Porella platyphylla | |
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Scientific classification | |
Kingdom: | Plantae |
Division: | Marchantiophyta |
Class: | Jungermanniopsida |
Order: | Porellales |
Family: | Porellaceae |
Genus: | Porella |
Species: | P. platyphylla |
Binomial name | |
Porella platyphylla | |
Synonyms [1] | |
Porella platyphylla is a species of liverwort belonging to the family Porellaceae. It has a Holarctic distribution, occurring across Eurasia and North America, [2] where it typically grows on tree bark and rocks in areas with adequate rainfall. The species is most common in regions receiving at least 600 millimetres of annual precipitation. The species forms part of a complex taxonomic group that includes several closely related species and hybrids, with populations showing distinct genetic differences between continents despite their morphological similarity.
The species is characterised by its obtuse-rounded leaf lobes with flat margins, and irregular teeth at the perianth mouth when mature. It reproduces both sexually, with separate male and female plants, and asexually through leaf fragments. P. platyphylla is known for its ability to survive long periods of desiccation, being able to recover normal photosynthetic function within hours of rehydration. The species produces unique chemical compounds, including pinguisane sesquiterpenoids and sacculatane diterpenoids, which help distinguish it from related species. Recent molecular studies have revealed significant genetic variation within the species, particularly in European populations, though this variation is not always reflected in physical characteristics.
The species was first described by Carl Linnaeus, with early descriptions and illustrations provided by Pier Antonio Micheli (1729) and Johann Jacob Dillenius (1741). P. platyphylloidea was later described by Lewis David von Schweinitz in 1821 from a North Carolina collection, who distinguished it primarily by its occurrence on rocks rather than bark, larger plants with often tripinnate branching, quadrate leaves with inflexed margins, and differences in the relative sizes of the plant's structural features. [3]
Historical taxonomic treatments varied in their interpretation of these plants. While some researchers like Marshall Avery Howe (1897) and Theodore Christian Frye and Lois Clark (1946) treated them as a single variable species, others such as Alexander William Evans (1916), Karl Müller of Freiburg (1915), and Rudolf Mathias Schuster (1980) maintained them as separate species. The perianth mouth structure and elater spiral patterns were particularly emphasised as diagnostic features by later authors. [3]
A comprehensive 1998 study examining both morphological and genetic variation in Porella platyphylla and P. platyphylloidea led to a significant taxonomic revision. The research, which analysed 35 populations across North America and Europe, revealed three distinct genetic groups within what had previously been considered two separate species. To help stabilize the taxonomy of this complex group, researchers later designated a DNA voucher specimen from Germany (Heinrichs and Feldberg 4600) as an epitype for P. platyphylla. [4]
Porella platyphylla forms part of a complex of closely related taxa, with its closest relative being P. cordaeana . DNA analyses have revealed two main clades within P. platyphylla: one predominantly North American and another widespread in Europe but also present in North America and Asia. Both species maintain distinct European and North American lineages that form sister groups in phylogenetic analyses. [4]
Porella cordaeana can be distinguished from P. platyphylla by its crenulate -sinuous perianth mouth and acute, ventrally reflexed and twisted lobules. While both species have leaf lobules that fold under, in P. cordaeana these are twisted and taper to a point, whereas in P. platyphylla they are merely folded under without twisting and have rounded or obtuse tips. [5]
These species can sometimes be found growing together in the same habitat, and multiple species may even grow intermixed within a single colony. This close physical association, combined with their ability to reproduce sexually, has led to hybridisation between species. [5] A hybrid between these species, P. × baueri has been documented in Europe. This hybrid is an allopolyploid, meaning it originated through hybridisation between two species and underwent chromosome doubling. While P. platyphylla and P. cordaeana are typically haploid with eight chromosomes (occasionally nine in P. platyphylla), their hybrid derivative P. × baueri is polyploid. Genetic evidence suggests this hybridisation event was relatively recent, and the hybrid may have originated multiple times. [4]
Analyses of chloroplast and nuclear DNA have produced conflicting evolutionary trees, suggesting that the North American populations formerly known as P. platyphylloidea may be the result of ancient hybridisation between P. cordaeana and P. platyphylla, though through a different and much older hybridisation event than that which produced P. × baueri. [4]
Porella platyphylla shows several distinctive morphological features that help identify it, though these can show considerable variation both within single populations and between different geographical regions. The leaves have tips (known as lobes) that are obtuse-rounded in shape, with edges that lie flat rather than being wavy or folded. The cells in the middle portion of these lobes measure between 28.5 and 36.1 μm in width. [5]
