BFNA Title: Splachnaceae
Author: P. Marino 
Date: October 22, 2009
Edit Level: R 
Version: 1

Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden

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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 North America.

 

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

 

Voitia hookeri Mitten

 

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., Alaska, Colo.; Europe; ne Asia.

 

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 Rocky Mountains.  Voitia hyperborea is mainly Arctic in distribution.

 

Greenland; N.W.T., Nunavut; Alaska; Arctic Eurasia.

 

SELECTED REFERENCES Steere, W. C. 1974. The status and geographical distribution of Voitia hyperborea in North America (Musci: Splachnaceae). Bull. Torrey Bot. Club. 101: 55--63.

 

2.  TAYLORIA Hooker,  J. Sci. and Arts (London). 2(3): 144. 1816 * [For Thomas Taylor, 1775--1848, British muscologist and coauthor (with William Jackson Hooker) of the Muscologia Britannica]

 

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 North America. Bryologist 88: 82--93.

 

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 Labr., N.W.T., Nunavut, Ont., Que., Yukon; Alaska, Wash.; n, c Europe, Asia (Siberia); Atlantic Islands (Iceland).

 

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., Yukon; Alaska, Colo., Utah; n, c Europe; w, s Asia.

 

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 North America. Bryologist 88: 82--93.

 

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 North America.  Bryologist 88: 82--93.

 

5.  Tayloria splachnoides (Schleicher ex Schwägrichen) Hooker,  J. Sci. and Arts (London) 2(3): 144. 3.  1816

 

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.), Nunavut, Que.; Alaska; c, n Europe, Asia (Japan).

 

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 & W. Gümbel, Bryol. Eur. 3: 211 (fasc. 23--24. Mon.1). 1844. * [ Greek tetraplo, fourfold, and odous, a tooth, alluding to the arrangement of the teeth] 

 

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. Berlin. Pp. 241—264.

 

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) I. Hagen, Nyt Mag. Naturvidensk. 38: 332. 1901

 

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., Nunavut, Yukon; Alaska; n Europe (Arctic Russia).

 

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 North America. Brittonia 29: 353--367.

 

2.  Tetraplodon angustatus (Hedwig) Bruch & Schimper in P. Bruch, W. P. Schimper & W. Gümbel, Bryol. Eur. 3: 214 (fasc. 23--24. Monogr. 4). 1844

 

Splachnum angustatum Hedwig,  Sp. Musc. Frond., 51. 1801; Splachnum setaceum Michaux

 

Plants 2--8 cm, in compact tufts, yellow-green to brown, stems rather loosely foliate. Leaves erect-flexuose when dry, erect when moist, lanceolate-acuminate, narrowing to a very slender, long, flexuose acumen, 3--4 x 0.5 mm, reaching to the capsule or beyond, large teeth proximally to the middle, teeth larger distally or occasionally almost entire, costa nearly filling the subula; distal cells rectangular or oblong-hexagonal. Sexual condition autoicous or dioicous. Seta brownish, stout, not twisted, 2--4 mm. Capsule 1.5--2.2 mm, brown becoming dark brown with age; apophysis green becoming brown but longer and wider than the urn; stomata numerous in distal part of apophysis; annulus none; operculum hemispheric or bluntly conic. Calyptra short, conic-mitrate. Spores 9--10 \um smooth.

 

Droppings of carnivores, old bones, owl pellets, mainly in dry boreal habitats; Greenland; Alta, B.C., Man, N.B., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont., Que., Sask., Yukon; Alaska, Maine, Mich., Minn., Mont., N.H., N.Y.; n, c Europe; Asia (China, Japan, Russia in Siberia).

 

Tetraplodon angustatus, like T. mnioides, is mainly boreal in distribution and occurs on similar substrates. It is easily distinguished from T. mnioides, however, by lanceolate-acuminate, irregularly serrate leaves that narrow to a slender, elongate acumen, shorter seta, and green (brown with age) apophysis.  In Alberta, sporophytes mature in spring prior to the maturation of T. mnioides sporophytes, resulting in the temporal separation of spores of these two species on fresh droppings and thus the physical separation of T. angustatus and T. mnioides on droppings where these two species occur together regionally.

