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Sporophytes of Marchantia. (A) Sporophyte (hanging downward) with foot (F) embedded in the archegoniophore (ARP). The seta (S) supports the capsule (C) containing spores. (A) Enlargement of the capsule showing thread-like elaters (arrows). Images were obtained from slides and used with permission from Carolina Biological Supply, Whitsett, North Carolina, Copyright Carolina Biological Supply. 

Sporophytes of Marchantia. (A) Sporophyte (hanging downward) with foot (F) embedded in the archegoniophore (ARP). The seta (S) supports the capsule (C) containing spores. (A) Enlargement of the capsule showing thread-like elaters (arrows). Images were obtained from slides and used with permission from Carolina Biological Supply, Whitsett, North Carolina, Copyright Carolina Biological Supply. 

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Context 1
... are 5000 to 7500 species of liverworts world- wide (S€ oderstr€ om et al., 2016;von Konrat et al., 2010) making this the second largest phylum of bryophytes. Morphologically the liverwort gametophytes are diverse with both simple and complex thalloid species that are flattened dorsoventrally ( Figure 6) and leafy liverworts that have a three-ranked caulid and phyllid arrangement, which are somewhat similar to mosses. Marchantia, belongs to the complex, thalloid liver- worts, which include about 5% of all liverworts and has often been used to represent the morphology of liverworts. In Marchantia photosynthesis only occurs in a defined cell layer on the dorsal or upper surface bounded by epidermal cells with pores for gas exchange (Figure 7). Rhizoids and scales are located on the lower surface and serve to absorb water and anchor the plant to the substrate. The cells between the upper and lower epidermis are mostly thin-walled and used for storage of oils and other products. In Marchantia, male and female gametangia are pro- duced on umbrella-like antheridiophores and archego- niophores (Figure 8) on separate plants. The archegonia are inverted and hang downward from the bottom of the structure (Figure 9A), whereas the male gametangia are located on the upper surface of the gametophores ( Figure 9B). Sperm dispersal is similar to mosses in that they are facilitated by water, though no tests have been made to determine whether ani- mals may contribute to their dispersal. Marchantia sporophytes ( Figures 10A,B) are composed of a foot embedded in the female gametophyte, setae, and a capsule and as in the mosses the sporophyte is dependent on the gametophyte for water and nutri- tion. The capsules of some, but not all, liverworts con- tain elaters (Figures 10B), which are diploid structures intermixed with the haploid spores. Elaters are long, narrow cells with secondary wall thickenings thought to have two functions. First, they may serve as nutri- tion for the developing spores and second, may be involved in spore dispersal in response to changes in moisture and humidity (Schuster, 1966;Kremer and Drinnan, ...
Context 2
... are 5000 to 7500 species of liverworts world- wide (S€ oderstr€ om et al., 2016;von Konrat et al., 2010) making this the second largest phylum of bryophytes. Morphologically the liverwort gametophytes are diverse with both simple and complex thalloid species that are flattened dorsoventrally ( Figure 6) and leafy liverworts that have a three-ranked caulid and phyllid arrangement, which are somewhat similar to mosses. Marchantia, belongs to the complex, thalloid liver- worts, which include about 5% of all liverworts and has often been used to represent the morphology of liverworts. In Marchantia photosynthesis only occurs in a defined cell layer on the dorsal or upper surface bounded by epidermal cells with pores for gas exchange (Figure 7). Rhizoids and scales are located on the lower surface and serve to absorb water and anchor the plant to the substrate. The cells between the upper and lower epidermis are mostly thin-walled and used for storage of oils and other products. In Marchantia, male and female gametangia are pro- duced on umbrella-like antheridiophores and archego- niophores (Figure 8) on separate plants. The archegonia are inverted and hang downward from the bottom of the structure (Figure 9A), whereas the male gametangia are located on the upper surface of the gametophores ( Figure 9B). Sperm dispersal is similar to mosses in that they are facilitated by water, though no tests have been made to determine whether ani- mals may contribute to their dispersal. Marchantia sporophytes ( Figures 10A,B) are composed of a foot embedded in the female gametophyte, setae, and a capsule and as in the mosses the sporophyte is dependent on the gametophyte for water and nutri- tion. The capsules of some, but not all, liverworts con- tain elaters (Figures 10B), which are diploid structures intermixed with the haploid spores. Elaters are long, narrow cells with secondary wall thickenings thought to have two functions. First, they may serve as nutri- tion for the developing spores and second, may be involved in spore dispersal in response to changes in moisture and humidity (Schuster, 1966;Kremer and Drinnan, ...
