Peltigera Cinnamomea
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''Peltigera cinnamomea'', commonly known as the cinnamon-pelt lichen, is a muscicolous (
moss Mosses are small, non-vascular plant, non-vascular flowerless plants in the taxonomic phylum, division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Wilhelm Philippe Schimper, Schimp. 1879) may also refer to the parent group bryo ...
-dwelling), leafy lichen in the family
Peltigeraceae The Peltigeraceae are a family of lichens in the order Peltigerales. The Peltigeraceae, which contains 15 genera and about 600 species, has recently (2018) been emended to include the families Lobariaceae and Nephromataceae. Many Peltigeraceae sp ...
. The Canadian lichenologist Trevor Goward
formally described A species description is a formal scientific description of a newly encountered species, typically articulated through a scientific publication. Its purpose is to provide a clear description of a new species of organism and explain how it differ ...
the species in 1995. The lichen is found in northwestern North America's forested regions, particularly in the unique montane and subalpine forest communities of the northern
Rocky Mountains The Rocky Mountains, also known as the Rockies, are a major mountain range and the largest mountain system in North America. The Rocky Mountains stretch in great-circle distance, straight-line distance from the northernmost part of Western Can ...
. ''Peltigera cinnamomea'' grows under prolonged snow cover, surviving well into spring. This trait distinguishes it from many other ''
Peltigera ''Peltigera'' is a genus of approximately 100 species of foliose lichens in the family Peltigeraceae. Commonly known as the dog or pelt lichens, species of ''Peltigera'' are often terricolous (growing on soil), but can also occur on moss, trees ...
'' species in similar North American forest ecosystems. ''Peltigera cinnamomea'' forms a specialised
symbiotic Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...
relationship with specific strains of the cyanobacterium ''
Nostoc ''Nostoc'', also known as star jelly, troll's butter, spit of moon, fallen star, witch's butter (not to be confused with the fungi commonly known as witches' butter), and witch's jelly, is the most common genus of cyanobacteria found in a variety ...
'', enabling it to fix atmospheric nitrogen, crucial for survival in nutrient-poor environments. This lichen is part of the broadly defined
species complex In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
centred around the widely distributed ''
Peltigera canina ''Peltigera canina'', commonly known as the dog lichen, is a widely distributed species of foliose lichen in the family Peltigeraceae. It was originally described by Carl Linnaeus in his 1753 work ''Species Plantarum''. German botanist Carl Ludw ...
''. Within this complex, ''P. cinnamomea'' falls under the CICADE group, indicating a preference for moist,
woodland A woodland () is, in the broad sense, land covered with woody plants (trees and shrubs), or in a narrow sense, synonymous with wood (or in the U.S., the '' plurale tantum'' woods), a low-density forest forming open habitats with plenty of sunli ...
habitats. Characterised by its leafy
thallus Thallus (: thalli), from Latinized Greek (), meaning "a green shoot" or "twig", is the vegetative tissue of some organisms in diverse groups such as algae, fungi, some liverworts, lichens, and the Myxogastria. A thallus usually names the entir ...
, which is loosely attached to its , ''P. cinnamomea'' typically measures in diameter. The thallus has a distinctive appearance, with a dull, billowed upper surface covered in a soft, dense growth of hairs. The
species name In taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, altho ...
''cinnamomea'' is reflected in the pale tan undersurface of the thallus, featuring rusty-brown, cinnamon-coloured . The lichen produces
apothecia An ascocarp, or ascoma (: ascomata), 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. As ...
(
fruiting bodies The sporocarp (also known as fruiting body, fruit body or fruitbody) of fungi is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne. The fruitbody is part of the sexual phase of a fungal life cyc ...
) that bear clear with three internal partitions.