 

SELECTED REFERENCES   Marino, P. C. 1997.  Competition, dispersal and coexistence of Splachnaceae in patchy habitats. In: R. E. Longton, ed. 1997. Advances in Bryology. Berlin.  Pp. 241--264.

 

3.  Tetraplodon urceolatus (Hedwig) Bruch & Schimper in P. Bruch, W. P. Schimper & W. Gümbel,  Bryol. Eur. 3: 217 (fasc. 23--24. Monogr. 7). 1844

 

Splachnum urceolatum Hedwig, Sp. Musc. Frond., 52. 1801; Tetraplodon mnioides var. cavifolius Schimper

 

Plants 1--4 cm, in very dense, compact tufts, slender yellow-green distally, densely radiculose proximal to the distal 2 - 5 mm. Leaves imbricate, entire, subcochleariform and very concave, ovate, about 1.5 x 0.8 mm with a flexuous yellow costa ending before the subula; leaf cells distally thick-walled, rectangular, 25--30 x 15--18 \um, those at the leaf base long and thin-walled, 65--70 x 18 \um. Sexual condition autoicous or dioicous. Seta stout, not twisted, straw-colored usually becoming dark red with age, 3--7 mm. Capsule dark reddish brown, cylindric, somewhat oblong; apophysis about the same length as the urn, stomata  over whole of apophysis; peristome teeth reflexed orange-red at the tips, inserted proximal to the mouth, in 4’s then 2’s; operculum hemispheric. Calyptra cucullate, about 2 mm. Spores 10--11 \um, smooth or slightly papillose.

 

Droppings of carnivores, old bones, owl pellets in dry usually very exposed places, open tundra, mountain summits;  Greenland; Alta, B.C., Man, N.W.T., Nunavut, Que., Yukon; Alaska; c Europe Alps, e, s Asia.

 

Tetraplodon urceolatus can be difficult to distinguish from T. mnioides in the northern part of the latter species’ range.  In general, T. urceolatus has leaves firm, concave, with stiff, rounded apices, and setae yellow, generally less than 2 cm.

 

SELECTED REFERENCE   Marino, P. C. 1988. The North American distribution of the circumboreal species of Splachnum and Tetraplodon. Bryologist 91: 161--166.

 

4.  Tetraplodon mnioides (Hedwig) Bruch & Schimper in P. Bruch, W. P. Schimper & W. Gümbel,  Bryol. Eur. 3: 215 (fasc. 23--24. Monogr. 5).  1844

 

Splachnum mnioides Swartz ex Hedwig, Sp. Musc. Frond., 51. 1801

 

 

Plants 1--4 (--8) cm, in dense light green or yellow-green tufts. Leaves crowded, erect, somewhat contorted when dry, 1--2 x 3--5 mm, concave, entire, oblong-ovate, abruptly narrowed to a ± long, often yellowish subula; costa vanishing in the subula; distal cells rectangular and oblong-hexagonal. Sexual condition autoicous. Seta stout, not twisted, straw-colored, usually becoming dark red with age, 10--50 mm. Capsule red, becoming dark or almost black, urn 1--1.5 mm, apophysis somewhat wider than the urn distally, gradually tapered to the seta, about twice as long as or somewhat longer than the urn, 2--3.5 mm, sometimes lighter than the urn but becoming the same color or darker, somewhat wrinkled longitudinally when dry, stomata large, numerous, over whole of aphophysis; annulus poorly developed; operculum bluntly conic. Calyptra short, conic-mitrate. Spores 9--12 \um smooth or slightly papillose.

 

Droppings of carnivores, old bones, owl pellets in dry alpine, boreal or arctic habitats, Greenland; Alta, B.C., Man, N.B., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont., Que., Sask., Yukon; Alaska, Maine, Mich., Minn., N.H., N.J., N.Y., Ore., Vt., Wash., W.Va.; n Europe; Asia.

