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'''''Sphagnum papillosum''''', the '''papillose peatmoss''', is a species of [[peat moss]] with a [[Nearctic]] distribution.<ref>{{BioRef|Tropicos|title=''Sphagnum papillosum'' Lindb.|id=35174721|access-date=11 April 2020}}</ref>
'''''Sphagnum papillosum''''', the '''papillose peatmoss''', is a species of [[peat moss]] distributed throughout the [[northern hemisphere]].<ref>{{BioRef|Tropicos|title=''Sphagnum papillosum'' Lindb.|id=35174721|access-date=11 April 2020}}</ref> Although sometimes confused with ''Sphagnum imbricatum'' and ''[[Sphagnum magellanicum]]'', it is distinguished by its yellow-green to brown short, blunt branches and papillose chlorophyllose cells.<ref name=H.M.S.O>{{cite book |last1=H.M.S.O |title=Handbook of European sphagna |date=1990 |location=London}}</ref>

''S. papillosum'' possesses a [[pectin]]-like [[polysaccharide]] called Sphagnan, Sphagnum acid, and phenolic compounds in its leaves that have been observed to contribute to known [[antimicrobial]] and preservative properties of [[Sphagnum]] mosses. <ref>{{cite journal |last1=Painter |first1=Terence J. |title=Lindow man, tollund man and other peat-bog bodies: The preservative and antimicrobial action of Sphagnan, a reactive glycuronoglycan with tanning and sequestering properties |journal=Carbohydrate Polymers |date=1991 |volume=15 |issue=2 |pages=123–142 |doi=10.1016/0144-8617(91)90028-b}}</ref><ref>{{cite journal |last1=Mellegård |first1=H. |last2=Stalheim |first2=T. |last3=Hormazabal |first3=V. |last4=Granum |first4=P.E. |last5=Hardy |first5=S.P. |title=Antibacterial activity of sphagnum acid and other phenolic compounds found in against food-borne bacteria |journal=Letters in Applied Microbiology |date=July 2009 |volume=49 |issue=1 |pages=85–90 |doi=10.1111/j.1472-765x.2009.02622.x}}</ref>

==Geographic Distribution==
S. papillosum is widely distributed throughout North America, Europe, and Asia.<ref name=H.M.S.O /> It is especially common in western and northern Europe and can extend as far south as the [[Himalaya mountains]].<ref name=H.M.S.O />

==Habitat and Ecology==

''S. papillosum'' is oligo-mesotrophic, commonly found in shaded, nutrient-poor [[fen]] habitats to open, acidic peatlands.<ref name=H.M.S.O /> Tends to form hummocks and dense carpets.<ref name=H.M.S.O /> It can also be found at low to mid-elevations near streams, flushes or transitional [[mires]].<ref name=H.M.S.O /> These types of habitats are mainly supplied by [[precipitation]]; therefore, nutrients such as [[nitrogen]] are particularly low and water availability is dependent on rates of rainfall, which both limits the growth and [[productivity]] of ''S. papillosum''.<ref>{{cite journal |last1=Aerts |first1=Rien |last2=Wallen |first2=Bo |last3=Malmer |first3=Nils |title=Growth-Limiting Nutrients in Sphagnum-Dominated Bogs Subject to Low and High Atmospheric Nitrogen Supply |journal=The Journal of Ecology |date=March 1992 |volume=80 |issue=1 |pages=131 |doi=10.2307/2261070}}</ref><ref>{{cite journal |last1=Vitousek |first1=Peter M. |last2=Aber |first2=John D. |last3=Howarth |first3=Robert W. |last4=Likens |first4=Gene E. |last5=Matson |first5=Pamela A. |last6=Schindler |first6=David W. |last7=Schlesinger |first7=William H. |last8=Tilman |first8=David G. |title=HUMAN ALTERATION OF THE GLOBAL NITROGEN CYCLE: SOURCES AND CONSEQUENCES |journal=Ecological Applications |date=August 1997 |volume=7 |issue=3 |pages=737–750 |doi=10.1890/1051-0761(1997)007[0737:HAOTGN]2.0.CO;2}}</ref> Although S. papillosum favours habitats with consistent precipitation, it is relatively tolerant to water stress.<ref>{{cite journal |last1=Clymo |first1=R. S. |last2=Hayward |first2=P. M. |title=The Ecology of Sphagnum |journal=Bryophyte Ecology |date=1982 |page=229-289 |doi=10.1007/978-94-009-5891-3_8}}</ref>

