length; cells rhombic to
oblong-hexagonal, 1-papillose on both surfaces. Branch leaves smaller than stem leaves, ovate, acute to broadly
acuminate; margins as in stem leaves but mostly plane; costa as in stem
leaves; cells rounded-quadrate to short-rectangular, 1-papillose on both
surfaces except terminal laminal cell pluripapillose. Sexual condition dioicous. Perichaetial
leaves strongly differentiated, erect, oblong-lanceolate, long-acuminate;
margins serrate to serrulate above; costa subpercurrent; cells elongate,
thick-walled, porose. Seta elongate,
slender, smooth. Capsule inclined,
strongly curved, cylindric; annuli differentiated in 2--3 rows; annuli not
seen; opercula obliquely short-rostrate; peristomes hypnoid with endostomial
segments narrowly perforate and cilia 1--2. Calyptra cucullate, naked. Species
3 (1 in the flora): North America, Europe, Abietinella
is an easy genus to recognize in the flora. It is characterized by relatively
robust, stiff, more or less erect, once-pinnate, paraphylliate stems, often golden-brown,
growing most often on fairly dry, mostly calcareous sands. 1.
Abietinella abietina (Hedwig) M. Fleischer, Musci Fl.
Buitenzorg 4: 1497. 1923 F Hypnum
abietinum Hedwig, Sp. Musc. 353. 1801; Thuidium abietinum (Hedwig) Schimper Plants
dark-green, yellowish or dark brown, sometimes blackish-tinged. Stems to 12 cm, erect-ascending;
branches short, unequal, tapered; paraphyllia abundant. Stem leaves erect when dry, erect-spreading when moist, 1.2--1.8
mm, orange at the insertion; cells stoutly 1-papillose over the lumina. Branch leaves erect when dry,
erect-spreading when moist, 0.6--0.7 mm. Perichaetial
leaves to 4 mm, long-acuminate, distal margins strongly serrate. Seta 2--2.5 cm; capsules 2--3 mm,
yellow-brown. Spores 9--11 \um,
finely papillose. Capsules very
rare, maturing late summer. Dry, exposed calcareous rocks and soil, sand of
partially stabilized dunes, among talus at base of cliffs, and humus in open,
coniferous forests; low to moderate elevations; Greenland; Alta., B.C., Man.,
N.B., Nfld. and Labr. [Nfld.], N.W.T., N.S., Nunavut, Ont., Que., Sask.,
Yukon; Alaska, Ariz., Colo., Conn., Iowa, Maine, Mich., Minn., Mo., Mont.,
N.H., N.J., N.Mex., N.Y., N.Dak., Pa., S. Dak., Va., Vt., Wis., Wyo.; Europe;
Asia. Abietinella
abietina is the easiest of the Thuidium-like
plants to recognize in our flora, in part because of the
distinctive habitat requirements. The stems are once-pinnate, erect-ascending
and often occur in extensive, lax mats. Paraphyllia are abundant on stems and
branches, but are usually not nearly as strongly branched as in Thuidium s.str. It is widespread
throughout the northern part of our range. Unlike Thuidium, Abietinella
is papillose on both leaf surfaces rather than just the back. 2. CYRTO-HYPNUM
(Hampe) Hampe & Lorentz in G. Hampe, Bot. Zeit. 27: 455. 1869 * [Greek cyrto, curved or arched, and hypnum,
lichen, or by usage, pleurocarpous moss] William
R. Buck Hypnum
subg. Cyrto-hypnum Hampe, Flora 50:
78. 1867; Thuidium sect. Minutula Schimper; T. subg. Microthuidium Limpricht Plants
small and slender, 1--2-pinnate; paraphyllia filamentous,
unbranched; axillary hairs 2-celled. Stem leaves broadly triangular to
broadly ovate, acuminate; margins crenulate-papillose, often recurved at
least basally; costa strong, extending 4/5\x leaf length to subpercurrent;
cells ± quadrate, pluripapillose [1-papillose] on both surfaces, usually with
the papillae arranged around the periphery of the lumina. Branch
leaves typically smaller than stem leaves, ovate to oblong-ovate, obtuse
to short-acuminate; margins as stem leaves but plane; costa strong, ending
3/4--7/8\x leaf length, sometimes crested or the apex projecting as a small
spine; cells similar to those of stem leaves.
