Erioderma Pedicellatum
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''Erioderma pedicellatum'' is a medium-sized, foliose
lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.Pannariaceae The Pannariaceae are a family of lichens in the order Peltigerales (suborder Collematineae). Species from this family have a widespread distribution, but are especially prevalent in southern temperate regions. Genera According to a recent (202 ...
, commonly called the boreal felt lichen. It grows on trees in damp
boreal Boreal may refer to: Climatology and geography *Boreal (age), the first climatic phase of the Blytt-Sernander sequence of northern Europe, during the Holocene epoch *Boreal climate, a climate characterized by long winters and short, cool to mild ...
forests along the Atlantic coast in
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...
, as well as in southcentral
Alaska Alaska ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U.S., ...
and in the
Kamchatka Peninsula The Kamchatka Peninsula (russian: полуостров Камчатка, Poluostrov Kamchatka, ) is a peninsula in the Russian Far East, with an area of about . The Pacific Ocean and the Sea of Okhotsk make up the peninsula's eastern and we ...
.Nelson, P, J Walton and C Roland. 2009. ''Erioderma pedicellatum'' (Hue) P.M.Jorg, new to the United States and western North America, discovered in Denali National Park and Preserve and Denali State Park, Alaska. Evansia 26: 19 – 23.


Description

''Erioderma pedicellatum'' is a foliose
cyanolichen Cyanolichens are lichens that apart from the basic fungal component ("mycobiont"), contain cyanobacteria, otherwise known as blue-green algae, as the photosynthesizing component ("photobiont"). Overall, about a third of lichen photobionts are cyan ...
with lobes 2–5 cm across, and occasionally reaching 12 cm in diameter. It has a distinctively fuzzy upper surface that is greyish-brown when dry, and slate-blue when moist. Its underside is white, and its edges usually curl upwards, giving it the appearance of having a white fringe.COSEWIC Status Report for ''Erioderma pedicellatum''
/ref> It differs from the two other North American species of '' Erioderma'' by lacking
soredia Soredia are common reproductive structures of lichens. Lichens reproduce asexually by employing simple fragmentation and production of soredia and isidia. Soredia are powdery propagules composed of fungal hyphae wrapped around cyanobacteria or gr ...
, and by having small, reddish-brown
apothecia An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are mo ...
on its upper surface.


Taxonomy and naming

''Erioderma pedicellatum'' was first collected in 1902 from
Campobello Island Campobello Island (, also ) is the largest and only inhabited island in Campobello, a civil parish in southwestern New Brunswick, Canada, near the border with Maine, United States. The island's permanent population in 2021 was 949. It is the si ...
,
Charlotte County, New Brunswick Charlotte County (2016 population 25,428) is the southwest-most List of counties of New Brunswick, county of New Brunswick, Canada. It was formed in 1784 when New Brunswick was partitioned from Nova Scotia. Once a layer of local government, th ...
, Canada, by
William Gilson Farlow William Gilson Farlow (December 17, 1844 – June 3, 1919) was an American botanist, born in Boston, Massachusetts, and educated at Harvard (A.B., 1866; M.D., 1870), where, after several years of European study, he became adjunct professor of ...
.Jørgensen, Per M. 2000. Survey of the Lichen Family Pannariaceae on the American Continent, North of Mexico. The Bryologist: 103(4): 670-704. It was originally identified as a species of ''
Pannaria ''Pannaria'' is a genus of lichen-forming fungi in the family Pannariaceae. The widespread genus contains an estimated 51 species, found primarily in tropical regions. Species *''Pannaria aenea'' *'' Pannaria andina'' *'' Pannaria aotearoana' ...
'' and named ''Pannaria pedicellata'' by French botanist
Auguste-Marie Hue Father Auguste-Marie Hue (15 August 1840– 22 June 1917) was a French lichenologist. Biography Hue was born on 15 August 1840, in Saint-Saëns, Seine-Maritime. He was Ordination, ordained as a priest in 1865. From 1890 to 1915 he was a chaplain ...
.Jørgensen, M. 2001. The present status of the names applicable to species and infraspecific taxa of ''Erioderma''(lichenised Ascomycetes) included in Zahlbruckner's "Catalogus". Taxon 50(2): 525-541. It remained in this genus until 1972 when it was reexamined by the Norwegian botanist
Per Magnus Jørgensen Per Magnus Jørgensen (born 1944) is a Norwegian botanist and lichenologist, and Professor Emeritus of systematic botany at the University of Bergen. He is known for his work on the lichen families Pannariaceae and Collemataceae. Jørgensen wa ...
and placed in the genus '' Erioderma'' as ''Erioderma pedicellatum''. It is an unusual species within that genus, both because of its laminal (upper surface)
apothecia An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are mo ...
(lacking in other ''Erioderma'') and its boreal distribution. ''Erioderma pedicellatum'' has also been incorrectly called ''E. boreale''.