Like other Sphagnum mosses, ''S. papillosum'' is sensitive to minerals and nutrients in the water and has been observed to [[bioaccumulate]] metals such as [[cadmium]], [[chromium]], and [[zinc]] in its tissues.<ref name=Gstoettner>{{cite journal |last1=Gstoettner |first1=E. M. |last2=Fisher |first2=N. S. |title=Accumulation of cadmium, chromium, and zinc by the mossSphagnum papillosum Lindle |journal=Water, Air, & Soil Pollution |date=January 1997 |volume=93 |issue=1-4 |pages=321–330 |doi=10.1007/bf02404764}}</ref> 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]] of metal concentrations present in the environment.<ref name=Gstoettner />

==Description==
===Morphology===
''S. papillosum'' is distinguished by its robust, golden brown capitulum.<ref name=H.M.S.O /> It usually has 4 branches comprised of 2 short and blunt divergent (spreading) branches and 2, sometimes 3, short [[pendent]] branches. <ref name=H.M.S.O /><ref name=eflora>{{cite web |title=Sphagnum papillosum in Flora of North America @ efloras.org |url=http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=200000821 |website=www.efloras.org}}</ref> The central cylinder or “wood” of the [[stem]] is usually dark brown to almost black in colour and sometimes green.<ref name=H.M.S.O /> 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.<ref name=H.M.S.O /><ref name=eflora /> The branch leaves are spreading and can be up to 2 mm long and 1 mm wide.<ref name=H.M.S.O /><ref name=eflora /> They are ovate to broadly ovate in shape and usually strongly concave.<ref name=H.M.S.O /><ref name=eflora />

===Cellular Structure===
The stem [[cortical]] cells of ''S. papillosum'' are fibrillose with 1-2 pores.<ref name=H.M.S.O /><ref name=eflora /> The branch cortical cells are also strongly fibrillose.<ref name=H.M.S.O /> The hyaline cells that make up the leaves are never ornamented, fairly wide, and often divided.<ref name=H.M.S.O /> 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.<ref name=H.M.S.O /><ref name=eflora /> 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.<ref name=H.M.S.O /> The chlorophyllose cells are rather small compared to other species and are trapezoidal to barrel-shaped in cross section.<ref name=H.M.S.O /> They are usually equally exposed on both sides or slightly more exposed on the adaxial side of the leaf.<ref name=H.M.S.O /><ref name=eflora />

==Life Cycle==
===Gametophyte===
''S. papillosum'' is [[dioicous]].<ref name=H.M.S.O /> Its antheridial and archegonial branches are morphologically similar to the vegetative branches.<ref name=H.M.S.O /> The formation of [[antheridia]] and [[archegonia]] occur in late summer to early autumn.<ref name=Cronberg>{{cite journal |last1=Cronberg |first1=N. |title=Reproductive biology of Sphagnum |journal=Lindbergia |date=1993 |issue=17 |page=69-82}}</ref> [[Fertilization]] occurs in spring. <ref name=Cronberg />

===Sporophyte===
The [[sporangium]] mature in mid to late summer and are relatively spherical with numerous pseudostomata.<ref name=eflora /> The [[spores]] are dispersed all at once when the sporangium matures through an explosive, [[hygroscopic]] mechanism triggered by warm and dry conditions.<ref name=Cronberg /> 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.<ref name=Cao>{{cite journal |last1=Cao |first1=T. |last2=Vitt |first2=D.H. |title=Spore surface structure of Sphagnum |journal=Nova hedwigia |date=1986 |volume=43 |issue=1-2 |page=191-220}}</ref> They are finely papillose with rounded, triradiate ridges on the distal surface.<ref name=H.M.S.O /><ref name=Cao />

===Asexual Reproduction===
''S. papillosum'' can also establish through [[asexual reproduction]] by the [[fragmentation]] of its stems and branches.<ref>{{cite journal |last1=Longton |first1=R.E. |title=Reproductive biology and life-history strategies |journal=Bryology for the Twenty-First Century |date=2018 |page=369–369 |doi=10.1201/9781315138626-26}}</ref><ref name=Gaudig>{{cite journal |last1=Gaudig |first1=G. |last2=Krebs |first2=M. |last3=Joosten |first3=H. |title=Sphagnum farming on cut-over bog in NW Germany: Long-term studies on Sphagnum growth |journal=Mires and Peat |date=2017 |volume=20 |issue=4 |page=1-19 |doi=10.19189/MaP.2016.OMB.238}}</ref>

==Classification==
''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.<ref name=Isoviita>{{cite journal |last1=Isovitta |first1=P. |title=Studies on Sphagnum L. I. Nomenclatural revision of the European taxa |journal=Annales Botanici Fennici |date=1966 |volume=3 |issue=2 |pages=199-264}}</ref> 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 of ''S. papillosum'' and was an isotype specimen.<ref name=Isoviita /> Other synonyms include ''S. immersum'' Nees & Hornsch and ''S. waghornei'' Warnst.<ref name=Isoviita />