A key identifying feature of the species is found in its perianth—a protective tube-like structure that surrounds the developing reproductive organs in female plants. When fully mature, the mouth of this perianth has irregularly spaced teeth that are 2–6 cells wide at their base. The female reproductive structures (called bracts) typically have smooth, unbroken edges, unlike some related species which may have toothed or spiny margins. [5]
Features previously used to distinguish P. platyphylla from P. platyphylloidea, such as leaf lobule width and elater spiral patterns, have shown no consistent correlation with genetic groupings. Even populations that were genetically identical sometimes displayed different morphological characteristics traditionally used to separate the species. More recent molecular studies have revealed the existence of cryptic species—genetically distinct lineages that cannot be reliably distinguished through morphological examination alone. While some tendencies exist, such as larger leaf lobules being more common in North American populations, these characteristics are not consistent enough for reliable identification. This challenge is further complicated by evidence that some populations may represent ancient hybrid lineages, explaining the high degree of morphological variability observed within populations and the unreliability of traditionally used diagnostic features. [4]
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The life cycle and development of Porella platyphylla was first studied in detail in the early 20th century. Like other liverworts, this species displays distinct stages in its life cycle, including both a dominant gametophyte (the main plant body) and a smaller sporophyte phase. [6]
The main plant body (gametophyte) grows flat against its substrate in a pattern known as dorsiventral (having distinct upper and lower surfaces). It develops from a pyramid-shaped growing tip called an apical cell, which continually divides to form new tissue. The plant produces three rows of leaves: two rows on the upper ( dorsal ) surface and one row on the lower ( ventral ) surface. The ventral leaves, technically called amphigastria, are supplemented by special flaps or lobes from the dorsal leaves that fold underneath, giving the appearance of five rows of leaves when viewed from below. [6]
Porella platyphylla reproduces sexually, with male and female reproductive organs developing on short side branches. The female organs (archegonia) develop on particularly short branches and consist of a protective jacket of cells surrounding a central channel containing an egg cell. The male organs (antheridia) form on slightly longer branches and produce sperm cells. Each antheridium develops on a long, slender stalk made up of two rows of cells. [6] Individual colonies are often multiclonal, with male and female plants growing intermixed, indicating establishment from multiple spores. While female plants are relatively common (about 49% of shoots), male plants are less frequently found (around 10% of shoots). [5]
After fertilisation, the embryo develops into a sporophyte—the spore-producing generation. The mature sporophyte consists of a capsule containing spores mixed with elongated cells called elaters, which help disperse the spores. The sporophyte base forms a foot that anchors it to the parent plant, with the shape of this foot varying from club-like to anchor-shaped. [6]
One main characteristic of P. platyphylla is its ability to survive long periods of drying out (desiccation). The plant can remain dormant in a dried state for months or even years, reviving quickly when moisture becomes available again. This adaptation allows it to survive in environments with irregular water availability. [6]
Studies using chlorophyll fluorescence measurements have shown that the species can recover rapidly and completely after spending a week in an air-dry state. When rehydrated, the plant's photosynthetic apparatus returns to normal functioning within just two hours, regardless of whether recovery takes place in light or darkness. This rapid recovery suggests that P. platyphylla has "constitutive" protection mechanisms—meaning the protective systems are always present rather than being produced in response to drying. [7]
During drying, the plant's photosynthetic efficiency remains near optimal until water content falls below about half of the fully hydrated state, after which it declines steeply. However, this decline is fully reversible, and the period of desiccation appears as only a temporary interruption of normal photosynthetic function. The plant shows increased photoprotection during recovery in light conditions, with these protective processes being essentially complete within 24 hours. [7]
Porella platyphylla produces a wide range of terpenoid metabolites, including mono-, sesqui-, di- and triterpenoids. The species belongs to the "nonpungent" chemotype of Porella species (referring to taste rather than smell), producing primarily pinguisane sesquiterpenoids and sacculatane diterpenoids – carbon skeletons that remain unique to liverworts. [8] The species can be distinguished chemically from pungent Porella species, which produce intensely pungent drimane compounds and related substances. P. platyphylla belongs to the pinguisane-sacculatane chemotype, characterised by high amounts of pinguisane-type sesquiterpenoids alongside sacculatane-type diterpenoids. It is known for the presence of monoterpenoids, with α-terpinene as the most abundant component. [9]
