Ulota

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Ulota
Ulota coarctata (a, 135714-474438) 7767.JPG
Ulota coarctata
Scientific classification Red Pencil Icon.png
Kingdom: Plantae
Division: Bryophyta
Class: Bryopsida
Subclass: Bryidae
Order: Orthotrichales
Family: Orthotrichaceae
Genus: Ulota
Mohr
U. crispa growing as an epiphyte on a tree. Ulota crispa (30083650062).jpg
U. crispa growing as an epiphyte on a tree.

Ulota is a genus of mosses comprising 69 species with a worldwide distribution, though most species are found in the southern hemisphere.

Contents

They have commonly been grouped within the genus Orthotrichum , though recently they have been placed in their own genus. Their phylogeny and taxonomy have been an area of debate ever since they were placed in their own genus, though it has been largely accepted to place Ulota as a genus of their own. They can be found growing on trees close to the coast, though some species are also found on rocks. Their shoots form small tufts to large mats across habitats, often intertwined with other mosses, including Orthotrichum species.

Ulota reproduce using sexual structures, sporangium (plural: sporangia), that are terminal on the shoot. The calyptra covering the developing sporangium can be hairy or not hairy depending on the species, but the hairs extend from the sporangium base to the apex as opposed to the calyptra hairs of Polytrichum mosses which extend from the apex to the base.

Taxonomy

Due to morphological similarities, this genus has often been taxonomically clustered with the genus Orthotrichum , and has also been considered as a subgenus of Orthotrichum. [1] [2] Often, these two genus are found growing together. Mohr first segregated Ulota from Orthotrichum in 1806, though the segregation has been debated. The two distinguishing features were the splitting of the calyptra at the base and the curled edges of the leaves. However, some Orthotrichum species also have calyptrae which split at the base. [3] Molecular data has made some progress as to differentiating the taxa, though certain morphological features can be distinguishing. [4] Ulota have superficial stomata, whereas Orthotrichum have cryptoporous stomata. [5] Other distinguishing features of Ulota include being monoicous, and the absence of asexual reproductive structures. [5] There is one dioicous species U. phyllantha, and according to molecular studies it is closer related to other Ulota species than Orthotrichum species which can be dioicous. [6] The hyaline cells at the base of the leaf margin also helps to distinguish between Orthotrichum and most Ulota species. [3]

Recent genetic analysis of the mitochondrial genomes of species within Orthotrichum suggests the presence of seven genera within the taxon. [7] The seven genera include: the genus Orthotrichum, which groups together the three previous subgenera Callistoma, Pulchella, and Orthotrichum; [7] the genus Leratia, comprising the previous subgenus Exiguifolium; [8] the genus Lewinskya, which includes the two previous subgenera Phaneroporum and Gymnoporus; [9] the genus Pulvigera, comprising only one species, Orthotrichum lyelli ; [5] the genus Nyholmiella, composed of the previous subgenus Orthophyllum; [6] the genus Plenogemma, composed of the single species U. phyllantha; [5] and the genus Ulota, comprising all previous species classified as Ulota. [7]

Species list

There are currently 69 recognized species of Ulota, over half of which occur only in the southern hemisphere. This number also includes U. phyllantha which has been proposed to reside within its own genus. [2] [5]

Description

Light microscopy of a leaf of U. bruchii showing the rectangular hyaline cells along the margin of the leaf. Ulota bruchii (f, 144651-474826) 8609.JPG
Light microscopy of a leaf of U. bruchii showing the rectangular hyaline cells along the margin of the leaf.
A mature sporangium of U. crispa showing the ribs along the sporangium wall. Ulota crispa (b, 144647-474823) 0442.JPG
A mature sporangium of U. crispa showing the ribs along the sporangium wall.