==Uses and Economic Importance==
''S. papillosum'' is a major peat-forming moss, dominant in peatlands that are mined for [[horticultural]] purposes, as peat is a popular growing medium for [[ornamental plants]].<ref>{{cite journal |last1=Emmel |first1=M. |title=GROWING ORNAMENTAL PLANTS IN SPHAGNUM BIOMASS |journal=Acta Horticulturae |date=January 2008 |issue=779 |pages=173–178 |doi=10.17660/actahortic.2008.779.20}}</ref><ref name=Gaudig /> An alternative to peat mining is Sphagnum farming, which is the commercial practice of cultivating and harvesting Sphagnum moss.<ref name=Gaudig /> In particular, ''S. papillosum'' has been observed to establish well on restored [[bog]] grasslands that could potentially be used for Sphagnum farming.<ref name=Gaudig /> ''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.<ref name=Gaudig /><ref>{{cite journal |last1=Krebs |first1=M. |last2=Gaudig |first2=G. |last3=Matchutadze |first3=I. |last4=Joosten |first4=H. |title=Sphagnum regrowth after cutting |journal=Mires and Peat |date=2018 |volume=20 |issue=12 |pages=1-20 |doi=10.19189/MaP.2017.OMB.298}}</ref>


==References==
==References==

Revision as of 21:50, 12 April 2020

Sphagnum papillosum
Scientific classification Edit this classification
Kingdom: Plantae
Division: Bryophyta
Class: Sphagnopsida
Subclass: Sphagnidae
Order: Sphagnales
Family: Sphagnaceae
Genus: Sphagnum
Species:
S. papillosum
Binomial name
Sphagnum papillosum

Sphagnum papillosum, the papillose peatmoss, is a species of peat moss distributed throughout the northern hemisphere.[1] Although sometimes confused with Sphagnum imbricatum and Sphagnum magellanicum, it is distinguished by its yellow-green to brown short, blunt branches and papillose chlorophyllose cells.[2]

S. papillosum possesses a pectin-like polysaccharide called Sphagnan, Sphagnum acid, and phenolic compounds in its leaves that have been observed to contribute to known antimicrobial and preservative properties of Sphagnum mosses. [3][4]

Geographic Distribution

S. papillosum is widely distributed throughout North America, Europe, and Asia.[2] It is especially common in western and northern Europe and can extend as far south as the Himalaya mountains.[2]

Habitat and Ecology

S. papillosum is oligo-mesotrophic, commonly found in shaded, nutrient-poor fen habitats to open, acidic peatlands.[2] Tends to form hummocks and dense carpets.[2] It can also be found at low to mid-elevations near streams, flushes or transitional mires.[2] These types of habitats are mainly supplied by precipitation; therefore, nutrients such as nitrogen are particularly low and water availability is dependent on rates of rainfall, which both limits the growth and productivity of S. papillosum.[5][6] Although S. papillosum favours habitats with consistent precipitation, it is relatively tolerant to water stress.[7]

Like other Sphagnum mosses, S. papillosum is sensitive to minerals and nutrients in the water and has been observed to bioaccumulate metals such as cadmium, chromium, and zinc in its tissues.[8] 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 of metal concentrations present in the environment.[8]

Description

Morphology

S. papillosum is distinguished by its robust, golden brown capitulum.[2] It usually has 4 branches comprised of 2 short and blunt divergent (spreading) branches and 2, sometimes 3, short pendent branches. [2][9] The central cylinder or “wood” of the stem is usually dark brown to almost black in colour and sometimes green.[2] 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.[2][9] The branch leaves are spreading and can be up to 2 mm long and 1 mm wide.[2][9] They are ovate to broadly ovate in shape and usually strongly concave.[2][9]

Cellular Structure

The stem cortical cells of S. papillosum are fibrillose with 1-2 pores.[2][9] The branch cortical cells are also strongly fibrillose.[2] The hyaline cells that make up the leaves are never ornamented, fairly wide, and often divided.[2] 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.[2][9] 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.[2] The chlorophyllose cells are rather small compared to other species and are trapezoidal to barrel-shaped in cross section.[2] They are usually equally exposed on both sides or slightly more exposed on the adaxial side of the leaf.[2][9]

Life Cycle

Gametophyte

S. papillosum is dioicous.[2] Its antheridial and archegonial branches are morphologically similar to the vegetative branches.[2] The formation of antheridia and archegonia occur in late summer to early autumn.[10] Fertilization occurs in spring. [10]

Sporophyte

The sporangium mature in mid to late summer and are relatively spherical with numerous pseudostomata.[9] The spores are dispersed all at once when the sporangium matures through an explosive, hygroscopic mechanism triggered by warm and dry conditions.[10] 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.[11] They are finely papillose with rounded, triradiate ridges on the distal surface.[2][11]