Context 3
... dispersal is similar to mosses in that they are facilitated by water, though no tests have been made to determine whether ani- mals may contribute to their dispersal. Marchantia sporophytes ( Figures 10A,B) are composed of a foot embedded in the female gametophyte, setae, and a capsule and as in the mosses the sporophyte is dependent on the gametophyte for water and nutri- tion. The capsules of some, but not all, liverworts con- tain elaters (Figures 10B), which are diploid structures intermixed with the haploid spores. ...

Citations

... Among the bryophytes, mosses are the most diverse in terms of species richness (Crandall-Stotler & Bartholomew-Began, 2007), with about 12,000 (Vanderpoorten & Goffinet, 2009) and 14,000 (Asakawa & Ludwiczuk, 2017;Das et al., 2022) species worldwide. Moreover, bryophytes are ubiquitous plants (Hodgetts et al., 2019) that can occupy a variety of habitats (Crandall-Stotler & Bartholomew-Began, 2007;Budke et al., 2018;Taha, 2020;Dziwak et al., 2022) compared to vascular plants (Sabovljević et al., 2022). Due to their structure, bryophytes are very sensitive to environmental conditions (Sabovljević et al., 2022), which makes them good indicators for environmental changes (Hallingback & Hodgetts, 2000;Mishler, 2001;Das et al., 2022), especially the epiphytic bryophytes which are predictably sensitive to global warming because of their direct connection to atmospherical conditions (Gradstein, 2008). ...
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The bryoflora of the Setifian High Plains, which situated in the Northeastern region of Algeria is poorly known. Megriss Mountain is one of the most important ecosystems in this region. The examination of Orthotrichaceae samples collected during bryophytes diversity investigations in the northern part of Megriss resulted in the identification of the epiphytic species; Orthotrichum macrocephalum F. Lara, Garilleti & Mazimpaka. The identification was made based on its leaves with rounded or obtuse apices, recurved margins, pluripapillose laminal cells, the presence of propagules, emergent capsule with eight longitudinal orange ribs and star-shaped mouth when dry, scarcely constricted below the mouth, immersed stomata in the capsule wall and the papillose endostome segments. This acrocarpous moss with a marked affinity for the Mediterranean climate has never been recorded in the Algerian bryoflora. A comprehensive description, microphotographs, and distribution map in Algeria of this species are provided. Moreover, its ecology is also discussed.
... Bryophytes are small plants that are essential elements of the vegetation in a diverse range of habitats, including moist forests, wetlands, mountain, and tundra ecosystems (von Konrat et al., 2010), growing on soils, rocks, and trees as epiphytes. Although they are most abundant in relatively moist areas (Schofield, 1985;Gradstein et al., 2001;Taylor et al., 2009), most species can survive periods of desiccation (Budke et al., 2018). They are indicators of environmental health due to their sensitivity to air/rainwater pollution (Pescott et al., 2015;Gatziolis et al., 2016). ...
Article
We describe fossil liverworts, mosses, and thallose forms of unknown affinity from the Barreal and Cortaderita formations (Sorocayense Group, Middle Triassic) in the Barreal depocenter, southwestern San Juan Province. The specimens are impressions-compressions of gametophytes and sporophytes, preserved in claystones, siltstones, and tuffaceous and bentonitic silty sandstones. The thallose forms consist of dichotomous simple thalli lacking evidence of a midrib and are assigned to the genus Thallites. Within the liverworts, we identified a new genus and two new species. Ricciellites unsaltoensis gen. et sp. nov. is characterized by thalli forming partial rosettes with distinct midrib, polygonal cells, air pores surrounded by a ring of cells, enclosed sporophytes, and ventral scales arranged in two rows. Ricciopsis cortaderitaensis sp. nov. has thalli forming rosettes, with distinct midrib, polygonal cells, and rhizoids. We also describe two types of acrocarpous mosses assignable to the genus Muscites. They are characterized by delicate unbranched axes with lanceolate unicostate leaves. Muscites sp. 1 is represented by several specimens arranged in a loose cushion, while Muscites sp. 2 corresponds to isolated specimens. Together, these fossils represent the first unequivocal evidence of mosses and liverworts for the Middle Triassic of Argentina. Given the scarcity of bryophyte megafossils of Triassic age worldwide (31 worldwide, three confirmed records for Argentina), these findings add significantly to the bryophyte fossil record.
