Conocephalum

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Conocephalum
Conocephalum conicum (f, 145046-474908) 3102.JPG
Conocephalum conicum
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Division: Marchantiophyta
Class: Marchantiopsida
Order: Marchantiales
Family: Conocephalaceae
Müll.Frib. ex Grolle
Genus: Conocephalum
Hill, 1773, corr. Wiggers 1780, >nom. et orth. cons.

Conocephalum is a genus of complex thalloid liverworts in the order Marchantiales and is the only extant genus in the family Conocephalaceae. [1] [2] Some species of Conocephalum are assigned to the Conocephalum conicum complex, which includes several cryptic species. [1] Conocephalum species are large liverworts with distinct patterns on the upper thallus, giving the appearance of snakeskin. [3] The species Conocephalum conicum is named for its cone-shaped reproductive structures, called archegoniophores. [4] Common names include snakeskin liverwort, great scented liverwort [5] [6] and cat-tongue liverwort. [7]

Contents

Species of Conocephalum are relatively common [5] and widely distributed throughout North America, Europe and East Asia. [8] Conocephalum often occurs in moist and shaded habitats [1] [5] [9] [10] and are also found in open woodlands, sandy banks, wet rocks and cliffs and moist soils. [4] Species of Conocephalum are also often associated with calcareous substrates. [9]

Conocephalum has a relatively large thallus [4] [5] with irregular branching. [4] Plants grow by overlapping lobes, often creating large mats. [5] Regarding reproduction, species of Conocephalum are dioicous. [11] Species of Conocephalum produce different terpenes and aromatic compounds. [12] [13] Considerable variation in species have been identified based on chemical composition [14] and different species have been identified based on their unique compounds. A unique sesquiterpene alcohol known as conocephalenol was identified and extracted form C. conicum. [12] [13]

Classification and taxonomy

Some species of Conocephalum are placed in the Conocephalum conicum complex, which includes several cryptic species. [1] Consequently, it has been challenging to identify the exact number of species in this genus. [15] Cryptic species refers to a species which demonstrates a genetic difference but lacks morphological differences. Within liverworts, cryptic species are suggested to be related to both geographical disjunction and to reproductive biology in combination with isolation and habitat differentiation. [14]

Molecular research has indicated that Conocephalum comprises a complex of six cryptic species (A, C, F, J, L and S). [14] [16] In 2005 C. conicum cryptic species S was described as a separate species, C. salebrosum. [1] [9] [14] Conocephalum salebrosum has a wider distribution and is present in North America, in contrast to C. conicum. [1] [3] [9] More recent examinations of the Conocephalum conicum complex in Japan and Taiwan have identified three new species within Conocephalum, C. orientalis, C. purpureorubum and C. toyotae, which were formerly described as C. conicum J, F, and R respectively. [2]

Species

Distribution

Conocephalum conicum growing in a moist and shady habitat Conocephalum conicum jagoke04.jpg
Conocephalum conicum growing in a moist and shady habitat

Species of Conocephalum are distributed throughout North America, Europe and East Asia. [8] [11] [15] Conocephalum salebrosum displays the widest distribution and is found throughout North America, [5] [9] [15] Europe and Asia. [17] In North America, C. salebrosum occurs throughout Canada and parts of the United States and has also been reported from Russia. [9] In contrast to C. salebrosum, C. conicum is found throughout Europe [11] [15] and has been recorded in Norway, Finland, Great Britain, Ireland, Belgium, France, Germany, Czech Republic, Austria, Hungary, Romania, Italy, Portugal, Spain, Croatia, Bulgaria, Greece, Ukraine, Poland and Russia. [18]

The species C. supradecompositum is more restricted in its distribution and is mainly found in China and Japan. [19] [20] Regarding the most recently described species of Conocephalum, C. purpureorubum has been observed in Japan, China, Taiwan and South Korea and C. orientalis has been found in Japan and Taiwan. [2] [11]

Habitat

Species of Conocephalum often occur in moist and shaded habitats. [1] [5] [9] [10] Conocephalum species also grow in specialized micro-habitats near both running and standing water. [17] Conocephalum conicum is often found in open woodlands, sandy banks, wet rocks and cliffs and moist soils. [4] Both C. conicum and C. salebrosum are strongly associated with calcareous substrates. [9] It has also been suggested that C. salebrosum is likely more tolerant of desiccation than C. conicum. [9] [10] [11] [17] [21]

