Pleopeltis polypodioides

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Pleopeltis polypodioides
GreenPolypods.jpg
Pleopeltis polypodioides on an oak limb after a brief rain
PoliypodiumPolypoioides.jpg
Fronds on the same limb in desiccated state
Status TNC G5.svg
Secure  (NatureServe) [1]
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Division: Polypodiophyta
Class: Polypodiopsida
Order: Polypodiales
Suborder: Polypodiineae
Family: Polypodiaceae
Genus: Pleopeltis
Species:
P. polypodioides
Binomial name
Pleopeltis polypodioides
(L.) E.G.Andrews & Windham
Synonyms

Acrosticum polypodioidesL.

Pleopeltis polypodioides (syn. Polypodium polypodioides), also known as the resurrection fern, is a species of creeping, coarse-textured fern native to the Americas and Africa.

Contents

Description

The evergreen fronds of this fern are 25 cm high by 5 cm wide and monomorphic. The leathery, yellow-green pinnae (leaflets) are deeply pinnatifid, oblong to narrowly lanceolate, usually widest near middle, occasionally at or near base. It attaches to the limbs of its host plant with a branching, creeping, slender rhizome, which grows to 2 mm in diameter. The scales are lanceolate, with light brown base and margins, and having a dark central stripe.

Upper and lower sides of fronds PolypodFronds2.jpg
Upper and lower sides of fronds

The gametophytes (the haploid gamete producers) of this plant develop from very small spores that float in the air and are deposited on moist tree branches. These spores are produced in sporangia that develop on the leaves of the fern's sporophyte. The fern can also reproduce by the division of its rhizomes.

On the underside of the blades, the sori (reproductive clusters) are round, discrete, and sunken. Their outline can be seen as raised dimples on the upper surface. They are typically near the outer edge, and occur on all but the lowest pinnae of fertile fronds. Indusia are absent. Sporangia are yellow to brown at maturity. Spores are produced from summer to fall.

Habitat

Sori on the underside of the leaf Pleopeltis-polypodioides-Frond-Rear-Narrow-2014-10-19-v1.jpg
Sori on the underside of the leaf

This fern is an epiphyte, or air plant, which means it attaches itself to other plants and gets its nutrients from the air and from water and nutrients that collect on the outer surface of bark. The resurrection fern lives on the branches of large trees such as cypresses and can often be seen carpeting the shady areas on limbs of large oak trees repeatedly exposed to rainfall. However, it is known to grow on the surfaces of rocks and dead logs as well. It is often found in the company of other epiphytic plants, such as Spanish moss.

The P. polypodioides resides in the hardwood forests of southeastern United States in areas including Delaware, Maryland, the Carolinas, Virginia, West Virginia, Illinois, Ohio, Alabama, Georgia, Florida, Mississippi, Arkansas, Texas and Oklahoma. This fern can also be found in areas such as subtropical America, parts of southern Africa and other humid or sub-humid climate areas.[ citation needed ]

Physiology and rehydration

The resurrection fern gets its name because it can survive long periods of drought by curling up its fronds and appearing desiccated, grey-brown and dead. However, when just a little water is present, the fern will uncurl and reopen, appearing to "resurrect". It has been estimated that these plants could last 100 years without water and still revive after a single exposure. [2] [3]

Pleopeltis polypodioides or resurrection ferns in their desiccated, curled state. Pleopeltis polypodioides Cumberland Island 3.jpg
Pleopeltis polypodioides or resurrection ferns in their desiccated, curled state.