Ulota mosses tend to grow in a tuft on the substrate, with the shoots growing in an acrocarpous fashion with the shoots standing up. [10] The tufts can vary in colour, ranging from a deep green at the top of the tuft to a darker brown/red at the base of the cushion. [4] The oblong to linear-lanceolate leaves are attached to an erect stem, and can be slightly flexed to fully curled when dry. [11] [5] The genus is characterized by the slightly to fully curled leaves when dry. The leaves have a costa, or midrib, which runs the entire length of the leaf. [5] The leaves have hyaline cells at the basal margins of the leaves, which are typically square or rectangular shaped. [11] The leaf base can be expanded or gradually tapered as it connects to the stem. [12] At the tip of the leaves, there are cells with very thick cell walls. [13] At maturity, the sporangia can be ribbed or unribbed. [4] The sporangia sit on top of a seta, which holds it above the gametophyte. Between species of Ulota, there is variation in the length of the seta which can be a useful trait in classifying species. [12] The sporangia have peristome teeth are diplolepidous, with 8 to 16 exostome teeth and 8 endostome teeth. [11] [5] The sporangia sit on top of a seta which attaches to the gametophyte at the apex of the shoot. [3] The calyptra covers the developing sporangium and is typically conic, split at the base, and can be naked or hairy. [5] The rhizoids are placed at the base of them stem, occasionally appearing along the stem, and are smooth. [4]

Habitat and distribution

Ulota species occur typically in temperate climates, growing as an epiphyte on tree trunks. Some species have also been known to extend into the tropics and subarctic environments. [11] Multiple species can grow together in a habitat, often intertwined with each other. [14] The genus is typically found in temperate, humid, oceanic coasts, though some species are also found in montane environments in the tropics and subtropics. [4] Their habitat is found to be heavily determined by the available substrate, but also on the dispersal ability of the species and the dispersal obstacles that are present. [15] The species which tend to be correlated with the presence of Ulota species include: Salix caprea, Populus tremula, Fraxinus excelsior, Sorbus aucuparia, and Acer pseudoplatanus . [15] They can also be found growing on rocks, though this is reportedly less common. [4] Two species, U. crispa and U. bruchii, tend to be found together, though U. crispa tends to be more common along the coast. [16]

Within their distribution, four areas have the highest rates of endemism for Ulota species: Papua New Guinea (75%), Subtropical China (77.8%), Southern Australasia (78.6%), and South America (81.8%). South America also has the highest richness of Ulota species with 22 recorded species. The next richest area is Australasia with 14 recognized species. [2]

Lifecycle

Lifecycle of a typical moss. Ulota species follow this general lifecycle. Lifecycle moss svg diagram.svg
Lifecycle of a typical moss. Ulota species follow this general lifecycle.

Like other bryophytes, show an alternation of generations. [17] The vegetative gametophyte generation is haploid, and is dominant for most of the life history. [18] Most plants are dioicous except for U. phyllantha, which is monoicous. [6] Asexual reproduction is uncommon, except in U. phyllantha, in which asexual propagules can commonly be found at the leaf apex. [19]

Sexual reproduction involves the transfer of sperm from the antheridium (plural: antheridia) via water. Drops of water carrying sperm allow for the fertilization of the egg housed within the archegonium (plural: archegonia). The heads of the archegonia have a mucilage which attracts the sperm, and the degeneration of the neck canal allows for the passage of sperm to the egg, resulting in fertilization. The fertilized egg develops into a sporophyte, which is diploid, and protected by a remnant of the archegonium now known as the calyptra. Inside the developing sporangium, haploid spores are generated via meiosis. The seta raises the sporangium into the air, and once the sporangium is mature, the operculum falls off and the spores are released into the wind. The peristome teeth are hygroscopic and move in relation to the moisture content of the air, which allows for spores to be released over a longer period of time. [17] Once mature, the sporangia are typically 8-ribbed, though in some species the ribs are only clearly visible right at the base of the peristome mouth. [4]

Spores can be unicellular, although there five known species in which multicellular spores have been identified. Endosporic germination is the process in which mitosis occurs within the sporeling before the spore is released, and once released, it is considered a multicellular spore. [14] This kind of germination is known in all three lineages of bryophytes: liverworts, hornworts, and mosses. [20] Once the spores have landed on suitable substrata, protonematal threads will begin to grow, followed by rhizoids which will help attach the developing plant to the substrate. [14] The leafy shoot develops soon after, reaching upwards towards light. [14]

Conservation

U. bruchii Ulota bruchii (c, 144704-474820) 5284.jpg
U. bruchii

Of the 69 recognized species, three of the species have been assessed by the IUCN. [21] Native to northwestern Europe and Serbia, U. calvescens shows an increasing population trend and is listed "least concerned". There is one protected area within its range, and it is listed as vulnerable in Portugal, Spain, and Serbia, and data deficient in Norway. [22] U. bruchii is found throughout northern and western Europe, with a few pockets in eastern Europe and is listed as "least concerned". [23] U. macrospora exists in a small area within central Europe, and is listed as "endangered". [24] It has a small population size and the current threats it faces includes a decline in habitat area and lowering of habitat quality. [24] Declines in air quality have also been correlated with declines in the size of the tufts of this genus found on trees in urban areas. [25]