Morphology

Conocephalum conicum, the thallus is distinctly shiny in appearance Conocephalum conicum (g, 145025-474547) 6046.JPG
Conocephalum conicum, the thallus is distinctly shiny in appearance
Conocephalum salebrosum, the thallus is dull in appearance Conocephalum salebrosum (a, 145855-475042) 7776.JPG
Conocephalum salebrosum, the thallus is dull in appearance

Conocephalum conicum and C. salebrosum share some similarities in morphological characteristics, in addition to having their own unique traits which help distinguish the two species. [21]

Conocephalum salebrosum, air pore present in the upper surface of the thallus Conocephalum salebrosum (a, 145855-475042) 7806.JPG
Conocephalum salebrosum, air pore present in the upper surface of the thallus

Gametophyte

The vegetative structure of Conocephalum is a thallus which has the appearance of a flattened body of plant tissue. [5] [22] The thallus is irregularly branched [4] and relatively large, reaching lengths of roughly 20-24 cm. [4] [5] In contrast to C. conicum and C. salebrosum, the thallus of C. supradecompositum is relatively small, measuring 2-3 cm long. [20] The thallus grows by developing lobes [22] which wither away as the plant matures. [23] Plants of C. salebrosum often grow by overlapping lobes, sometimes creating large mats. [5]

Species of Conocephalum have a thallus that is either dull in appearance, such as C. salebrosum, or distinctly shiny, such as C. conicum. [1] [11] The upper surface of the thallus has characteristic hexagonal outlines formed by shallow grooves around each air chamber. [4] Photosynthetic tissue and chloroplasts are located within the air chambers. [24] In the middle of each air chamber is a white-ringed pore. [4] [23] The upper walls of large air chambers are often visible on the surface of the thallus. [23] The air chamber pore remains open, in contrast to the stomata of vascular plants where the pores can open and close. [5]

The underside of the thallus has both rhizoids and scales. [4] The scales are purple in colour and are arranged along the middle of the underside of the thallus. [4] Rhizoids are also present on the underside of the thallus. There are two types of rhizoids, both long smooth rhizoids and short pegged rhizoids. [4] [22] [25] The short rhizoids are thought to play a role in absorbing water and nutrients. In contrast, the longer rhizoids help anchor the thallus to the underlying substrate. The rhizoids are single-celled, in contrast to the multicellular rhizoids found in mosses. [22]

Complex oil bodies

Liverworts cells often contain complex oil bodies. [14] [22] [25] [26] The oil bodies are intracellular organelles bounded by a single membrane. [24] [27] The oil bodies have been known to contain a variety of unique phytochemicals, such as terpenes and flavonoids. [26] The function of oil bodies is still poorly understood. [24] [25] [27] It has been suggested that oil bodies might function as a deterrent to herbivory or could protect from cold temperatures or harmful ultraviolet radiation. [24] [25]

Sporophyte

The sporophyte consists of an unbranched stalk called a seta, which bears a terminal spore capsule called a sporangium. [25] The sporangia of Conocephalum are borne beneath stalked gametophytic structures called archegoniophores. [4] In contrast to mosses, the sporophyte matures before the seta elongates. [25] [28] Unlike mosses, liverwort sporophytes lack stomata, a columella and peristome teeth. [24] [25]

Life cycle

General life cycle of Marchantia liverwort Cycle de reproduction de Marchantia polymorpha.jpg
General life cycle of Marchantia liverwort

The life cycles of liverworts involves alternating haploid gametophyte and diploid sporophyte generations. The gametophyte generation is more dominant, while the sporophyte generation is relatively short-lived. The gametophyte produces haploid gametes, egg and sperm, which fuse to form a diploid zygote. The zygote then develops into a sporophyte which ultimately produces haploid spores through meiosis. The sporophyte requires nutrients supplied by the gametophyte to sustain growth and development.