The resurrection fern roots have evolved to function as anchors for the epiphytes, holding them to their host tree’s bark and have been found to be inefficient in absorption. When in clustered groups of ferns on the bark of trees, the increased transpiration of the ferns and bark allow the middle ferns of the cluster to remain open longer than ferns on the outside. The relative humidity of the air and the closing rate of the leaves of the ferns is an inverse relationship. The greater the humidity, the slower the leaves of the ferns close because they have prolonged exposure to moisture in the air. [4]

The resurrection fern can severely desiccate and lose almost all of its water. Experiments have shown they can lose up to 97% and remain alive, though more typically they lose only around 76% in dry spells. [5] For comparison, most other plants would die after losing only 8–12%. This fern can lose almost all the water not hydrating the cells in its leaves and survive. Upon rainfall or even minute exposures to water, they can fully rehydrate and return to a normal state in 24 to 48 hours. [2] [6] Following substantial exposures to moisture, there is an immediate increase in its water content of up to 50% after the first hour and 65–70% after three hours. [6] When it regains moisture, the fern can once again become photosynthetically active, increasing its metabolism and release of organic compounds that provide nutrients for symbiotic bacteria that allow them both to thrive.  

At least one study has shown association between P. polypodioides and moss, [7] indicating that this fern may rely on moss for some of its water needs.

Resurrection ferns rehydrated in their uncurled state. Polypodium polypodioides, Loxahatchee.JPG
Resurrection ferns rehydrated in their uncurled state.

The resurrection ferns accumulate sugar in their dehydrated cells in order to protect them and stabilize the membranes and proteins in the dry state. By accumulating sugar, the cells can maintain hydrogen bonds between necessary macromolecules needed for their structure. An interesting feature of the resurrection fern is it does not immediately take up water after having been in a desiccated state. The resurrection ferns have porous cell structure networks that can expand and rearrange to accommodate absorbing three to four times their own weight in water after being exposed. [8] Initially, the water flows via capillary pressure into the spongy layer of the epidermis and the porous network of the fern’s leaves, without altering them significantly. Over time, pressure builds up in the fern cell network, changing the turgor pressure and strain in the plant, initiating the unfolding of the leaves. The fern stops unfolding its leaves only when the cell network of the fern is entirely filled with water. The uncurling of the leaves was believed to be an entirely osmotic phenomenon, but when the fern is again exposed to water, it absorbs the water through solid colloids, causing an increase in volume and capillary pressure of the cells of the fern.

Timelapse of a resurrection fern (Pleopeltis polypodioides) absorbing a drop of water.

When the fronds “dry,” the mesophyll deforms, which causes the lamina to curl and exposes the underside and shows the peltate scales. The peltate scales help prevent photooxidation and mechanical damage, as well as assist in quick rehydration. [9] The fern fronds contain canal cells in the center of their surface scales that direct water to the epidermis of the fern leaves, allowing absorption. The ability of the fern fronds to unroll after exposure to desiccation is attributed to the large cells of the upper epidermis along the midrib of the frond that increase in width more than any other epidermal cell, which forces the fronds to uncurl and flatten out. [4]

One way the resurrection fern has adapted to survive desiccation is the ability of its cell walls to deform and reform, without bursting or rupturing. It has been observed that dehydrins may allow the cell walls of the fern fronds and leaves to deform and reform in times of extreme drought followed by water exposure, due to large fluctuations in water content. Dehydrins were found to be expressed only when the fern was drying or in the desiccated state, with the dehydrins located on the outsides of the cells near the cell walls, allowing the leaves and fronds of the ferns to deform and reform accordingly. [10]

Thermoluminescence of the resurrection fern has been tested to observe at what temperatures it occurs and therefore shows what temperatures and amount of desiccation the ferns can tolerate before their chloroplasts lose the energy they have stored for the plant, in the form of light emissions. One study found the resurrection fern required high temperatures before thermoluminescence occurred at around 50 °C, whereas other desiccation-tolerant plants showed thermoluminescence at 40 °C. [11] The thermoluminescent activity of the resurrection fern increased as it was exposed to six separate but consecutive flashes of green safelight at its leaves. Because thermoluminescence from the fern fronds was not observed until higher temperatures, this hints that the fern’s chloroplasts may have a mechanism to withhold and store energy for the fern as it desiccates, allowing it to survive higher temperatures and extreme periods of desiccation. [11]

Reproduction

The fern has spores on the bottom of the fronds, contained in sori. Sori can be found aligned in rows on the underside of fertile fronds. They start as yellow color, but as they mature, they turn a brown color and split. [12] The fern sporulates in Summer and early Fall. Rhizome sections are also viable offspring and can root themselves in new medium.