Sexual condition autoicous. Perichaetial
leaves strongly differentiated, erect, oblong-lanceolate to oblong-ovate,
acuminate; margins entire to ciliate; costa percurrent to excurrent. Seta
slender, smooth or papillose. Capsule
inclined to horizontal, cylindric; annulus differentiated in 2--3 rows;
operculum obliquely long-rostrate; peristome hypnoid with endostomial
segments not or narrowly perforate and cilia 1--3. Calyptra
cucullate, naked. Species 30--40 (4 in the flora): mostly tropical. SELECTED REFERENCE
Buck, W. R. and H. A. Crum. 1990. An evaluation of familial limits
among the genera traditionally aligned with the Thuidiaceae and
Leskeaceae. Contr. Univ. Michigan
Herb. 17: 55--69. 1. Stems 1-pinnate; branch and stem leaves of about the
same size, more than 0.4 mm, strongly incurved when dry . . . . 1. Cyrto-hypnum involvens 1. Stems 2-pinnate (sometimes inconspicuous); branch
leaves smaller than stem leaves, less than 0.4 mm, erect to somewhat but not
strongly incurved when dry. 2.
Stems and branches as well as the paraphyllia papillose . . . . 2. Cyrto-hypnum pygmaeum 2. Stems and branches with papillose
paraphyllia, but not papillose themselves. 3. Stems weakly and irregularly
2-pinnate; cells of paraphyllia mostly twice as long as wide; seta smooth;
perichaetial leaves serrulate . . . .
3. Cyrto-hypnum minutulum 3. Stems regularly 2-pinnate; cells
of paraphyllia mostly quadrate; seta rough; perichaetial leaves ciliate . . .
. 4. Cyrto-hypnum schistocalyx 1. Cyrto-hypnum involvens
(Hedwig) W. R. Buck & H. A. Crum, Contr. Univ. Michigan Herb. 17: 66.
1990 Leskea
involvens Hedwig, Sp. Musc. Frond,
218. 1801; Thuidium involvens (Hedwig) Mitten Stems
1-pinnate, smooth; paraphyllia to ca. 8 cells in length,
cells ca. 1:1, all sparingly papillose.
Stem leaves
flexuose-incurved when dry, ovate-triangular, (0.3--)0.5--0.6 mm, ± abruptly
short-acuminate; costa disappearing in the acumen but not filling it. Branch
leaves strongly incurved when dry, ± 2-ranked, often laxly disposed, ca.
0.5 mm, obtuse to acute; costa ca. 7/8\x leaf length, often ± flexuose, not
covered with quadrate cells. Perichaetial leaves laxly serrulate
distally, subentire proximally or with a few coarse, blunt teeth. Seta
papillose throughout. Capsule with
endostomial cilia single, rarely with rudimentary additional cilium. Capsules mature Nov.--Dec.(--Mar.). Lowland, moist forests, growing over
limestone, soil, humus, rotten wood; 0 m; s The once-pinnate stems, strongly incurved leaves,
non-ciliate perichaetial leaves and rough seta distinguish Cyrto-hypnum involvens. The aspect is
quite different from the other species in the genus. It occurs between 0 and
1000 m south of the flora area. 2.
Cyrto-hypnum pygmaeum (Schimper) W. R. Buck & H. A.
Crum, Contr. Univ. Michigan Herb. 17: 67. 1990 Thuidium
pygmaeum Schimper in P. Bruch, W.
P. Schimper & W. Gümbel, Bryol. Eur. 5(fasc. 49/51): 162. 1852 Stems
2-pinnate, stems and branches papillose; paraphyllia
(1--)2--4(--5) cells in length, cells ca. 1:1, all strongly papillose. Stem
leaves slightly incurved when dry, ovate-triangular, 0.2--0.35 mm,
acuminate; costa about 3/4\x leaf length.
Branch leaves
erect-incurved when dry, 0.2--0.3 mm on primary branches, ca. 0.15 mm on secondary branches, acute to
short-acuminate; costa 1/2--3/5\x leaf length, straight, not covered with
quadrate cells. Perichaetial leaves serrulate distally, subentire
proximally. Seta smooth. Capsule with endostomial cilia in pairs. Capsules mature July--Sept. Moist woods, growing over moist rocks,
usually limestone but less often sandstone and quartzite; mostly at moderate
to low elevations; Ont., Que.; Ala., Ark., Conn., Fla., Ga., Ill., Ind.,
Iowa, Ky., Mass., Mo., N.C., N.J., N.Y., Ohio, Okla., Pa., Tenn., Va., Vt.
