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''Sphagnum papillosum'', the papillose peatmoss, is a species of
peat moss ''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store water, since ...
distributed throughout the
northern hemisphere The Northern Hemisphere is the half of Earth that is north of the equator. For other planets in the Solar System, north is defined by humans as being in the same celestial sphere, celestial hemisphere relative to the invariable plane of the Solar ...
. 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.


Taxonomy

''S. papillosum'' is classified under section Sphagnum of the genus, which also includes the species ''S. magellanicum'', ''S. palustre'', and ''S. austinii''. It was first described by Lindberg in 1872 and typified in 1913. It has previously been described in 1907 from specimens in Japan as ''S. hakkodense'' Warnst. & Card.; however, this was later identified to be a synonym and an isotype specimen of ''S. papillosum''. Other synonyms include ''S. immersum'' Nees & Hornsch and ''S. waghornei'' Warnst.


Description


Morphology

''S. papillosum'' is distinguished by its robust, golden brown capitulum. It usually has 4 branches: 2 short and blunt divergent (spreading) branches and 2, sometimes 3, short
pendent Pendent is an adjective that describes the condition of hanging, either literally, or figuratively, as in undecided or incomplete. The word is to be distinguished from the spelling "pendant", which is the noun. * In botany and anatomy the term ...
branches. The central cylinder or “wood” of the
stem Stem or STEM most commonly refers to: * Plant stem, a structural axis of a vascular plant * Stem group * Science, technology, engineering, and mathematics Stem or STEM can also refer to: Language and writing * Word stem, part of a word respon ...
is usually dark brown to almost black in colour and sometimes green. The stem leaves are approximately 1.3 mm long and 0.7 mm wide, usually rectangular or spatulate in shape with a resorbed or fringed abaxial surface, inconspicuous on the adaxial surface. The branch leaves are spreading and can be up to 2 mm long and 1 mm wide. They are ovate to broadly ovate in shape and usually strongly concave. ''S. papillosum'' possesses a
pectin Pectin ( ': "congealed" and "curdled") is a heteropolysaccharide, a structural polymer contained in the primary lamella, in the middle lamella, and in the cell walls of terrestrial plants. The principal chemical component of pectin is galact ...
-like
polysaccharide Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long-chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with wat ...
called Sphagnan, Sphagnum acid, and phenolic compounds in its leaves that have been observed to contribute to known
antimicrobial An antimicrobial is an agent that kills microorganisms (microbicide) or stops their growth (bacteriostatic agent). Antimicrobial medicines can be grouped according to the microorganisms they are used to treat. For example, antibiotics are used aga ...
and preservative properties of
Sphagnum ''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store water, since ...
mosses.


Cellular structure

The stem cortical cells of ''S. papillosum'' are fibrillose with 1-2 pores. The branch cortical cells are also strongly fibrillose. The hyaline cells that make up the leaves are never ornamented, fairly wide, and often divided. There is usually a single pore present at the upper apex of the cell on the adaxial side, while there are many round to elliptic pores along the commissures of the abaxial side. The internal commissural walls appear to be rough or papillose due to cell wall projections that form papillae, a distinguishing feature of this Sphagnum moss. The chlorophyllose cells are rather small compared to other species and are trapezoidal to barrel-shaped in cross section. They are usually equally exposed on both sides or slightly more exposed on the adaxial side of the leaf.


Distribution

''S. papillosum'' is widely distributed throughout North America, Europe, and Asia. It is especially common in western and northern Europe and can extend as far south as the
Himalaya mountains The Himalayas, or Himalaya ( ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the Earth's highest peaks, including the highest, Mount Everest. More than 100 peak ...
.


Habitat and ecology

''S. papillosum'' is oligo-mesotrophic, commonly found in shaded, nutrient-poor
fen A fen is a type of peat-accumulating wetland fed by mineral-rich ground or surface water. It is one of the main types of wetland along with marshes, swamps, and bogs. Bogs and fens, both peat-forming ecosystems, are also known as mires ...
habitats to open, acidic peatlands. Tends to form hummocks and dense carpets. It can also be found at low to mid-elevations near streams, flushes or transitional mires. These types of habitats are mainly supplied by
precipitation In meteorology, precipitation is any product of the condensation of atmospheric water vapor that falls from clouds due to gravitational pull. The main forms of precipitation include drizzle, rain, rain and snow mixed ("sleet" in Commonwe ...
; therefore, nutrients such as
nitrogen Nitrogen is a chemical element; it has Symbol (chemistry), symbol N and atomic number 7. Nitrogen is a Nonmetal (chemistry), nonmetal and the lightest member of pnictogen, group 15 of the periodic table, often called the Pnictogen, pnictogens. ...
are particularly low and water availability is dependent on rates of rainfall, which both limits the growth and
productivity Productivity is the efficiency of production of goods or services expressed by some measure. Measurements of productivity are often expressed as a ratio of an aggregate output to a single input or an aggregate input used in a production proce ...
of ''S. papillosum''. Although ''S. papillosum'' favours habitats with consistent precipitation, it is relatively tolerant to water stress. Like other Sphagnum mosses, ''S. papillosum'' is sensitive to minerals and nutrients in the water and has been observed to
bioaccumulate Bioaccumulation is the gradual accumulation of substances, such as pesticides or other chemicals, in an organism. Bioaccumulation occurs when an organism absorbs a substance faster than it can be lost or eliminated by catabolism and excretion. Th ...
metals such as
cadmium Cadmium is a chemical element; it has chemical symbol, symbol Cd and atomic number 48. This soft, silvery-white metal is chemically similar to the two other stable metals in group 12 element, group 12, zinc and mercury (element), mercury. Like z ...
,
chromium Chromium is a chemical element; it has Symbol (chemistry), symbol Cr and atomic number 24. It is the first element in Group 6 element, group 6. It is a steely-grey, Luster (mineralogy), lustrous, hard, and brittle transition metal. Chromium ...
, and
zinc Zinc is a chemical element; it has symbol Zn and atomic number 30. It is a slightly brittle metal at room temperature and has a shiny-greyish appearance when oxidation is removed. It is the first element in group 12 (IIB) of the periodic tabl ...
in its tissues. The metal concentrations in ''S. papillosum'' has been observed to be a direct reflection of concentrations in its natural waters; therefore, it presents itself as an effective
bioindicator A bioindicator is any species (an indicator species) or group of species whose function, population, or status can reveal the qualitative status of the environment. The most common indicator species are animals. For example, copepods and other sma ...
of metal concentrations present in the environment.