 

Tetraplodon mnioides is much more common than other species of the genus, but is readily distinguished.  It differs from T. angustatus in larger tufts and longer sporophytes, which are dark red, becoming almost black with age.  Sterile material of T. mnioides differs from that of T. angustatus in leaves oblong-ovate, rather abruptly subulate, and entire, whereas those of T. angustatus are oblong-lanceolate, gradually subulate, and serrate.  Tetraplodon pallidus has straw-colored setae, and capsules with a much more delicate capsule wall such that when the teeth become reflexed, they tend to make the capsule mouth square.  Tetraplodon paradoxus, like T. pallidus, has a yellowish setae and capsules.  It has, however, a narrow, tapered apophysis and is cleistocarpous.  Tetraplodon urceolatus is an Arctic or alpine moss very similar to reduced northern forms of T. mnioides; it has, however, a short yellowish seta, small strongly concave appressed leaves, and a short apophysis only about as long as the urn and with relatively fewer stomata generally confined to the distal portion of the apophysis. 

 

SELECTED REFERENCES Bequaert, J. 1921.  On the dispersal by flies of the spores of certain mosses of the family Splachnaceae.  Bryologist. 24: 1--4.    Marino, P. C. 1988. The North American distribution of the circumboreal species of Splachnum and Tetraplodon. Bryologist 91: 161--166.     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.   

 

5.  Tetraplodon pallidus I. Hagen , Kongel. Norske Vidensk. Selsk. Skr. (Trondheim) 1893: 75.  1894

 

Tetraplodon mnioides subsp. pallidus (I. Hagen) Kindberg

 

Plants in dense light green, yellow-green tufts. Leaves 2--3.5 mm, ovate, concave with acuminate apex; costa thin, ending in subula; cells thin-walled, distal cells mostly hexagonal, about 30 \um wide, in proximal part elongate, rectangular.  Sexual condition autoicous. Seta 1--2 cm, clear pale-yellow to straw-colored. Capsule clear pale-yellow to straw-colored; capsule wall delicate such that the reflexed peristome teeth tend to make the capsule mouth square; apophysis generally not broader than the urn and often narrower; stomata confined to distal portion of apophysis; operculum large, bluntly conic. Spores ca. 8 \um, smooth.

 

Caribou or musk ox dung; Greenland; B.C., Nfld. and Labr. (Labr.), N.W.T., Nunavut, Yukon; Alaska; n Europe, Arctic Siberia.

 

Tetraplodon pallidus  and T. paradoxus often grow intermixed and can be distinguished by several characters (W. C. Steere 1977).  Tetraplodon paradoxus is cleistocarpous, the setae are longer (up to twice as long) and leaves smaller and more narrow than in T. pallidus, and the capsule is darker and spindle-shaped or club-shaped, with the apophysis noticeably narrower than the urn.  In contrast, T. pallidus has a large operculum that falls off exposing a broad, square-shaped mouth of the capsule and the well-developed reflexed teeth. 

 

SELECTED REFERENCES Steere, W. C. 1977. Tetraplodon paradoxus and T. pallidus (Musci: Splachnaceae) in northern North America. Brittonia. 29: 353--367.

 

 

4. SPLACHNUM Hedwig,  Sp. Musc., Frond., 51.  1801 *  [Used by Dioscorides for a lichen with a lung-like appearance, like the shrunken apophysis of Splachnum]

 

Plants in soft, lose, light to emerald green, often shiny tufts, radiculose. Stem 1--4 cm, loosely foliate, simple or forked. Stem leaves large, usually more crowded toward the stem tip, obovate or obovate-lanceolate, obtuse, acute or acuminate; erect or erect-spreading, soft, ± contorted when dry; entire or serrate distally; costa ending in or before the apex; cells large, oblong-hexagonal distally, rectangular proximally. Sexual condition autoicous or dioicous; perichaetia terminal on stem or branches with fewer and smaller leaves; perigonia large and cupulate. Seta 1--20 cm, usually erect, smooth; apophysis usually much wider than the capsule sometimes skirt-like or umbrella-like; growth usually continued after spore maturation; annulus none. Capsule smaller than apophysis, erect, reddish-brown, cylindric; exothecial cells transversely elongated in distal rows, quadrate-hexagonal and rectangular at the base; columella shortly exserted; peristome teeth 16, inserted at or proximal to the mouth, sometimes fused, reflexed when dry, inflexed when moist; finely papillose; composed of 3 layers of cells and chambered; operculum conic to hemispheric and apiculate. Calyptra small, reaching to the apophysis, conic-mitrate, not constricted distal to the base, epapillose. Spores small, about 11 \um, yellow, smooth.