Wis.; Asia (China, Japan, Korea). Cyrto-hypnum
pygmaeum is the flora’s smallest species of Cyrto-hypnum. The
diminutive stature coupled with papillose stems and branches is
distinctive. It is most likely
confused with Heterocladium macounii, which also has
pluripapillose leaf cells and papillose stems and branches. However, that species is larger and has a
costa ending near or before mid leaf and is without paraphyllia; although
pseudoparaphyllia are present, these have pointed terminal cells, not
truncate as are the paraphyllia of C.
pygmaeum. 3.
Cyrto-hypnum minutulum (Hedwig) W. R. Buck & H. A.
Crum, Contr. Univ. Michigan Herb. 17: 66. 1990 Hypnum
minutulum Hedwig, Sp. Musc. Frond.,
260. 1801; Thuidium minutulum (Hedwig) Schimper Stems
weakly and irregularly 2-pinnate, smooth; paraphyllia
2--3(--5) cells in length, cells mostly ca. 2:1, usually only the terminal
one papillose. Stem leaves erect-incurved when dry, broadly ovate,
0.3--0.45(--0.55) mm, ± abruptly shortly and broadly acuminate; costa ending
about 4/5\x leaf length. Secondary branch leaves
erect-incurved when dry, 0.15--0.25 mm, acute; costa ending ca. 3/4\x leaf
length, straight, apical portions often covered with quadrate cells and thus
projecting as low crests. Perichaetial leaves serrulate
distally, subentire basally. Seta smooth. Capsule with endostomial cilia 1--2(--3). Capsules mature Feb.--Aug.(--Oct.). Moist woods, usually growing on rotten
logs, sometimes on rock, especially limestone, and bases of trees; mostly
moderate to low altitudes; Ont., N.B.,; Ala., Ark., Conn., Fla., Ga., Ill.,
Ind., Iowa, La., Maine., Md., Mich., Minn., Mo., N.J., N.Y., N.C., Ohio, Pa.,
S.C., Tenn., Va., W.Va.; Mexico, Central America, n,c South America; West Indies; Europe. Because of its wide range in eastern 4.
Cyrto-hypnum schistocalyx (Müller Hall.) W. R. Buck & H.
A. Crum, Contr. Univ. Michigan Herb. 17: 67. 1990 Hypnum
schistocalyx Müller Hall., Syn.
Musc. Frond. 2: 691. 1851; Thuidium
schistocalyx (Müller Hall.) Mitten Stems
regularly 2-pinnate, smooth; paraphyllia 2--8 cells in
length, cells mostly 1:1, all papillose.
Stem leaves appressed when
dry, broadly triangular, (0.2--)0.25--0.32(--0.5) mm, gradually and broadly
acuminate, sometimes piliferous; costa ending ca. 5/6 the leaf length. Secondary
branch leaves incurved-appressed when dry, 0.15--0.25(--0.3) mm, acute to
obtuse; costa ending ca. 3/4\x leaf length, straight, apical portions often
covered with quadrate cells and thus projecting as low crests. Perichaetial
leaves strongly flexuose-ciliate. Seta papillose throughout;
endostomial cilia 1(--2). Capsule maturity not determined. Moist forests, usually
growing on limestone or bases of trees; 0 m; The ciliate perichaetial leaves and roughened seta
characterize Cyrto-hypnum schistocalyx,
an essentially tropical taxon. Cyrto-hypnum schistocalyx occurs from sea level to 200 m south of the flora
area. 3. Rauiella
Reimers, Hedwigia 76: 287. 1937 * [For Eugene Abraham Rau, 1848--1932,
American bryologist, and Latin -ella,
diminutive] William
R. Buck Plants
mostly medium-sized, 1-pinnate; stems creeping, the branches prostrate to ascending;
paraphyllia ± filamentous, mostly unbranched, abundant on stems, sparse on
branches; axillary hairs 3-celled. Stem leaves broadly ovate, abruptly
long-acuminate, concave, somewhat decurrent; margins crenulate to
crenulate-papillose, plane or recurved; costa strong, ending near base of the
acumen, the apical portion not covered with short cells; cells ± quadrate,
bulging on both surfaces, pluripapillose (or rarely 1-papillose) on both
surfaces, with the papillae clustered over the lumina or arranged around the
periphery. Branch leaves smaller than stem leaves, ovate to, gradually
broadly acuminate; margins as stem leaves; costa strong, ending well beyond
midleaf, covered distally with short laminal cells; cells similar to those of
stem leaves. Sexual condition autoicous.
