Trichocolea Tomentella
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''Trichocolea tomentella'' is a species of
liverwort Liverworts are a group of non-vascular land plants forming the division Marchantiophyta (). They may also be referred to as hepatics. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry ...
belonging to the family Trichocoleaceae. It forms loose, pale green to yellowish-white mats and is characterised by its highly divided leaves that give it a feathery or fuzzy appearance. The species has a wide distribution across
temperate In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
regions of the Northern Hemisphere, particularly in oceanic and suboceanic areas, occurring in Europe, Asia, North Africa, and eastern North America. It typically grows in moist, shaded locations, especially near springs and streams in
deciduous In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...
and
coniferous forest Conifers () are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All e ...
s. While capable of
sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...
, with male and female structures on separate plants, it reproduces predominantly through vegetative means via branching and fragmentation. The species shows considerable morphological variation between populations but maintains stable taxonomic characteristics in its cell structure. Though it can form extensive pure patches and effectively compete with other
bryophyte Bryophytes () are a group of embryophyte, land plants (embryophytes), sometimes treated as a taxonomic Division (taxonomy), division referred to as Bryophyta ''Sensu#Common qualifiers, sensu lato'', that contains three groups of non-vascular pla ...
s in suitable habitats, ''T. tomentella'' faces threats from habitat destruction, particularly through logging and drainage of its preferred moist forest habitats.


Taxonomy

''Trichocolea tomentella'' was first documented in 1700 by
Joseph Pitton de Tournefort Joseph Pitton de Tournefort (5 June 165628 December 1708) was a French botanist, notable as the first to make a clear definition of the concept of genus for plants. Botanist Charles Plumier was his pupil and accompanied him on his voyages. Li ...
under the name . In 1783,
Jakob Friedrich Ehrhart Jakob Friedrich Ehrhart (4 November 1742, Holderbank, Aargau – 26 June 1795) was a German botanist, a pupil of Carl Linnaeus at Uppsala University, and later director of the Botanical Garden of Hannover, where he produced several major botanical ...
recognised it as distinct from '' Jungermannia ciliaris'' and named it ''Jungermannia tomentella''. The genus ''Trichocolea'' was proposed by
Barthélemy Charles Joseph Dumortier Barthélemy Charles Joseph Dumortier (; 3 April 1797 – 9 July 1878) was a Belgians, Belgian who conducted a parallel career of botanist and Member of Parliament and is the first discoverer of biological cell division. Over the course of his lif ...
in 1822 (initially spelled as ''Thricholea'') to accommodate this species and ''T. tomentosa''. After several spelling variations,
Christian Gottfried Daniel Nees von Esenbeck Christian Gottfried Daniel Nees von Esenbeck (14 February 1776 – 16 March 1858) was a prolific Germany, German botanist, physician, zoologist, and natural philosopher. He was a contemporary of Goethe and was born within the lifetime of Carl Li ...
standardised the spelling to ''Trichocolea'' in 1838, which was later conserved as the official genus name. The species shows considerable morphological variation between populations, particularly in growth form, plant size, branching frequency, stem thickness, density of stem paraphyllia (small leaf-like structures), and leaf characteristics. Plants growing in very wet, shaded habitats often have a more ascending habit, smaller size, and sparser branches compared to those in merely humid conditions, which tend to grow
prostrate Prostrate may refer to:- *Prostration, a position of submission in religion etc. *Prone position, a face-down orientation of the body *Prostrate shrub A prostrate shrub is a woody plant, most of the branches of which lie upon or just above the ...
and form more compact mats with denser branching. However, certain characteristics remain stable and taxonomically significant, including the thin-walled leaf cells, the - to weakly -verrucose surface of the (hair-like projections), and non-dilated
septa SEPTA, the Southeastern Pennsylvania Transportation Authority, is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly four million people througho ...
between cells.
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 based on ''
rbcL Ribulose-1,5-bisphosphate carboxylase/oxygenase, commonly known by the abbreviations RuBisCo, rubisco, RuBPCase, or RuBPco, is an enzyme () involved in the Photosynthesis#Light-independent reactions, light-independent (or "dark") part of photosyn ...
'' gene sequences have revealed that populations of ''T. tomentella'' show different DNA
haplotype A haplotype (haploid genotype) is a group of alleles in an organism that are inherited together from a single parent. Many organisms contain genetic material (DNA) which is inherited from two parents. Normally these organisms have their DNA orga ...
patterns between European and Southeast Asian populations. European populations show genetic homogeneity across their range, while Southeast Asian populations exhibit three distinct haplotypes. This greater genetic diversity in Asian populations suggests that the species may have originated in Southeast Asia before spreading to Europe, as ancestral populations typically show higher genetic diversity than more recently established ones.


