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BFNA Title: Splachnaceae |
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(for illustrations CLICK HERE) XX. SPLACHNACEAE
Greville & Arnott Paul
Marino Plants
small to medium-sized, green or yellowish sometimes brownish. Stems acrocarpous, forked by
subfloral innovations; minute claviform filaments in axils of distal leaves
common. Stem leaves soft, homogeneous along stem or larger and
crowded at stem apex, mostly broad, ovate-lanceolate to oblong or spatulate,
entire to dentate, sometimes bordered; costa single, strong, mostly ending
before apex; cells rhomboidal, large, oblong or oblong-hexagonal in distal
portion of leaf; basal cells oblong. Perichaetial
leaves like stem leaves but often larger. Seta thin or thick, usually elongate, erect. Capsule mostly erect, exerted and symmetrical; often with an
elongate neck or with a wide inflated or long and narrow apophysis proximal
to the urn; annulus mostly lacking; operculum convex to conic; stomata
abundant, with 2 guard cells; peristome mostly present, single, teeth 16,
densely and finely papillose, approximate in 2’s and 4’s, entire or rarely forked;
columella sometimes exerted. Calyptra
mitrate, rarely cucullate, smooth or sometimes hairy. Genera 6, species 73 (5 genera, 17 species in the flora):
tropical to sub-polar regions worldwide. Almost half the species of Splachnaceae possess three
noteworthy ecological features: First,
their gametophytes are “coprophilous,” growing on feces and occasionally old
bones and other animal matter. Second, their spores are small and sticky
making them suitable for insect dispersal.
Third, the sporophytes of all entomophilous Splachnaceae examined to
date produce complex, species-specific odors that are thought to promote the
attraction of flies (Diptera). Although
the leaves are soft textured and similar in shape to those of the
Funariaceae, recent phylogenetic studies (C. J. Cox and T. A. Hedderson 1999;
C. J. Cox et al. 2000) suggest that the Splachnaceae are closely related to
the Meesiaceae rather than to the Funariales as previously proposed (V. F.
Brotherus, 1924; D. H. Vitt, 1984).
Like many Meesiaceae most Splachnaceae grow in moist habitats such as
peatlands in temperate and boreal forests.
Splachnaceae differ from the Meesiaceae in the structure of the
capsule, which in Splachnaceae is erect with a mitrate calyptra whereas in
the Meesiaceae the capsule is curved with a cucullate calyptra (B. Goffinet
et al. 2004). SELECTED REFERENCES
Goffinet, B., A. J. Shaw, and C. Cox. 2004. Phylogenetic inferences in
the dung moss family Splachnaceae from analysis of cpDNA sequence data and
implications for the evolution of entomophily. Am. J. Bot. 91: 748--759. Koponen, A. 1978. The peristome and
spores in Splachnaceae and their evolutionary and systematic significance.
Bryophyt. Biblioth. 13: 535--567.
Koponen, A. 1982. The classification of the Splachnaceae. Nova Hedw.
Beih. 71: 237--245. Marino, P. C., R. Raguso, and B. Goffinet. 2008.
The ecology and evolution of fly dispersed dung mosses (Family
Splachnaceae): manipulating insect behaviour through odor and visual cues.
Symbiosis 47: 61-76 1. Capsule
cleistocarpous; coprophilous. 2. Peristome not differentiated;
apophysis absent; capsule dark red...........1. Voitia, p. xx 2. Peristome differentiated,
apophysis present; capsule yellowish………………………………………...……….4. Tetraplodon (in part), p. xx 1. Capsule not
cleistocaropous, color various; apophysis clearly differentiated;
coprophilous or not. 3.
Apophysis short to elongate or narrowly pyriform similar in color or
darker than the urn; calyptra either constricted or not constricted distal to
the base; coprophilous or not. 4.
Apophysis short to elongate, as wide as or narrower than the urn and
similar to it in color; calyptra constricted distal to the base; peristome
teeth 16; calyptra constricted distal to the base; not coprophilous (North
American species) …………………………………………………………………....2. Tayloria, p. xx 4.
Apophysis narrowly pyriform, somewhat broader and usually longer than
the urn, the same color or darker than the urn; peristome teeth single with 8
well-developed exostome teeth, calyptra not constricted distal to the base;
coprophilous or grows on old bones …………….……………………..….3. Tetraplodon (in part), p. xx 3.