Perichaetial leaves
enlarged, mostly erect, mostly lanceolate, long-acuminate, ± plicate; margins
crenulate, never ciliate; costa subpercurrent; cells more elongate than in
vegetative leaves, often smooth. Seta elongate, smooth. Capsule suberect to horizontal,
short-cylindric, arcuate; annuli differentiated in 2--3 rows; opercula
stoutly and obliquely apiculate to rostrate; peristomes hypnoid with
endostomial segments not or narrowly perforate and cilia 1--3. Calyptra
cucullate, naked. Species 8 (2 in the flora): North
America, Central and South America, Africa, Rauiella
has frequently been misunderstood, being confused with Thuidium and Haplocladium.
It is characterized in the flora region by leaf pluripapillose on both sides
of the leaf, by well-developed papillose paraphyllia of oblate cells on the
stems (sparse on the branches), a smooth seta, and autoicous sexuality. The
distal part of the costa on branch leaves are covered by laminal cells (as
seen in Cyrto-hypnum), but in Rauiella the cells are also bulging on
both surfaces (but flat in Cyrto-hypnum).
SELECTED REFERENCE Reimers, H. 1937. Die europäischen Haplocladium-Arten mit besonderer
Berücksichtigung ihrer aussereuropäischen Verbreitung und ihrer Verwandtschaft.
Hedwigia 76: 191--298. 1. Apical cell of branch leaves pointed, not papillose;
southwestern ... 1. Rauiella praelonga 1. Apical cell of branch leaves pluripapillose;
eastern........... 2. Rauiella scita 1.
Rauiella praelonga (Bescherelle) Wijk & Margadant,
Taxon 11: 222. 1962 F Pseudoleskea
praelonga Schimper ex Bescherelle, Mém. Soc. Sci. Nat. Cherbourg
16: 233. 1872; Bryohaplocladium
praelongum (Bescherelle) H. A. Crum;
Haplocladium praelongum (Bescherelle) H. A. Crum; Rauia praelonga (Bescherelle) Brotherus Stems
2--3(--6) cm, ± regularly 1-pinnate, the branches to ca. 1 cm, mostly
shorter; paraphyllia mostly filamentous, 1-seriate distally and
2(--3)-seriate proximally, mostly unbranched, the cells oblate,
papillose. Stem leaves erect when dry, spreading to wide-spreading when
moist, broadly ovate, 0.5--0.6 mm, abruptly acuminate; margins plane in the
acumen, mostly recurved proximally, rarely erect; costa ending near base of
the acumen, pellucid; cells quadrate to isodiametric, ca. 5--6 \um wide,
pluripapillose on both surfaces, with 3--5 papillae mostly around the
periphery of the lumina. Branch leaves erect when dry,
spreading to wide-spreading when moist, ovate, ca. 0.4 mm, gradually
broad-acuminate; margins plane to irregularly recurved; costa ca. 4/5\x leaf
length, pellucid proximally, obscure distally by overlying laminal cells;
cells quadrate to isodiametric, with 4--6 papillae mostly around the
periphery of the lumina, ca. 5--6 \um wide, the apical cell enlarged,
pellucid and smooth, pointed. Perichaetial leaves ca. 2.5 mm,
lanceolate, long-acuminate, plicate; costa subpercurrent; cells rectangular
to long-rectangular, smooth. Seta smooth, 1--1.5 cm; operculum
stoutly apiculate; segments not or narrowly perforate, cilia in groups of
1--2. Spores ca. 17 \um, finely
papillose. Dead branches and rocks, probably in
mostly exposed habitats; ca. 1200 m; Ariz.; Mexico; West Indies; Central
America; South America; e,s Africa. Rauiella praelonga, only occurring
rarely, in the southwestern part of our flora, differs from R. scita by the shorter acumen of the
stem leaves and the nonpapillose apical cell of the branch leaves. 2.
Rauiella scita (Palisot de Beauvois) Reimers,
Hedwigia 76: 287. 1937. E Hypnum
scitum Palisot de Beauvois, Prodr. Aethéog. 69. 1905; Rauia scita (Palisot de Beauvois) Stems
4--6 cm, ± regularly 1-pinnate, the branches 2--3 mm; paraphyllia mostly
filamentous, 1-seriate distally and 2(--3)-seriate proximally, weakly
branched, the cells oblate, papillose.