Asexual Reproduction

S. papillosum can also establish through asexual reproduction by the fragmentation of its stems and branches.[12][13]

Classification

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.[14] 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 of S. papillosum and was an isotype specimen.[14] Other synonyms include S. immersum Nees & Hornsch and S. waghornei Warnst.[14]

Uses and Economic Importance

S. papillosum is a major peat-forming moss, dominant in peatlands that are mined for horticultural purposes, as peat is a popular growing medium for ornamental plants.[15][13] An alternative to peat mining is Sphagnum farming, which is the commercial practice of cultivating and harvesting Sphagnum moss.[13] In particular, S. papillosum has been observed to establish well on restored bog grasslands that could potentially be used for Sphagnum farming.[13] 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.[13][16]

References

  1. ^ "Sphagnum papillosum Lindb". Tropicos. Retrieved 11 April 2020.
  2. ^ a b c d e f g h i j k l m n o p q r s t u v H.M.S.O (1990). Handbook of European sphagna. London.{{cite book}}: CS1 maint: location missing publisher (link)
  3. ^ Painter, Terence J. (1991). "Lindow man, tollund man and other peat-bog bodies: The preservative and antimicrobial action of Sphagnan, a reactive glycuronoglycan with tanning and sequestering properties". Carbohydrate Polymers. 15 (2): 123–142. doi:10.1016/0144-8617(91)90028-b.
  4. ^ Mellegård, H.; Stalheim, T.; Hormazabal, V.; Granum, P.E.; Hardy, S.P. (July 2009). "Antibacterial activity of sphagnum acid and other phenolic compounds found in against food-borne bacteria". Letters in Applied Microbiology. 49 (1): 85–90. doi:10.1111/j.1472-765x.2009.02622.x.
  5. ^ Aerts, Rien; Wallen, Bo; Malmer, Nils (March 1992). "Growth-Limiting Nutrients in Sphagnum-Dominated Bogs Subject to Low and High Atmospheric Nitrogen Supply". The Journal of Ecology. 80 (1): 131. doi:10.2307/2261070.
  6. ^ Vitousek, Peter M.; Aber, John D.; Howarth, Robert W.; Likens, Gene E.; Matson, Pamela A.; Schindler, David W.; Schlesinger, William H.; Tilman, David G. (August 1997). "HUMAN ALTERATION OF THE GLOBAL NITROGEN CYCLE: SOURCES AND CONSEQUENCES". Ecological Applications. 7 (3): 737–750. doi:10.1890/1051-0761(1997)007[0737:HAOTGN]2.0.CO;2.
  7. ^ Clymo, R. S.; Hayward, P. M. (1982). "The Ecology of Sphagnum". Bryophyte Ecology: 229-289. doi:10.1007/978-94-009-5891-3_8.
  8. ^ a b Gstoettner, E. M.; Fisher, N. S. (January 1997). "Accumulation of cadmium, chromium, and zinc by the mossSphagnum papillosum Lindle". Water, Air, & Soil Pollution. 93 (1–4): 321–330. doi:10.1007/bf02404764.
  9. ^ a b c d e f g h "Sphagnum papillosum in Flora of North America @ efloras.org". www.efloras.org.
  10. ^ a b c Cronberg, N. (1993). "Reproductive biology of Sphagnum". Lindbergia (17): 69-82.
  11. ^ a b Cao, T.; Vitt, D.H. (1986). "Spore surface structure of Sphagnum". Nova hedwigia. 43 (1–2): 191-220.
  12. ^ Longton, R.E. (2018). "Reproductive biology and life-history strategies". Bryology for the Twenty-First Century: 369–369. doi:10.1201/9781315138626-26.
  13. ^ a b c d e Gaudig, G.; Krebs, M.; Joosten, H. (2017). "Sphagnum farming on cut-over bog in NW Germany: Long-term studies on Sphagnum growth". Mires and Peat. 20 (4): 1-19. doi:10.19189/MaP.2016.OMB.238.
  14. ^ a b c Isovitta, P. (1966). "Studies on Sphagnum L. I. Nomenclatural revision of the European taxa". Annales Botanici Fennici. 3 (2): 199–264.
  15. ^ Emmel, M. (January 2008). "GROWING ORNAMENTAL PLANTS IN SPHAGNUM BIOMASS". Acta Horticulturae (779): 173–178. doi:10.17660/actahortic.2008.779.20.
  16. ^ Krebs, M.; Gaudig, G.; Matchutadze, I.; Joosten, H. (2018). "Sphagnum regrowth after cutting". Mires and Peat. 20 (12): 1–20. doi:10.19189/MaP.2017.OMB.298.


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