Bryoria | |
---|---|
Bryoria nadvornikiana | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Fungi |
Division: | Ascomycota |
Class: | Lecanoromycetes |
Order: | Lecanorales |
Family: | Parmeliaceae |
Genus: | Bryoria Brodo & D.Hawksw. (1977) |
Type species | |
Bryoria trichodes (Michx.) Brodo & D.Hawksw. (1977) | |
Synonyms [1] | |
Bryoria is a genus of lichenized fungi in the family Parmeliaceae. Many members of this genus are known as horsehair lichens. [2] The genus has a widespread distribution, especially in boreal and cool temperate areas.
Bryoria was circumscribed in 1977 by lichenologists Irwin Brodo and David Hawksworth, with Bryoria trichodes as the type species. [3]
Molecular studies have revealed that the genus originated approximately 11.5 million years ago during the Miocene period. The genus comprises several sections, including a recently revised understanding of section Bryoria, which is now known to be polyphyletic and divided into two distinct clades. The first group (Bryoria clade 1) has been placed in an emended section Americanae, which is primarily restricted to western North America and the Himalayan region, with B. americana being the only widely distributed species in this section. [4]
Molecular studies by Velmala et al. (2014) [5] and Boluda et al. (2019) [6] showed that several North American Bryoria taxa, including B. pikei (formerly identified as North American "B. capillaris"), B. pseudofuscescens , B. friabilis, and B. inactiva, belong to a single genetic clade with minimal genetic variation. While these taxa were originally distinguished by their secondary metabolites, ecology, and geographic distributions, DNA analysis did not support maintaining them as separate species. However, Bruce McCune and Daphne Stone (2022) noted that these chemical variants have distinct ecological preferences – for instance, B. friabilis is associated with inland wet forests and is common at low elevations in western Oregon and Washington, while B. pseudofuscescens dominates at higher elevations in the Cascades. To preserve this valuable ecological information while reflecting genetic reality, these taxa were reclassified as varieties of B. pseudofuscescens, distinguished by their chemical composition: var. pseudofuscescens (norstictic acid), var. friabilis (gyrophoric acid), var. inactiva (acid-deficient), and var. pikei (alectorialic acid). [7]
Of particular interest is section Divaricatae, which molecular studies indicate is relatively young (approximately 5 million years old) and is currently undergoing active diversification. This diversification is particularly evident in South-East Asia and western North America, where new species continue to be discovered and described. [4]
Bryoria species are fruticose, slender and hair-like, tending to grow hanging (pendent) or like a small bush. They range in colour from dark brown to pale greyish brown to grey in some species. Ascospores are colourless, ellipsoid, numbering eight per ascus. [8]
The genus shows considerable morphological variation, particularly well demonstrated in species like B. tenuis. Within this species alone, five distinct growth forms have been documented: cobwebby (finely threadlike and pliant throughout), threadlike- spinulose (with short, stiff branchlets ), threadlike- flexuose (with longer, more flexible branchlets), thickening-spinulose (thickened and brittle in older parts with stiff branchlets), and thickening-flexuose (thickened stems with flexible branchlets). These variations can occur within the same species, suggesting significant morphological plasticity. [4] Main stems in Bryoria species can vary from uniformly thin throughout to distinctly thickened, particularly in basal portions. Third-order branchlets, when present, may be short and stiff or longer and more flexuous, with their arrangement varying from sparse to abundant. These morphological characteristics, while useful for identification, should be considered alongside other features as they can vary considerably within species. [4]
Other hair-like lichens that may be confused with Bryoria include dark brown species of Bryocaulon , Nodobryoria , Pseudephebe , Alectoria , or Cetraria . [8]