Taxonomy

The
type specimen In biology, a type is a particular wikt:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to ancho ...
of ''Peltigera cinnamomea'' was collected by the Canadian lichenologist Trevor Goward on 13 April 1985, within the Clearwater River Basin in
British Columbia British Columbia is the westernmost Provinces and territories of Canada, province of Canada. Situated in the Pacific Northwest between the Pacific Ocean and the Rocky Mountains, the province has a diverse geography, with rugged landscapes that ...
(Canada), at an elevation of . The lichen was found growing over a mossy boulder situated in a boulder bed in an open,
mixed forest Temperate broadleaf and mixed forest is a temperate climate terrestrial habitat type defined by the World Wide Fund for Nature, with broadleaf tree ecoregions, and with conifer and broadleaf tree mixed coniferous forest ecoregions. These ...
. ''Peltigera cinnamomea'' was first scientifically documented by Goward in a 1994 report on the ''
Peltigera ''Peltigera'' is a genus of approximately 100 species of foliose lichens in the family Peltigeraceae. Commonly known as the dog or pelt lichens, species of ''Peltigera'' are often terricolous (growing on soil), but can also occur on moss, trees ...
'' species in
Alberta Alberta is a Provinces and territories of Canada, province in Canada. It is a part of Western Canada and is one of the three Canadian Prairies, prairie provinces. Alberta is bordered by British Columbia to its west, Saskatchewan to its east, t ...
, published by the
Provincial Museum of Alberta The Royal Alberta Museum (RAM) is a museum of human history, human and natural history in Downtown Edmonton, Alberta, Canada, located north of Edmonton City Hall, City Hall. The museum is the largest in western Canada with more than exhibition ...
. It did not, however, meet the specific standards for
valid publication In botanical nomenclature, a validly published name is a name that meets the requirements in the ''International Code of Nomenclature for algae, fungi, and plants'' (''ICN'') for valid publication. Valid publication of a name represents the minim ...
of a new species, due to non-compliance with Articles 39.1 and 40.1 of the nomenclatural rules, which require a clear
description Description is any type of communication that aims to make vivid a place, object, person, group, or other physical entity. It is one of four rhetorical modes (also known as ''modes of discourse''), along with exposition, argumentation, and narr ...
and the designation of a type specimen. Goward published it formally a year later in 1995, as part of a synopsis of ''Peltigera'' occurring in British Columbia. ''Peltigera cinnamomea'', although readily recognisable in its natural habitat, was historically misidentified as '' P. praetextata'', because both species share some superficial similarities. However, their distinct ecological preferences, alongside more detailed morphological differences, led to the formal recognition of ''P. cinnamomea''. The
species epithet Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In Taxonomy (biology), taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany) ...
refers to the
cinnamon Cinnamon is a spice obtained from the inner bark of several tree species from the genus ''Cinnamomum''. Cinnamon is used mainly as an aromatic condiment and flavouring additive in a wide variety of cuisines, sweet and savoury dishes, biscuits, b ...
-coloured veins on the thallus underside. Goward called it the "dog pelt" in a previous publication before it was officially named. It has also been referred to as the "cinnamon-pelt lichen". Early (2003)
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
studies of taxa within the ''P. canina''
species complex In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
delineated two major
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
groups, meaning they consist of all the descendants of a
common ancestor Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. According to modern evolutionary biology, all living beings could be descendants of a unique ancestor commonl ...
, each with distinct ecological preferences. The two groups are known as CICADE and PORUDI. The acronym CICADE stands for "Cinnamomea, Canina, Degelii, and their close relatives", encompassing species that are primarily
mesophytic Mesophytes are terrestrial plants which are adapted to neither particularly dry nor particularly wet environments. An example of a mesophytic habitat would be a rural temperate meadow, which might contain goldenrod, clover, oxeye daisy, and '' Ros ...
, meaning they thrive in moderately moist environments, to subhygrophytic, indicating they prefer slightly wetter conditions. While these species typically occupy humid, woodland habitats, they can also be found in drier locations. ''Peltigera cinnamomea'' is a part of the CICADE group, reflecting its preference for environments that maintain moderate to high humidity. This categorisation is supported by genetic data that confirms its close relationship with other species in this group.