Uses

Ulota crispa and U. megalospora have both been found to be intolerant of high levels of sulfur dioxide, which may indicate both species' use as an air quality indicator. [26] [25]

Related Research Articles

Sporangium Enclosure in which spores are formed

A sporangium is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their life cycle. Sporangia can produce spores by mitosis, but in nearly all land plants and many fungi, sporangia are the site of meiosis and produce genetically distinct haploid spores.

Moss Division of non-vascular land plants

Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophytasensu stricto. Bryophyta may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hornworts. Mosses typically form dense green clumps or mats, often in damp or shady locations. The individual plants are usually composed of simple leaves that are generally only one cell thick, attached to a stem that may be branched or unbranched and has only a limited role in conducting water and nutrients. Although some species have conducting tissues, these are generally poorly developed and structurally different from similar tissue found in vascular plants. Mosses do not have seeds and after fertilisation develop sporophytes with unbranched stalks topped with single capsules containing spores. They are typically 0.2–10 cm (0.1–3.9 in) tall, though some species are much larger. Dawsonia, the tallest moss in the world, can grow to 50 cm (20 in) in height. There are approximately 12,000 species.

Bryophyte Terrestrial plants that lack vascular tissue

Bryophytes are a proposed taxonomic division containing three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. They are characteristically limited in size and prefer moist habitats although they can survive in drier environments. The bryophytes consist of about 20,000 plant species. Bryophytes produce enclosed reproductive structures, but they do not produce flowers or seeds. They reproduce sexually by spores and asexually by fragmentation or the production of gemmae. Though bryophytes were considered a paraphyletic group in recent years, almost all of the most recent phylogenetic evidence supports the monophyly of this group, as originally classified by Wilhelm Schimper in 1879. The term bryophyte comes from Ancient Greek βρύον (brúon) 'tree moss, liverwort', and φυτόν (phutón) 'plant'.

Sporophyte Diploid multicellular stage in the life cycle of a plant or alga

A sporophyte is the diploid multicellular stage in the life cycle of a plant or alga. It develops from the zygote produced when a haploid egg cell is fertilized by a haploid sperm and each sporophyte cell therefore has a double set of chromosomes, one set from each parent. All land plants, and most multicellular algae, have life cycles in which a multicellular diploid sporophyte phase alternates with a multicellular haploid gametophyte phase. In the seed plants, the largest groups of which are the gymnosperms and flowering plants (angiosperms), the sporophyte phase is more prominent than the gametophyte, and is the familiar green plant with its roots, stem, leaves and cones or flowers. In flowering plants the gametophytes are very reduced in size, and are represented by the germinated pollen and the embryo sac.

Hornwort One of the three Divisions of bryophytic plants

Hornworts are a group of bryophytes constituting the division Anthocerotophyta. The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, the flattened, green plant body of a hornwort is the gametophyte plant.

<i>Fissidens adianthoides</i> Species of moss

Fissidens adianthoides, the maidenhair pocketmoss, is a moss in the family of Fissidentaceaea. It was first collected by Hedwig in 1801.

<i>Funaria</i> Genus of mosses

Funaria is a genus of approximately 210 species of moss. Funaria hygrometrica is the most common species. Funaria hygrometrica is called “cord moss” because of the twisted seta which is very hygroscopic and untwists when moist. The name is derived from the Latin word “funis”, meaning "a rope". In funaria root like structures called rhizoids are present.

Monoicy is a sexual system in haploid plants where both sperm and eggs are produced on the same gametophyte, in contrast with dioicy, where each gametophyte produces only sperm or eggs but never both. Both monoicous and dioicous gametophytes produce gametes in gametangia by mitosis rather than meiosis, so that sperm and eggs are genetically identical with their parent gametophyte.

Operculum (botany)


In botany, an operculum or calyptra is a cap-like structure in some flowering plants, mosses, and fungi. It is a covering, hood or lid, describing a feature in plant morphology.