Biology and ecology

''Erioderma pedicellatum'' grows on mossy trunks and branches of trees on slopes in areas that have a constant supply of moisture, and are rich in ''
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 ...
'' moss. It is usually found on
balsam fir ''Abies balsamea'' or balsam fir is a North American fir, native to most of eastern and central Canada (Newfoundland west to central Alberta) and the northeastern United States (Minnesota east to Maine, and south in the Appalachian Mountains to ...
, occasionally on
black spruce ''Picea mariana'', the black spruce, is a North American species of spruce tree in the pine family. It is widespread across Canada, found in all 10 provinces and all 3 territories. It is the official tree of the province of Newfoundland and Labra ...
, and rarely on
white spruce White spruce is a common name for several species of spruce (''Picea'') and may refer to: * ''Picea glauca'', native to most of Canada and Alaska with limited populations in the northeastern United States * ''Picea engelmannii'', native to the Ro ...
,
red maple ''Acer rubrum'', the red maple, also known as swamp maple, water maple, or soft maple, is one of the most common and widespread deciduous trees of eastern and central North America. The U.S. Forest Service recognizes it as the most abundant nativ ...
, or
white birch ''Betula papyrifera'' (paper birch, also known as (American) white birch and canoe birch) is a short-lived species of birch native to northern North America. Paper birch is named for the tree's thin white bark, which often peels in paper like ...
. It does not appear to grow directly on bare bark, and is usually found growing in association with the
epiphytic An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
liverwort The Marchantiophyta () are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of g ...
'' Frullania asagrayana''.
Old growth An old-growth forestalso termed primary forest, virgin forest, late seral forest, primeval forest, or first-growth forestis a forest that has attained great age without significant disturbance (ecology), disturbance, and thereby exhibits un ...
balsam fir boreal forests in wet areas of eastern Canada regenerate by gap replacement, which creates many forest stands of different ages, while still maintaining a full or partial canopy.Thompson ID, DJ Larson, and WA Montevecchi. 2003. Characterization of old “wet boreal” forests, with an example from balsam fir forests of western Newfoundland. Environmental Reviews 11: S23-S4

/ref> It appears that this diversity in forest stands age is necessary for a viable population of ''E. pedicellatum''. Natural dispersal of ''E. pedicellatum'' is evidently possible within these old-growth forests, but there are no known examples of ''E. pedicellatum'' establishing in stands previously Clearcutting, clear-cut. A healthy, mature specimen of ''E. pedicellatum'' can grow at a rate of 11 to 14 mm per year, and populations of this lichen have a generation time of about 30 years. The species sexually reproduces. However, as with most lichen, considerable information in understanding ''E. pedicellatum'' biology is missing, especially information about its life stages. ''
Scytonema ''Scytonema'' is a genus of photosynthetic cyanobacteria that contains over 100 species. It grows in filaments that form dark mats. Many species are aquatic and are either free-floating or grow attached to a submerged substrate, while others s ...
''
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
photobionts of this lichen make it particularly sensitive to acid rain and other atmospheric pollutants. It requires relatively cool and moist oceanic climates, and an open canopy. It deteriorates rapidly on dead trees, or if habitat succession occurs that reduces, or increases light availability. Altered micro-climatic conditions caused by extensive logging also causes ''E. pedicellatum'' to deteriorate.


