Pogonatum Classification Essay

1. The Plant Cuticle

Plants colonized land some 500 million years ago [1]. The evolution of molecular barriers formed from lipid-based polyesters was essential for the long-term success of land plants. These barriers protect the plant and also control the fluxes of gases, water, and solutes. The cuticle forms a waterproof layer on the epidermis that is important for protecting plants from biotic and abiotic stresses, such as herbivore attacks, dehydration and radiation [2]. Further, the cuticle is also involved in controlling the morphology of plants [3]. The cuticle is formed from cutin and waxes [2]. Cutin is a lipid polyester, formed mainly from glycerol and long chain (C16 and C18) hydroxy fatty acids, that in the cuticle are interspersed and covered with cuticular waxes [2,4]. Cuticular wax is a complex mixture of straight-chain C20 to C60 hydrocarbons, and may include secondary metabolites such as triterpenoids, phenylpropanoids, and flavonoids [5].

Synthesis of lipid polymers and cuticular waxes require de novo synthesis of precursors followed by transfer of the precursors through the plasma membrane to the apoplastic compartment [6]. Once exported, the hydrophobic polymer precursors and wax compounds are delivered to the polymerization sites outside the cell wall. This last step is probably the least understood in cuticle biosynthesis. It seems to require that the largely hydrophobic cuticle monomers traffic through the hydrophilic polysaccharide wall to reach the site of cuticle assembly. One family of proteins hypothesized to be involved in this trafficking is the non-specific lipid transfer proteins (LTPs) [4,7].

2. Lipid Transfer Proteins Could Have a Key Role in Cuticle Biosynthesis

LTPs are soluble, cysteine-rich, and small proteins with a molecular size usually below 10 kDa [8]. They are translated with an N-terminal signal peptide that localizes the protein to the apoplastic space. An LTP protein possesses four or five α-helices, which are stabilized by four conserved disulfide bridges, formed by an eight-Cys motif (8 CM) with the general form C-Xn-C-Xn-CC-Xn-CXC-Xn-C-Xn-C, where X specifies any amino acid and n is an unspecified number of amino acids. The disulfide bridges promote the folding of the helices around a central hydrophobic cleft (Figure 1), which is suitable for binding of hydrophobic ligands [9,10]. LTPs are compact structures that, to a high degree, are insensitive to heat and denaturing agents [11,12,13]. LTPs are abundant in all investigated land plants, but have not been detected in any other organisms [14]. They are encoded by large gene families in many flowering plants, while in bryophytes and ferns the gene families are significantly smaller [14,15,16,17]. LTPs are classified to one of five major types (LTP1, LTP2, LTPc, LTPd and LTPg) or four minor types (LTPe, LTPf, LTPh, LTPj and LTPk) [14]. The classification is based on the spacing between the Cys residues in the 8CM, the polypeptide sequence identity and the position of evolutionary conserved introns. The classification also reflects post-translational modifications, e.g., LTPs with a glycosylphosphatidylinositol (GPI)-anchor belong to LTPg. LTPd and LTPg were possibly the first LTP types that evolved in land plants, whereas LTP1 and LTP2, the most abundant LTP types in flowering plants, are not found in liverworts, mosses, or other non-seed plants [7,14].

There are a number of features that support the LTPs as stron candidates for delivering hydrophobic cuticle compounds to the apoplastic space: LTPs are synthesized with a signaling peptide and are secreted into the apoplast [7]. They are also abundantly expressed in the epidermis [18,19], small enough to traverse the pores of the cell walls, and their hydrophobic pocket is capable of binding long-chain fatty acids [20]. There is also some experimental evidence supporting a role for the LTPs in cuticular biosynthesis; when gene expression data from rice and Arabidopsis was investigated for co-expression patterns, the LTPgs could be arranged in three co-expressed clusters [21]. For the first cluster (I), expression was observed in aerial parts of the plant. The second cluster (II), was the only one with expression in roots, while expression of the third cluster (III) was restricted to reproductive tissues. Gene ontology analyses of genes coexpressed with the three Arabidopsis LTPg-clusters showed for cluster I an enrichment of genes involved with cuticular wax accumulation, for cluster II an enrichment of genes involved with suberin synthesis or deposition, and for cluster III an enrichment for genes acting in sporopollenin accumulation [21]. These coexpression patterns suggest that the LTPgs in the three clusters are involved in the assembly of the cuticle, suberin and sporopollenin, respectively.

In Arabidopsis AtLTPg1 and AtLTPg2, which both encode GPI-anchored LTPs, are highly expressed in the epidermis of inflorescence stems and silique walls [22,23], suggesting a role in cuticle development. Knock-down of AtLTPg1 resulted in reduced wax load on stem surfaces [22]. In Atltpg1 and Atltpg2 knock-out mutants, there was a 4–20% reduction in stems and siliques of the C29 alkane (nonacosane) component of cuticular wax, while an Atltpg1 Atltpg2 double mutant showed even stronger reductions [23,24]. There was also less total wax load in the stems and siliques of the double mutant and in the siliques of the ltpg2 single mutant [23,24]. Overexpression of the Brassica rapa BrLTPd1 gene in Brassica napus caused reduced wax deposition on leaves and morphological changes of leaves and flowers [25]. It was speculated that overexpression of BrLTPd1 leads to disordered secretion of wax, which was then lost from the surface, or to inhibition of other LTPs [25].