Apophysis clearly differentiated, elongate and somewhat broader to
much broader than the urn; calyptra not constricted distal to the base;
coprophilous or grows on old bones. 5.
Apophysis globose to turbinate, sometimes becoming umbrella-like,
greatly differentiated in size, shape, color, and texture; seta smooth and
slender; peristome teeth double and fused in pairs, chambered …….....4. Splachnum, p. xx 5.
Apophysis rounded, not or only slightly wider than the urn; seta
hyaline and slender; peristome single with 8--12 rudimentary outer teeth, not
chambered………………………………………………….…….5. Aplodon,
p. xx 1. VOITIA Hornschuch, De Voitia et Systylio 5, plate
1. 1818 * [For Johann
Gottlob Wilhelm Voit, 1787--1813, German bryologist] Plants in tufts 2--6 cm, light green to yellow
green, densely radiculose proximally the distal 5 mm. Stems 1--6 cm, often
branched. Stem leaves slightly contorted when
dry erect-spreading when wet, lanceolate, long-acuminate, margins entire, ±
incurved; costa excurrent; distal leaf cells rectangular, 35--55 x 20--25
\um; basal cells 55--95 \um in length. Specialized
asexual reproduction none. Sexual
condition autoicous; perigonia
and perichaetia terminal on separate branches. Seta 1--3 cm, yellowish
to dark red, darkest in the distal portion, straight or slightly twisted
basally, usually strongly twisted immediately proximal to the capsule;
apophysis absent. Capsule cleistocarpous, dark red, symmetric or
slightly curved; annulus and peristome lacking; operculum not differentiated.
Calyptra enlarged and
completely covering the capsule, sometimes split at the base. Spores
7--18 \um, smooth. Species 2 (2 in the flora): circumboreal and mountainous
regions of Europe and western Voitia
species are coprophilic, but, as they are cleistocarpous, do not have the
spores dispersed by flies (Diptera).
Spores are released by the disintegration of the sporangial wall. The dark red capsule is quite distinct and
the two species are easily distinguished by the shape of the capsule. SELECTED REFERENCES
Goffinet, B. and A. J. Shaw. 2002. Independent origins of cleistocarpy
in the Splachnaceae: analyses of cpDNA sequences reveals polyphyly of the
Voitioideae. Syst. Bot. 27: 203--208.
Goffinet, B., A. J. Shaw, and C. Cox. 2004. Phylogenetic inferences in
the dung moss family Splachnaceae from analysis of cpDNA sequence data and
implications for the evolution of entomophily. Am. J. Bot. 91: 748--759. 1. Capsule
oblong-ovate, narrowing gradually to the seta without a ridge formation; seta
usually strongly twisted immediately proximal to the capsule
………...………………………..1. Voitia nivalis 1. Capsule
ovate-globose, base abruptly narrowing to the seta forming a definite ridge
along the base; seta not twisted proximal to the capsule................................................................2. Voitia hyperborea 1. Voitia nivalis
Hornschuch, Voitia et Systylio, 5, plate 1. 1818 Plants
3--6 cm. Leaves , 3--5.5 mm, ovate-lanceolate, long-acuminate,
concave. . Capsule dark red, oblong-ovate, erect and symmetric or
slightly curved, 3 mm, broader at the base, drawn out to an oblique point
distally about 1 mm, narrowing gradually to seta without ridge; outer wall of
capsule smooth or nearly so. Calyptra cucullate, constricted and
sometimes split at the base, often sliding down on the seta. Spores
7--11 \um,. Alta., N.W.T., 2. Voitia hyperborea Greville & Arnott,
Tent. Meth. Musc. 4: 149, plate 7, figs. 19--22. 1822 Voitia
nivalis var. hyperborea (Greville & Arnott)
Schimper Plants
in tufts 2--5 cm, , tufts deeper and thicker than in Voitia nivalis. Leaves lanceolate, long-acuminate,
narrower with a much longer acumination than Voitia nivalis. Capsule
dark red, ovate-globose, broadly conic, symmetric and almost round, very
abruptly narrowing to the seta forming a definite ridge along the base; outer
wall of capsule irregularly but strongly wrinkled, pustulate; or even ridged.