Description

''Peltigera cinnamomea'' is characterised by its foliose, loosely attached
thallus Thallus (: thalli), from Latinized Greek (), meaning "a green shoot" or "twig", is the vegetative tissue of some organisms in diverse groups such as algae, fungi, some liverworts, lichens, and the Myxogastria. A thallus usually names the entir ...
, which typically spans in diameter. The structure of the thallus facilitates various ecological functions. The of this lichen are somewhat leathery and stiff, measuring approximately in width, elongating and loosely overlapping with irregular branches. The tips of the lobes are rounded and may either lie flat or curve downwards, with generally smooth margins. The upper surface varies in colour from pale bluish-grey to pale brownish-grey, and may be tinged with shades of cinnamon brown. This surface is dull, often billowed broadly, and covered with a tomentum—a short, soft, dense growth of hairs—that tends to fade near the thallus centre. The lichen lacks
soredia Soredia are common reproduction, reproductive structures of lichens. Lichens asexual reproduction, reproduce asexually by employing simple fragmentation and production of soredia and isidia. Soredia are powdery propagules composed of fungus, fung ...
(granular reproductive structures) and
isidia An isidium (plural: isidia) is a tiny, wart- or finger-like outgrowth on the thallus surface of certain lichen species. It is one of two principal types of vegetative reproduction, vegetative reproductive structures in lichens, the other being ...
(outgrowths containing both the algae and the fungus of the lichen), indicative of its specific reproductive and structural characteristics. The margins are devoid of (small lobes). The lower surface features pale tan that darken to rusty brown or cinnamon brown towards the centre. These veins are narrow, occasionally raised, and smooth (), while the areas between them (interstices) are whitish and moderately deep. The
rhizine In lichens, rhizines are multicellular root-like structures arising mainly from the lower surface. A lichen with rhizines is termed rhizinate, while a lichen lacking rhizines is termed erhizinate. Rhizines serve only to anchor the lichen to their s ...
s, root-like structures, match the colour of the veins and range from to intricately branched (), enhancing the lichen's attachment to its substrate. The of the thallus is 50–80 
μm The micrometre (Commonwealth English as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American English), also commonly known by the non-SI term micron, is a unit of length in the International System ...
thick, and beneath this lies a layer, 20–80 μm thick, hosting the cyanobacterium ''
Nostoc ''Nostoc'', also known as star jelly, troll's butter, spit of moon, fallen star, witch's butter (not to be confused with the fungi commonly known as witches' butter), and witch's jelly, is the most common genus of cyanobacteria found in a variety ...
''. The
medulla Medulla (Latin for "marrow") or medullary may refer to: Science * Medulla oblongata, a part of the brain stem * Renal medulla, a part of the kidney * Adrenal medulla, a part of the adrenal gland * Medulla of ovary, a stroma in the center of the ...
, a loosely packed layer of fungal
hypha 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 o ...
e located beneath the photobiont layer, is white and measures 70–180 μm in thickness.
Apothecia An ascocarp, or ascoma (: ascomata), 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. As ...
(fruiting bodies) are commonly found at the margins on narrow, elongate lobes. These bear a medium brown, longitudinally folded , averaging 6–10 mm in length and typically erect. The are clear (
hyaline A hyaline substance is one with a glassy appearance. The word is derived from , and . Histopathology Hyaline cartilage is named after its glassy appearance on fresh gross pathology. On light microscopy of H&E stained slides, the extracellula ...
), three-septate, and measure 40–49 μm in length by 3–5 μm in width, with eight spores per
ascus An ascus (; : asci) is the sexual spore-bearing cell produced in ascomycete fungi. Each ascus usually contains eight ascospores (or octad), produced by meiosis followed, in most species, by a mitotic cell division. However, asci in some gen ...
(the spore-bearing sacs).
Conidiomata Conidiomata (singular: Conidioma) are blister-like fruiting structures produced by a specific type of fungus called a coelomycete. They are formed as a means of dispersing asexual spores call conidia, which they accomplish by creating the blister- ...
(asexual reproductive structures) have not been observed to occur in this species. Other than trace amounts of unidentified substances, no
secondary metabolite Secondary metabolites, also called ''specialised metabolites'', ''secondary products'', or ''natural products'', are organic compounds produced by any lifeform, e.g. bacteria, archaea, fungi, animals, or plants, which are not directly involved ...
s (
lichen product Lichen products, also known as lichen substances, are organic compounds produced by a lichen. Specifically, they are secondary metabolites. Lichen products are represented in several different chemical classes, including terpenoids, orcinol deri ...
s) have been detected in this species.


Photobiont

''Peltigera cinnamomea'' forms a specialised
symbiotic Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...
relationship with the
cyanobacterium Cyanobacteria ( ) are a group of autotrophic gram-negative bacteria that can obtain biological energy via oxygenic photosynthesis. The name "cyanobacteria" () refers to their bluish green (cyan) color, which forms the basis of cyanobacteria' ...
''Nostoc'', which serves as its primary
photosynthetic Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...
partner (photobiont). Unlike many lichens that may associate with various ''Nostoc'' strains, ''P. cinnamomea'' specifically partners with certain genetic clusters of ''Nostoc''. Genetic studies have identified specific clusters of ''Nostoc'', indicating a closely adapted relationship where the lichen's fungal component and its photosynthetic partner have co-evolved to maximise mutual survival benefits. This partnership benefits ''P. cinnamomea'' by enabling it to efficiently fix atmospheric nitrogen, vital for thriving in nutrient-sparse environments. This tailored relationship suggests that the distribution of ''P. cinnamomea'' may be closely tied to the presence of its specific ''Nostoc'' partners, reflecting a delicate balance between the lichen and its preferred photobiont environments.