 

Species 11 (6 in the flora): tropical to subarctic worldwide except Antarctic.

 

Splachnum are entomophilous and coprophilous, and all circumboreal species appear to be restricted to growing on herbivore dung (e.g., moose, caribou).  The genus is characterized by the chambered peristome teeth as well as several species having greatly enlarged, colorful, differentiated apophyses.

 

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.   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. Berlin. Pp. 241--264.    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. Apophysis more or less globose or turbinate, much broader than the urn; wrinkled when dry.

 

2. Apophysis yellow or pink, rarely reddish; leaves ± oblanceolate, with large teeth, slenderly acuminate………………….……….……………1. Splachnum ampullaceum

 

2. Apophysis purple when fresh, leaves ovate to obovate, always entire, apex blunt and rounded or with an obtuse point………………………………2. Splachnum vasculosum

 

1. Apophysis either large and skirt- or umbrella-like or not much broader than the urn and globose or narrowly pyriform.

 

3. Apophysis large, discoid or convex umbrelliform.

 

4. Apophysis discoid umbrelliform, yellow; leaves toothed only in distal portion…………………………………...….3. Splachnum luteum

 

4. Apophysis convex umbrelliform, red-purple; leaves toothed nearly to the base ............................................................... 4. Splachnum rubrum

 

3. Apophysis not much broader than the urn, globose or narrowly pyriform.

 

5. Setae less than 1 cm; capsules shortly exserted; apophysis green with red-brown tip; leaves long-lanceolate, slenderly long-acuminate …… 6. Splachnum pensylvanicum

 

5. Setae considerably longer than 1 cm (to 7.5 cm); capsules long-exserted; apophysis green turning red-brown upon drying; leaves broadly obovate, abruptly short- to long-acuminate ………………5. Splachnum sphaericum

 

1.  Splachnum ampullaceum Hedwig, Sp. Musc. Frond., 55.  1801

 

Plants soft, loose, pale- or yellow-green, somewhat shiny tufts, radiculose proximally, 1--2 cm. Leaves somewhat longer and crowded at stem tips, 3.5--4 mm, long-lanceolate to narrowly oblong-obovate, slenderly acuminate; margins deeply spinose-dentate in the distal half; costa disappearing in the acumen somewhat before the apex. Sexual condition dioicous, male and female plants mixed.  Seta 15--65 mm, flexuose, red or red brown. Capsule urn distally yellow-brown, 1--1.2 mm; apophysis much broader than the urn, 2--6 mm, top-shaped, rugose, color yellow to pinkish or pale purple; annulus none; operculum hemispheric, blunt; peristome teeth 16, inserted near the mouth, united in pairs, pale-brown or orange-brown, reflexed when dry, inflexed when moist, lanceolate, densely and minutely papillose on the outer surface; operculum hemispheric, blunt. Spores 7--9.5 \um subspherical, smooth, yellow-green, sticky.

 

Dung of large boreal herbivores (e.g., moose), mainly muskeg and other boggy habitats;  Greenland; Alta, B.C., Man, N.B., Nfld. and Labr. (Nfld.), N.W.T., N.S., Nunavut, Ont., Que., Sask., Yukon; Alaska, Maine, Mich., Minn., Mont., N.H., N.Y., Ore.; n, c Europe; Asia (China, Japan, Siberia).

 

Splachnum ampullaceum is the most common species of the genus in boreal North America. It sometimes grows with S. luteum in the western North America and S. pensylvanicum in eastern North America.  Splachnum ampullaceum is easily recognized by both sporophyte and gametophyte morphology.  The name of the species is derived from its ampulla-shaped capsules with abruptly dilated bases.  The apophysis is broad and top-shaped and ranges in color from yellow to pink and almost dark-red with age.  The leaves, unlike other North American species in the genus, have deeply spinose-dentate margins. 

 

SELECTED REFERENCES Cameron, R. G., D. Troilo. 1982. Fly-mediated spore dispersal in Splachnum ampullaceum  (Musci). Michigan Botanist 21: 59--65.  Whitmire, R. S. 1965.  Ecological observations on Splachnum ampullaceum.  Bryologist. 68:342--343.