The life cycle of Marchantia liverworts also applies to Conocephalum, with the exception that Conocephalum lacks a stalked antheridiophore and instead has small flat antheridial heads on the surface of the thallus. [25]

Reproduction

Conocephalum conicum, sporophytes (black) hanging beneath the umbrella-shaped, stalked archegoniophores Conocephalum conicum (c, 144707-474823) 1912.jpg
Conocephalum conicum, sporophytes (black) hanging beneath the umbrella-shaped, stalked archegoniophores

Liverworts reproduce through both sexual and asexual reproduction. [5] In natural populations, the high genetic variation observed suggests that sexual reproduction might dominate. Species of Conocephalum are dioicous, [11] meaning that the male and female reproductive structures are produced on separate plants. [29]

Sexual reproduction

In Conocephalum the male and female reproductive parts are embedded in receptacles on separate plants. On male plants, the receptacle is slightly raised, lacking a stalk, and often circular or oval shaped. Antheridia are embedded in the receptacle [5] and at maturity the sperm is released into the air. [5] [25] [30] In contrast, on female plants the receptacles are dome-shaped, with several drooping lobes at the end of an erect stalk. The receptacles are often described as a tiny umbrella, with the archegonia beneath. [5]

Gametophytes produce eggs and sperm in the archegonia and antheridia, respectively. [22] Fertilization occurs when the sperm reach the egg within the archegonia of a female plant. [23] Once fertilization occurs, the ovule within an archegonium develops into a sporophyte. [22] Mature sporangia on the underside of the receptacle resemble black capsules. These capsules split open to release both spores and elaters, [23] which are dispersed mainly by wind. [22] [23] The elaters function to propel spores during dispersal. [28]

Conocephalum elaters are unique and display a wide range of variability in shape, size and number. Often the abundance of elaters within a capsule are 2-3 times more abundant than spores. [31] Elaters form from an initial mother cell which develops into a diploid cell with spiral thickenings. In contrast, spores develop from an initial diploid mother cell that ultimately forms haploid spores by meiosis. [24]

Conocephalum conicum, thallus with scales at the tips covering new spring buds Conocephalum conicum (e, 144739-474742) 1022.jpg
Conocephalum conicum, thallus with scales at the tips covering new spring buds

Asexual reproduction

The production of gemmae is a common method of asexual reproduction in liverworts. [5] [28] [32] Gemmae are small packets of tissue consisting of haploid cells that are genetically identical with those of the parent plant. They are dispersed by rainfall and ultimately grow into new individuals. [32] In C. conicum, gemmae are located on the lower layers of the thallus and are released as the thallus degrades. [4] In contrast, C. salebrosum does not produce gemmae.

Vegetative reproduction can occur when a piece of the thallus breaks off and is transported away from the parent plant. [5] The individuals resulting from vegetative reproduction are genetically identical to the parent plant and therefore clonal colonies often exist as either all male or all female. [28]

Conocephalum species are perennial, meaning that they can overwinter and produce new growth in the spring. These new buds are covered and protected by small scales. [11]

Biochemistry

Chemical structure of Cubebol Cubebol.png
Chemical structure of Cubebol

Many liverworts produce different terpenes and aromatic compounds. [12] [13] Terpenoids and aromatic compounds are often accumulated within the oil bodies of many liverworts, including Conocephalum. Within Conocephalum, considerable variation in species have been identified based on chemical composition and these compounds been used to identify different cryptic species. [14]

Three different groups of Conocephalum were identified baed on their unique primary volatile compounds. For example, the compound cubebol, a sesquiterpene alcohol, is characteristic of C. salebrosum. [14] It has also been noted that C. supradecompositum has a distinct chemical composition compared to C. conicum as well, mainly that the Monoterpenoid content in C. supradecompositum is much less than observed in C. conicum. [20]

A unique sesquiterpene alcohol known as conocephalenol was identified and extracted from C. conicum. [12] [13] Conocephalenol has a unique chemical skeleton that is characteristic of a sesquiterpenes present in red algae. [33]

Associations with other species

Fungal interactions

Conocephalum can form associations with fungi that are similar to the mycorrhizal associations observed vascular plants. Molecular analyses demonstrated that Conocephalum contained fungal endophytes from the group of fungi known as the Glomeromycota. [34]

