Liverwort Family Research Articles

A Taxonomic Revision of Aneuraceae (Marchantiophyta) from Eastern Africa with an Interactive Identification Key

A taxonomic revision of the liverwort family Aneuraceae in eastern Africa is presented based on the results of an integrative approach. Molecular and morphological data lead to the recognition of fourteen species in eastern Africa, in three genera: eleven species in Riccardia Gray, two in Aneura Dumort. and one in Afroriccardia Reeb & Gradst. One further species, R. multifida, may be expected in eastern Africa and is included in this treatment. Keys, descriptions and illustrations are provided for the accepted species together with data on types, synonymy, geographical distribution, habitat and differentiating characters. A knowledge database was built using Xper3 and an interactive key was generated using the Xper3 platform tools. Riccardia gasparii sp. nov., R. martinii sp. nov. and R. vohimanensis sp. nov. are described as new to science. A comparison with other regions of the world shows that Africa has relatively few Aneuraceae taxa; only the Holarctic region, with the exception of East Asia, has fewer species. Moreover, dendroid taxa and species with cuticular ornamentation are lacking in Africa. On the other hand, Africa stands out by the presence of the endemic genus Afroriccardia. More collecting is needed in future to clarify the species ranges, the status of questionable taxa and the knowledge of local floras.

The checklist of Hungarian bryophytes – second update

According to the checklist presented here, the bryophyte flora of Hungary consists of 2 species in 2 genera and 2 families of hornworts, 146 species and 3 additional infraspecific taxa (1 subspecies and 2 varieties) in 60 genera and 34 families of liverworts, and 521 species and 17 additional infraspecific taxa (5 subspecies and 12 varieties) in 186 genera and 64 families of mosses, in total 669 species, 6 subspecies and 14 varieties or 689 taxa. During the last decade, 40 bryophyte taxa (6 liverworts and 34 mosses) were added to the Hungarian bryophyte flora (Appendix 1.1), and an additional 19 taxa (4 liverworts and 15 mosses) that were already part of the Hungarian flora but not recognised in the previous checklist, are now incorporated in the list (Appendix 1.2). On the other hand, 29 taxa (7 liverworts and 22 mosses) must be excluded according to our present state of knowledge (Appendix 2). Another 13 taxa (2 liverworts and 11 mosses) that are at present not considered members of the Hungarian bryoflora are annotated in Appendix 3. They were not recognised in the previous checklist (with one exception), but are reported from Hungary in some literature and with five exceptions are also recognised in the European checklist. In many cases we have insufficient information to definitely decide upon their status. Annotations to all taxa listed in the three appendices provide additional information. In Appendix 4 to all names of bryophyte taxa that appeared in the previous checklist (except excluded taxa) the presently accepted names are assigned.

<p><strong><em>Cololejeunea manilaliana </em></strong><strong>a new synonym of <em>Cololejeunea nilgiriensis </em>(Lejeuneaceae: Marchantiophyta)</strong></p>

Lejeuneaceae, the largest family of liverworts with over 1800 species in 74 genera, is considered taxonomically very difficult (Ye & Zhu 2018; Zhu & Shu 2018; Zhu et al. 2017; 2018a, b; Zhu et al. 2019; Asthana & Srivastava 2019; Asthana et al. 2019). Cololejeunea (Spruce 1884: 291) Stephani (1891: 208) is the largest genus in the family (He et al. 2012; Zhu et al. 2018b), and represented in India by 63 taxa (Asthana & Srivastava 2003; Asthana & Srivastava 2015; Dey & Singh 2016a,b; Singh & Singh 2016; Manju et al. 2017; Singh 2018; Asthana & Srivastava 2019; Asthana et al. 2019). Among these, one of the recently described species, Cololejeunea manilalia Manju, Chandini & Rajesh (2017: 262) appears to be conspecific with Cololejeunea nilgiriensis Asthana & Srivastava (2003: 27).