Calyptra with a hyaline collar-like base clasping the seta proximal to
the capsule, expansion of the capsule splitting the calyptra from the base
nearly to the tip. Spores 10--18 \um. The principle difference between Voitia hyperborea
and V. nivalis is the shape of the
capsule, it being oblong-ovate in V. nivalis and ovate-globose in V.
hyperborea. The base of the
capsule in V. hyperborea is abruptly much narrowed to the seta,
forming a definite ridge along the base.
In V. nivalis, the more elongate capsule narrows gradually to
the seta without any ridge formation.
Both species grow on herbivore droppings and, because the capsules are
cleistocarpous, it is unclear how they are dispersed. W. C. Steere (1974) suggested that spore
dispersal to fresh dung may occur when the sporophytes are ingested by
caribou or musk-oxen, and the spores subsequently dropped by them at some
other location. The species do not,
for the most part, have overlapping distributions. Voitia
nivalis has a more southerly distribution than V. hyperborea and, in North America, is mainly a species
of the Greenland; N.W.T., SELECTED REFERENCES Steere, W. C. 1974. The status and geographical distribution of Voitia hyperborea in 2. TAYLORIA Hooker, J. Sci. and Arts ( Hookeria
Schwägr. Sp. Musc. Frond., Suppl.
1(2): 340. 1816 (replaced name) Plants loosely or densely tufted green or
yellowish sometimes brownish green. Stems
0.5--3 cm, rarely to 6 cm,
sometimes branched. Stem leaves erect to wide-spreading,
imbricate to somewhat contorted when dry, small proximally, larger and more
crowded distally, margins toothed to entire, oblong-lanceolate to obovate,
blunt or acute to acuminate; costa ending well before the apex to excurrent;
cells large oblong-hexagonal distally, longer proximally. Sexual condition autoicous, rarely synoicous, often
dioicous. Seta 1--4 cm,
apophysis narrower than the capsule and usually colored the same and
generally tapered and constricted when dry. Capsule almost always
exerted, erect, ovoid to oblong-cylindric or claviform; peristome teeth 16,
erect or reflexed, equally spaced, sometimes fused in pairs or rarely
becoming split, consisting of two layers, inserted at or proximal to the
mouth of the capsule; operculum convex or conic to hemispheric. Calyptra short, constricted at the base, mitrate,
smooth or rough, naked or hairy. Spores 15--45 \um, smooth or
slightly roughened. Species 45 (6 in the flora): Tropical to subarctic
worldwide. Tayloria
species are the only Splachnaceae that include both
anemophilous and entomophilous taxa, and they are also the most polymorphic
morphologically. All North American Tayloria species are anemophilous and
none are coprophilous although they often grow on nutrient-enriched
substrates. SELECTED
REFERENCE LaFarge-England and C., D.
H. Vitt 1985. A Taxonomic Study of Tayloria hornschuchii and T.
froelichiana in 1. Leaves lingulate, obtuse to broadly rounded at the
apex, entire. 2. Preperistome present; seta not
noticeably stout, 25--40 mm……………………1. Tayloria
lingulata 2. Preperistome never present; seta
stout, less than 10 mm. 3. Leaves ± uniform along the stem,
lax, slightly contorted when dry; columella included; apophysis about as long
as the urn; peristome teeth lanceolate, persistent, yellow to tan; operculum
deciduous ……………………………………….………..……3. Tayloria
froelichiana 3 Leaves clustered at stem and
branch tips; concave, turgid when dry, columella exserted; apophysis longer
than the urn; peristome teeth truncate, fragile, red to red-brown; operculum
systylious…………4. Tayloria hornschuchii 1. Leaves obovate to oblanceolate, acute or acuminate,
not broadly rounded at the apex, serrate distally. 4. Peristome teeth not divided or
recurved…………………..…………2. Tayloria
serrata 4. Peristome teeth split at base to
form long narrow segments tightly revolute inside the urn when moist,
reflexed and somewhat contorted when dry. 5. Robust plants up to 4 cm, leaves
rounded-obtuse with a short acumen, 2--5 mm; seta 10--30 mm; urn of capsule
2--3.5 mm, shrinking on drying to expose the columella; apophysis; operculum
high-conic, 1--1.25 mm ..…………5. Tayloria
splachnoides 5. Slender plants rarely more than 1
cm, leaves long-acuminate, 2--4 mm; seta 6--15 mm; urn of capsule 0.8--2 mm,
not shrinking greatly when drying and either not or barely exposing the
columella; operculum low-conic 0.5--0.6 mm…....6. Tayloria acuminata 1. Tayloria
lingulata (Dickson) Lindberg., Musci Scand. 19. 1879 Splachnum
lingulatum Dickson, Fasc. Pl. Crypt. Brit. 4: 4, plate 19, fig.
