Sphagnum squarrosum var. semi-squarrosumRussow ex Warnst.
Sphagnum cymbifolium var. squarrosum(Crome) Nees & Hornsch.
Sphagnum latifolium var. squarrosum(Crome) Wahlenb.
Sphagnum squarrosum var. immersumBeckett ex Warnst.
Sphagnum teres subsp. squarrosum(Crome) Warnst.
Sphagnum teres var. squarrosum(Crome) Warnst.
Sphagnum imbricatum var. brachycladum(Warnst.) Röll
Sphagnum obtusifolium var. crassisetum(Brid.) Sw.
Sphagnum squarrosum f. brachycladumWarnst.
Sphagnum squarrosum var. capitatumRöll
Sphagnum squarrosum var. compactumWarnst.
Sphagnum squarrosum f. cuspidatum(Warnst.) Warnst.
Sphagnum squarrosum f. densumRussow ex Warnst.
Sphagnum squarrosum f. elegans(Röll) Warnst.
Sphagnum squarrosum f. flagellare(Röll) Warnst.
Sphagnum squarrosum var. fuscolutescensC.E.O.Jensen
Sphagnum squarrosum f. gracileRussow ex Warnst.
Sphagnum squarrosum f. humile(Schlieph. ex Röll) Röll
Sphagnum squarrosum f. molle(Röll) Warnst.
Sphagnum squarrosum f. patulum(Röll) Warnst.
Sphagnum squarrosum f. pycnocladumWheldon
Sphagnum squarrosum f. robustum(Röll) Warnst.
Sphagnum squarrosum f. silvaticumWheldon
Sphagnum squarrosum var. spectabileRussow
Sphagnum squarrosum var. strictiformeRöll
Sphagnum squarrosum var. submersumRöll
Sphagnum squarrosum f. subsquarrosum(Russow) C.E.O.Jensen
Sphagnum teres var. cuspidatum(Warnst.) Cardot
Sphagnum teres var. imbricatum(Schimp.) Warnst.
Sphagnum teres f. immersumWarnst.
Sphagnum teres f. strictumWarnst.
Sphagnum squarrosum var. brachycladum(Warnst.) Grav. ex Röll
Sphagnum squarrosum var. cuspidatumWarnst.
Sphagnum squarrosum f. densum(Röll) Warnst.
Sphagnum squarrosum var. densumRöll
Sphagnum squarrosum f. elegansRussow ex Warnst.
Sphagnum squarrosum var. elegansRöll
Sphagnum squarrosum var. flagellareRöll
Sphagnum squarrosum var. gracile(Russow ex Warnst.) Röll
Sphagnum squarrosum var. humileSchlieph. ex Röll
Sphagnum squarrosum f. imbricatum(Schimp.) C.E.O.Jensen
Sphagnum squarrosum f. immersum(Warnst.) C.L.Beckm. ex Warnst.
Sphagnum squarrosum var. molleRöll
Sphagnum squarrosum var. patulumRöll
Sphagnum squarrosum var. robustumRöll
Sphagnum squarrosum var. strictum(Warnst.) Röll
Sphagnum squarrosum var. subsquarrosumRussow
Sphagnum squarrosum var. tenellumRöll
Sphagnum teres var. compactum(Warnst.) Cardot
Sphagnum squarrosum, commonly known as the spiky bog-moss or spreading-leaved bog moss, is a peat moss species found in nutrient-rich, damp soils and wetlands across the Northern Hemisphere, with isolated populations in South America. Its spiky appearance, resulting from strongly spreading branch leaves, distinguishes it from other peat moss species. Playing an important role in wetland succession, the species is one of the first Sphagnum mosses to colonise developing wetlands. It shows considerable tolerance to mineral-rich conditions and actively modifies its habitat through cation exchange processes.
Unlike many other peat mosses that require highly acidic conditions, S.squarrosum thrives in areas with moderate calcium levels, particularly along stream banks, in fens, and in wet woodlands dominated by Salix (willows) and Betula (birches). It disperses efficiently, producing more spores per capsule and per patch than other Sphagnum species. It shows high genetic diversity across its range, having survived the Last Glacial Maximum in multiple European refugia. The species serves as the specific host for the fungal parasite Discinella schimperi, which infects about half of all documented populations.