Spores of Plagiochila (Dumort.) Dumort.: the taxonomic relevance of morphology and ultrastructure

ABSTRACT Plagiochilaceae is a family of leafy liverworts that are distributed worldwide. It is of great importance due to its taxonomic and ecological implications among bryophytes. Most species of the family belong to the genus Plagiochila, but there is no consensus regarding its infrageneric circumscription. There have been few palynological studies involving Plagiochilaceae and Plagiochila. Here, we describe the spore morphology of seventeen species of Plagiochila and discuss the taxonomic value of palynological characters for these taxa. The spores were processed by standard palynological techniques and analyzed using light and electron microscopy. The spores were found to be apolar, spheroidal, released monads that vary in size from 13µm to 58µm (small to large size). The sporoderm comprises an intine (stratified), a nexine, and a sexine. The spore surface is ornamented with granules that vary in shape and morphology, thus allowing the studied species to be grouped into four spore types: regular and delicate granulate, irregular and coarse granulate, long granules with flattened apices, and long and straight granules. Hierarchical cluster analysis revealed five different groups of species, evidencing the importance of spore information for taxonomic and phylogenetic studies.

Mohamedia, a new genus of Lejeuneaceae (Marchantiophyta) from Oceania and tropical Asia

Lejeuneaceae is the largest family of liverworts with an estimated 1800 species in about 73 genera. The systematic position of a number of taxa in the family remains ambiguous owing to the lack of molecular data and morphological observations especially on oil bodies and ocelli in fresh samples. Drepanolejeunea brunnea Mizut. and Lepidolejeunea borneensis (Steph.) R.M.Schust. are two examples of these taxa. Here we investigate the phylogenetic relationships of these two species by using the nuclear ribosomal ITS region and two chloroplast regions (rbcL, trnL–trnF). The molecular data show that Drepanolejeunea brunnea and Lepidolejeunea borneensis form a well-supported clade unrelated to Drepanolejeunea (Spruce) Steph. and Lepidolejeunea R.M.Schust. On the basis of the combined molecular-phylogenetic and morphological data, a new genus in the subtribe Lepidolejeuneinae, Mohamedia R.L.Zhu et L.Shu, is described to accommodate Drepanolejeunea brunnea and Lepidolejeunea borneensis. Mohamedia is closely related to Otolejeunea Tixier endemic to Madagascar, but differs in the thinner stem with only 3(–4) rows of medullary cells, perianths with 4–5 crenulate to weakly dentate keels and without auricles, and lack of an unbroken row of ocelli in leaf lobes. Two new subgenera, Mohamedia subg. Mohamedia and Mohamedia subg. Piippolejeunea R.L.Zhu et L.Shu, are described and the new combinations (Mohamedia brunnea (Mizut.) R.L.Zhu et L.Shu and M. borneensis (Steph.) R.L.Zhu et L.Shu) are proposed.

ATLANTIC EPIPHYTES: a data set of vascular and non-vascular epiphyte plants and lichens from the Atlantic Forest.

Epiphytes are hyper-diverse and one of the frequently undervalued life forms in plant surveys and biodiversity inventories. Epiphytes of the Atlantic Forest, one of the most endangered ecosystems in the world, have high endemism and radiated recently in the Pliocene. We aimed to (1) compile an extensive Atlantic Forest data set on vascular, non-vascular plants (including hemiepiphytes), and lichen epiphyte species occurrence and abundance; (2) describe the epiphyte distribution in the Atlantic Forest, in order to indicate future sampling efforts. Our work presents the first epiphyte data set with information on abundance and occurrence of epiphyte phorophyte species. All data compiled here come from three main sources provided by the authors: published sources (comprising peer-reviewed articles, books, and theses), unpublished data, and herbarium data. We compiled a data set composed of 2,095 species, from 89,270 holo/hemiepiphyte records, in the Atlantic Forest of Brazil, Argentina, Paraguay, and Uruguay, recorded from 1824 to early 2018. Most of the records were from qualitative data (occurrence only, 88%), well distributed throughout the Atlantic Forest. For quantitative records, the most common sampling method was individual trees (71%), followed by plot sampling (19%), and transect sampling (10%). Angiosperms (81%) were the most frequently registered group, and Bromeliaceae and Orchidaceae were the families with the greatest number of records (27,272 and 21,945, respectively). Ferns and Lycophytes presented fewer records than Angiosperms, and Polypodiaceae were the most recorded family, and more concentrated in the Southern and Southeastern regions. Data on non-vascular plants and lichens were scarce, with a few disjunct records concentrated in the Northeastern region of the Atlantic Forest. For all non-vascular plant records, Lejeuneaceae, a family of liverworts, was the most recorded family. We hope that our effort to organize scattered epiphyte data help advance the knowledge of epiphyte ecology, as well as our understanding of macroecological and biogeographical patterns in the Atlantic Forest. No copyright restrictions are associated with the data set. Please cite this Ecology Data Paper if the data are used in publication and teaching events.