Symbiosis

''Erioderma pedicellatum'', like all lichens, is a
symbiotic Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
, in this instance between an
ascomycete Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The defi ...
fungus A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...
and
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
of the genus ''
Scytonema ''Scytonema'' is a genus of photosynthetic cyanobacteria that contains over 100 species. It grows in filaments that form dark mats. Many species are aquatic and are either free-floating or grow attached to a submerged substrate, while others s ...
'', and is therefore capable of
fixing nitrogen Nitrogen fixation is a chemical process by which molecular nitrogen (), with a strong triple covalent bond, in the air is converted into ammonia () or related nitrogenous compounds, typically in soil or aquatic systems but also in industry. Atmos ...
. ''E. pedicellatum'' may also be part of second symbiosis with the
epiphytic An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
liverwort The Marchantiophyta () are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of g ...
'' Frullania asagrayana''.Cornejo, C., P.R. Nelson, I. Stepanchikova, D. Himelbrant, P.-M. Jorgensen and C. Scheidigger. 2016. Contrasting pattern of photobiont diversity in the Atlantic and Pacific populations of Erioderma pedicellatum (Pannariaceae). The Lichenologist 48(4): 275 – 291 The symbiosis between the free-living ''
Scytonema ''Scytonema'' is a genus of photosynthetic cyanobacteria that contains over 100 species. It grows in filaments that form dark mats. Many species are aquatic and are either free-floating or grow attached to a submerged substrate, while others s ...
'' and the germinating ascomycete
spore 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 ...
s of ''E. pedicellatum'' is hypothesized to begin within the water sacs of ''Frullania asagrayana'', where the fungal
hyphae A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
assimilates a
cyanobacterium Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blue ...
, and needs to develop for 5 to 10 years before it reaches a visible size. The liverwort may also benefit from the nitrogen that is being fixed by the
cyanolichen Cyanolichens are lichens that apart from the basic fungal component ("mycobiont"), contain cyanobacteria, otherwise known as blue-green algae, as the photosynthesizing component ("photobiont"). Overall, about a third of lichen photobionts are cyan ...
growing within it. This complex relationship means that the ecological balance between ''E. pedicellatum'' and its cyanobacterial symbiont (''Scytonema''), its host tree, and (potentially) its liverwort nursemaid (''Frullania asagrayana''), is very delicate and easily impacted by logging, air pollution, and other factors. More information is needed to fully understand the symbiosis within ''E. pedicellatum''.


Distribution

''Erioderma pedicellatum'' is distributed in 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 ...
. It can be found on both sides of the Atlantic and Pacific coastal regions.


Atlantic populations

''Erioderma pedicellatum'' was once prevalent in Norway, Sweden, and the provinces of
New Brunswick New Brunswick (french: Nouveau-Brunswick, , locally ) is one of the thirteen provinces and territories of Canada. It is one of the three Maritime provinces and one of the four Atlantic provinces. It is the only province with both English and ...
,
Nova Scotia Nova Scotia ( ; ; ) is one of the thirteen provinces and territories of Canada. It is one of the three Maritime provinces and one of the four Atlantic provinces. Nova Scotia is Latin for "New Scotland". Most of the population are native Eng ...
, and
Newfoundland Newfoundland and Labrador (; french: Terre-Neuve-et-Labrador; frequently abbreviated as NL) is the easternmost province of Canada, in the country's Atlantic region. The province comprises the island of Newfoundland and the continental region ...
in
Atlantic Canada Atlantic Canada, also called the Atlantic provinces (french: provinces de l'Atlantique), is the region of Eastern Canada comprising the provinces located on the Atlantic coast, excluding Quebec. The four provinces are New Brunswick, Newfoundlan ...
. It has disappeared from Norway and Sweden (thus all of Europe). In Canada, it is no longer found in New Brunswick, and as of 2009 there were fewer than 200 individuals known in Nova Scotia.Cameron, R, C Hanel, I Goudie, and N. Neily. 2009. Boreal felt lichen; Current status, conservation issues and future prospects. Botanical Electronic News, no. 42