Chemical composition shows strong correlation with molecular phylogenetic data in the genus Porella. The chemical profile of P. platyphylla places it in a clade with P. navicularis, both sharing the pinguisane-sacculatane chemotype. The absence of pungent drimane compounds distinguishes this clade from the P. vernicosa complex, which forms a distinct section within the genus characterised by pungent taste and glossy appearance. [9] The chemical composition of Porella species, particularly their terpenoid profiles, has proven valuable for taxonomic investigations, helping to resolve uncertainties in this morphologically challenging genus through correlation with molecular data. [9] [8]
Porella platyphylla is widely distributed in forest ecosystems that receive at least 600 millimetres of annual precipitation. It is particularly abundant in northern and central Europe and along the Atlantic coast and islands, though it can also be found scattered throughout Mediterranean regions. [5] The liverwort has a Holarctic distribution pattern, occurring across Eurasia and North America. The species shows distinct geographical structuring in its genetic makeup, with European and North American populations forming separate sister clades. While the species is capable of long-distance dispersal—as evidenced by its wide distribution—molecular evidence suggests that successful establishment and genetic mixing between distant populations remains relatively rare. [4] Whilst the species is widespread in Europe, the typical European form (P. platyphylla sensu stricto) has been confirmed from only a single locality in New Mexico in North America. [4]
The species typically grows on tree bark and rocks, though habitat preferences may vary between different genetic lineages. The North American populations formerly known as P. platyphylloidea were historically noted to occur more frequently on rock substrates, whilst European populations were more commonly found on bark, though this distinction has proven unreliable for taxonomic purposes. [3] In the southern part of the Russian Far East, it primarily occurs on limestone and other basic rocks in partly shaded conditions. However, in northern regions such as the Commander Islands (around 55°N), it can be found on temporarily inundated stream beds, showing habitat preferences more similar to those seen in northern Europe. [10]
Within Russia, most occurrences lie between 43° and 45°N, though the species penetrates northward to several locations including the Bolshoy Anyuy (49°N), Komsomolsk Nature Reserve (51°N), and the Dzhagdy Range (~54°N). Its presence in the Commander Islands (at almost 55°N) appears to be relictual, particularly notable as the species is absent from the intervening Kamchatka Peninsula. [10]
Botany, also called plant science, plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek word botanē (βοτάνη) meaning "pasture", "herbs" "grass", or "fodder"; Botanē is in turn derived from boskein, "to feed" or "to graze". Traditionally, botany has also included the study of fungi and algae by mycologists and phycologists respectively, with the study of these three groups of organisms remaining within the sphere of interest of the International Botanical Congress. Nowadays, botanists study approximately 410,000 species of land plants, including some 391,000 species of vascular plants and approximately 20,000 bryophytes.
Alternation of generations is the predominant type of life cycle in plants and algae. In plants both phases are multicellular: the haploid sexual phase – the gametophyte – alternates with a diploid asexual phase – the sporophyte.
Bryophytes are a group of land plants (embryophytes), sometimes treated as a taxonomic division, that contains three groups of non-vascular land plants: the liverworts, hornworts, and mosses. In the strict sense, the division Bryophyta consists of the mosses only. Bryophytes are characteristically limited in size and prefer moist habitats although some species 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.
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 genetic information.
A sporophyte is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase.
Hornworts are a group of non-vascular Embryophytes constituting the division Anthocerotophyta. The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, hornworts have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of genetic information; the flattened, green plant body of a hornwort is the gametophyte stage of the plant.
Plant reproductive morphology is the study of the physical form and structure of those parts of plants directly or indirectly concerned with sexual reproduction.
Dioecy is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly or indirectly. Dioecious reproduction is biparental reproduction. Dioecy has costs, since only the female part of the population directly produces offspring. It is one method for excluding self-fertilization and promoting allogamy (outcrossing), and thus tends to reduce the expression of recessive deleterious mutations present in a population. Plants have several other methods of preventing self-fertilization including, for example, dichogamy, herkogamy, and self-incompatibility.
