Sphagnum Papillosum
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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, peat moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store wa ...
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 as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
. 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 may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
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 ( grc, πηκτικός ': "congealed" and "curdled") is a heteropolysaccharide, a structural acid contained in the primary lamella, in the middle lamella, and in the cell walls of terrestrial plants. The principal, chemical component of ...
-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 wa ...
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 or stops their growth. Antimicrobial medicines can be grouped according to the microorganisms they act primarily against. For example, antibiotics are used against bacteria, and antifungals ar ...
and preservative properties of
Sphagnum ''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, peat moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store wa ...
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 planet's highest peaks, including the very highest, Mount Everest. Over 100 ...
.


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 wetlands along with marshes, swamps, and bogs. Bogs and fens, both peat-forming ecosystems, are also known as mires. T ...
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 under gravitational pull from clouds. The main forms of precipitation include drizzle, rain, sleet, snow, ice pellets, graupel and hail. ...
; therefore, nutrients such as
nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
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 proces ...
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 at a rate faster than that at which the substance is lost or eliminated ...
metals such as
cadmium Cadmium is a chemical element with the symbol Cd and atomic number 48. This soft, silvery-white metal is chemically similar to the two other stable metals in group 12, zinc and mercury. Like zinc, it demonstrates oxidation state +2 in most of ...
,
chromium Chromium is a chemical element with the symbol Cr and atomic number 24. It is the first element in group 6. It is a steely-grey, lustrous, hard, and brittle transition metal. Chromium metal is valued for its high corrosion resistance and hardne ...
, and
zinc Zinc is a chemical element with the symbol Zn and atomic number 30. Zinc 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 periodi ...
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 where archegonia and antheridia are produced on separate gametophytes. It is one of the two main sexual systems in bryophytes. Both dioicous () and monoicous gametophytes produce gametes in gametangia by mitosis rathe ...
. 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. Androecium is also t ...
and
archegonia An archegonium (pl: 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 ga ...
occur in late summer to early autumn.
Fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proce ...
occurs in spring.


Sporophyte

The
sporangium A sporangium (; from Late Latin, ) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their life cy ...
mature in mid to late summer and are relatively spherical with numerous pseudostomata. The
spores In biology, a spore is a unit of sexual 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 plants, algae, f ...
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 or adsorption from the surrounding environment, which is usually at normal or room temperature. If water molecules become suspended among the substance ...
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 fu ...
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 is the branch of agriculture that deals with the art, science, technology, and business of plant cultivation. It includes the cultivation of fruits, vegetables, nuts, seeds, herbs, sprouts, mushrooms, algae, flowers, seaweeds and no ...
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 that i ...
. 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 muskeg; a ...
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; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
classifies the species as
Least concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...
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