See also

Related Research Articles

<span class="mw-page-title-main">Fern</span> Class of vascular plants

The ferns are a group of vascular plants that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissues that conduct water and nutrients, and in having life cycles in which the branched sporophyte is the dominant phase.

<i>Platycerium</i> Genus of ferns

Platycerium is a genus of about 18 fern species in the polypod family, Polypodiaceae. Ferns in this genus are widely known as staghorn or elkhorn ferns due to their uniquely shaped fronds. This genus is epiphytic and is native to tropical and temperate areas of South America, Africa, Southeast Asia, Australia, and New Guinea.

<i>Onoclea sensibilis</i> Species of fern

Onoclea sensibilis, the sensitive fern, also known as the bead fern, is a coarse-textured, medium to large-sized deciduous perennial fern. The name comes from its sensitivity to frost, the fronds dying quickly when first touched by it. It is sometimes treated as the only species in Onoclea, but some authors do not consider the genus monotypic.

<i>Alsophila brevipinna</i> Species of fern

Alsophila brevipinna, synonym Cyathea brevipinna, is a species of tree fern endemic to the higher parts of Mount Gower on Lord Howe Island, where it grows in exposed areas at an altitude of about 790 m. The trunk is erect and may reach 3 m in height. It is often covered with reddish brown scales and stipe bases. This species may produce stolons at ground level. Fronds are tripinnate, densely crowded, and up to about 3 m long. The stipe is brown and sometimes warty after scales fall off. The scales are long, glossy dark brown, with a distinctly narrow apex and fragile paler edges. Sori are attached to deeply divided fertile pinnules that may uncurl over the sori. Indusia are firm and large. A. brevipinna is a stunted plant with short pinnae.

<i>Phlebodium aureum</i> Species of fern

Phlebodium aureum is an epiphytic fern native to tropical and subtropical regions of the Americas.

<i>Selaginella lepidophylla</i> Species of spore-bearing plant

Selaginella lepidophylla, also known as a resurrection plant, is a species of desert plant in the spikemoss family (Selaginellaceae). It is native to the Chihuahuan Desert of the United States and Mexico. S. lepidophylla is renowned for its ability to survive almost complete desiccation. Resurrection plants are vascular rooted plants capable of surviving extreme desiccation, then resuming normal metabolic activity upon rehydration. The plant's hydro-responsive movements are governed by stem moisture content, tissue properties and a graded distribution of lignified cells affecting concentric stem stiffness and spiraling. During dry weather in its native habitat, its stems curl into a tight ball, uncurling only when exposed to moisture.

<i>Pleopeltis</i> Genus of ferns

Pleopeltis is a genus of ferns in the family Polypodiaceae, subfamily Polypodioideae, according to the Pteridophyte Phylogeny Group classification of 2016 (PPG I). The genus widely distributed in tropical regions of the world, and also north into temperate regions in eastern North America and eastern Asia. Several species are known by the common name scaly polypody and resurrection fern.

<i>Polypodium vulgare</i> Species of ferns in the family Polypodiaceae

Polypodium vulgare, the common polypody, is an evergreen fern of the family Polypodiaceae. The name is derived from Greek poly- ("many") and pous, podos ("foot"). Polypody has traditional uses in cooking for its aroma and sweet taste, and in herbal medicine as a purgative and vermifuge.

Poikilohydry is the lack of ability to maintain and/or regulate water content to achieve homeostasis of cells and tissue connected with quick equilibration of cell/tissue water content to that of the environment. The term is derived from Ancient Greek ποικίλος.

<i>Polypodium glycyrrhiza</i> Species of ferns in the family Polypodiaceae

Polypodium glycyrrhiza, commonly known as licorice fern, many-footed fern, and sweet root, is a summer deciduous fern native to western North America, where it is found in shaded, damp locations.

<i>Myriopteris gracillima</i> Species of fern

Myriopteris gracillima, formerly known as Cheilanthes gracillima, is a species of lip fern known by the common name lace lip fern. It is native to western North America, where it grows in rocky habitat from British Columbia to California to Montana.