Description

''Trichocolea tomentella'' is a distinctive liverwort that forms loose, pale green to yellowish-white mats. Individual plants grow to in length and have a characteristic feathery or fuzzy appearance due to their highly divided leaves. The plant grows in a branching pattern, with the main stem producing regular side branches that themselves branch two to three times, creating a fern-like appearance. The main stem is relatively robust, measuring about half a millimetre in width, and is covered with tiny leaf-like structures called paraphyllia. The leaves are divided into 6–8 delicate finger-like segments, giving the plant its characteristic fuzzy texture. These segments are made up of elongated cells containing 4–8 oil bodies - small
organelle In cell biology, an organelle is a specialized subunit, usually within a cell (biology), cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as Organ (anatomy), organs are to th ...
s that are characteristic of liverworts. Unlike many other liverworts, ''T. tomentella'' rarely produces root-like structures (
rhizoid Rhizoids are protuberances that extend from the lower epidermal cells of bryophytes and algae. They are similar in structure and function to the root hairs of vascular land plants. Similar structures are formed by some fungi. Rhizoids may be un ...
s). The species is
dioicous Dioicy () is a sexual system in non-vascular plants where archegonia (female organs) and antheridia (male organs) are produced on separate plants in the gametophyte phase. It is one of the two main sexual systems in bryophytes, the other being ...
, meaning male and female reproductive structures occur on separate plants. The male structures (androecia) grow along the main stem, while the female structures (
gynoecia Gynoecium (; ; : gynoecia) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ...
) develop into a club-shaped structure covered in small leaves. When
fertilisation Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a zygote and initiate its development into a new individual organism or of ...
occurs, this develops into a long stalk bearing a narrow, dark brown to black spore capsule. However,
sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...
is rare in many parts of its range, with the plant primarily spreading through
vegetative growth Vegetative phase change is the juvenile-to-adult transition in plants. This transition is distinct from the reproductive transition and is most prolonged and pronounced in woody species. Manipulating phase change may be an important avenue for p ...
. The spores, when produced, are reddish-brown and very small, measuring just in diameter. The plant's distinctive appearance, particularly its pale colour and highly divided leaves, makes it relatively easy to identify even for novice botanists.