6. 1801; Weissia turbinata Drummond Plants in tufts, green distally, brown
proximally, 3--4 cm, sometimes branched with red to purple rhizoids; radicles
not bearing brood bodies. Leaves somewhat contorted when dry,
erect-spreading when wet, 2--4 x 1--1.4 mm, lingulate, often broadest beyond
the middle, apex obtuse to rounded; margins plane or recurved proximally,
entire or nearly so, sometimes with blunt teeth; cost ending before the apex.
Specialized asexual reproduction none. Sexual condition autoicous or sometimes apparently
dioicous, the perigonia terminal. Seta 1.5--4 cm, slender, flexuose,
bright red. Capsule ± ovoid, 1.5--2.5 mm including the apophysis,
wide-mouthed when empty; apophysis about as long as the urn, stomata
numerous; columella included; annulus none; operculum convex (nearly flat
when dry), bluntly and obliquely umbonate to short-rostrate; peristome teeth
inserted somewhat proximal to the mouth, erect, evenly spaced, broadly
lanceolate, deep-yellow or orange-yellow, densely and finely papillose, with
small, brownish, papillose preperistome fragments at the base. Calyptra
smooth, naked, constricted proximally. Spores smooth, 26--45 \um. Damp soil or humus or on mud rich in organic matter such
as insect exuviae or bird droppings;
Greenland; Alta, B.C., Man., Nfld. and Tayloria
lingulata is distinguished from the other species by tongue-shaped
leaves. Also, the setae are relatively
slender. The 16 peristome teeth are separate and
erect and the short, broad urn is about as long as the apophysis. 2.
Tayloria serrata
(Hedwig) Bruch & Schimper in P. Bruch,
W. P. Schimper & W. Gümbel, Bryol. Eur. 3: 204. 284 (fasc. 23--24 Monogr.
6. 1.) 1844 Splachnum
serratum Hedwig, Sp. Musc. Frond., 53. 1801 Plants in loose, clear-green, ± radiculose
tufts about 0.5--3 cm; small, dark, red-brown, narrowly ellipsoidal brood
bodies sometimes produced on the radicles; infrequently branched. Leaves
soft, loosely erect, somewhat contorted when dry, imbricate to spreading when
wet, 2--5 x 1--1.5 mm, obovate to oblong-obovate, tapering proximally,
acuminate or acute, sometimes obtuse, apex often reflexed, margins plane and
toothed distally, entire and usually recurved proximally; costa ending before
the apex. Specialized asexual
reproduction none. Sexual
condition autoicous or
sometimes apparently dioicous; paraphyses lacking in the perichaetia, those
of the perigonia stoutly filiform. Seta 1--3 cm, yellow to dark red or
brown, flexuose, ± stout. Capsule cylindric, 2.5--5 mm including the
apophysis; urn oblong-cylindric, 1--2 mm when dry, brown or red-brown,
gradually narrowed to a somewhat darker apophysis about as long as the urn or
up to twice as long (1--2 mm); columella somewhat exserted; annulus none;
operculum hemispheric, sometimes bluntly apiculate or short-rostrate;
peristome teeth inserted proximal to the mouth, in pairs or separate, dark
red or red-brown, reflexed when dry, densely papillose. Calyptra
smooth, naked, constricted proximally. Spores light-yellow, smooth,
round 9--12 \um. Organic material of animal origin and also on humus;
Alta, B.C., N.B., Ont., Nfld. and Labr. (Labr.), Que.; Alaska, Maine, N.Y., Ore., Vt., Wash.;
n, c Europe. Tayloria
serrata can be distinguished from other species of the genus in
North American by several leaf characters, including obovate to
oblong-obovate leaves with an acuminate or acute apex, and unbordered,
serrate margins. In addition, the
peristome teeth are not divided or recurved as in T. acuminata and T.