<i>Buxbaumia</i> Genus of mosses

Buxbaumia is a genus of twelve species of moss (Bryophyta). It was first named in 1742 by Albrecht von Haller and later brought into modern botanical nomenclature in 1801 by Johann Hedwig to commemorate Johann Christian Buxbaum, a German physician and botanist who discovered the moss in 1712 at the mouth of the Volga River. The moss is microscopic for most of its existence, and plants are noticeable only after they begin to produce their reproductive structures. The asymmetrical spore capsule has a distinctive shape and structure, some features of which appear to be transitional from those in primitive mosses to most modern mosses.

Funariidae Subclass of mosses

The Funariidae are a widespread group of mosses in class Bryopsida. The majority of species belong to the genera Funaria and Physcomitrium.

<i>Tortilicaulis</i> Extinct genus of Devonian plants

Tortilicaulis is a moss-like plant known from fossils recovered from southern Britain, spanning the Silurian-Devonian boundary. Originally recovered from the Downtonian of the Welsh borderlands, Tortilicaulis has since been recovered in the famous Ludlow Lane locality.

Orthotrichum scanicum is a species of moss in the Orthotrichaceae family. It is found in Austria, the Czech Republic, Denmark, France, Germany, Greece, Hungary, Italy, Norway, Poland, Russia, Serbia and Montenegro, Sweden, and Switzerland. Its natural habitat is temperate forests. It is threatened by habitat loss.

Splachnaceae Family of mosses

Splachnaceae is a family of mosses, containing around 70 species in 6 genera. Around half of those species are entomophilous, using insects to disperse their spores, a characteristic found in no other seedless land plants.

<i>Pogonatum urnigerum</i> Species of moss

Pogonatum urnigerum is a species of moss in the family Polytrichaceae, commonly called urn haircap. The name comes from "urna" meaning "urn" and "gerere" meaning "to bear" which is believed to be a reference made towards the plant's wide-mouthed capsule. It can be found on gravelly banks or similar habitats and can be identified by the blue tinge to the overall green colour. The stem of this moss is wine red and it has rhizoids that keep the moss anchored to substrates. It is an acrocarpous moss that grows vertically with an archegonium borne at the top of each fertilized female gametophyte shoot which develops an erect sporophyte.

<i>Climacium dendroides</i> Species of moss

Climacium dendroides, also known as tree climacium moss, belongs in the order Hypnales and family Climaciaceae, in class Bryopsida and subclass Bryidae. It is identified as a "tree moss" due to its distinctive morphological features, and has four species identified across the Northern Hemisphere. The species name "dendroides" describes the tree-like morphology of the plant, and its genus name came from the structure of the perforations of peristome teeth. This plant was identified by Weber and Mohr in 1804. They often have stems that are around 2-10 cm tall and growing in the form of patches, looking like small palm-trees. They have yellow-green branches at the tip of stems. The leaves are around 2.5-3 mm long, with rounder stem leaves and pointier branch leaves. Their sporophytes are only abundant in late winter and early spring, and appears as a red-brown shoot with long stalk and cylindrical capsules.

<i>Orthotrichum lyellii</i> Species of moss

Orthotrichum lyellii, also known as Lyell's bristle moss, is a species of acrocarpous moss belonging to the family Orthotrichaceae. O. lyellii can be found throughout western North America and Europe. It is found most commonly growing epiphytically on a variety of trees, and less commonly on rocks or boulder substrates.

<i>Andreaea rupestris</i> Species of moss

Andreaea rupestris is a species of moss in the class Andreaeopsida, are commonly referred to as the "lantern mosses" due to the appearance of their dehisced sporangia. It is typically found on smooth, acidic, exposed rock in the Northern hemisphere. It exhibits the common features of the genus Andreaea such as being acrocarpous, having dark pigmentation, lacking a seta, and bearing 4 lines of dehiscence in its mature sporangia, but can be further identified upon careful examination of its gametophytic leaves which have an ovate base to a more blunt apex compared to other similar species.

<i>Octoblepharum</i> Genus of haplolepideous mosses

Octoblepharum is a genus of haplolepideous mosses (Dicranidae) in the monotypic family Octoblepharaceae . The genus Octoblepharum was previously placed in family Calymperaceae.