Similar species

''Peltigera cinnamomea'' is often recognisable in its natural habitat but was frequently misidentified as ''P. praetextata'' in taxonomic classifications. Unlike ''P. praetextata'', which has brownish or tan-coloured veins that are often tomentose and partially erect, ''P. cinnamomea'' has paler veins. ''P. praetextata'' has somewhat crisped lobe margins with lobules, particularly on older parts of the thallus. Both species occur throughout inland British Columbia, but ''P. cinnamomea'' predominantly inhabits areas with prolonged snow cover, persisting until May or June, conditions under which ''P. praetextata'' is absent. ''Peltigera cinnamomea'' closely resembles the lichen '' P. evansiana'', suggesting they might form a , where the species is fertile, and the reproduces
vegetatively Vegetative reproduction (also known as vegetative propagation, vegetative multiplication or cloning) is a form of asexual reproduction occurring in plants in which a new plant grows from a fragment or Cutting (plant), cutting of the parent pl ...
. This similarity warrants careful observation for accurate identification. Within the genus ''Peltigera'', ''P. cinnamomea'' is grouped with species that feature ''Nostoc'' as a primary photobiont and have an appressed laminal tomentum. It shares similar lobe widths of with ''P. canina'', '' P. membranacea'', and '' P. retifoveata'', unlike the narrower lobes of '' P. didactyla'' and '' P. ponojensis''. The veins of ''P. cinnamomea''—pale to cinnamon brown and low to partly raised—differ markedly from the darker, more elevated veins of ''P. praetextata'' and the broad, strongly raised veins of ''P. retifoveata''. Apothecia are frequently observed in ''P. cinnamomea'', similar to ''P. canina'', ''P. membranacea'', and ''P. rufescens'', but less common in ''P. retifoveata''. Unlike most species in this comparison, only ''P. retifoveata'' consistently demonstrates chemical traits, whereas ''P. cinnamomea'' does not. ''P. cinnamomea'' prefers mesic to hygric environments, particularly snowy locales, unlike the
xeric Deserts and xeric shrublands are a biome defined by the World Wide Fund for Nature. Deserts and xeric (Ancient Greek 'dry') shrublands form the largest terrestrial biome, covering 19% of Earth's land surface area. Ecoregions in this habita ...
to mesic habitats preferred by ''P. canina'', ''P. didactyla'', and ''P. ponojensis''. These distinctions in ecological adaptations and morphological features help identify and differentiate ''P. cinnamomea'' from closely related species.