Frullania grabenhorstii sp. nov., a fossil liverwort (Jungermanniopsida: Frullaniaceae) with perianth from Bitterfeld amber

Frullaniaceae are the most diverse family of leafy liverworts preserved in amber and are known from several deposits dating from the Miocene to the Early Cretaceous. In the fossil record, Frullania is represented by 15 species as well as the extinct genera Pseudofrullania, Protofrullania and Kaolakia. Here, we describe another species of Frullania from Bitterfeld amber (Germany) as Frullania grabenhorstii sp. nov. A combination of characters associated with the leaf lobe, leaf lobule, underleaf, branching patterns, and perianth distinguishes it from all other known extant and extinct taxa. Many characters of the new fossil are shared with F. subg. Frullania sect. Australes, especially the morphologically similar extant species F. incumbens and F. subincumbens-both of Australasia. Another similar species, Fullania densiloba, occurs in Japan. These distribution patterns reinforce previously described affinities of the Baltic and Bitterfeld bryophyte floras to the extant flora of Asia and Australasia. This pattern has been found in several taxa, e.g., Notoscyphus, Nipponolejeunea, and Metacalypogeia. The new fossil is compared with other species from Bitterfeld, Baltic, and Rovno amber, which show significant morphological differences. In addition to the description, we provide an overview of the family Frullaniaceae in different amber deposits.

Species diversity of the genus Riccia L. (Marchantiales, Ricciaceae) in Maranhão state, Brazil

Ricciaceae is a little-known liverwort family in northeastern Brazil. Fieldwork in 4 localities in Maranhão state yielded 4 species of Riccia, with 2 taxa, R. mauryana and R. weinionis, representing new state records. This paper describes the species diversity of the genus Riccia in Maranhão state, and provides descriptions, ecological notes, and illustrations for each species.

Soella (Marchantiophyta: Lejeuneaceae), a new genus from China and Japan

Lejeuneaceae is the largest family of liverworts with over 1800 species distributed mostly in moist tropical and subtropical regions. Drepanolejeunea (Spruce) Steph. is the fourth largest genus of this family with over 120 species currently listed in the updated checklist of liverworts and hornworts, however no phylogenetic studies of this genus based on an extensive taxon sampling have been done. Here we for the first time elucidate phylogenetic relationships of Drepanolejeunea by using the nuclear ribosomal ITS region and two chloroplast regions (rbcL, trnL−trnF). Our molecular phylogenetic results reveal that Drepanolejeunea as currently circumscribed is not monophyletic. On the basis of the combined molecular phylogenetic and morphological data, a new genus in the subtribe Lepidolejeuneinae, Soella, is proposed to include the remarkable and rare species, Drepanolejeunea obtusifolia T.Yamag. previously known from Japan. Drepanolejeunea subg. Acantholejeunea R.M.Schust. (≡ Acantholejeunea (R.M.Schust.) R.M.Schust.), one of five subgenera currently accepted in Drepanolejeunea, is suggested to be synonymous with Ceratolejeunea (Spruce) J.B.Jack et Steph. Soella obtusifolia (T.Yamag.) R.L.Zhu et al. comb. nov. and Ceratolejeunea spinistipula (Herzog) R.L.Zhu et al. comb. nov. are proposed. Drepanolejeunea tenera K.I.Goebel is new to the liverwort flora of China. With the description of Soella, the liverworts (Marchantiophyta) so far contain 369 genera, including 73 genera belonging to Lejeuneaceae.