References

  1. ↑ Mitten, William (1869). Musci Austro-Americani: Sive Enumeratio Muscorum Omnium Austro-Americanorum Mihi Hucusque Cognitorum, Eorum Praecipue in Terris Amazonicis Andinisque Ricardo Spruceo Lectorum. Longmans, Green, Reader, & Dyer.
  2. 1 2 3 Garilleti, Ricardo; Mazimpaka, Vicente; Lara, Francisco (2015). "Ulota larrainii (Orthotrichoideae, Orthotrichaceae, Bryophyta) a new species from Chile, with comments on the worldwide diversification of the genus". Phytotaxa. 217 (2): 133–144. doi:10.11646/phytotaxa.217.2.3 – via Biotaxa.
  3. 1 2 3 Wang, Qing-Hua; Jia, Yu (November 2012). "A taxonomic revision of the Asian species of Ulota Mohr (Orthotrichaceae)". The Bryologist. 115 (3): 412–443. doi:10.1639/0007-2745-115.3.412. ISSN   0007-2745. S2CID   86387122.
  4. 1 2 3 4 5 6 7 Caparrós, Rut; Lara, Francisco; Draper, Isabel; Mazimpaka, Vicente; Garilleti, Ricardo (2016-04-01). "Integrative taxonomy sheds light on an old problem: the Ulota crispa complex (Orthotrichaceae, Musci)". Botanical Journal of the Linnean Society. 180 (4): 427–451. doi: 10.1111/boj.12397 . ISSN   0024-4074.
  5. 1 2 3 4 5 6 7 8 9 Plášek, Vítězslav; Sawicki, Jakub; Ochyra, Ryszard; Szczecińska, Monika; Kulik, Tomasz (2015). "New taxonomical arrangement of the traditionally conceived genera Orthotrichum and Ulota (Orthotrichaceae, Bryophyta)". Acta Musei Silesiae, Scientiae Naturales. 64 (2): 169–174. doi: 10.1515/cszma-2015-0024 . ISSN   2336-3207.
  6. 1 2 3 Sawicki, Jakub; Plášek, Vítězslav; Szczecińska, Monika (2010). "Molecular studies resolve Nyholmiella (Orthotrichaceae) as a separate genus". Journal of Systematics and Evolution. 48 (3): 183–194. doi:10.1111/j.1759-6831.2010.00076.x. ISSN   1759-6831. S2CID   84318916.
  7. 1 2 3 Sawicki, Jakub; Plášek, Vítězslav; Ochyra, Ryszard; Szczecińska, Monika; Ślipiko, Monika; Myszczyński, Kamil; Kulik, Tomasz (2017-06-30). "Mitogenomic analyses support the recent division of the genus Orthotrichum (Orthotrichaceae, Bryophyta)". Scientific Reports. 7 (1): 4408. Bibcode:2017NatSR...7.4408S. doi:10.1038/s41598-017-04833-z. ISSN   2045-2322. PMC   5493672 . PMID   28667304.
  8. ↑ Goffinet, B.; Shaw, A. J.; Cox, C. J.; Wickett, N. J.; Boles, S. (2004). "Phylogenetic inferences in the Orthotrichoideae (Orthotrichaceae: Bryophyta) based on variation in four loci from all genomes". Monographs in Systematic Botany from the Missouri Botanical Garden. 98: 270–289.
  9. ↑ Lara, Francisco; Garilleti, Ricardo; Goffinet, Bernard; Draper, Isabel; Medina, Rafael; Vigalondo, Beatriz; Mazimpaka, Vicente (2016). "Lewinskya, a New Genus to Accommodate the Phaneroporous and Monoicous Taxa of Orthotrichum (Bryophyta, Orthotrichaceae)". Cryptogamie, Bryologie. 37 (4): 361–382. doi:10.7872/cryb/v37.iss4.2016.361. ISSN   1290-0796. S2CID   88561982.
  10. ↑ Caparrós, Rut; Lara, Francisco; Long, David G.; Mazimpaka, Vicente; Garilleti, Ricardo (2011-09-01). "Two new species of Ulota (Orthotrichaceae, Bryopsida) with multicellular spores, from the Hengduan Mountains, Southwestern China". Journal of Bryology. 33 (3): 210–220. doi:10.1179/1743282011Y.0000000008. ISSN   0373-6687. S2CID   84279281.
  11. 1 2 3 4 Wang, Qinghua; Jia, Yu (2016-04-01). "A taxonomic revision of Ulota Mohr (Orthotrichaceae) in South and Central America". Plant Diversity. 38 (2): 65–88. doi:10.1016/j.pld.2016.03.003. ISSN   2468-2659. PMC   6112220 . PMID   30159451.