The species is the type for sectionSquarrosa of the genus Sphagnum. Although closely related to Sphagnum teres, the two are morphologically and ecologically distinct, justifying their classification as separate species. This distinction is further supported by cytological evidence, with S.squarrosum reported to have double the chromosome number of S.teres.[3]
Over time, the species has accumulated numerous synonyms and infraspecific taxa, reflecting its morphological variability across its wide geographical range. Notable among these are S.semisquarrosum (Russow ex Warnst.) Lepage and various forms and varieties such as var. squarrosulumG.Müller and var. semisquarrosumRussow ex Warnstorf.[3]
Description
Sphagnum squarrosum is readily identified by its distinctive spiky appearance, which sets it apart from other peat mosses. This large, robust species forms loose carpets in pale to yellow-green colours, though it may develop pale brown colouration in exposed alpine or arctic habitats. Its common name "spiky bog-moss" comes from its most striking feature: branch leaves that spread outward at sharp angles from the branches like tiny stars, a growth pattern botanists term squarrose .[4][5][6]
The plant's structure follows the typical Sphagnum pattern, but with distinctive characteristics. At its crown, it possesses a very large and prominent apical bud. The stem is strong, measuring 0.7–1.3mm in diameter, with a dark brown internal cylinder that becomes paler in shade forms. The stem cortex comprises 2–3 layers of hyaline (colourless and translucent) cells, with the superficial layer showing indistinct thinnings known as 'shadow pores'. The branch cortex contains specialised pore-bearing cells called retort cells (flask-shaped cells that aid in water conduction), which are often relatively indistinct from other cortical cells and occurring in groups of one to four, with an internal cylinder that is pale brown or yellowish. The branches are arranged in fascicles (clusters) of 4–6, with 2–3 spreading branches measuring 20–30mm or more, and 2–3 pendent branches of 8–30mm.[4][5][6]
The stem leaves are shorter than the branch leaves, measuring 1.6–1.8mm long, and have an ovate- lingulate to oblong-lingulate shape with broadly rounded tips. These leaves possess short-lived borders 2–3 cells wide that are often lost in older leaves, though they persist longer than in S.teres. Branch leaves are larger, measuring 1.9–3.3mm long, with a broadened base that abruptly contracts at the middle to form pronounced 'shoulders' before tapering to a sharply reflexed , acuminate limb. The lower half remains erect and concave, forming a sheath around the branch.[4][5][6]
Morphology
Close-up of branches showing the distinctive squarrose (outward-spreading) branch leaves that give Sphagnum squarrosum its common name 'spiky bog-moss'
Microscope view of leaf showing the network of photosynthetic (chlorocysts) and water-storage (leucocysts) cells. Scale bar: 0.5 mm
Mature sporophytes (reddish-brown capsules) elevated on pale pseudopodia above the characteristic spiky gametophyte
At the cellular level, S.squarrosum shares the specialised cell structure common to all Sphagnum species. It possesses two types of cells: hyaline cells (leucocysts), which store water and give the plant its impressive water-holding capacity, and photosynthetic cells (chlorocysts). In the squarrose limb, hyaline cells are relatively small (70–100 × 15–22μm) but become larger towards the lower margins (up to 200 × 30–50μm). They bear 2–6 large, distinctly ringed pores on their adaxial surface, with similar or slightly fewer pores on the abaxial surface. The photosynthetic cells appear narrowly oval-triangular to trapezoidal in cross-section and reach both surfaces, though they are more widely exposed on the abaxial surface.[4][6]
At the reproductive level, S.squarrosum produces both male and female structures on the same plant. The antheridial bracts are densely imbricated (overlapping) and often show yellowish or pale brown colouration; they resemble branch leaves but are smaller and have less divergent apices. The inner perichaetial bracts are large with relatively narrow insertion points, becoming broad above with a retuse (slightly notched) apex, with abaxial resorption gaps and eroded apex. When present, the spore capsules contain yellow-brown, papillose spores. The branch anatomy includes retort cells, which are often indistinct from other cortical cells and occur in groups of one to four. The internal cylinder is pale brown or yellowish.[6]
The species shows some phenotypic plasticity in response to environmental conditions. In temperate regions, it typically appears more robust and develops fuller growth in shaded habitats, particularly along brook margins and in wet parts of forest floors. However, in subarctic and arctic regions, it commonly achieves similar robust growth in open, treeless areas.[5]
Diagnostic features and similar species