 

2.  Splachnum vasculosum Hedwig,  Sp. Musc. Frond., 53.  1801

 

Plants loosely tufted, light to dark green, 1--3 cm. Leaves ± contorted when dry, spreading when wet, ovate to obovate with narrow base, 2.5 x 4 mm, entire; apex blunt, rounded or obtusely pointed; proximal leaves obtuse, distal apiculate; basal cells rectangular, others ± rhomboidal; costa ending before the apex. Sexual condition dioicous. Seta 1--3 cm, thin, reddish orange, twisted. Capsule dark orange, 1 mm, cylindric; apophysis red-purple when moist and ovoid, broader distally, tapering proximally and as long as or longer and much broader than the urn, when dry rugose and narrower; operculum convex and apiculate to low-conic. Spores 8--12.5 \um smooth, greenish yellow.

 

Dung in bogs and other wet places; Greenland; Man., Nfld. and Labr., Que., N.W.T., Nunavut, Yukon; Alaska; n Europe; Asia (Siberia).

 

Splachnum vasculosum has a more northerly distribution than S. ampullaceum, a species with which it could be confused.  The leaves of S. vasculosum are entire and ovate to obovate with a blunt or blunt-rounded apex.  The apophysis is dark-purple, and compared to the apophysis of S. ampullaceum is similar in size but more globose and narrows more abruptly to the seta.

 

SELECTED REFERENCES   Brassard, G. R. 1971. The mosses of northern Ellesmere Island, Arctic Canada. II. Annotated list of the taxa.  Bryologist 74: 282--311.

 

3.  Splachnum luteum Hedwig,  Sp. Musc. Frond.,  56.  1801

 

Plants soft, loose, pale or yellow-green, somewhat shiny tufts, radiculose proximally, 1.5--3.5 cm. Leaves somewhat crowded at stem tips, 5--6 mm, oblong-obovate, abruptly narrowed to a slender acumen; margins bluntly serrate to subentire distally, indistinctly bordered; costa disappearing in the acumen somewhat before the apex. Sexual condition dioicous, male and female plants mixed. Seta 2--15.5 cm, greenish yellow to orange-red. Capsule urn orange-brown, 1--1.5 mm; apophysis discoid-umbrelliform, 4.5--11 mm across, bright yellow, smooth; annulus none; peristome teeth 16, inserted proximal to the mouth, approximate in pairs, orange-brown; reflexed when dry, inflexed when moist, lanceolate, densely and minutely papillose on the outer surface; operculum hemispheric, blunt.. Spores 7--9 \um subspherical, smooth, yellow-green, sticky.

 

Dung of large boreal herbivores (e.g., moose) mainly in muskeg and other boggy habitats; Alta., B.C., Man., Nfld. and Labr. (Labr.), N.W.T., Ont., Que., Sask.; Alaska; n Europe; Asia.

 

The bright yellow apophysis on its long seta of Splachnum luteum gives this moss the appearance of a flower. The apophysis is flattened and umbrelliform, unlike the more convex umbrelliform apophysis of S. rubrum.  Immature sporophytes look much like the mature sporophytes of S. sphaericum or the immature sporophytes of the other North American species of Splachnum.  The oblong-obovate leaves that narrow to a slender acumen and the bluntly serrate leaves, however, help distinguish this species. Splachnum luteum is generally rare although locally common in central Alaska; it is often found growing on the same patches of dung with S. sphaericum and sometimes S. rubrum. 

 

4.  Splachnum rubrum Hedwig,  Sp. Musc. Frond., 56.  1801

 

Plants soft, loose, pale or yellow-green, somewhat shiny tufts, radiculose proximally, 15--35 mm. Leaves somewhat crowded at stem tips, 5--7.5 mm, obovate, long-acuminate; margins coarsely serrate distally, indistinctly bordered; costa disappearing in the acumen somewhat before the apex. Sexual condition dioicous, male and female plants mixed. Seta 5--13 cm, orange-red. Capsule urn of capsule orange-brown, 1--1.5 mm; apophysis convex-umbrelliform, 6--11 mm across, bright magenta red, smooth; annulus none; peristome teeth 16 inserted proximal to the mouth, brownish, not fused or approximate in pairs, orange-brown; reflexed when dry, inflexed when moist, lanceolate, densely and minutely papillose on the outer surface; operculum hemispheric, blunt.. Spores 11--13 \um subspherical, smooth, yellow-green, sticky.