Found almost exclusively on conifers, or on tundra soil, [8] Bryoria collectively has a widespread distribution, especially in boreal and cool temperate areas. [9] The genus shows distinct regional diversity patterns, with particularly high species richness and ongoing diversification in South-East Asia and western North America. Some species show restricted distributions – for example, several recently described species from the Hengduan Mountains of China are known only from that region, while others like B. americana have broad circumpolar distributions. The genus appears to have undergone significant recent diversification in these areas, particularly within the last 5 million years. [4]
Studies have shown that species in the genus Bryoria, characterized by their dark melanic pigments, show distinct ecological preferences compared to pale yellow-green usnic acid-containing hair lichens. Research on coniferous forests in British Columbia demonstrated that Bryoria species strongly favor sun-exposed, well-ventilated locations, particularly dominating south-facing slopes and upper canopy positions. This distribution pattern reflects an evolutionary specialization where melanic pigments provide crucial protection against high light exposure, allowing these lichens to thrive in dry, sun-exposed environments while shielding their photobionts from radiation damage. The melanin in their cortex effectively absorbs most incident solar radiation, enabling Bryoria species to tolerate temperatures up to 70 °C (158 °F) when desiccated. This adaptation contrasts with usnic acid-containing hair lichens, which typically prefer more shaded and humid conditions. The strong correlation between Bryoria abundance and canopy openness suggests that these lichens are particularly well-adapted to early-successional forest conditions where light exposure is high. [10]
The Parmeliaceae is a large and diverse family of Lecanoromycetes. With over 2700 species in 71 genera, it is the largest family of lichen-forming fungi. The most speciose genera in the family are the well-known groups: Xanthoparmelia, Usnea, Parmotrema, and Hypotrachyna.
Irwin Murray Brodo is an emeritus scientist at the Canadian Museum of Nature, in Ottawa, Ontario, Canada. He is an authority on the identification and biology of lichens.
Pseudephebe is a genus of fruticose lichens in the family Parmeliaceae. It contains three species that grow on rocks.
Hypogymnia is a genus of foliose lichens in the family Parmeliaceae. They are commonly known as tube lichens, bone lichens, or pillow lichens. Most species lack rhizines that are otherwise common in members of the Parmeliaceae, and have swollen lobes that are usually hollow. Other common characteristics are relatively small spores and the presence of physodic acid and related lichen products. The lichens usually grow on the bark and wood of coniferous trees.
Melanohalea is a genus of foliose lichens in the family Parmeliaceae. It contains 30 mostly Northern Hemisphere species that grow on bark or on wood. The genus is characterised by the presence of pseudocyphellae, usually on warts or on the tips of isidia, a non-pored epicortex and a medulla containing depsidones or lacking secondary metabolites. Melanohalea was circumscribed in 2004 as a segregate of the morphologically similar genus Melanelia, which was created in 1978 for certain brown Parmelia species. The methods used to estimate the evolutionary history of Melanohalea suggest that its diversification primarily occurred during the Miocene and Pliocene epochs.
Nodobryoria is a genus of medium to large, reddish-brown lichens that are hair-like to shrubby in shape and grow on conifer trees. The genus contains three species, distributed in North America and Greenland, which were previously included in the genus Bryoria. Nodobryoria is similar in appearance to Bryoria, but is differentiated because it does not contain the polysaccharide lichenin, and it has a unique cortex composed of interlocking cells that look like pieces of a jigsaw puzzle when viewed under a light microscope.
Gowardia is a genus of medium-sized, greyish hair lichens in the family Parmeliaceae. It is a circumpolar genus, mainly restricted to arctic-alpine habitats in northern Canada, Europe, and Russia.
Alectoria is a genus of fruticose lichens belonging to the family Parmeliaceae.
Bryoria hengduanensis is a species of lichen of the genus Bryoria. It was described as new to science in 2003 by lichenologists Li-Song Wang and Hiroshi Harada. It is found in the Hengduan Mountains of southern China, where it grows on twigs and branches in coniferous forests at elevations of 3,000–4,000 metres (9,800–13,100 ft). The Hengduan Mountains is a region of high Bryoria biodiversity, as 24 species are known from this area.