Stem leaves erect when dry,
erect-spreading when moist, broadly ovate to cordate-deltoid, 0.5--0.6 mm,
narrowly long-acuminate, not to moderately 2-plicate; margins plane distally,
loosely revolute proximally; costa ending near base of the acumen, pellucid;
cells rounded hexagonal, ca. 5--6 \um wide, pluripapillose on both surfaces,
with 3--6 papillae mostly around the periphery of the lumina. Branch
leaves appressed when dry, erect-spreading when moist, ovate, ca. 0.4 mm,
acute; margins plane to irregularly recurved; costa ca. 4/5\x leaf length,
pellucid below, obscure distally by overlying laminal cells; cells quadrate
to isodiametric, with 4--6 papillae mostly around the periphery of the
lumina, ca. 5--6 \um wide, the apical cell truncate, pluripapillose. Perichaetial
leaves ca. 1.5 mm, lanceolate, long-acuminate, erect, plicate; costa
subpercurrent; cells rectangular to long-rectangular, smooth. Seta
smooth, 1--2 cm; operculum obliquely stoutly short-rostrate; segments
narrowly perforate, cilia in groups of 2--3. Spores 10--13 \um, finely papillose. Capsules
mature fall--early winter. Bark, especially at bases of hardwoods, mature
forests; mostly moderate elevations; N.B., N.S., Ont., Que.; Conn., D.C.,
Del., Ind., Iowa, Ky., Maine, Mass., Md., Mich., N.C., N.H., N.J., N.Y.,
Ohio, Pa., R.I., Tenn., Va., Vt., Wis., W.Va. Rauiella scita is most likely confused
with a Leskea rather than the other
Thuidiaceae. However, the abundant paraphyllia and pluripapillose laminal
cells will immediately separate it from that genus. Like R. praelonga, the distal portion of the costa of the branch
leaves is covered with laminal cells, but in R. scita the apical cell of the branch leaves is truncate and
pluripapillose, unlike the smooth, pointed apical cell in R. praelonga. 4. THUIDIUM Bruch & Schimper in P. Bruch, W. P.
Schimper & Howard
A. Crum Plants
usually relatively large, in dull, green to yellow-brown mats. Stems creeping to arched-ascending,
2--3-pinnate, and usually frondose; paraphyllia abundant, especially on
stems, lanceolate or filiform-branched, papillose. Stem and branch leaves dimorphic. Stem leaves ovate, acuminate, generally +/- plicate at base;
margins usually revolute to the base of the acumen; margins
papillose-serrulate proximally, often serrulate distally by projecting cell
ends; costa ending before the apex; cells mostly +/- uniform throughout,
rounded- to oblong-hexagonal, thick-walled, coarsely 1-papillose at back (or
less commonly with low, forked papillae and appearing to be pluripapillose). Primary branch leaves often rather
similar to stem leaves but shorter. Secondary
and tertiary branch leaves much smaller, concave, ovate, acute, with
erect margins and usually ending in a truncate, pluripapillose cell; costa
shorter and weaker. Sexual condition
dioicous. Perichaetial leaves +/- elongate, erect, pale, lanceolate or
ovate- lanceolate,
long-acuminate, sometimes ciliate-margined; costa ending near the apex; cells
elongate, smooth or somewhat papillose. Seta
elongate, smooth or rarely papillose. Capsule inclined to horizontal, curved-cylindric, asymmetric,
smooth; annulus of 2--4 irregular layers of small cells, often tardily
deciduous; operculum conic or rostrate from a convex-conic base; stomata present
in the short neck; peristome teeth yellow, yellow-brown, or red-brown,
densely cross-striolate basally; endostome consisting of a high basal
membrane, keeled segments, and nodulose cilia in groups of 2--4. Calyptra cucullate, smooth, naked. Species 229 (4 in the flora); almost worldwide. SELECTED REFERENCES
Buck, W. R. and H. A. Crum. 1990. An
evaluation of familial limits among the genera traditionally aligned
with the Thuidiaceae and Leskeaceae. Contr. Univ. Mich. Herb. 17: 55--69.