Marchantia quadrata is a species of liverwort, a simple non-flowering plant that grows as a flat, green, leaf-like structure (thallus) typically found on damp rocks and soil along stream banks in the Northern Hemisphere. The species was originally classified in its own genus Preissia due to its distinctive features, including larger spores and lack of the specialised cup-like reproductive structures common in other liverworts, but genetic studies have shown it belongs within the genus Marchantia. Like most liverworts, it reproduces both sexually, through separate male and female plants that produce umbrella-like reproductive structures, and through regeneration from fragments. The species prefers slightly drier habitats than its relatives and shows significant genetic variation across its range, suggesting it may comprise several distinct but closely related species.
Conocephalum is a genus of complex thalloid liverworts in the order Marchantiales and is the only extant genus in the family Conocephalaceae. Some species of Conocephalum are assigned to the Conocephalum conicum complex, which includes several cryptic species. Conocephalum species are large liverworts with distinct patterns on the upper thallus, giving the appearance of snakeskin. The species Conocephalum conicum is named for its cone-shaped reproductive structures, called archegoniophores. Common names include snakeskin liverwort, great scented liverwort and cat-tongue liverwort.
Porella is a large, common, and widespread genus of liverworts in order Porellales. It is a member of the family Porellaceae within that order.
Fulfordianthus evansii is a species of liverwort in the family Lejeuneaceae. It is found in Belize, Costa Rica, Guatemala, and Panama. Its natural habitat is subtropical or tropical moist lowland forests.
This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary of leaf morphology. For other related terms, see Glossary of phytopathology, Glossary of lichen terms, and List of Latin and Greek words commonly used in systematic names.
Ptilidium is a genus of liverwort, and is the only genus in family Ptilidiaceae. It includes only three species: Ptilidium californicum, Ptilidium ciliare, and Ptilidium pulcherrimum. The genus is distributed throughout the arctic and subarctic, with disjunct populations in New Zealand and Tierra del Fuego. Molecular analysis suggests that the genus has few close relatives and diverged from other leafy liverworts early in their evolution.
Lejeunea hodgsoniana is a species of liverwort in the family Lejeuneaceae. Endemic to New Zealand, it was first recognized in 1980 but not formally described until 2013. The plant forms bright green mats up to 7 centimetres in diameter on tree bark and occasionally on rocks. The species is found from the Kermadec Islands in the north to the Chatham Islands in the south, primarily in coastal and lowland areas below 100 metres elevation. It is distinguished from related species by its relatively large size, multi-celled tooth on the leaf lobule, and deeply divided underleaves with pointed tips. While showing a particular affinity for mahoe trees, it grows on various native and introduced trees and is considered "Not Threatened" under the New Zealand Threat Classification System due to its abundance within its range and ability to grow in both pristine and disturbed habitats.
Radula demissa is a species of liverwort in the family Radulaceae. It occurs in southeastern Australia and New Zealand, where it grows as an epiphyte in temperate rainforest environments.
Buxbaumia viridis, also known as the green shield-moss, is a rare bryophyte found sporadically throughout the northern hemisphere. The gametophyte of this moss is not macroscopically visible; the large, distinct sporophyte of B. viridis is the only identifying structure of this moss. This moss can be found singularly or in small groups on decaying wood, mostly in humid, sub-alpine to alpine Picea abies, Abies alba, or mixed tree forests. This moss is rare and conservation efforts are being made in most countries B. viridis is found in.
Trichocolea tomentella is a species of liverwort 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 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 and coniferous forests. While capable of sexual reproduction, 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 bryophytes in suitable habitats, T. tomentella faces threats from habitat destruction, particularly through logging and drainage of its preferred moist forest habitats.
Nowellia curvifolia is a species of liverwort belonging to the family Cephaloziaceae. It is a small, delicate plant that forms prostrate growths on rotting logs in forests throughout the northern temperate and subarctic regions. The species is distinguished by its distinctive billowed leaves arranged in two rows, each divided into two pointed lobes with the lower lobe forming a specialised water-holding pocket called a water-sac. The plant shows seasonal colour variation from rose to purple to brown to spring green, developing reddish-purple pigmentation in autumn and winter. It reproduces both sexually, through spores produced in reddish-brown capsules, and asexually via single-celled gemmae. N. curvifolia serves as an indicator species in forest ecosystems, particularly in montane spruce-fir forests, where it often forms pure mats on decorticated logs and is associated with the slime mould species Barbeyella minutissima.
Polytrichastrum formosum, commonly known as the bank haircap moss, is a species of moss belonging to the family Polytrichaceae.