... was established, bryological research is now embracing a golden era, propelled by new technologies for data management, molecular biology, genomics and ecological modelling. Such advances have been reflected by recent special issues addressing the state of the art of different fields of bryological research (Budke et al. 2018;Stech et al. 2021). However, key questions have only started to be largely resolved, most notably those concerning phylogenetic relationships among the three main bryophyte lineages and with respect to the vascular plants de Sousa et al. 2019;Harris et al. 2020;Su et al. 2021). ...
... Indeed, along with other important recent initiatives (e.g. Renzaglia et al. 2007;Budke et al. 2018;Câmara et al. 2021;Stech et al. 2021), we have sought to contribute and advance the bryophyte research agenda. Despite our large and ambitious list of research foci, much of hypothesis-driven and wellexecuted research discussed here has the full potential to inspire theoretical and empirical research in the near future. ...
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Introduction Half a century since the creation of the International Association of Bryologists, we carried out a review to identify outstanding challenges and future perspectives in bryology. Specifically, we have identified 50 fundamental questions that are critical in advancing the discipline. Methods We have adapted a deep-rooted methodology of horizon scanning to identify key research foci. An initial pool of 258 questions was prepared by a multidisciplinary and international working group of 32 bryologists. A series of online surveys completed by a broader community of researchers in bryology, followed by quality-control steps implemented by the working group, were used to create a list of top-priority questions. This final list was restricted to 50 questions with a broad conceptual scope and answerable through realistic research approaches. Key results The top list of 50 fundamental questions was organised into four general topics: Bryophyte Biodiversity and Biogeography; Bryophyte Ecology, Physiology and Reproductive Biology; Bryophyte Conservation and Management; and Bryophyte Evolution and Systematics. These topics included 9, 19, 14 and 8 questions, respectively. Conclusions Although many of the research challenges identified are not newly conceived, our horizon-scanning exercise has established a significant foundation for future bryological research. We suggest analytical and conceptual strategies and novel developments for potential use in advancing the research agenda for bryology.
... One important difference between bryophytes and most other plants is that bryophytes are poikilohydric, i.e., their cell water content varies with ambient water supply. They have to cope with drought and recover from drought-induced damages (Oliver et al. 2005;Budke et al. 2018). Our data suggested that bryophyte species living in wet habitats with reduced need to cope with desiccation had higher mutation rate. ...
Article
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Plants commonly referred to as “bryophytes” belong to three major lineages of non-vascular plants: the liverworts, the hornworts and the mosses. They are unique among land plants in having a dominant haploid generation and a short-lived diploid sporophytic generation. The dynamics of selection acting on a haploid genome differs from those acting on a diploid genome: new mutations are directly exposed to selection. The general aim of this paper is to investigate the diversification rate of bryophytes - measured as silent site substitution rate representing neutral evolution (mutation rate) and the nonsynonymous to synonymous substitution rate ratio (dN/dS) representing selective evolution - and compare it with earlier studies on vascular plants. Results show that the silent site substitution rate is lower for liverworts as compared to angiosperms, but not as low as for gymnosperms. The selection pressure, measured as dN/dS, is not remarkably lower for bryophytes as compared to other diploid dominant plants as would be expected by the masking hypothesis, indicating that other factors are more important than ploidy.
... The bryophytes consisting of more than 20,000 species are widespread covering every part of the world with exception of sea 9,10 . However, despite their wide coverage, the mosses have been unable to maintain their rightful place as a preferred research object and therefore most plant mor-phogenesis studies have been carried out in vascular plants 11 . The most widely used application of bryophytes includes its use for biomonitoring/bioindicator of waters and air pollution in addition to its use for determination of heavy metal accumulation for 12,13 . ...