Lunularia cruciata (L.) Dumort. ex Lindb.: A New Plant Record for the Bryoflora of Kashmir Himalaya, India

Lunularia cruciata (L.) Dumort. ex Lindb., the only species of the monotypic liverwort family Lunulariaceae (Lunulariales), is reported for the first time for the bryoflora of Kashmir Himalaya. It is easily recognizable by crescent-shaped gemma cups on the surface of the thalli.

Lejeunea apiahyna, a new synonym of Lejeunea cristuliflora (Lejeuneaceae, Marchantiophyta)

Lejeuneaceae is the largest liverwort family with more than 1800 species in about 70 genera (Zhu et al. 2017). Lejeunea Libert (1820: 373) is the second largest genus of Lejeuneaceae with many taxa poorly known (He et al. 2012). Lejeunea apiahyna (Stephani 1914: 514) Singh (2013: 63) is one of those poorly known species and is known only from Brazil. It was described by Stephani (1914) as Otigoniolejeunea apiahyna Stephani (1914: 514) based on J. Puiggari’s collection from the State of São Paulo, Brazil, and transferred to the genus Lejeunea by Singh (2013). According to Wei et al. (2014), Otigoniolejeunea Spruce (1884: 77) is represented by only two ocellus containing species: Otigoniolejeunea huctumalcensis (Lindenberg & Gottsche in Gottsche et al. 1847: 762) Wei et al. (2014: 237), and Otigoniolejeunea portoricensis (Hampe & Gottsche 1853: 352) Wei et al. (2014: 237).

New or little known epiphyllous liverworts, XXII. Reinerantha (Lejeuneaceae), a genus new to Venezuela

Reinerantha foliicola was recently described as a new epiphyllous genus and species in the Cololejeuneinae subtribe of the liverwort family Lejeuneaceae, from the montane rainforest region of Ecuador. A second locality of this unusual plant was detected in Venezuela in a rich Andean montane rainforest near Mérida, at an elevation of 2,300 m.

Reinerantha foliicola, a new genus and species of Lejeuneaceae subtribe Cololejeuneinae (Marchantiophyta) from Ecuador

AbstractCololejeuneinae is the largest subtribe of the liverwort family Lejeuneaceae and the crown group of the family. The species often grow on the surfaces of living leaves and twigs or in swiftly running waters and many of them possess neotenic gametophytes, considered an adaptation to these extreme habitats. We describe a new, epiphyllous genus and species in Cololejeuneinae from the Western Cordillera of Ecuador, Reinerantha foliicola Gradst. & R.L.Zhu. Analysis of combined molecular datasets with parsimony, maximum likelihood and Bayesian methods revealed Reinerantha in a sister relationship to the large pantropical genus Diplasiolejeunea. Reinerantha differs from Diplasiolejeunea in leaf segmentation, epidermal wall thickening, stylus, male bracts and perianth keels, and is morphologically more similar to the genus Tuyamaella. In the molecular analysis, however, Reinerantha and Tuyamaella were resolved in different clades and not closely related. Tuyamaella is shown to be restricted to Asia; the report of Tuyamaella from South America (Peru) was erroneous and a case of mislabeling.

Towards a monophyletic classification of Lejeuneaceae V: the systematic position of Pictolejeunea

With more than 1,000 species in some 70 genera, Lejeuneaceae represent the largest family of liverworts. Although much progress has been made in recent years, the supraspecific classification of Lejeuneaceae has not yet been satisfactorily solved. Maximum likelihood and Bayesian analyses of a three marker dataset (nrITS, cp DNA rbcL and trnL-trnF) derived from 85 accessions of Lejeuneeae and 13 outgroup species revealed Pictolejeunea picta in a sister relationship to the remainder of Lejeuneeae, the most speciose tribe of Lejeuneaceae. Based on the outcome of the phylogenetic analyses, we establish subtribe Pictolejeuneinae subtr. nov.