Conocephalum conicum often colonizes bare soils or rocky substrates, where mineral nutrients can often be limiting. The fungal endophyte establishes a complex relationship with C. conicum, which is characterized by the formation of arbuscules. These fungi form a highly branched mycelium outside of the plant which then colonize the outside of the rhizoids and pass into the gametophyte. The fungal infection induces grown of fungal hypha within the host cells of C. conicum. This association of the fungal hypha with the hosts plastids suggests that photosynthates produced through photosynthesis in C. conicum are likely transferred to the fungus. A similar situation regarding this fungal association has also been identified in the thalloid liverwort Pellia epiphylla . Although these associations are common in vascular plants, they have rarely been described in non-vascular plants. [35]

Animal interactions

Herbivory

Approximately 25 species of moths which are endemic to East Asia associate exclusively with Conocephalum. [11] The larval stage of Epimartyria pardella moths feed on C. conicum. [36] [37] In addition, the fungal species Loreleia marchantiae also feeds on C. conicum. [11]

Pathogens

The fungal pathogen belonging to the genus Pythium has often been isolated from infected rhizoids and thallus of Conocephalum. [11] Bryoscyphus conocephali is another fungal pathogen that has been associated with C. conicum. [38]

Human applications

Conocephalum as a bioindicator for pollution

Conocephalum conicum has been identified as being tolerant of heavy metals and has therefore been suggested to have a possible role as a bioindicator for pollution. Conocephalum conicum takes up ions from both the soil and the atmosphere. Therefore, heavy metals contamination of C. conicum is related not only to air pollution, but environmental contamination from different sources. [39] Recent research has also examined C. conicum as a bioindicator for cadmium pollution. Cadmium a toxic metal and considered the third highest contaminant, after mercury and lead. Conocephalum conicum was shown to respond to cadmium stress by changing its biological activity. These biological changes could be used as biomarkers for cadmium pollution. [40]

Chemical structure of aflatoxin B1 found in Aspergillus fungi (-)-Aflatoxin B1 Structural Formulae V.1.svg
Chemical structure of aflatoxin B1 found in Aspergillus fungi

Anti-fungal activity

Conocephalum conicum has been suggested to have a possible role in the management of food borne disease caused by species of Aspergillus fungi. Aspergillus produces highly potent toxins, carcinogens, referred to as aflatoxins. Aflatoxins are harmful both plants and animals. Aspergillus can cause disease in many important crops, which can ultimately cause disease in humans. Conocephalum has been shown to have a variety of bioactive compounds which promote anti-fungal property against Aspergillus. [26]

Ethnomedicine

Throughout North America, China and India, liverworts such as Conocephalum have been used for ethnomedical purposes. Conocephalum is known to be important to Bhotia, Raji, Tharus and Boxas tribes in Pithoragarh district of Kumaon Himalaya. Conocephalum conicum is used to treat burns, and the extract of C. conicum is also used in treating gallstones. Conocephalum has also demonstrated antidote activity against venomous snake bites. The role of Conocephalum regarding modern medicine has yet to be investigated. [26]

Cosmetic industry

Liverworts often contain highly pungent compounds. [41] Conocephalum species often exude an odour that is characteristic of turpentine. The odour is thought to be related to the presence of monoterpenoids. The compound conocephalenol is widely used in the cosmetic industry for its odourant properties. [33]

Related Research Articles

<span class="mw-page-title-main">Bryophyte</span> Terrestrial plants that lack vascular tissue

Bryophytes are a group of land plants, sometimes treated as a taxonomic division, that contains three groups of non-vascular land plants (embryophytes): 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. 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'.

<span class="mw-page-title-main">Marchantiophyta</span> Botanical division of non-vascular land plants

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.

<span class="mw-page-title-main">Hornwort</span> Division of non-vascular land plants with horn-shaped sporophytes

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.

<span class="mw-page-title-main">Marchantiales</span> Order of non-vascular plants known as liverworts

Marchantiales is an order of thallose liverworts that includes species like Marchantia polymorpha, a widespread plant often found beside rivers, and Lunularia cruciata, a common and often troublesome weed in moist, temperate gardens and greenhouses.

<i>Marchantia</i> Genus of plants in the liverwort family Marchantiaceae

Marchantia is a genus of liverworts in the family Marchantiaceae and the order Marchantiales.