/ref> In Newfoundland, it is estimated that there between 15,000 to 20,000 individuals. The remaining habitat in Newfoundland is therefore critical for the global survival of this species. Lockyer's Waters and Hall’s Gullies on the
Avalon Peninsula The Avalon Peninsula (french: Péninsule d'Avalon) is a large peninsula that makes up the southeast portion of the island of Newfoundland. It is in size. The peninsula is home to 270,348 people, about 52% of Newfoundland's population, according ...
to the southeast, and
Bay d'Espoir Bay d'Espoir ( ) is an arm of Hermitage Bay in the Gulf of Saint Lawrence, located on the south coast of Newfoundland. Communities in Bay d'Espoir include: Milltown-Head of Bay d'Espoir, Morrisville, St. Alban's, St. Joseph's Cove, St. Veronica ...
to the south, are three of the Newfoundland's most prolific rare lichen habitats, and are important for the conservation of ''E. pedicellatum''.


Pacific populations

Recently, populations were discovered in
Denali Denali (; also known as Mount McKinley, its former official name) is the highest mountain peak in North America, with a summit elevation of above sea level. With a topographic prominence of and a topographic isolation of , Denali is the thir ...
area of
Alaska Alaska ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U.S., ...
, and in the
Kamchatka Peninsula The Kamchatka Peninsula (russian: полуостров Камчатка, Poluostrov Kamchatka, ) is a peninsula in the Russian Far East, with an area of about . The Pacific Ocean and the Sea of Okhotsk make up the peninsula's eastern and we ...
of
Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...
, increasing the known range of the species, and is the first time the species has been observed in the Pacific region. While little is known about the population in Russia, the Alaskan population has been projected to be able to increase the global population to about 100,000 individuals based on
statistical model A statistical model is a mathematical model that embodies a set of statistical assumptions concerning the generation of Sample (statistics), sample data (and similar data from a larger Statistical population, population). A statistical model repres ...
ing (ie. projecting the population size based on the known number of individuals counted in a certain area of the site, quantifying known variables, and the actual size of the site.) However, more research is needed on the Alaskan population to determine its population stability, as well as whether it is genetically distinct from the Atlantic populations, since the Alaskan population were found on
white spruce White spruce is a common name for several species of spruce (''Picea'') and may refer to: * ''Picea glauca'', native to most of Canada and Alaska with limited populations in the northeastern United States * ''Picea engelmannii'', native to the Ro ...
twigs, a different type of tree usually associated with ''E. pedicellatum'' in Canada. The Canadian populations were also found mostly on trunks, as opposed to twigs.


Conservation status

Lichen conservation is a growing field. ''E. pedicellatum'' is currently listed as endangered by the
Committee on the Status of Endangered Wildlife in Canada The Committee on the Status of Endangered Wildlife in Canada (COSEWIC, French: Comité sur la situation des espèces en péril au Canada, COSEPAC) is an independent committee of wildlife experts and scientists whose "raison d'être is to identify s ...
(COSEWIC) and critically endangered by the
International Union for Conservation of Nature 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 ...
(IUCN) since its last assessment in 2003. No other lichen is listed by the IUCN.