Sphagnum squarrosum is typically recognisable by its robust habit, bright green to yellow-green colouration, and distinctive "bottle brush" appearance created by its strongly squarrose branch leaves. This characteristic spiky appearance is most pronounced in well-developed plants, though it may be less obvious in some forms.[6]
Several other Sphagnum species may be confused with S.squarrosum under certain conditions. Its closest relative, S.teres, can appear similar, particularly when S.squarrosum produces weaker growth forms with less pronounced leaf squarrosity. In such cases, microscopic examination becomes necessary for definitive identification, though even this may be challenging due to gradation of characters in extreme forms of both species.[6]
Shade forms of S.palustre may superficially resemble S.squarrosum, but can be distinguished by two key features: the roughened, cucullate (hood-shaped) apices of their branch leaves and the presence of spiral fibrils in their branch cortex. While S.compactum can also produce squarrose-leaved forms, it is readily distinguished by its notably minute stem leaves.[6]
In exposed alpine or arctic habitats, S.squarrosum may develop a pale brown colouration rather than its typical green hues. The species shows some morphological plasticity in response to environmental conditions, with shade forms often developing a paler internal stem cylinder compared to the typically dark brown colouration of plants from more exposed sites.[6]
Habitat, distribution, and ecology
Sphagnum squarrosum has a broad geographical range spanning much of the Northern Hemisphere, occurring throughout North America and Eurasia.[7] While it is widely distributed, the species shows particular concentration in northern regions of Europe.[7] Its circumpolar range has been well-documented through specimens from Canada (including the Northwest Territories and Yukon), the United States (including Alaska), Greenland, multiple European countries, Japan, and various regions of Russia.[8] The 2017 report of populations in southeastern Brazil has extended its known range into South America, representing its southernmost occurrence globally.[4]
Sphagnum squarrosum forming characteristic green carpets at the edge of standing water in Torfowisko pod Zieleńcem Nature Reserve, Poland
Unlike many other peat mosses that require highly acidic conditions, S.squarrosum thrives in mineral-rich environments with moderate calcium levels. This tolerance is supported by laboratory studies showing the species maintains consistent photosynthetic rates across a wide pH range (5.2–7.5).[9] The species actively shapes its environment through cation exchange, absorbing calcium and releasing protons to gradually alter water chemistry. This ability makes it an important pioneer species in wetland development, often appearing alongside S.fimbriatum in early successional stages before more acid-loving bog species can establish.[10][11]
The species shows distinct habitat preferences across its range. In temperate eastern Canada, it is more commonly found in shaded habitats, particularly along brook margins and wet parts of forest floors. In contrast, in subarctic to arctic regions, it grows in open, treeless areas.[5] It typically inhabits mesotrophic to slightly eutrophic conditions, growing in wet habitats such as river banks, pond edges, fens, and woodland carr dominated by Salix, Betula or Alnus species.[7] In North America, it is particularly associated with woodlands subject to seasonal flooding, swamps, and stream margins, while in montane regions it can also colonise wet rock ledges.[4]
The species reaches its northernmost extent in Svalbard (Norway), where it is the most prevalent Sphagnum species. Here, at about 80°32.5'N at Nordkapp, at the northern extreme of Chermsideøya on Nordaustlandet, it grows from sea level to approximately 300m (980ft) elevation, forming low mats or cushions on moist, gently sloping moss-rich tundra and mineral-rich water seepages. Unlike its growth patterns elsewhere, in Svalbard it typically occurs in pure stands rather than mixing with other Sphagnum species. It develops a distinctly yellowish-green to pale yellowish-brown colouration, with brown, pale brown or yellowish-brown stems.[12]
Sphagnum squarrosum is ecologically adaptable throughout its range. It can withstand periods of dryness and grows successfully in both rain-fed and ground water conditions. The species shows particularly vigorous growth when nutrients are abundant, responding positively to enrichment.[7] This adaptability is further evidenced by its wide altitudinal range, occurring from sea level to elevations of at least 1,100m (3,600ft).[4]
Reproduction and dispersal
Historical illustration of Sphagnum reproductive structures: (C) mature sporophyte showing capsule and pseudopodium; (A) longitudinal section of developing archegonium; (B) cross-section of mature capsule showing spore chamber and air space.