Article
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The present study aimed to evaluate the chemical analysis and antimicrobial effects of the Thamnobryum alopecurum (Hedw.) Gangulee essential oils were obtained by different distillation methods (Clevenger apparatus hydrodistillation (HD) and microwave-assisted hydrodistillation (MD)). In this study obtained volatile oils were analyzed by gas chromatography-mass spectrometry which revealed the presence of forty-five compounds present in the essential oil obtained by HD method representing 99.48 % of the total identified compounds and thirty-four compounds were identified in the essential oil extracted by MD method representing 95.39 % of the total identified compounds. 3-octanone was the major constituent of the essential oils extracted by both HD (14.28 %) and MD (20.07 %). Terpenes were the major constituents in the essential oil extra-cted by both methods (54.41 % in HD and 52.21 % in MD) followed by oxygenated terpenoids (17.16 % in HD and 14.94 % in MD). The other major compounds of the essential oil in HD was limonene (9.65 %) followed by β-pinene (9.13 %), cuparene (6.04 %), gymnomitrone (5.72 %) and 4-epi-abietal (5.59 %). While in MD essential oil the other major compounds were β-pinene (12.20 %) followed by limonene (8.56 %), α-pinene (9.01 %), cupa-rene (6.44 %), abietadiene (5.97 %). The antimicrobial effects of the essential oils were determined tested quantitatively in respective broth media by using double microdilution and the minimal inhibition concentration (MIC) values (μg/mL). The isolated essential oils of T. alopecurum showed effective antimicrobial activities against Escherichia coli, Yersinia pseudotuberculosis, Staphylococcus aureus, Enterococcus faecalis, Bacillus cereus, Mycobacterium smegmatis and Candida albicans. However, no activity was not observed against Pseudomonas aeruginosa. The hydrodistillation method was found to be more effective than the microwave method in iden-tifying the chemical composition of the structure of T. alopecurum essential oil. While for antimicrobial study MD method was found to be more effective in terms of antimicrobial effect. The T. alopecurum thus is a potential source of an antimicrobial agent with a wide range of chemical compounds that could be responsible for their activity.
... This is the second largest phylum of bryophytes as there are 5000 to 7500 species of liverwort all over the world (Soderstrom et al., 2016;Von Konrat et al., 2010). Common liverwort belongs to the thalloid complex of liverworts, which includes about 5% of all liverworts, and has often been used to represent the model morphology of liverworts (Budke et al., 2018). The thickness of the thallus is 0.3 to 0.6 mm at the midrib region and gradually becomes thinner toward the margin (Shimamura, 2015). ...
... The thallus is dorsiventral with a broad laminar surface for maximum interception of light (Raven et al., 1999). In common liverwort, photosynthesis occurs in a defined cell layer on the dorsal surface (Budke et al., 2018). On the lower surface there are rhizoids and scales, which absorb moisture and anchor the plant body to the substrate (Budke et al., 2018). ...
... In common liverwort, photosynthesis occurs in a defined cell layer on the dorsal surface (Budke et al., 2018). On the lower surface there are rhizoids and scales, which absorb moisture and anchor the plant body to the substrate (Budke et al., 2018). The rhizoids are in localized areas over the whole ventral surface of the thallus (McConaha, 1941), and they also grow down the grooves in the gametophore stalks (Bell, 1992). ...
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Common liverwort ( Marchantia polymorpha ) is a primitive, spore-bearing bryophyte that thrives in containerized ornamental crop propagation and production environments. It is one of the major weed problems in container nurseries and greenhouses because it competes with ornamental plants for soil/growing medium, nutrients, water, space, and oxygen within the container. As a result, its presence can reduce the overall quality and market value of the ornamental crop. Once established in nurseries and greenhouses, it spreads rapidly because of its ability to propagate both asexually and sexually. Currently, no effective methods of controlling common liverwort in container production systems are available because a significant knowledge gap exists. Therefore, research is needed to determine whether organic mulches (types, depths, moisture holding capacity, and particle size), biopesticides, and strategic placement of fertilizers within containers suppress or inhibit common liverwort growth and development. In addition, newer chemicals (both synthetic and organic) and combinations need to be tested on different growth stages of common liverwort. The objective of this review was to summarize previous and current research related to common liverwort control in container production, and to identify areas where additional research is needed either to improve current control methods or to develop new ones.
... Hornwort consists of about 200 species in the world (Budke et al., 2018); (Cole and Hilger, 2016). Bryophyte of this class is similar to some liverwort, differ from only in thalloid gametophyte in the form of discs and chiseled edges. ...
Article
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Bryophyte is a division of plants that lives on land, generally it is green and reproduces through spores, has ecological and economic functions, and plays an important role in forest ecosystems. It spreads out almost in all parts of the earth with different characters in each group. It is divided into 3 groups, namely liverwort, true moss, and hornwort, which are phylogeny and true liverwort is in the same lineage. The number of bryophytes species is around 18000 with the largest distribution area of bryophyte diversity in tropical and subtropical latitudes, such as the Malesia region which includes Malaysia, Indonesia, Timor Leste, Papua New Guinea, the Philippines, and Brunei. Various studies were carried out related to the diversity of bryophyte, especially in the Malesia region, and found various types including new species, new records, and new characters.