Crum, H. & L. E. Anderson. 1981. Mosses of Thuidium
is characterized by abundant, 2--3-pinnate branching
paraphyllia; dioicous sexuality; dimorphic stem and branch leaves with short,
thick-walled cells that are papillose abaxially and nearly always
1-papillose; 3-celled axillary hairs with a single, short, brown basal cell
and two elongate, hyaline ones; terminal cells of branch leaves mostly
truncate and pluripapillose; and setae nearly always smooth . The genus is treated here in a sense
exclusive of Abietinella, Rauiella, and Cyrto-hypnum (cf. W. R. Buck and H. A. Crum 1990). Abietinella is relatively robust and
has 1-pinnate branching, abundant paraphyllia, dioicous inflorescences, leaf cells
1-papillose on both surfaces, and 3-4-celled, brown axillary hairs. Rauiella and Cyrto-hypnum are small plants with autoicous inflorescences,
paraphyllia short and not much branched, and leaf cells papillose on both
surfaces (most commonly pluripapillose). Rauiella
has 1-pinnate branching, 3-celled axillary hairs, leaves not incurved when
dry, strongly bulging leaf cells, and smooth setae. Cyrto-hypnum has mostly 2-pinnate branching, as well as 2-celled
axillary hairs, leaves incurved when dry, leaf cells plane, and setae often
roughened. 1. Branching mostly 3-pinnate; terminal cells of branch
leaves acute, smooth. . . . . 1. Thuidium
tamariscinum 1. Branching mostly 2-pinnate; terminal cells of branch
leaves truncate and pluripapillose. 2. Stem leaves incurved at base and
spreading at the tips when dry, distinctly plicate; costa nearly filling the
acumen; paraphylla papillose at cell ends. . . . 2. Thuidium recognitum 2. Stem leaves ± erect, less
markedly plicate; costa not filling the acumen; paraphylla papillose at cell
midpoints. 3. Stem leaves short-pointed, with
plane margins or sometimes revolute at base; cells with low, forked papillae
and appearing 1--3-papillose. . . . 3. Thuidium
allenii 3. Stem leaves acuminate, revolute-margined;
cells coarsely 1-papillose, sometimes forked but not appearing
pluripapillose. . . 4. Thuidium
delicatulum 1.
Thuidium tamariscinum (Hedwig) Bruch & Schimper in P.
Bruch, W. P. Schimper & W. Gümbel, Bryol. Eur. 5: 163 (fasc. 49--51 Mon.
7. 2, 3). 1852 Hypnum
tamariscinum Hedwig, Sp. Musc.
Frond., 261. 1801 Plants
vivid-green to yellowish. Stems up
to 15 cm, creeping or arched, (2--)3-pinnately branched and frondose;
paraphyllia papillose near the middle of cells. Stem leaves loosely erect when dry, erect-spreading when moist,
1.5--2 mm, broadly ovate, gradually to rather abruptly acuminate, plicate;
margins revolute to the base of the acumen, denticulate distally; costa
strong, ending near the leaf apex; cells rounded to elliptic, thick-walled,
13--20 x 8--10 \um, stoutly 1-papillose. Branch
leaves ending in a sharp, smooth cell. Primary branch leaves somewhat resembling stem leaves, ca. 0.8
mm, 2-plicate; margins somewhat recurved proximally; costa delicate, ending
before the apex. Secondary and
tertiary branch leaves to about 0.4 mm, ovate, acute, with erect margins
and a weak costa ending near leaf middle; cells stoutly 1-papillose, the
papillae +/- curved. Perichaetial leaves pale yellow, ca.
3 mm, oblong-lanceolate, long-subulate, serrate distally, long-ciliate
proximally; costa ending in the subula. Seta
25--45 mm. Capsule 3--4 mm;
operculum 1.8 mm, obliquely long-rostrate; cilia of endostome 3--4. Spores ca. 12 \um, smooth or very
finely papillose. Moist soil or soil over rocks, sometimes in stream beds;
Nfld.; West Indies ( Thuidium
tamariscinum is a woodland species of relatively large and elegantly
frondose plants. The branching is typically tripinnate, but it is sometimes bipinnate
or imperfectly tripinnate. The leaf cells are stoutly 1-papillose, except
that the terminal cell of branch leaves is sharp and smooth. 2.
Thuidium recognitum (Hedwig) Lindberg, Not. Sällsk. F.
Fl. Förh. 13: 416. 1874 Hypnum
recognitum J. Hedwig, Sp. Musc.