Sphagnum squarrosum is a monoecious species (having both male and female reproductive organs on the same plant) and frequently produces spores throughout its range.[7] The species has efficient dispersal capabilities. Among studied Sphagnum species, it produces both the highest number of spores per capsule and the highest overall spore output per patch of growth.[7]
Large capsules and small spores enhance the species' long-distance dispersal efficiency. Compared to other Sphagnum species, Sphagnum squarrosum retains fewer spores within its capsules after dispersal, resulting in greater spore release and wider distribution.[7]
Research on island colonization reveals that S.squarrosum is an effective long-distance disperser, successfully establishing populations as far as 40km (25mi) from parent sources. The species is notable for being able to colonise habitats closer to the sea than other Sphagnum species, demonstrating higher tolerance to salt spray and ionic concentrations. Like other Sphagnum mosses, spore production in natural populations is relatively rare, with studies finding only about 2% of patches producing spore capsules. However, as a bisexual species (i.e. monoecious–having both male and female reproductive structures on the same plant), S.squarrosum has greater potential for successful spore production compared to unisexual Sphagnum species that require separate male and female plants in close proximity.[13] With approximately 243,000 spores per capsule, S.squarrosum produces the highest recorded spore output among Sphagnum species. This high spore output is achieved through a combination of large capsules (averaging about 6 cubic millimetres in volume) and relatively small spores (averaging 25.8 micrometres (μm) in diameter, with individual spores ranging from 19–30μm). The size of spores shows a curvilinear relationship with capsule size, with larger capsules generally producing more spores.[14]
The species uses an 'air gun' dispersal mechanism, where spores are explosively discharged from the capsule by built-up air pressure (approximately 5 x 105Pascals) during warm, dry conditions. S.squarrosum's large capsules are particularly effective at this dispersal method, shooting spores higher into the air than smaller-capsuled species. Studies have shown that only about 7% of spores remain in the capsule after discharge, with another 7% landing within the parent colony. The fact that only about 11% of released spores are found within 3.2m (10ft) of the parent plant suggests the majority of spores are capable of long-distance dispersal.[15]
As a bisexual species, S.squarrosum produces both male and female reproductive structures on the same plant. Studies have found no evidence of reduced fitness in self-fertilised plants.[16]
Evolutionary history
Genetic studies of European populations suggest that S.squarrosum likely survived the Last Glacial Maximum in multiple scattered refugia across Europe. Analysis of chloroplast DNA variation indicates that the species maintained relatively stable historical population sizes and experienced less severe population bottlenecks during glacial periods compared to some other Sphagnum species.[7]
The species has high genetic diversity, with widely distributed haplotypes and weak geographic structure in Europe. This pattern suggests the species spread from multiple refugial populations after the last ice age. Some genetic variants found at higher altitudes may represent ancient lineages that were once more widely distributed across European tundra following the LGM.[7]
Fossil history
Sphagnum squarrosum appears in late Holocene subfossil records, notably from peat deposits in northwestern Iran dating back approximately 2,000 years. These subfossil remains were found in the Tuska Tchal peat bog at an elevation of 1,034m (3,392ft), where evidence suggests the species was continuously present from around 2,000 years ago through to recent times. The presence of associated sphagnophilous (Sphagnum-loving) organisms in these deposits, including the fungus Geoglossum sphagnophilum and the rotifer species Habrotrocha angusticollis, further confirms the long-term presence of S.squarrosum at this site.[17]
These Iranian subfossil populations may be glacial relicts from the last ice age, when colder conditions allowed the species to expand its range southward. Such findings suggest that highland wetland sites in Iran served as interglacial refugia for boreal species like S.squarrosum.[17]
Associated species
Sphagnum squarrosum is a specific host to the fungal parasite Discinella schimperi (formerly Helotium schimperi), which infects specialised mucilaginous cells in the moss. The fungus is highly host-specific, infecting only S.squarrosum and not other Sphagnum species. Herbarium studies indicate that about half or more of examined S.squarrosum samples were infected with D.schimperi, suggesting a widespread relationship between the two species. The fungus has been documented throughout the moss's circumpolar range, including North America, Europe, and Asia.[8]
Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophytasensu stricto. Bryophyta may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hornworts. Mosses typically form dense green clumps or mats, often in damp or shady locations. The individual plants are usually composed of simple leaves that are generally only one cell thick, attached to a stem that may be branched or unbranched and has only a limited role in conducting water and nutrients. Although some species have conducting tissues, these are generally poorly developed and structurally different from similar tissue found in vascular plants. Mosses do not have seeds and after fertilisation develop sporophytes with unbranched stalks topped with single capsules containing spores. They are typically 0.2–10 cm (0.1–3.9 in) tall, though some species are much larger. Dawsonia, the tallest moss in the world, can grow to 50 cm (20 in) in height. There are approximately 12,000 species.