<i>Zealandia pustulata</i> Species of fern

Zealandia pustulata is a species of fern native to eastern Australia and New Zealand. It is commonly referred to as 'kangaroo fern' or 'kangaroo paw fern' as its native range includes Australia and the shape of its mature foliage tends to resemble the shape of a kangaroo's foot. It is also referred to as 'hound's tongue', and as kōwaowao and pāraharaha in the Māori language.

<i>Polystichum vestitum</i> Species of fern

Polystichum vestitum, commonly known as the prickly shield fern or pūnui (Māori), is a hardy, evergreen or semi-evergreen ground fern.

<i>Myrothamnus flabellifolius</i> Species of plant of Southern Africa

Myrothamnus flabellifolius is a plant species in the family Myrothamnaceae native to central and southern Africa. It is also called the resurrection plant for the appearance of dead leaves reviving during rain.

<i>Aglaomorpha</i> (plant) Genus of ferns

Aglaomorpha is a genus of ferns in the subfamily Drynarioideae of the family Polypodiaceae. The Pteridophyte Phylogeny Group classification of 2016 uses this genus name, while other sources use Drynaria to include Aglaomorpha. Species are commonly known as basket ferns. As circumscribed in PPG I, the genus contains around 50 species.

<i>Lygodium japonicum</i> Species of fern

Lygodium japonicum is a species of fern that is known by the common names vine-like fern and Japanese climbing fern. It is native to eastern Asia, including Taiwan, Japan, Korea, southeastern Asia, and India, and eastern Australia. The fern is present in the southeastern United States and Puerto Rico as an introduced species.

<i>Gleichenia polypodioides</i> Species of fern

Gleichenia polypodioides (L.) Sm., commonly known as coral fern, kystervaring or ystervaring due to its glabrous, brown, wiry stipes. The species is widespread in south- and east tropical Africa, southern Africa and the western Indian Ocean region. It occurs naturally in a broad coastal belt in South Africa, Lesotho, Eswatini, Angola, Malawi, Burundi, Tanzania, Mozambique, Zimbabwe, Mauritius, Réunion, Amsterdam Island and Madagascar, and was first described by Carl Linnaeus in 1771 under the name Onoclea polypodioides. Often forming dense and impenetrable thickets, sometimes over large areas, this rhizomatous perennial is an important pioneer in disturbed areas such as pine plantations. It is often mistakenly seen as an exotic invader rather than as a useful rehabilitation plant, a source of peat and growing medium, while showing exceptional resistance to herbicides.

Rhizome brown, 1–2.5 mm. in diam., creeping, with long-spined dark-brown scales up to 0.5 mm. in diam., with fronds spaced 2–20 cm. apart. Stipe castaneous, up to 60 cm. long and up to 1.5 mm. in diam., glabrous or with a few scales similar to those on the rhizome, shallowly sulcate. Frond bifurcate to reniform-lunate in outline, with 1 level of false dichotomy in each lateral branch system arising from each side of the terminal bud; all branches bearing distant foliar segments. Aborted apical buds up to 1.2 mm. long, clothed in dark-brown lanceolate laciniate scales. Pinnules linear, up to 7 x 0.75 cm., pinnate, usually glabrous, divided into sessile rounded entire triangular lobes, 3 x 2 mm., green to glaucous below. Sori partially immersed in the lamina, consisting of 2–4 sporangia, each in a separate but adjoining pit.

<i>Vandenboschia boschiana</i> Species of fern

Vandenboschia boschiana, synonym Trichomanes boschianum, also known as the Appalachian bristle fern or Appalachian filmy fern, is a small delicate perennial leptosporangiate fern which forms colonies with long, black creeping rhizomes.

<i>Hymenophyllum tunbrigense</i> Species of fern

Hymenophyllum tunbrigense, the Tunbridge filmy fern or Tunbridge filmy-fern, is a small, fragile perennial leptosporangiate fern which forms large dense colonies of overlapping leaves from creeping rhizomes. The common name derives from the leaves which are very thin, only a single cell thick, and translucent, giving the appearance of a wet film. The evergreen fronds are bipinnatifid, deeply and irregularly dissected, about 3 to 6 cm long, 2 cm across with dark winged stipes. In contrast to the similar H. wilsonii the fronds are more divided, flattened, appressed to the substrate and tend to have a bluish tint.