... Liverworts (Marchantiophyta) are a relatively small group of plants with an estimated 5-6000 species in the world [21,22], but they are plants that produce a wide array of biologically active secondary metabolites [1]. These compounds are accumulated in the oil bodies, which are a prominent and highly distinctive organelle uniquely found in liverworts [23,24]. ...
... Liverworts are plants in which one can find relatively rare groups of naturally occurring compounds. These are seco-africanes (10)(11)(12), noraristolanes (13,14), 1,10-seco-aromadendranes (15, 16), 2,3-seco-aromadendranes (17,18), seco-cuparanes (19), tridensanes (20), ricciocarpanes (21,22), modified pacifigorgianes (23,24), pinguisanes (25,26), neotrifaranes (27), chenopodanes (28,29), and riccardiphanes (30, 31), among others ( Fig. 3) [1,4]. ...
Article
Bryophytes, phylogenetically placed between algae and pteridophytes, are divided into three classes, mosses, liverworts, and hornworts. Traditional system of medicine throughout the world has been utilizing this group of plants to treat various ailments. One of the outstanding features of these spore forming plants is their chemistry, especially that of the liverworts. Liverworts, and other bryophytes have yielded of rich array of secondary metabolites. Many of these compounds are characterized by unprecedented structures, and some have not been found in any other plants, fungi or marine organisms. Among the bryophytes, the chemical constituents of liverworts and their biological activity have been studied in the most detail. In this review, we demonstrate the chemistry of the liverworts, mosses and hornworts, and also neuroptotective activity of dimeric herbertane-type sesquiterpenoids, mastigophorenes and secoaromadendrane-type sesquiterpenoids, marchantin-type cyclic bisbibenzyls with muscle-relaxant activity, or ent-longipinane-type sesqiterpenoids with acetylcholinesterase inhibitory activity, among others.
... Bryophytes are the one of the most important group of plants consisting of about 24,000 species worldwide with a wide range of distribution. These cosmopolitan species are distributed from the polar region to the tropics (1) . They grow especially in hills and forests throughout the year. ...
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A leafy liverwort, Jungermannia exertifolia Steph. (Hepaticae, Jungermanniales,) has been reported from the Adampur Forest of Moulvibazar district as a new species record for Bangladesh. Jungermannia exertifolia is characterized by its flattened leaves and rhizoids arising ventrally all along the stem with alternate leaves, which are obliquely inserted and succubous in arrangement. It grows in the shady habitats of Adampur forest on the surface of the hilly rocks. Detailed taxonomy of the newly reported species along with its photographs is provided. Dhaka Univ. J. Biol. Sci. 29(1): 133-136, 2020 (January)
... These are classified under three groups: the mosses (14,000 species), the liver-worts (6000 species), and the hornworts (300 species). Mosses are considered to have the most significant number of species among green plants, next to angiosperms (Budke et al., 2018). The potential for medical use in bryophytes is one of the reasons for exploring biological compounds. ...
Article
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This study was conducted to assess the efficacy of SCoT and DAMD molecular markers in genetic diversity of three moss species and compare them with ISSR marker. Another objective was to evaluate the suitability of these DNA markers in species identification (delimitation) in three moss species, including Neckera complanata, Homalothecium sericeum and Neckera crispa. To prevent ecological impact on results, all samples were collected from one locality. PIC and MI in three markers showed closely ranged values. Our results revealed that the average values of Rp and the number of species-specific bands in SCoT primers were more than those of DAMD and ISSR. It showed considerable capability of SCoT marker in discriminating individuals. The highest value of genetic parameters Na (1.08), He (0.15) and I (0.23) were obtained with SCoT marker showing the power of this marker in genetic diversity analysis. Moreover, dendrograms produced from SCoT and DAMD data indicated similar results in the placement of closely related species. SCoT markers were shown to be more efficient than DAMD and ISSR markers based on the multiplex ratio (EMR), Rp, genetic diversity parameters (Na, He, I) and the number of species-specific bands. The findings demonstrated that the SCoT and DAMD markers could be applied for the estimation of genetic relationships and separation of two closely related genera. This is the first report of its type on the genetic diversity of mosses by application of SCoT and DAMD.