splachnoides. 3. Tayloria froelichiana
(Hedwig) Mitten, J. Linn. Soc. London (Suppl.) 1: 57. 1859 Splachnum
froelichianum Hedwig, Sp. Musc. Frond., 52. 1801 Plants small in loose, shiny tufts, yellow-green
10--20 mm high, unbranched. Leaves dense, imbricate, slightly keeled,
obovate to oblong, bluntly acute, obtuse, 1--2.8 x 0.8--1.2 mm, ± entire;
margins plane or slightly revolute before the middle; proximal leaves smaller
and somewhat radiculose; costa stout, subpercurrent, scarcely narrowed
distally, ending 6 cells or less
proximal to the apex. Specialized
asexual reproduction none. Sexual
condition cladautoicous or synoicous. Seta twisted, stout, 1 cm;
with a long, slender, dark-orange apophysis, 1 mm. Capsule pyriform,
conic, or globose-conic, symmetric, when old consisting of a wide reddish
mouth and a constriction from which the apophysis tapers to the seta;
apophysis usually of a lighter color and much narrower than the urn, 1--1.5
mm; operculum flat to conic, often with a blunt oblique tip, deciduous;
peristome teeth yellow, in pairs, lanceolate, acute to obtuse, erect to
slightly reflexed when dry; columella included. Calyptra mitrate, constricted at base, split singly when mature. Spores
27--48 \um coarsely papillose. Mesic semi-disturbed sites; higher latitudes and
elevations; Alta, B.C., Tayloria
froelichiana can be distinguished most readily by several sporophytic
traits that help distinguish this species from T. horschuchii: apophysis
twisted and conic, tapering quickly to the seta; operculum deciduous, with an
immersed columella; exostome reticulate to striate, exostome teeth paired, 8
in number; sexual condition autoicous or synoicous. The main gametophytic difference between T. froelichiana and the
gametophytically similar T.
hornschuchii are the lack of axillary propagula in T. froelichiana and the longer basal (125--195 \um) and apical
(75--115 \um) leaf cells. SELECTED
REFERENCES LaFarge-England, C. and
D. H. Vitt 1985. A Taxonomic Study of Tayloria hornschuchii and T. froelichiana in 4. Tayloria hornschuchii
(Greville & Arnott) Brotherus, A.
Engel and K. Prantl, Natur. Pflanzenfam. 1(3): 502. 1903; Dissodon
hornschuchii Grevile & Arnott,
Tent. Meth. Musc. 5(2): 468.
1826 Plants
in short, shiny tufts, 3--7 mm high, yellow-green distally, brown proximally;
unbranched or with 6 or less subterminal inovations. Leaves crowded at
the apex of stems and innovations, imbricate, lingulate and obtuse, strongly
keeled, stiffly erect imbricate when dry, entire, 1--2.5 x 0.7--1.2 mm
margins plane or slightly revolute in the proximal portion, basal leaves
smaller; costa often brown to red-brown, ending 3--4 cells before the apex. Specialized asexual reproduction propagula
6 cells or fewer, red to red-brown at maturity abundant to rare predominately
in proximal leaf axils. Sexual
condition dioicous. Seta stout, straw-colored, not or scarcely twisted,
mostly less than 1 cm. Capsule oblong to cylindric, symmetric, about 1
x 1 mm, quadrate at the middle of the capsule, exserted; apophysis about 1.5
mm, slender and tapering, as wide as the urn, straw-colored to dark red-brown
when mature; calyptra constricted at the base, 2 mm; operculum remaining
attached to the exserted columella, flat and with an obtuse, usually long
apiculus; peristome orange-red, erect, inserted at the mouth, 16 teeth
partially split into 32, truncate to obtuse, densely papillose. Calyptrae mitrate, constricted at
base, splitting by ± 2--3 slits, apiculate tip, naked. Spores 30--40 \um papillose.
Humus and humic soil and more exposed soils; Alta,
Nunavut, Yukon; Alaska, Colo., Wyo.; c, n Europe. Tayloria
hornschuchii is distinguished by the
well-developed, often wrinkled apophysis, gradually tapering to the seta;
operculum systylious, with exserted columella; exostome papillose of 16
unpaired teeth; and sexual condition dioicous. The main gametophytic difference between T. hornschuchii and the similar T. froelichiana are the presence of