Habitat, distribution, and ecology

''Trichocolea tomentella'' has a wide distribution across
temperate In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
regions of the Northern Hemisphere, particularly in oceanic and suboceanic areas. Its range extends from northern Portugal and Spain northwards to southern Norway, Sweden and Finland, and from Ireland and Great Britain eastward to western Russia. The species is also found in Asia, North Africa and eastern North America. The species typically grows in moist, well-shaded locations, with a particular preference for
deciduous forest In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, after flo ...
habitats. It is frequently found near springs,
stream A stream is a continuous body of water, body of surface water Current (stream), flowing within the stream bed, bed and bank (geography), banks of a channel (geography), channel. Depending on its location or certain characteristics, a strea ...
s and
gullies A gully is a landform created by running water, mass movement, or both, which erodes soil to a sharp angle, typically on a hillside or in river floodplains or terraces. Gullies resemble large ditches or small valleys, but are metres to t ...
where there is a constant water supply and moderately diffuse light. In western Norway, it occurs in two distinct habitat types: ''
Alnus glutinosa ''Alnus glutinosa'', the common alder, black alder, European alder, European black alder, or just alder, is a species of tree in the family (biology), family Betulaceae, native plant, native to most of Europe, southwest Asia and northern Africa. ...
'' forests with '' Carex remota'' understory, and on poorer mineral soils in narrow river gorges dominated by either ''
Betula pubescens ''Betula pubescens'' (syn. ''Betula alba''), commonly known as downy birch and also as moor birch, white birch, European white birch or hairy birch, is a species of deciduous tree, native and abundant throughout northern Europe and northern Asia ...
'' or ferns. In other regions, it can be found in coniferous
swamp forest Freshwater swamp forests, or flooded forests, are forests which are inundated with freshwater, either permanently or seasonally. They normally occur along the lower reaches of rivers and around freshwater lakes. Freshwater swamp forests are foun ...
s. ''Trichocolea tomentella'' can form extensive pure patches covering up to , though it often grows mixed with other
bryophyte Bryophytes () are a group of embryophyte, land plants (embryophytes), sometimes treated as a taxonomic Division (taxonomy), division referred to as Bryophyta ''Sensu#Common qualifiers, sensu lato'', that contains three groups of non-vascular pla ...
s. Common associate species include ''
Brachythecium rivulare ''Brachythecium'' is a genus of mosses belonging to the family Brachytheciaceae. The genus was first described by Wilhelm Philippe Schimper. Species The following species are recognised in the genus ''Brachythecium'': * '' Brachythecium albicans ...
'', ''
Thuidium tamariscinum ''Thuidium tamariscinum'' is a species of moss belonging to the family Thuidiaceae. It has an almost cosmopolitan distribution. In a study of the effect of the herbicide Asulam on moss growth, ''Thuidium tamariscinum'' was shown to have intermed ...
'', '' Calliergonella cuspidata'', and various species of ''
Rhytidiadelphus ''Rhytidiadelphus'' is a genus of mosses belonging to the family Hylocomiaceae. The genus was first described by Karl Gustav Limpricht in 1906. The genus has cosmopolitan distribution. Species: * '' Rhytidiadelphus japonicus'' T. Koponen, 197 ...
''. The species reproduces predominantly through
asexual Asexual or Asexuals may refer to: *Asexual reproduction **Asexual reproduction in starfish *Asexuality, the lack of sexual attraction to anyone or lack of interest in or desire for sexual activity. **Gray asexuality, the spectrum between asexualit ...
means via clonal
regeneration Regeneration may refer to: Science and technology * Regeneration (biology), the ability to recreate lost or damaged cells, tissues, organs and limbs * Regeneration (ecology), the ability of ecosystems to regenerate biomass, using photosynthesis ...
and branching, with sexual reproduction (sporophyte production) being rare in much of Europe, though more common in eastern North America. Its dispersal ability appears to be limited, particularly in areas where it reproduces only asexually, though it may occasionally spread through water, mammals, or birds carrying detached
shoot Shoot most commonly refers to: * Shoot (botany), an immature plant or portion of a plant * Shooting, the firing of projectile weapons * Photo shoot, a photography session; an event wherein a photographer takes photographs Shoot may also refer t ...
fragments. The species faces threats from
habitat destruction Habitat destruction (also termed habitat loss or habitat reduction) occurs when a natural habitat is no longer able to support its native species. The organisms once living there have either moved elsewhere, or are dead, leading to a decrease ...
, particularly through
logging Logging is the process of cutting, processing, and moving trees to a location for transport. It may include skidder, skidding, on-site processing, and loading of trees or trunk (botany), logs onto logging truck, trucks ''Trichocolea tomentella'' shows high clonal persistence in undisturbed habitats, with individual plants reproducing primarily through vegetative means via branching and fragmentation rather than sexually through spores. Studies have shown the species maintains relatively high
genetic diversity Genetic diversity is the total number of genetic characteristics in the genetic makeup of a species. It ranges widely, from the number of species to differences within species, and can be correlated to the span of survival for a species. It is d ...
within populations despite limited sexual reproduction, likely due to long-term accumulation of genetic variants through
somatic mutation A somatic mutation is a change in the DNA sequence of a somatic cell of a multicellular organism with dedicated reproductive cells; that is, any mutation that occurs in a cell other than a gamete, germ cell, or gametocyte. Unlike germline muta ...
s and occasional
recruitment Recruitment is #Process, the overall process of identifying, sourcing, screening, shortlisting, and interviewing candidates for Job (role), jobs (either permanent or temporary) within an organization. Recruitment also is the process involved in ...
. The species grows in dense
colonies A colony is a territory subject to a form of foreign rule, which rules the territory and its indigenous peoples separated from the foreign rulers, the colonizer, and their '' metropole'' (or "mother country"). This separated rule was often or ...
that can effectively exclude other bryophytes – as ''T. tomentella'' shoot density increases, the presence of other bryophyte species decreases significantly.
Gene flow In population genetics, gene flow (also known as migration and allele flow) is the transfer of genetic variation, genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent ...
between populations appears to be limited even at small spatial scales (1 km), suggesting the species has restricted dispersal abilities. This combination of traits – clonal persistence, competitive ability in suitable microsites, but limited dispersal – makes the species vulnerable to habitat fragmentation while also allowing it to maintain stable populations in undisturbed conditions. '' Penicillium concentricum'' is an
endophytic An endophyte is an endosymbiont, often a bacterium or fungus, that lives within a plant for at least part of its life cycle without causing apparent disease. Endophytes are ubiquitous and have been found in all species of plants studied to date; ...
fungus that associates with ''Trichocolea tomentella'' and that produces several biologically active
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.


References

{{Taxonbar, from=Q1139601 Jungermanniales Flora of Asia Flora of North Africa Flora of Europe Flora of Northern America Taxa named by Jakob Friedrich Ehrhart Plants described in 1783