Habitat and distribution

''Peltigera cinnamomea'' inhabits various forested areas across western North America. It predominantly grows on moss, moss-covered rocks, and logs, in open and somewhat sheltered inland forests at various elevations. This species adapts well to mesophytic to hygrophytic conditions and is particularly prevalent in areas with prolonged snow cover, often surviving under snow until late spring or early summer (May or June). This resilience makes it one of the most snow-tolerant ''Peltigera'' species that occur in the local lichen
funga Funga is all the fungi of a particular region, habitat, or geological period. In life sciences, "funga" is a recent term (2000s) for the kingdom fungi similar to the longstanding ''fauna'' for animals and ''flora'' for plants. The term seeks to ...
. ''Peltigera cinnamomea'' is extensively documented and is locally widespread in suitable forested habitats. It is most abundant in the
Rocky Mountains The Rocky Mountains, also known as the Rockies, are a major mountain range and the largest mountain system in North America. The Rocky Mountains stretch in great-circle distance, straight-line distance from the northernmost part of Western Can ...
. In Canada,
collections Collection or Collections may refer to: Computing * Collection (abstract data type), the abstract concept of collections in computer science * Collection (linking), the act of linkage editing in computing * Garbage collection (computing), autom ...
have been made in Alberta at
Swan Hills Swan Hills is a town in northern Alberta, Canada. It is in the eponymous Swan Hills (Alberta), Swan Hills, approximately north of Whitecourt and northwest of Fort Assiniboine. The town is at the junction of Alberta Highway 32, Highway 32 and Alb ...
and in British Columbia at multiple locations, including the Clearwater River Basin,
Skeena River The Skeena River is the second-longest river entirely within British Columbia, Canada (after the Fraser River). Since ancient times, the Skeena has been an important transportation artery, particularly for the Tsimshian and the Gitxsan—whose na ...
Basin,
Skagit River The Skagit River ( ) is a river in southwestern British Columbia in Canada and northwestern Washington (state), Washington in the United States, approximately 150 mi (240 km) long. The river and its tributaries drain an area of 1.7&nbs ...
Basin,
Fraser River The Fraser River () is the longest river within British Columbia, Canada, rising at Fraser Pass near Blackrock Mountain (Canada), Blackrock Mountain in the Rocky Mountains and flowing for , into the Strait of Georgia just south of the City of V ...
Basin, and
Nechako River The Nechako River (Carrier language, Dakelh: ᘅᐪᙠᗶᑋ) arises on the Nechako Plateau east of the Kitimat Ranges of the Coast Mountains of British Columbia, Canada, and flows north toward Fort Fraser, British Columbia, Fort Fraser, then east ...
Basin. In the United States, it has been identified in
Montana Montana ( ) is a landlocked U.S. state, state in the Mountain states, Mountain West subregion of the Western United States. It is bordered by Idaho to the west, North Dakota to the east, South Dakota to the southeast, Wyoming to the south, an ...
at
Lake Country Lake Country is a district municipality with a population of approximately 15,000 in the Okanagan Valley region of British Columbia, Canada. It is a part of the Central Okanagan Regional District, and of the Kelowna metropolitan area. The city o ...
and in
Pend Oreille County, Washington Pend Oreille County ( ) is a county located in the northeast corner of the U.S. state of Washington, along the Canada–US border. As of the 2020 census, the population was 13,401. The county seat and largest city is Newport. The county w ...
. It also occurs in
Oregon Oregon ( , ) is a U.S. state, state in the Pacific Northwest region of the United States. It is a part of the Western U.S., with the Columbia River delineating much of Oregon's northern boundary with Washington (state), Washington, while t ...
. In
Alaska Alaska ( ) is a non-contiguous U.S. state on the northwest extremity of North America. Part of the Western United States region, it is one of the two non-contiguous U.S. states, alongside Hawaii. Alaska is also considered to be the north ...
, most records of the lichen are from the southeast region of the state. ''P. cinnamomea'' is one of 29 ''Peltigera'' species in the
Pacific Northwest The Pacific Northwest (PNW; ) is a geographic region in Western North America bounded by its coastal waters of the Pacific Ocean to the west and, loosely, by the Rocky Mountains to the east. Though no official boundary exists, the most common ...
region of North America.


Ecology

''Peltigera cinnamomea'' grows in specific
ecological communities In ecology, a community is a group or association of populations of two or more different species occupying the same geographical area at the same time, also known as a biocoenosis, biotic community, biological community, ecological community ...
within
montane Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures lapse rate, fall as elevation increases, causing the ecosystem to stratify. This stratification is ...
and subalpine forests in the northern Rocky Mountains, particularly in the northern Whitefish Range, Montana, USA. These communities are classified in a unique forest order called ''Calamagrostio-Pseudotsugetalia glaucae'', characterised by its dominant plant species and unique environmental conditions. Though termed an "order", this classification groups ecosystems with similar species and interactions, typical of valley bottoms and lower slopes across northwestern North America. These forests are distinct from the coastal forests of the ''Vaccinio-Piceetea'' class, filling niches similar to those of broad-leaved forests in other temperate regions. Influenced by Pacific
air mass In meteorology, an air mass is a volume of air defined by its temperature and humidity. Air masses cover many hundreds or thousands of square miles, and adapt to the characteristics of the surface below them. They are classified according to ...
es, these areas benefit from moist conditions and extensive snow cover, providing a stable environment for the growth of species like ''P. cinnamomea''.


Conservation

''Peltigera cinnamomea'' has an "S5" provincial
conservation status The conservation status of a group of organisms (for instance, a species) indicates whether the group still exists and how likely the group is to become extinct in the near future. Many factors are taken into account when assessing conservation ...
in British Columbia, meaning it is "demonstrably widespread, abundant, and secure". It is also listed as "yellow" in the B.C. List Status, indicating that it is "apparently secure and not at risk of extinction". It has not been assessed 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 ...
. In Oregon, it was listed as a "
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
of concern" by the Oregon Biodiversity Information Center in 2016.


References

{{Taxonbar, from=Q10622568 cinnamomea Lichen species Lichens described in 1995 Lichens of Western Canada Lichens of the Northwestern United States Lichens of Subarctic America Taxa named by Trevor Goward