 

Dung of large boreal herbivores (e.g., moose) mainly in muskeg and other boggy habitats.  Alta., B.C., N.B., N.S., Nfld. and Labr. (Labr.), N.W.T., Ont., Que.; Alaska, Mich.; n Eurasia.

 

The magenta red, almost iridescent apophysis on a long seta gives Splachnum rubrum the appearance of a flower.  The apophysis is convex-umbrelliform, unlike the discoid-umbrelliform apophysis of S. lutem.  Also, like S. luteum, the young sporophytes of S. rubrum resemble the mature sporophytes of S. sphaericum and the immature sporophytes of S. ampullaceum.  The leaves of S. rubrum are less broadly obovate than those of S. sphaericum or S. luteum.  Of the mainly boreal North American species of Splachnum, this species is by far the rarest.  It is often found growing intermixed with S. luteum. 

 

5.  Splachnum sphaericum Hedwig,  Sp. Musc. Frond., 55.  1801

 

Splachnum ovatum var. sphaericum (Hedwig) Dixon, Rev. Bryol. Lichénol. 6: 98 1933; S. ovatum Dickson ex Hedwig

 

Plants in light or yellow-green tufts, often branched, 5--30 mm. Leaves somewhat crowded at stem tips, ± contorted when dry, spreading when wet, 2.5--3.5 mm, broadly obovate, often narrowing abruptly to an acuminate tip; margins entire or toothed distally, yellowish; costa almost as wide as leaf base at insertion, disappearing in the acumen just before the apex; proximal leaves smaller, narrower. Sexual condition dioicous, male and female plants mixed. Seta 1.5--10 cm, thin, flexuose, yellow, becoming reddish. Capsule urn of capsule orange-red, capsule and apophysis 2--4 mm; apophysis green when first mature, becoming purplish and wrinkled upon aging and drying, slightly wider than the urn, ± spherical to obovoid; peristome teeth inserted proximal to the mouth, fused in pairs, orange-brown; operculum hemispheric, apiculate. Spores 7--13 \um, spherical, smooth, yellow-green, sticky.

 

Dung of large boreal herbivores (e.g., moose), mainly in muskeg and other boggy habitats;  Greenland; Alta., B.C., Man., Nfld. and Labr. (Labr.), N.W.T., Ont., Que.; Alaska, Colo., Mich., Mont., Wash., Wyo.; n, c Europe; n Asia.

 

Splachnum sphaericum often grows intermixed with S. luteum in western Canada and central Alaska.  Unlike other North American species of Splachnum, its apophysis is scarcely wider than the capsule and is green upon maturity, only turning purplish as it ages and dries.  Immature sporophytes of the other North American species of Splachnum, however, resemble the mature sporophytes of S. sphaericum.  The gametophyte of S. sphaericum can be distinguished from the other North American species by its broadly obovate, narrowly acuminate leaves.  The apophysis of S. sphaericum is much more strongly scented than that of S. luteum.  Because of the size, shape, and color of the apophysis, S. sphaericum could be confused with Tetraplodon mnioides, but T. mnioides is usually found growing on carnivore droppings in drier habitats and has oblong-obovate leaves that narrow abruptly. 

 

5.  Splachnum pensylvanicum (Bridel) Grout ex H. A. Crum, Bryologist 69:206.  1966

 

Mnium pensylvanicum Bridel,  Muscol. Recent. Suppl. 3: 44.  1817 

 

Plants slender, in green tufts, often branched, 2--25 mm. Leaves erect; slightly contorted when dry, long-lanceolate and slenderly long-acuminate, 2.5--5 mm, margins not distinctly bordered, occasionally entire but mostly irregularly but sharply serrate, teeth in distal half often long and ciliate; costa stout, disappearing in the acumen near the apex. Sexual condition autoicous. Seta 2--10 mm, slender, almost hyaline, light green, flexuose. Capsule exerted or sometimes barely emergent, 1--2.5 mm; urn 0.6--0.7 mm, orange brown; apophysis greenish with dark red or purplish coloring distally, wrinkled when dry, as wide or slightly wider than the urn, tapered towards the seta, as long or longer than the urn; peristome teeth inserted proximal to the mouth, fused in 8 pairs, brown or orange-brown; operculum conic, bluntly apiculate. Spores 7--11 \um, subspherical, smooth, light green, sticky.