Estudo comparativo de comunidades de briófitas sujeitas a diferentes graus de inundação no município de São Domingos do Capim, PA, Brasil

RESUMO Foram analisadas 35 parcelas (10 × 10 m) para a listagem florística e, destas, 18 parcelas (seis para cada fitofisionomia: Floresta de Igapó (IG), Floresta de Várzea (VA) e Floresta de Terra Firme (TE)) foram sorteadas para a análise de comparação das comunidades de briófitas. Encontramos 118 espécies de briófitas: 37 musgos e 82 hepáticas, sendo que Lejeuneaceae (63 espécies) foi a família mais rica para hepáticas e Calymperaceae (oito espécies) para musgos. A estimativa de riqueza está abaixo da esperada, porém a curva de coletor estabilizada. O substrato mais colonizado foi tronco vivo (101 spp., 45 exclusivas) seguido de tronco em decomposição (66 spp., 16 exclusivas). Os resultados obtidos mostram que as fitofisionomias analisadas apresentam diferenças na riqueza (IG > TF > VA) e composição das comunidades de briófitas, devido à existência de espécies exclusivas e as análises de Agrupamento (UPGMA) e Ordenação (NMDS) indicaram estruturação das comunidades entre as três fitofisionomias.

Epiphyllous Liverworts of India: An Overview

Taxonomy of the epiphyllous liverworts in India has been reviewed and their diversity and distribution has been discussed. A total of 160 species, one subspecies and two varieties of epiphyllous liverworts belonging to 23 genera in eight families have been recognized in Indian bryoflora, distributed only in eastern Himalaya and the north-east, Western Ghats, and the Andaman & Nicobar Islands. Eastern Himalaya, including the north-east, with 133 taxa shows the maximum diversity of epiphylls, whereas Sikkim with 80 taxa is the richest amongst the States. Lejeuneaceae with 131 species belonging to 16 genera is the most prolific family of epiphyllous liverworts accounting for over 80 per cent of their total diversity in India, while Cololejeunea with 54 species is the most dominant genus. Twenty species are endemic to India, of which 11 are restricted to eastern Himalaya, three to Western Ghats, and one to Andaman & Nicobar, while five species are common between eastern Himalaya and the Western Ghats.

Survey of bryophytes in Serra da Canastra National Park, Minas Gerais, Brazil

Serra da Canastra National Park is located in southwestern Minas Gerais State and contains several phytophysiognomies of the Brazilian Cerrado. To date, no study on bryophytes has been conducted in this area and the present study is the first to be carried out in this major biological reserve of Cerrado located on the Brazilian Plateau. This study found 289 species of bryophytes, including mosses, liverworts, and one species of hornwort, representing fourty-one Brazilian endemic species and 56 new records for Minas Gerais State. Most species are widely distributed in Brazil, with only 16% having restricted distributions. Regarding worldwide distribution, 31% are Neotropical. Lejeuneaceae had the highest species richness among liverwort families with 53 species, whereas Sphagnaceae had the highest richness among moss families with 26 species. Phaeoceros laevis was the only hornwort species found in the park. This study contributes to the understanding of bryophyte species richness and distribution, and provides the worldwide and Brazilian distribution of the bryophyte flora of Serra da Canastra National Park, an important center for biodiversity conservation.

Liverworts to the rescue: an investigation of their efficacy as mycorrhizal inoculum for vascular plants