axillary propagula in T. hornschuchii
and the shorter basal (75--115 \um) and apical (40--65 \um) leaf cells. SELECTED
REFERENCES LaFarge-England, C. and
D. H. Vitt 1985. A Taxonomic Study of Tayloria hornschuchii and T. froelichiana in 5.
Tayloria splachnoides (Schleicher ex Schwägrichen)
Hooker, J. Sci. and Arts ( Hookeria
splachnoides Schleicher ex Schwägrichen, Sp. Musc. Frond., Suppl. 2:
340, plate 100. 1816 Plants
in lax tufts, 1--3 cm, clear green distally; sometimes branched. Leaves slightly
crisped, lingulate to obovate, blunt or apiculate, strongly serrate beyond the
middle, proximal portion smooth and recurved, ca. 3 x 0.7--1 mm, costa ending
4--5 cells before the apex. Specialized
asexual reproduction none. Sexual
condition synoicous or
autoicous. Seta 1.5--3 cm, slightly twisted, yellow, becoming red with
age. Capsule before maturity elongate-ovoid, cylindric and contracted
very suddenly to the apophysis, tough-walled, pale-brown, somewhat darker at
the mouth and sometimes slightly darker at the neck; urn 1--1.5 mm when dry,
2--3.5 mm when moist; columella short- to long-exerted when dry; annulus
none; operculum high-conic and acuminate, 1--1.25 mm, peristome inserted
proximal to the mouth, the 16 teeth split into 32 linear-lanceolate filaments
tightly rolled inside the urn and usually hidden when moist but loosely
rolled and reflexed when dry red, densely and minutely papillose, 0.6--1 mm;
apophysis elongate, to 3 mm, much more slender than the capsule, colored as
the capsule or somewhat more pale. Calyptra constricted below the
middle, smooth and naked. Spores 14--16 \um, slightly roughened. Humus covered rocks, decaying logs or soil; B.C., Nfld. and Labr. (Nfld.), Tayloria
splachnoides is uncommon and closely resembles T. acuminata but differs from this species by leaves blunt or
apiculate, operculum high-conic and acuminate, and columella exserted. Both T.
splachnoides and T. acuminata
have long, bifid red or red-brown peristome teeth that are reflexed and
tightly rolled inside the urn when dry.
6.
Tayloria acuminata Hornschuch, Flora 8: 78. 1825 Tayloria
splachnoides var. acuminata
(Hornschuch) Huebener, Muscol. Germ., 96. 1833; T.
splachnoides subsp. acuminata
(Hornschuch) Kindberg;
T. serrata subsp. acuminata
(Hornschuch) J. J. Amann Plants
bright green, loosely tufted, 5--10 mm, radiculose, radicles violet becoming
purple to dark-red with age; sometimes branched. Leaves erect and
moderately contorted when dry, erect-spreading when moist, ovate-lanceolate,
acuminate, margin in distal portion irregularly serrate, proximally smooth
and recurved; costa yellow-green to reddish, tapered, ending 5--9 cells
before the apex. Specialized asexual
reproduction small, clavate, papillose brood bodies 2--4 cells in length
born on radicles. Sexual condition autoicous. Seta 6--15 mm, slightly twisted when
dry, yellow, becoming red with age. Capsule suberect or often slightly
asymmetric, cylindrical, wide-mouthed and abruptly contracted to the slender
neck when dry, less abruptly narrowed when moist, tough-walled, pale brown,
sometimes slightly darker at the neck, varying in length, 0.5--1.5 mm when
dry, 0.8--2 mm when moist; apophysis gradually narrowed towards seta;
columella not exserted when dry; annulus none; operculum 0.5--0.6 mm,
short-conic with a blunt, often slightly oblique point; peristome inserted
proximal to the mouth, the 16 teeth 2-fid nearly to the base to form long,
linear-lanceolate divisions tightly rolled inside the urn and usually hidden
when moist but loosely rolled and ± reflexed when dry, red-brown, minutely
papillose. Calyptra constricted proximally to the middle, smooth and
naked. Spores 15--18 \um finely papillose. Damp places, humus, rotten logs, rock; Alta, B.C., Man, N.W.T., Nunavut, Ont.,
Que., Yukon; Alaska, Idaho, Mont., Nev., Wash., Wyo.; n, c Europe, n, c Asia.
In comparison with Tayloria
splachnoides, which has a close resemblance, T. acuminata has more slender gametophytes and sporophytes. The leaves are also more pointed, the
columella barely exerted upon drying of the capsule, and the operculum
low-conic 3.