Sphagnum is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, also bog moss and quacker moss. Accumulations of Sphagnum can store water, since both living and dead plants can hold large quantities of water inside their cells; plants may hold 16 to 26 times as much water as their dry weight, depending on the species. The empty cells help retain water in drier conditions.
Helodium blandowii, also known as Blandow's helodium moss, Blandow's tamarisk-moss, Blandow's bogmoss, and Blandow's feathermoss, is a common moss species in Europe. It is also found in North America, Central Asia and Greenland. It is considered rare plant in the Western U.S., including Oregon and California. It occurs all around the northern hemisphere in higher latitudes, and in some places is not as rare as in the Western U.S.
Meesia triquetra, the three-ranked hump-moss, is a moss that occurs all around the northern hemisphere in higher latitudes.
Meesia uliginosa, the broad-nerved hump-moss, is a rare moss of the Western U.S. It occurs all around the northern hemisphere in higher latitudes, and in some places is not as rare as in the Western U.S.
Aulacomnium palustre, the bog groove-moss or ribbed bog moss, is a moss that is nearly cosmopolitan in distribution. It occurs in North America, Hispaniola, Venezuela, Eurasia, and New Zealand. In North America, it occurs across southern arctic, subboreal, and boreal regions from Alaska and British Columbia to Greenland and Quebec. Documentation of ribbed bog moss's distribution in the contiguous United States is probably incomplete. It is reported sporadically south to Washington, Wyoming, Georgia, and Virginia.
Sphagnum palustre, the prairie sphagnum or blunt-leaved bogmoss, is a species of peat moss from the genus Sphagnum, in the family Sphagnaceae. Like other mosses of this type it can soak up water up to the 30-fold amount of its own dry weight thanks to its elastic spiral fibers. S. palustre is rather frequent and is spread almost all over the world. It mainly grows in wet forests and—compared to other specimens of this genus—rarely grows in moors.
Sphagnum capillifolium, the red bogmoss, northern peat moss, acute-leaved bog-moss, or small red peat moss, is a species of peat moss native to Canada, the northern United States, Greenland, and Europe. Small red peat moss can be distinguished by its sweeping, outward-curving branches that resemble tresses. Sphagnum moss can hold large amounts of water within its cells, up to 20 times its own weight. This capability is due to its dead, empty cells called hyaline cells that fill up with water. This allows the moss to survive in wet, boggy habitats around rivers and lakes.
Lysimachia asperulifolia is a rare species of flowering plant in the Primulaceae known by the common name rough-leaved loosestrife and roughleaf yellow loosestrife. It is endemic to the Atlantic coastal plain in North Carolina and northern South Carolina in the United States, where there are 64 known populations. It is a federally listed endangered species of the United States.
Inocybe saliceticola is a fungus found in moist habitats in the Nordic countries. The species produces brown mushrooms with caps of varying shapes up to 40 millimetres (1.6 in) across, and tall, thin stems up to 62 mm (2.4 in) long. At the base of the stem is a large and well-defined "bulb". The species produces unusually shaped, irregular spores, each with a few thick protrusions. This feature helps differentiate it from other species that would otherwise be similar in appearance and habit.
Polytrichastrum alpinum, also known as Alpine haircap, is a species of moss from the family Polytrichaceae. It is widely distributed and may be found growing among other moss species.
Sphagnum fuscum, the rusty bogmoss or rusty peat moss, is a peat moss found commonly in Norway and Sweden, and can be found scattered across North America, the United Kingdom, and in southern to eastern Europe.
Sphagnum cuspidatum, the feathery bogmoss, toothed sphagnum, or toothed peat moss, is a peat moss found commonly in Great Britain, Norway, Sweden, the eastern coast of the United States, and in Colombia.
Polytrichum strictum, commonly known as bog haircap moss or strict haircap, is an evergreen and perennial species of moss native to Sphagnum bogs and other moist habitats in temperate climates. It has a circumboreal distribution, and is also found in South America and Antarctica.