<i>Polypodium</i> Genus of ferns in the family Polypodiaceae

Polypodium is a genus of ferns in the family Polypodiaceae, subfamily Polypodioideae, according to the Pteridophyte Phylogeny Group classification of 2016 (PPG I). The genus is widely distributed throughout the world, with the highest species diversity in the tropics. The name is derived from Ancient Greek poly ("many") + podion, on account of the foot-like appearance of the rhizome and its branches. They are commonly called polypodies or rockcap ferns, but for many species unique vernacular names exist.

References

  1. "NatureServe Explorer 2.0 - Pleopeltis polypodioides Resurrection Fern". explorer.natureserve.org. Retrieved 9 October 2020.
  2. 1 2 North Carolina Cooperative Extension (17 Feb 2012). "Resurrection ferns make awe-inspiring comebacks". Wayne County Center--North Carolina Cooperative Extension. Goldsboro, North Carolina: North Carolina State University. Archived from the original on 30 March 2013. Retrieved 14 Jul 2015.
  3. Stevenson, Carrie (2 Sep 2013). "Resurrection Ferns". Gardening in the Panhandle. Gainesville, Florida: Institute of Food and Agricultural Sciences. Archived from the original on 14 July 2015. Retrieved 14 Jul 2015.
  4. 1 2 Potts, Roberta; Penfound, Wm. T. (1948). "Water Relations of the Polypody Fern, Polypodium Polypodioides (L.) A. S. Hitchcock". Ecology. 29 (1): 43–53. Bibcode:1948Ecol...29...43P. doi:10.2307/1930343. JSTOR   1930343.
  5. Moran 2004
  6. 1 2 Jackson, Evelyn F.; Echlin, Haley L.; Jackson, Colin R. (2006). "Changes in the phyllosphere community of the resurrection fern, Polypodium polypodioides, associated with rainfall and wetting: Changes in the phyllosphere community of the resurrection fern". FEMS Microbiology Ecology. 58 (2): 236–246. doi: 10.1111/j.1574-6941.2006.00152.x . PMID   17064265.
  7. "Proceedings of the Oklahoma Academy of Science". Archived from the original on 2005-03-17. Retrieved 2005-03-14.
  8. Helseth, L. E.; Fischer, T. M. (2005-06-09). "Physical mechanisms of rehydration in Polypodium polypodioides, a resurrection plant". Physical Review E. 71 (6): 061903. Bibcode:2005PhRvE..71f1903H. doi:10.1103/PhysRevE.71.061903. ISSN   1539-3755. PMID   16089761.
  9. Prats, Kyra A; Brodersen, Craig R (2021-08-02). "Desiccation and rehydration dynamics in the epiphytic resurrection fern Pleopeltis polypodioides". Plant Physiology. 187 (3): 1501–1518. doi:10.1093/plphys/kiab361. ISSN   0032-0889. PMC   8566288 . PMID   34618062.
  10. Layton, B. E.; Boyd, M. B.; Tripepi, M. S.; Bitonti, B. M.; Dollahon, M. N. R.; Balsamo, R. A. (2010), "Dehydration-induced expression of a 31-kDa dehydrin in Polypodium polypodioides (Polypodiaceae) may enable large, reversible deformation of cell walls", American Journal of Botany, 97 (4): 535–44, doi: 10.3732/ajb.0900285 , PMID   21622416
  11. 1 2 Maslenkova, T. L.; Homann, P. H. (2000). "2000CRABS..53d..99M Page D99". Comptes Rendus de l'Académie Bulgare des Sciences. 53: 4. Bibcode:2000CRABS..53d..99M.
  12. Webmaster, ANPS (2018-01-16). "Know Your Natives – Resurrection Fern". Arkansas Native Plant Society. Retrieved 2023-10-16.