  12. 1 2 Blockeel, Tom; Bosanquet, Sam (May 2017). "The Ulota crispa group in Britain and Ireland, with notes on other species of the genus" (PDF). Field Bryology. 117: 8–19 – via British Bryological Society.
  13. ↑ Vitt, Dale; Koponen, Timo; Norris, Daniel (1993). "Bryophyte flora of the Huon Peninsula, Papua New Guinea. LIII". Acta Botanica Fennica. 148: 5–25 – via ResearchGate.
  14. 1 2 3 4 Garilleti, Ricardo; Mazimpaka, Vicente; Lara, Francisco (December 2012). "New Ulota species with multicellular spores from southern South America". The Bryologist. 115 (4): 585–600. doi:10.1639/0007-2745-115.4.585. ISSN   0007-2745. S2CID   86367474.
  15. 1 2 Heegaard, Einar; Hangelbroek, Helen H. (1999). "The Distribution of Ulota crispa at a Local Scale in Relation to Both Dispersal- and Habitat-Related Factors". Lindbergia. 24 (2): 65–74. ISSN   0105-0761. JSTOR   20150006.
  16. ↑ Albertos, B.; Lara, F.; Garilleti, R.; Mazimpaka, V. (2000). "Distribution and abundance of corticolous species of the genus Ulota (Orthotrichaceae, Musci) in the NW Iberian Peninsula". Nova Hedwigia. 70 (3–4): 461–470. doi:10.1127/nova.hedwigia/70/2000/461. hdl: 11268/5759 .
  17. 1 2 McMillin, John (1966). "A Laboratory Investigation of the Mosses". The American Biology Teacher. 28 (10): 814–817. doi:10.2307/4441553. ISSN   0002-7685. JSTOR   4441553.
  18. ↑ During, Heinjo J. (1979). "Life Strategies of Bryophytes: A Preliminary Review". Lindbergia. 5 (1): 2–18. ISSN   0105-0761. JSTOR   20149317.
  19. ↑ Seppelt, R. D. (1978-03-01). "Studies on the bryoflora of Macquarie Island". New Zealand Journal of Botany. 16 (1): 21–23. doi: 10.1080/0028825X.1978.10429654 . ISSN   0028-825X.
  20. ↑ Schuette, Scott; Renzaglia, Karen S. (2010). "Development of multicellular spores in the hornwort genus Dendroceros (Dendrocerotaceae, Anthocerotophyta) and the occurrence of endospory in Bryophytes". Nova Hedwigia. 91 (3): 301–316. doi:10.1127/0029-5035/2010/0091-0301.
  21. ↑ "The IUCN Red List of Threatened Species". IUCN Red List of Threatened Species. Retrieved 2020-04-05.
  22. ↑ Consultant), Nick Hodgetts (Europe LIFE project-Freelance (2017-05-20). "IUCN Red List of Threatened Species: Balding Pincushion". IUCN Red List of Threatened Species. Retrieved 2020-04-05.
  23. ↑ Consultant), Nick Hodgetts (Europe LIFE project-Freelance (2017-05-20). "IUCN Red List of Threatened Species: Bruch's pincushion". IUCN Red List of Threatened Species. Retrieved 2020-04-05.
  24. 1 2 Vana, Jiri; Norbert Schnyder (University of Zürich, Switzerland); Consultant), Nick Hodgetts (Europe LIFE project-Freelance; History), Irene Bisang (Swedish Museum of Natural; Baudraz, Maude; Kučera, Jan; Ştefănuţ, Sorin; Lars Hedenäs (Natural History Museum, Stockholm); Steffen Caspari (Zentrum für Biodokumentation, Schiffweiler (2017-01-04). "IUCN Red List of Threatened Species: Ulota macrospora". IUCN Red List of Threatened Species. Retrieved 2020-04-05.
  25. 1 2 Hallingbäck, Tomas (1992-01-01). "The effect of air pollution on mosses in southern Sweden". Biological Conservation. Endangered Bryophytes in Europe-Causes and Conservation. 59 (2): 163–170. doi:10.1016/0006-3207(92)90577-A. ISSN   0006-3207.
  26. ↑ Ireland, Robert R. (2004). "Review of Grimmias of the World". The Bryologist. 107 (2): 274–276. doi:10.1639/0007-2745(2004)107[0275:R]2.0.CO;2. ISSN   0007-2745. JSTOR   3244857.
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