BFNA Title: Splachnaceae
(for illustrations CLICK HERE)
XX. SPLACHNACEAE Greville & Arnott
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,.
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
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.
REFERENCE LaFarge-England and C., D.
H. Vitt 1985. A Taxonomic Study of Tayloria hornschuchii and T.
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.
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.
REFERENCES LaFarge-England, C. and
D. H. Vitt 1985. A Taxonomic Study of Tayloria hornschuchii and T. froelichiana in
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
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
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.
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
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