Sphagnum papillosum, the papillose peatmoss, is a species of peat moss distributed throughout the northern hemisphere. Although sometimes confused with Sphagnum imbricatum and Sphagnum palustre, it is distinguished by its yellow-green to brown short, blunt branches and papillose chlorophyllose cells.
Trichocolea tomentella is a species of liverwort belonging to the family Trichocoleaceae. It forms loose, pale green to yellowish-white mats and is characterised by its highly divided leaves that give it a feathery or fuzzy appearance. The species has a wide distribution across temperate regions of the Northern Hemisphere, particularly in oceanic and suboceanic areas, occurring in Europe, Asia, North Africa, and eastern North America. It typically grows in moist, shaded locations, especially near springs and streams in deciduous and coniferous forests. While capable of sexual reproduction, with male and female structures on separate plants, it reproduces predominantly through vegetative means via branching and fragmentation. The species shows considerable morphological variation between populations but maintains stable taxonomic characteristics in its cell structure. Though it can form extensive pure patches and effectively compete with other bryophytes in suitable habitats, T. tomentella faces threats from habitat destruction, particularly through logging and drainage of its preferred moist forest habitats.
Sphagnum wulfianum, commonly known as Wulf's peatmoss, is a species of moss belonging to the family Sphagnaceae. It has a circumboreal distribution, occurring primarily in moist boreal forest environments across Eurasia and North America, with rare occurrences in Arctic tundra regions. The species is morphologically distinctive among peat mosses, characterised by having six to twelve branches per fascicle, a unique feature in the genus. First described in 1860 from Estonia, it typically grows in boggy mineral-rich spruce forests and at the borders of mires, forming small carpets and hummocks. While the species has a large geographic range and stable overall population trends in Europe, it faces regional conservation challenges, particularly at its range edges, where it is considered endangered in several countries due to habitat degradation from peatland drainage, forestry, and agriculture. Genetic studies indicate remarkably low genetic diversity throughout its range, suggesting high rates of gene flow across large distances despite infrequent spore production.
Dicranoloma billardierei is a species of bryophyte in the genus Dicranoloma. This moss is extremely common in wet rainforest habitats. In the field, Dicranoloma billardierei, is often confused with Dicranoloma robustum and Dicranoloma fasciatum. However, the short and obtuse nature of the leaves make this moss normally very distinctive.
Sphagnum australe is a species of Sphagnum found in southeastern Australia.
Sphagnum teres, or rigid bogmoss, is a species of moss from the Sphagnaceae family. Widely distributed in the Northern Hemisphere, it grows in mountainous areas in the southern part of its range. It thrives in fertile, minerotrophic peatlands. It is characterized by a clearly visible terminal bud in the middle part of the head and usually a dark brown stem.
↑ Crome, Georg Ernst Wilhelm (1803). Sammlung deutscher Laub-Moose (in German). Schwerin: Gedruckt mit Bärensprungschen Schriften. p.24.
1 2 3 Isoviita, Pekka (1966). "Studies on Sphagnum L. I. Nomenclatural revision of the European taxa". Annales Botanici Fennici. 3 (2): 199–264. JSTOR23724595.
↑ Sundberg, Sebastian; Hansson, Jesper; Rydin, Håkan (2006). "Colonization of Sphagnum on land uplift islands in the Baltic Sea: time, area, distance and life history". Journal of Biogeography. 33 (8): 1479–1491. doi:10.1111/j.1365-2699.2006.01520.x.
↑ Sundberg, Sebastian; Rydin, Håkan (1998). "Spore number in Sphagnum and its dependence on spore and capsule size". Journal of Bryology. 20 (1): 1–16. doi:10.1179/jbr.1998.20.1.1.
↑ Sundberg, Sebastian (2005). "Larger capsules enhance short-range spore dispersal in Sphagnum, but what happens further away?". Oikos. 108 (1): 115–124. doi:10.1111/j.0030-1299.2005.12916.x.
↑ Johnson, Matthew G.; Shaw, A. Jonathan (2015). "Genetic diversity, sexual condition, and microhabitat preference determine mating patterns in Sphagnum (Sphagnaceae) peat-mosses". Biological Journal of the Linnean Society. 115 (1): 96–113. doi:10.1111/bij.12497.
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