Protection

Two of the main populations of ''E. pedicellatum'' in Newfoundland are currently within protected areas: Jipujijkuei Kuespem Provincial Park, and the Lockyer’s Waters interim protected area, which was established specifically to protect this lichen. The Bay du Nord Wilderness Area in Newfoundland also includes some populations of the lichen. The Hall’s Gullies site is more in jeopardy, as it remains a designated cutblock (area allowed to be forested) under the current Forest Management Operating Plan. It has been noted that populations of this lichen can decline even in protected areas, which has been linked to air pollution and introduced herbivores, such as moose. The Atlantic population of the ''E. pedicellatum'' is protected in Canada under the Federal Species at Risk Act (SARA), and is the focus of an ongoing recovery strategy. Efforts are being made, through land purchases and agreements with landowners, to formally protect areas of forest that are home to this rare species. Furthermore, conservationists are engaging with private and government forest managers to encourage their participation in the mapping of ''E. pedicellatum'' habitat and the implementation of management plans that will prevent further habitat loss. For the Alaskan population, in the United States, ''E. pedicellatum'' appears not to be listed on the
United States Fish and Wildlife Service The United States Fish and Wildlife Service (USFWS or FWS) is an agency within the United States Department of the Interior dedicated to the management of fish, wildlife, and natural habitats. The mission of the agency is "working with othe ...
's Threatened and Endangered Species List. However, the population is in the
Denali National Park and Preserve Denali National Park and Preserve, formerly known as Mount McKinley National Park, is an American national park and preserve located in Interior Alaska, centered on Denali, the highest mountain in North America. The park and contiguous preserve e ...
, a protected area, and efforts have been made by researchers to not give away its location within the site so that they will not be tampered with.


Threats and efforts

The biggest threat to ''E. pedicellatum'' is deforestation and habitat loss. Through statistical modeling, it has been shown that ''E. pedicellatum'' has an increased chance of death when growing next to areas with deforestation. The habitat of ''E. pedicellatum'' was shown to be cut down faster than it can grow back, which is of concern given ''E. pedicellatum'''s narrow habitat
niche Niche may refer to: Science *Developmental niche, a concept for understanding the cultural context of child development *Ecological niche, a term describing the relational position of an organism's species *Niche differentiation, in ecology, the ...
. Current models predict that the Atlantic ''E. pedicellatum'' population will continue to decrease substantially over the next 25 years. Most current efforts in studying the status and conservation efforts of ''E. pedicellatum'' are through statistical modeling to predict its population and habitat quality. Through statistical modeling, more information has been developed on the habitat of ''E. pedicellatum'', including how it prefers older, established forests with poor soil drainage. However, statistical modeling has disadvantages, including making broad assumptions based on missing knowledge in lichen ecology. Efforts have been made to further understanding the effects of air quality. Air quality is shown to have a major impact on ''E. pedicellatum,'' most likely due to it being a cyanolichen. In particular,
acid rain Acid rain is rain or any other form of precipitation that is unusually acidic, meaning that it has elevated levels of hydrogen ions (low pH). Most water, including drinking water, has a neutral pH that exists between 6.5 and 8.5, but acid ...
is a major concern to ''E. pedicellatum'' because its habitat is already acidic. Research has also been conducted looking at impacts from invasive
gastropods The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. Ther ...
, which were shown to possibly be grazing on ''E. pedicellatum'' in Nova Scotia. More research is needed on their impacts. Researchers are trying to understand the genetic differences between each segmented population of ''E. pedicellatum''. Currently, initial genetic analysis hypothesizes that the Pacific populations of ''E. pedicellatum'' are more diverse than the Atlantic population. However, research is still needed on how the populations interact with one another, specifically through
gene flow In population genetics, gene flow (also known as gene migration or geneflow and allele flow) is the transfer of genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent a ...
.


Gallery

File:Erioderma pedicellatum2.JPG, ''Erioderma pedicellatum'' growing on a tree in Newfoundland File:Erioderma pedicellatum3.JPG, ''Erioderma pedicellatum'' growing on a tree in Newfoundland File:Erioderma pedicellatum4.JPG, ''Erioderma pedicellatum'' growing on a tree in Newfoundland


References

* Robertson, A. 1998. The boreal felt lichen (''Erioderma pedicellatum'' (Hue) P M. Jørg.) in Newfoundland. Government of Newfoundland and Labrador of Forest Resources and Agrifoods, Forestry, Wildlife and Inland Fish Branch, St. John's, NF.


External links

*
Pictures of ''Erioderma pedicellatum''
a
www.erioderma.com

Newfoundland Lichen Education and Research Group

Memorial University's NL Nature project, focusing primarily on Boreal Felt Lichens


* {{Authority control Peltigerales Lichen species Lichens described in 1911 Fungi of Asia Fungi of North America Taxa named by Auguste-Marie Hue