Summary Pezoloma ericae (D.J. Read) Baral, a widespread mycorrhizal fungus of plants in the Ericales, is known to form intracellular associations with several families of leafy liverworts (Schistochilaceae, Lepidoziaceae, Cephaloziaceae, Cephaloziellaceae) in vitro. The ecological significance of this link between vascular and non‐vascular plants is unknown. Fungal symbionts were isolated from rhizoids of the leafy liverworts Cephalozia connivens (Dicks.) Lindb. and C. bicuspidata (L.) Dum. (Cephaloziaceae), as well as from the hair roots of two dominant ericoid mycorrhiza‐forming species of European heathlands, Erica tetralix (L.) and Calluna vulgaris (L.). Using pure cultures of P. ericae, we resynthesized liverwort–fungus associations to use colonized liverworts as inoculum which was applied to substrates supporting the growth of heather seedlings and cuttings. Effects were quantified using germination, rooting, plant colonization, plant survival under waterlogging stress and growth in height in experimental systems with and without liverworts and/or fungi. Fungal symbionts growing from liverwort rhizoids readily colonized the hair roots of ericaceous plants to form typical ericoid mycorrhizas. The presence of inoculum‐bearing liverworts led to significant increases in plant growth. Erica tetralix was more responsive to inoculation than C. vulgaris. Ericaceous cuttings rooted and survived more successfully when they were coplanted with previously colonized liverwort stems. We demonstrate, under realistic ecological circumstances, that liverworts can deliver mycorrhizal inoculum and improve the establishment of vascular plants. We propose that by providing sources of mycorrhizal inoculum, symbiotic non‐vascular plants can contribute to the restoration of plant communities dominated by Ericales plants. This research leads to broader knowledge about the function of ericoid mycorrhizas in ecosystems.

Towards a monophyletic classification of Lejeuneaceae IV: reinstatement of Allorgella, transfer of Microlejeunea aphanella to Vitalianthus and refinements of the subtribal classification

The classification of Lejeuneeae, the most speciose tribe of the largely epiphytic leafy liverwort family Lejeuneaceae, is deemed notoriously difficult and hampered by extensive morphological homoplasy. We employed sequences of two chloroplast regions (trnL-trnF, rbcL) and the nuclear ribosomal ITS region of several critical taxa to explore their phylogenetic position. Based on our phylogenetic analyses and accompanying morphological investigations, we transfer Microlejeunea aphanella to Vitalianthus, reinstate the genus Allorgella, transfer Vitalianthus from Drepanolejeuneinae to Lepidolejeuneinae, identify Metalejeunea as another element of Lepidolejeuneinae, and place Allorgella and Haplolejeunea in Echinolejeuneinae. We map selected morphological characters on the reconstructed phylogenetic hypothesis to demonstrate the importance of subgynoecial innovations and ocelli for the circumscription of genera and subtribes of Lejeuneeae.

Preliminary Attempts to Conserve Bryophytes in Lowland Forests of Sri Lanka: A Case Study at Yagirala Forest Reserve

Bryophytes were the first dwellers of the land plant history and play a vital role in the dynamics of the forest ecosystems such as water flow regulation and nutrient cycling. Considerable efforts on surveying and taxonomic studies of bryoflora in Sri Lanka was done in the early and mid-19th century by foreign scientists and there is a huge gap for several decades till late 1990s and no proper systematic survey of bryoflora in lowland forests in Sri Lanka has been carried out in the recent past. Sri Lanka is a biodiversity hotspot and it has a rich and diverse tropical bryoflora in the lowland and highland natural wet -moist forests. Yagirala forest reserve is a lowland tropical rainforest situated in the low country wet zone. Many floristic researches on angiosperms have been carried out but there is no systematic survey has been carried out to study the cryptogamic plants in the forest. Hence this is the first attempt to study and identify distribution of the bryoflora at Yagirala forest reserve. The bryophyte diversity in three main ecosystems in the forest including the primary forest, secondary forest and the forest associated with the Pinus plantation was studied. Ten (10) different families of mosses (including 12 genera, 16 species), Six (06) families of leafy liverworts (including 12 genera, 14 species) and 03 families of simple thalloid liverworts (including 03 genera, 03 species) were found. All species records were mapped and seemed to be new species records to Sri Lanka. DNA samples were collected for further barcoding studies and voucher specimens were prepared. A comparison of the species diversity in each major habitat type was studied and some anthropogenic and natural threats to the native bryoflora in the forest were studied. Further intensive research work is needed to prepare a complete checklist of bryoflora in the Yagirala Forest reserve and the final output of the project will be developing a user-friendly pocket guide to study the common bryophytes in the forest and this will open a novel avenue to initiate the conservation and IUCN redlisting of lowland bryoflora in Sri Lanka. Keywords: Bryophytes, Preliminary survey, Yagirala Forest Reserve, Conservation