<i>Lunularia</i> Species of liverwort

Lunularia is a genus of liverworts whose only species is Lunularia cruciata, the crescent-cup liverwort. Lunularia is either the only genus in the order Lunulariales, or may be placed in the order Marchantiales. The name, from Latin luna, moon, refers to the moon-shaped gemma cups.

<i>Riccia</i> Genus of liverworts

Riccia is a genus of liverworts in the order Marchantiales.

<i>Anthoceros</i> Genus of hornworts

Anthoceros is a genus of hornworts in the family Anthocerotaceae. It is distributed globally. Species of Anthoceros are characterized by having a small to medium-sized, green thallus that is more or less lobed along the margins.

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.

<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.

<span class="mw-page-title-main">Prothallus</span> Gametophyte stage in the fern life cycle

A prothallus, or prothallium, is usually the gametophyte stage in the life of a fern or other pteridophyte. Occasionally the term is also used to describe the young gametophyte of a liverwort or peat moss as well. In lichens it refers to the region of the thallus that is free of algae.

<i>Cavicularia</i> Genus of liverworts

Cavicularia densa is the only species in the liverwort genus Cavicularia. The species was first described in 1897 by Franz Stephani, and is endemic to Japan, where it grows on fine moist soil.

<i>Pellia epiphylla</i> Species of liverworts in the family Pelliaceae

Pellia epiphylla is a species of thallose liverwort. It occurs in North America, Europe, North Africa and parts of Asia. It grows in patches in damp, sheltered places on neutral or acidic substrates. It is common on the banks of rivers, streams and ditches and also grows in wet woodland, marshes and on wet rocks.

<i>Conocephalum conicum</i> Species of liverwort

Conocephalum conicum, also known as the great scented liverwort or snakeskin liverwort, is a liverwort species in the genus Conocephalum. C. conicum is part of the Conocephalum conicum complex, which includes several cryptic species. The name C. conicum refers to the cone-shaped archegoniophore, which bear sporangia.

<i>Marchantia berteroana</i> Species of liverwort

Marchantia berteroana is a liverwort species in the genus Marchantia.

<i>Dawsonia superba</i> Species of moss

Dawsonia superba is a moss in the family Polytrichaceae that is found in Australia, New Guinea, Malaysia and New Zealand. D. superba is the tallest self-supporting moss in the world, reaching heights of 60 cm (24 in). It has analogous structures to those in vascular plants that support large size, including hydroid and leptoid cells to conduct water and photosynthate, and lamellae that provide gas chambers for more efficient photosynthesis. D. superba is a member of the class Polytrichopsida, although it has a sporophyte that is unique from other hair-cap mosses.

<i>Asterella californica</i> Species of plant

Asterella californica is a complex thallic liverwort in the phylum Marchantiophyta. A. californica often grows as colonies of flat rosettes of light green, rigid thalli, with undersides dark wine-red to nearly black. The receptacles are rounded, with four lobes each bearing a single sporangium sheathed by a white tattered skirt. A. californica is dioecious with separate male plants often intermingled with female plants. This species is found throughout California. See Distribution information below. Asterella californica is the commonest species of the three species of Asterella occurring in California; the other two species are A. bolanderi and A. palmeri.

<i>Conocephalum salebrosum</i> Species of liverwort

Conocephalum salebrosum, commonly known as snakewort, is a species of liverwort, a non-vascular land plant, with a broad, holarctic distribution. It is also known as snakeskin liverwort, cat-tongue liverwort, mushroom-headed liverwort, and great scented liverwort.

Conocephalum supradecompositum is a species of thalloid liverwort in the genus Conocephalum, of the order Marchantiales and the family Conocephalaceae. C. supradecompositum has a distribution that is mainly restricted to China and Japan. C. supradecompositum has very distinct chemical composition from the species Conocephalumconicum.

<i>Monoclea forsteri</i> Species of liverwort

Monoclea forsteri is one of the two species in the thallose liverwort family Monocleaceae. It is dioicous with the capsule dehiscing with a single longitudinal slit. Endemic and widely distributed throughout New Zealand, it is also the country's largest thalloid liverwort. Hooker described the species in 1820. The holotype is in the British Museum.

References

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