Teuvo ("Ted") Tapio Ahti is a Finnish botanist and lichenologist who has made significant contributions to the taxonomy and biogeography of lichens. Known particularly for his work on the lichen family Cladoniaceae, he has had a long career at the University of Helsinki beginning in 1963, and following his retirement in 1997, has continued his research at the Botanical Museum of the Finnish Museum of Natural History. His research output spans more than seven decades, comprising over 450 scientific publications across lichenology, mycology, and botanical science.
Melanohalea zopheroa is a species of lichen in the family Parmeliaceae. It was first formally described in 1977 by Ted Esslinger as Parmelia zopheroa. A year later, he transferred it to the new genus Melanelia, which he created to contain the brown Parmeliae species. In 2004, after early molecular phylogenetic evidence showed that Melanelia was not monophyletic, Melanohalea was circumscribed by lichenologists Oscar Blanco, Ana Crespo, Pradeep K. Divakar, Esslinger, David L. Hawksworth and H. Thorsten Lumbsch, and M. zopheroa was transferred to it. The lichen has a disjunct distribution, as it is found in South America (Chile) and in New Zealand.
Bryoria implexa is a species of horsehair lichen in the family Parmeliaceae.
Bryoria nadvornikiana, commonly known as the spiny grey horsehair lichen or the blonde horsehair lichen, is a species of horsehair lichen in the family Parmeliaceae.
Bryoria subcana is a species of horsehair lichen in the family Parmeliaceae. It is found in North America and Europe.
Bryoria kockiana is a species of horsehair lichen in the family Parmeliaceae. It is found in North America, where it grows from the branches of conifer trees.
Raesaenenia is a fungal genus in the large family Parmeliaceae. It is a monotypic genus, containing the single lichenicolous fungus Raesaenenia huuskonenii, which parasitises lichens of genus Bryoria in the Northern Hemisphere.
Gowardia zebrina is a rare species of fruticose lichen in the family Parmeliaceae. Found in Canada, it was formally described as a new species in 2020 by the lichenologists Trevor Goward and Leena Myllys. The type specimen was collected by Goward on Mount Cain at an altitude of 1,440 m (4,720 ft). Here the lichen was found growing on the lower branches of a hemlock tree. The specific epithet zebrina refers to the diagnostic alternating pale and dark banding pattern of the terminal branches. The lichen has a limited distribution, as it is only known to occur in coastal northwest North America, extending from the Insular Mountains of southern Vancouver Island north to Hazelton. Its preferred habitat is in open, wind-scoured old growth forests, and its preferred substrate is the trunks and lower branches of conifer trees like fir and hemlock.
Gowardia nigricans, commonly known as the gray hair lichen or gray witch's hair, is a species of fruticose lichen in the family Parmeliaceae.
Gowardia arctica is a species of terricolous (ground-dwelling), fruticose (bushy) lichen in the family Parmeliaceae. Found in arctic regions of Northern Canada and Russia, it was formally described as a new species in 2009 by Pekka Halonen, Leena Myllys, Saara Velmala, and Heini Hyvärinen. The type specimen was collected from Banks Island in Swan Lake ; here, at an elevation of 100 m (330 ft), it was found growing among mesic mountain heath. It also occurs along the Arctic Ocean coast of Russia. The lichen is richly branched, black to black-brown in colour, and reaches up to 13 cm (5.1 in) in diameter. It contains alectorialic acid and two other unknown lichen products.
Sulcaria spiralifera is a species of fruticose lichen in the family Parmeliaceae. First described in 1977 as a species of Bryoria, it was transferred to the genus Sulcaria in 2014 based on DNA analysis. The species comprises two chemical variants now recognized as varieties: the typical variety and var. pseudocapillaris, which differ in their chemical composition and spot test reactions. It is found in the northwestern United States, where it grows as an epiphyte, hanging from a variety of tree species in open or shaded maritime forests. The lichen is characterized by its pendulous brown thallus with spiral-arranged white pores (pseudocyphellae) on its surface, extending 4–12 cm in length.