TETRAPLODON Bruch & Schimper in P. Bruch, W. P. Schimper & Plants in dense cushions, bright green to
yellow-green distally, often brown and matted with rhizoids proximally. Stem 0.5--3(--8) cm, erect, Stem leaves slender
to oblong-lanceolate, or obovate and acuminate, margins toothed or entire;
costa usually ending in the subula; cells shortly rectangular to
oblong-hexagonal. Sexual condition
autoicous or rarely dioicous. Seta
0.3--5 cm, stout, erect; apophysis narrower to wider than the urn and usually
colored the same and wrinkled when dry. Capsule symmetric, cylindric
to ovoid; annulus mostly lacking; peristome teeth 16, at first ± coherent in
4’s, later in 2’s, composed of two layers of cells; in most species reflexed
when dry; inflexed when moist; operculum hemispheric to bluntly conic. Calyptra
short, conic-mitrate or
cucullate, not constricted distal to the base, smooth and naked. Spores
small, 8--12 \um, smooth. Species 10 (5 in the flora): alpine and subalpine and
temperate to subarctic worldwide. Tetraplodon
species are entomophilous and coprophilous although
apparently restricted to the dung of carnivores, bones, and owl pellets. The apophysis is well-developed and
elongate, and the peristome teeth are joined in 4’s when young but in 2’s as
they age. SELECTED REFERENCES
Goffinet, B., A. J. Shaw, and C. Cox. 2004. Phylogenetic inferences in
the dung moss family Splachnaceae from analysis of cpDNA sequence data and
implications for the evolution of entomophily. Amer. J. Bot. 91:
748--759. Marino, P. C. 1988. The
North American distribution of the circumboreal species of Splachnum and Tetraplodon. Bryologist 91: 161--166. Marino, P. C. 1997. Competition,
dispersal and coexistence of Splachnaceae in patchy habitats. In: R. E.
Longton, ed. Advances in Bryology. 1. Leaves long-lanceolate with large teeth. …………..…..……2.
Tetraplodon angustatus 1. Leaves ovate or obovate-lanceolate, entire or nearly
so. 2. Capsule cleistocarpous; capsule
yellowish, apophysis narrower than the urn ……… 1. Tetraplodon paradoxus 2. Capsule not cleistocarpous;
capsule mostly brown at maturity, apophysis usually as broad or broader than
the urn. 3. Seta less than 1 cm; leaves ±
imbricate, costa ending at the base of the acumination, distal leaf cells
thick-walled …………3. Tetraplodon
urceolatus 3 Seta 1--5 cm; leaves subulate or
concave acuminate, costa ending in the subula, distal leaf cells not
noticeably thickened. 4. Leaves abruptly subulate;
capsules red, becoming dark red almost black, mouth round …………… ……………4. Tetraplodon mnioides 4. Leaves concave with acuminate
apex; capsule pale-yellow to straw-colored, mouth square…………..……5. Tetraplodon pallidus 1.
Tetraplodon paradoxus
(R. Brown) Splachnum paradoxum R. Brown,
Chlor. Melvill., 44. 1823; Tetraplodon mnioides var. paradoxus
(R. Brown) Jensen Plants in dense light green or
yellow-green tufts. Leaves 2--5 mm, ovate, concave with acuminate
apex; costa thin, ending in subula; cells thin-walled, distal cells mostly
hexagonal, about 30 \um wide, in proximal portion elongate, rectangular. Sexual condition autoicous. Seta 2--3.5 cm,
clear pale-yellow to straw-colored. Capsule cleistocarpous; clear pale
yellow to straw-colored; spindle or club-shaped, apophysis conspicuously
narrower than the urn; stomata confined to upper part of apophysis; operculum
not developed. Spores
9 \um smooth. Caribou or musk ox dung; Greenland; N.W.T., Tetraplodon
paradoxus is cleistocarpous and much confused in the literature
with T. pallidus (W. C. Steere 1977) although easily
distinguished. Compared to T. pallidus, T. paradoxus is cleistocarpous, the setae are longer (up to
twice as long), leaves smaller and more narrow and the capsule is darker and
spindle-shaped or club-shaped, with the apophysis noticeably narrower than
the urn and having a smaller number of stomata. Steere (1977) suggested that spore
dispersal to fresh dung may occur when the sporophytes are ingested by
caribou or musk-oxen and the spores subsequently dropped in dung at some
other location. SELECTED REFERENCES Steere, W. C. 1977. Tetraplodon paradoxus and T. pallidus
(Musci: Splachnaceae) in northern
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