Frond., 261. 1801; H. protensum Michaux Plants
in light-green to yellowish or brownish mats. Stems 4--9 cm, 2-pinnately branched and frondose; paraphyllia
papillose mostly at or near cell ends, the terminal cell usually with 2--3
large, divergent papillae. Stem leaves
spreading with reflexed tips when moist, +/-
incurved at base and wide-spreading at the tips when dry, ca. 1 mm, broadly
ovate, abruptly acuminate, distinctly plicate; margins papillose-serrulate,
more strongly so in the acumen, plane at least when removed from the stem,
rarely +/- revolute proximally;
costa broadened distally, nearly filling the acumen but disappearing near the
apex; distal cells to 24 x 8--10 \um, rhombic to oblong-rhombic, coarsely
1-papillose. Branch leaves ending in a truncate,
pluripapillose cell. Primary branch
leaves smaller, with a shorter costa. Secondary branch leaveses ca. 0.2 mm, erect-spreading when wet or
dry, ovate, acute; costa 1/3--2/3 the leaf length; distal cells rhombic,
8--12 x 8--10 \um, thick-walled, stoutly 1-papillose, the papillae somewhat
curved, the terminal cell truncate, pluripapillose. Perichaetial leaves up to 4.2 mm; margins denticulate, sometimes
dentate or notched near base of the acumen. Seta 20--42 mm, reddish. Capsule
2--3.5 mm; operculum 0.7--1 mm, high-conic or bluntly short-rostrate; cilia
of endostome in 2's and 3's. Spores
11--16 \um, smooth or nearly so. Moist soil, humus, or rocks, infrequently on logs or bark
at base of trees, calcareous habitats, woodlands, often in clearings and
timber trails; Alta., B.C., Man.,
Ont., Que.; Alaska, D.C., Ga., Ill., Ind., Iowa, Kans., Ky., Maine, Md.,
Mass., Mich., Minn., Mo., N.H., N.J., N.Y., Ohio, Okla., Oreg., Pa., Tenn.,
Vt., Va., Wash., W.Va., Wis.; Europe. The specific epithet of Thuidium recognitum was applied by Hedwig because he recognized
the many differences that separate the species from T. delicatulum: stem
leaves that are shortly, broadly, and abruptly acuminate and plicate, with margins
plane or nearly so; and costa spreading out and nearly filling the acumen;
tips of the stem leaves wide-spreading from an arched and incurved base;
perichaetial leaves eciliate; paraphyllia papillose at or near cell ends; and
operculum not or only shortly and bluntly rostrate. 3.
Thuidium allenii Austin, Bull. Torrey Bot. Club 7:
16. 1880 Thuidium
glaucinum var. ludovicianum Cardot, Bryologist 8: 51. 1905 Plants
slender, light-green or yellow. Stems
loosely branched, irregularly to regularly 2-pinnate, loosely foliate;
paraphyllia papillose at the side of cells. Stem leaves
erect-spreading when moist, loosely erect or slightly incurved when dry, ca.
1 mm, ovate, broadly short-pointed, acute, not or somewhat plicate; margins +/- denticulate all around because of
projecting cell ends, plane or sometimes reflexed near the base; costa ending
near the apex; distal cells hexagonal, mostly about 7 \um, moderately
thick-walled, 1-papillose, the papillae low and usually forked thus appearing
to be 1--3 per cell. Branch leaves ending in a truncate,
pluripapillose cell. Primary branch
leaves strongly incurved, 0.5--0.6 mm, ovate, acute; costa 8/10--9/10 the
leaf length; cells 8--9 \um. Secondary
branch leaves similar to those of primary branches but smaller, 0.2--0.4
mm, acute or obtuse; costa 7/10--8/10 the leaf length. Perichaetial leaves
eciliate. Seta 25--30 mm. Capsule ca. 2.5 mm; operculum bluntly
conic. Spores not seen. Uncommon, soil, logs, exposed roots, tree bases in
swamps, often just above water line, characteristically in Taxodium-Nyssa-Chamaecyparis
swamps; Ala., Ark., Conn., Del., Fla., Ga., Ind., La., Md., Mass., Miss.,
N.Y., N.C., Pa., Tenn., Tex., Va. Thuidium
allenii is essentially restricted
to the 4.
Thuidium delicatulum (Hedwig) Bruch & Schimper in P.
Bruch, W. P. Schimper & W. Gümbel, Bryol. Eur. 5: 164, fig. 484. (fasc.
49--51 Mon. 8, fig. 4). 1852 Hypnum
delicatulum Hedwig, Sp. Musc.