Life cycle


Gametophyte

''S. papillosum'' is
dioicous Dioicy () is a sexual system in non-vascular plants where archegonia (female organs) and antheridia (male organs) are produced on separate plants in the gametophyte phase. It is one of the two main sexual systems in bryophytes, the other being ...
. Its antheridial and archegonial branches are morphologically similar to its vegetative branches. The formation of
antheridia An antheridium is a haploid structure or organ producing and containing male gametes (called ''antherozoids'' or sperm). The plural form is antheridia, and a structure containing one or more antheridia is called an androecium. The androecium is al ...
and
archegonia An archegonium (: archegonia), from the Ancient Greek ''ἀρχή'' ("beginning") and ''γόνος'' ("offspring"), is a multicellular structure or organ of the gametophyte phase of certain plants, producing and containing the ovum or female gamet ...
occur in late summer to early autumn.
Fertilization Fertilisation or fertilization (see American and British English spelling differences#-ise, -ize (-isation, -ization), spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give ...
occurs in spring.


Sporophyte

The
sporangium A sporangium (from Late Latin, ; : sporangia) is an enclosure in which spores are formed. It can be composed of a unicellular organism, single cell or can be multicellular organism, multicellular. Virtually all plants, fungus, fungi, and many ot ...
mature in mid to late summer and are relatively spherical with numerous pseudostomata. The
spores In biology, a spore is a unit of sexual (in fungi) or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plant ...
are dispersed all at once when the sporangium matures through an explosive,
hygroscopic Hygroscopy is the phenomenon of attracting and holding water molecules via either absorption (chemistry), absorption or adsorption from the surrounding Natural environment, environment, which is usually at normal or room temperature. If water mol ...
mechanism triggered by warm and dry conditions. The spores are discharged into the air and dispersed by the wind. Its spores are 26-36 μm in diameter and yellow brown in colour. They are finely papillose with rounded, triradiate ridges on the distal surface.


Asexual reproduction

''S. papillosum'' can also establish through
asexual reproduction Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the f ...
by the fragmentation of its stems and branches.


Uses and economic importance

''S. papillosum'' is a major peat-forming moss, dominant in peatlands that are mined for
horticultural Horticulture (from ) is the art and science of growing fruits, vegetables, flowers, trees, shrubs and ornamental plants. Horticulture is commonly associated with the more professional and technical aspects of plant cultivation on a smaller and mo ...
purposes, as peat is a popular growing medium for
ornamental plants Ornamental plants or ''garden plants'' are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars th ...
. An alternative to peat mining is Sphagnum farming, which is the commercial practice of cultivating and harvesting Sphagnum moss. In particular, ''S. papillosum'' has been observed to establish well on restored
bog A bog or bogland is a wetland that accumulates peat as a deposit of dead plant materials often mosses, typically sphagnum moss. It is one of the four main types of wetlands. Other names for bogs include mire, mosses, quagmire, and musk ...
grasslands that could potentially be used for Sphagnum farming. ''S. papillosum'' has also been observed to regenerate after cutting when tested for harvesting techniques; therefore, it may provide a potentially renewable and environmentally friendly alternative to peat mining.


Conservation

The
IUCN The International Union for Conservation of Nature (IUCN) is an international organization working in the field of nature conservation and sustainable use of natural resources. Founded in 1948, IUCN has become the global authority on the status ...
classifies the species as
Least concern A least-concern species is a species that has been evaluated and categorized by the International Union for Conservation of Nature (IUCN) as not being a focus of wildlife conservation because the specific species is still plentiful in the wil ...
due to its common occurrence across its wide geographic range, but several local populations (e.g. in Romania, Slovakia and Serbia) are considered threatened.


References

{{Taxonbar, from=Q958193 papillosum Taxobox binomials not recognized by IUCN