 

Dung of herbivores (e.g., moose, cattle); N.B.,Nfld. and Labr. (Nfld.), N.S.; Fla., Ga., Maine, Mass., Pa., N.J., N.Y., N.C., S.C., Tenn., Tex.

 

Splachnum pensylvanicum is endemic to eastern North America, and has a much more southerly distribution than other North American species. It has often been collected from cattle droppings from Florida through the Atlantic coastal plan north to New England.  It has become rare, however, within this range.  It is quite abundant on the Avalon Peninsula of eastern Newfoundland where it grows both in pure and in mixed populations with S. ampullaceum on moose droppings.  In both sporophytic and gametophytic characteristics S. pensylvanicum is quite distinct from the other North American species.  Its sporophyte is relatively diminutive, with a very short seta, and a small, barely inflated apophysis.  The leaves, like those of S. ampullaceum are long-lanceolate and slenderly acuminate but, unlike the spinose-dentate distal leaf margins of S. ampullaceum, the distal leaf margins of S. pensylvanicum range from entire to irregularly serrate. 

 

SELECTED REFERENCE    Bagnell, B. A. and S. R. Clayden. 1993. Splachnum pensylvanicum in Canada. Bryologist 96: 216--218.

 

5. APLODON R. Brown,  Chlor. Melvill., 299.  1823

 

Haplodon R. Brown ex I. Hagen

 

Plants slender lax tufts, light green distally, often with flagellate branches. Stem to 6 cm. Stem leaves concave, broadly ovate or obovate, margins entire; costa ending before apex; cells large, oblong-hexagonal distally, rounded-rectangular proximally. Sexual condition autoicous with perichaetia terminal on axillary branches. Seta hyaline, very slender, twisted, flexuose, hyaline, 1--2 cm. Capsule ovate, reddish; apophysis rounded, not or only slightly wider than the urn; peristome single with 8--12 rudimentary outer teeth; operculum hemispheric and sometimes apiculate; columella included.. Calyptra conical, mitrate or cucullate. Spores 10--15 \um smooth, yellow-green.

 

 

Species 1 (1 in Flora): subarctic to Arctic Northern Hemisphere.

 

Although Aplodon has an inflated apophysis, it is not, based upon the molecular data, well-resolved within the Splachnaceae, and its relationship to this family requires further study (B. Goffinet et. al. 2004).

 

1.  Aplodon wormskjoldii (Hornemann) R. Br.  Chlor. Melvill., 41.  1823

 

Splachnum wormskioldii Hornemann, Flora Danica 10(28): 8, 9, fig. 1659.  1819; T. wormskioldii (Hornemann) Lindberg 

 

Leaves 1--1.5 x 2--3 mm, shrunken when dry, erectopatent when moist, concave, broadly ovate or obovate, the proximal obtuse, the distal with frequently a reflexed obtuse acumen; costa sometimes forked. Capsule ovate when wet, widest at the mouth when dry, reddish becoming almost black with age; capsule and apophysis 1.5--2 x 0.5--1 mm; peristome teeth at first in pairs, then separate, of two layers, short, truncate, reflexed or erect.

 

Dung, animal remains; Greenland; Man, Nfld. and Labr. (Labr.), N.W.T., Nunavut, Que.; Alaska; n Eurasia.

 

Aplodon wormskjoldii can be distinguished by the hyaline, slender and twisted seta.  The gametophyte differs from other Splachnaceae in having shorter and much wider apical cells and a wider marginal row of cells on the leaves. 

 

SELECTED REFERENCES    Brassard, G. R. 1978. An extensive subfossil deposit of the arctic moss Aplodon wormskioldii.  Canad. J. Bot. 56: 1852--1859. Observations on the genus Aplodon (Musci: Splachnaceae) Bryologist. 76: 347--355.