Frond., 160. 1801; Thuidium recognitum var. delicatulum (Hedwig) Warnstorf Plants
green or yellowish. Stems 3--8 cm,
2--3-pinnate, ± frondose; paraphyllia mostly papillose at cell points. Stem leaves erect-spreading when
moist, appressed when dry, 0.6--1.5 mm, triangular-ovate, gradually or
abruptly narrowed to a broad acumen or ending in a hyaline point of 2--8
1-seriate cells, not plicate; margins papillose-serrulate, revolute from the
insertion to the acumen; costa ending well before the apex; distal cells
irregularly oblong-hexagonal, 6--10 x 8--12 \um, rhombic, moderately
thick-walled, coarsely 1-papillose. Branch
leaves ending in a truncate,
pluripapillose cell. Primary branch
leaves erect-spreading, to 0.5 mm, acute; costa 1/2--2/3 the leaf length;
cells 6--8 x 8--12 \um, rhombic, coarsely 1-papillose, the papillae curved,
sometimes forked. Secondary branch
leaves similar but smaller. Perichaetial
leaves to 5 mm, eciliate to ciliate proximally, often denticulate
distally. Seta 15--45 mm. Capsule 1.8--4 mm; operculum 0.7--2
mm, long-rostrate; cilia of endostome in 2's and 3's. Spores 12--24 \um, smooth. Common in woodlands, soil, humus, rocks, logs, and
stumps, infrequently bases of trees; Alta., B.C., Man., N.B., Nfld., N.S.,
Ont., Que., Yukon; Ala., Alaska, Ariz., Ark., Conn., Del., D.C., Fla., Ga.,
Ill., Ind., Iowa, Kans., Ky, La., Maine, Md., Mass., Mich., Minn., Mo., N.H.,
N.J., N.Mex., N.Y., N.C., Ohio, Okla., Pa., R.I., S.C., S.Dak., Tenn., Tex.,
Vt., Va., W.Va., Wis.; Mexico; Central America; n South America; Europe; Asia. Thuidium
delicatulum, needlessly confused with T. recognitum, can be
recognized by stem leaves erect or erect-spreading when moist, not plicate
and rather gradually acuminate, with margins recurved to the base of the acumen
and costa ending well before the apex. The paraphyllia have small papillae
along the cell midpoints. The leaf cells are stoutly 1-papillose, but often,
especially in the South, the papillae are forked. The operculum is
long-rostrate . 1. Stem leaves merely acute; perichaetial leaves ciliate.
. . 4a. Thuidium delicatulum var. delicatulum 1. Stem leaf apex extending by 2--8 hyaline cells in a
single row; perichaetial leaves not ciliate or rarely ± so. . . . 4b. Thuidium delicatulum var. radicans 4a.
Thuidium delicatulum var. delicatulum Stem
leaves somewhat shorter, 0.6--1.4 mm, averaging 0.9 mm, acute
and not ending in a hyaline point 1-seriate cells. Perichaetial leaves ciliate proximally, denticulate distally. Common in woodlands, soil, humus, rocks, logs, and
stumps, infrequently bases of trees; N.B., N.S., Ont., Que.; Ala., Ariz.,
Ark., Conn., Del., D.C., Fla., Ga., Ill., Ind., Iowa, Kans., Ky, La., Md.,
Mass., Mich., Minn., Mo., N.H., N.J., N.Y., N.C., Ohio, Okla., Pa., R.I.,
S.C., Tenn., Tex., Vt., Va., W.Va., Wis.; Mexico; Central America; n South
America; Europe. Thuidium
delicatulum var. delicatulum is
apparently more often in acid substrates than is T. recognitum and T. delicatulum
var. radicans. The perichaetial leaves are copiously ciliate. 4b.
Thuidium delicatulum var. radicans (Kindberg) H. A. Crum, Steere & L. E. Anderson,
Bryologist 68: 434. 1965 (1966) Thuidium
recognitum var. radicans Kindberg, Rev. Bryol. 19:
103. 1892; T. philibertii Limpricht; T. recognitum
ssp. philibertii (Limpricht) Stem
leaves somewhat longer, 1--1.5 mm, averaging 1.2 mm, ending in
a hyaline point of 2--8 1-seriate cells. Perichaetial
leaves usually not ciliate but sometimes sparsely so, rarely copiously
ciliate. Apparently a calciphile, soil, humus, rocks, or logs in
swampy places; Alta., B.C., Ont., Man., Nfld., N.S., Yukon; Alaska, Iowa,
Maine, Mich., Minn., N.J., N.Mex., N.Y., S.C., S.Dak., Va., Wis.; Mexico;
South America (Peru); Europe; Asia. The var. radicans intergrades
with the var. delicatulum. The
annulus of the var. radicans is of
2--3 rows of cells scarcely differentiated from suboral cells and is
sometimes of aid in distinguishing this variety from var. delicatulum.
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