It is still unclear how LTPs aid in the extracellular transport of building blocks for lipid polymer synthesis. In previously suggested models, ATP-binding cassette (ABC) transporters move the cuticle polymer compounds through the plasma membrane [26,27,28,29]. On the extracellular side of the plasma membrane, lipids are possibly transferred from the ABC transporters to LTPs [7]. Hypothetically, LTPs could stimulate the diffusion or transport of lipid polymer and wax components to the sites of cuticle accumulation on the extracellular side of the plasma membrane [30], which for instance could be the surfaces of leaves, stems or pollen. It is possible that the ABC transporters deliver the polymer building blocks to LTPgs, which are attached to the apoplastic side of the plasma membrane through their GPI-anchor. The cargo may then be transferred from an LTPg to an LTP of another type that may diffuse freely in the cell wall.

An alternative hypothesis for the role of LTPs in cuticle assembly comes from investigations of AtLTP2 [31]. We previously renamed AtLTP2 to AtLTP1.4 to emphasize that it is of type LTP1 [7], and will use AtLTP1.4 onwards in this review. AtLTP1.4 is expressed only in epidermal cells of aerial organs, and an Atltp1.4-mutant has an increased cuticle permeability. In comparison to wild type, the Atltp1.4-mutant shows only minor differences in cuticular wax composition. However, in this mutant there is a 30% increase in 18:2 dicarboxylic acid, which is a major cutin component. The Atltp1.4-mutant also shows structural defects at the cell wall–cuticle interphase [31]. It was therefore proposed that AtLTP1.4 could play a major structural role by maintaining the integrity of the adhesion between the mainly hydrophobic cuticle and the underlying hydrophilic cell wall [31]. In another study, seeds from several Arabidopsis LTPg loss-of-function mutant lines showed increased permeability to tetrazolium salt, which suggests a malfunctioning seed coat. Morphological seed coat alterations were also shown for several of these LTPg mutant lines, as well as seed coat lipid polyesters with increased levels of unsubstituted fatty acids and decreased levels of ω-hydroxy fatty acids [32].

Hence, there are rather different roles suggested for LTPs in cuticle assembly and biosynthesis. LTPs may facilitate the transport or diffusion of the hydrophobic cuticle monomers and waxes in the hydrophilic cell wall, such LTPs could be functionally classified as Transporter LTPs (Figure 2). LTPs may also stabilize the adhesion between the cuticle and the cell wall. We classify these LTPs as Adhesion LTPs (Figure 2). These different hypotheses on the function of LTPs do not necessarily contradict each other. In the light of the variety of different LTP-types and the large number of members in the LTP family, it is likely that distinct members of the LTP family could participate as Transfer LTPs or Adhesion LTPs. It is also possible that both functions in transfer and adhesion could be fulfilled by singular LTPs. Further studies will hopefully reveal whether particular LTPs are involved in separate and specific processes during cuticle biosynthesis.

3. The Cuticle in Mosses and Liverworts

Already in the first half of the 20th century, the Finnish botanist Hans Buch described cuticles and waxy surfaces of mosses and liverworts [33]. In 1975, Schönherr and Ziegler [34] used scanning and transmission electron microscopy (SEM and TEM) and histochemistry to show beyond doubt the cuticle at the air pores of the thalli of several liverworts, such as Marchantia polymorpha, M. paleacea, and Plagiochasma elongatum [34]. Four years later, the leaf surfaces of 43 species of mosses were examined with SEM. Twelve species showed well developed superficial wax comparable to the cuticular wax of flowering plants [35]. This study was followed by one of the first reports on the chemical nature of the surface waxes of mosses, as the gametophytes of Andreae rupestris, Pogonatum aloides, and Pogonatum urngerum were shown to contain surface waxes in amounts of 0.05–0.1% of dry weight [36]. The main components of the surface waxes in these mosses were esters, free fatty acids, alcohols, aldehydes, and alkanes. The carbon chain lengths for these compounds were mainly C20–C28 in length for free fatty acids, C20–C24 for fatty acid esters, and C24–C28 for free alcohols and ester alcohols.

Also, the sporophyte, at least in some mosses, is covered by a cuticle, as shown by surface analysis of the sporophytes of Buxbamia viridis [37]. The calyptra is the small cap of maternal gametophyte tissue that covers the top of the sporophyte during development. This structure is covered with a multi-layered cuticle, similar in structure to the cuticle in flowering plants, as shown by SEM and TEM of the calyptra from the moss Funaria hygrometrica [38]. Hence, it appears that most aerial surfaces of bryophytes such as the sporophytes and the thalli of liverworts and the gametophores of many leafy mosses are protected by a cuticle with a similar composition and structure as the cuticle in vascular plants.

In vascular plants, the cuticle is a dynamic structure that can be structurally and chemically modified according to environmental and development requirements [39]. There are not many published experiments that address the physiological function and regulation of the cuticle in mosses. However, removal of the calyptra cuticle had a negative impact on development and reproduction. Without the calyptra cuticle, dehydration disrupts sporophyte maturation resulting in decreased spore production [40]. Although it is clear that the cuticle forms a protective barrier on outer surfaces also in liverworts and mosses, we lack substantial information about any dynamic modifications of the bryophyte cuticle in response to developmental or environmental cues.

Lipid profiling experiments of gametophores of the moss model organism Physcomitrella patens have revealed the presence of both cutin and cuticular waxes [13,41]. We have previously reported that unsubstituted fatty acids, fatty alcohols and ω-hydroxylated fatty acids are represented among the moss cutin monomers [13]. Generally, these compounds had chain-lengths of C16 or C18. The unsubstituted fatty acids were the largest class, with 70% of the total monomer content, and among the hydroxylated fatty acids, only C16 were found. There were no dicarboxylated fatty acids found in the moss samples. This class constitutes about 50% of the total cutin monomer content in Arabidopsis, but is significantly lower in other plants [42,43,44].

Furthermore, Buda and coworkers [41] identified large amounts of phenolic monomers, such as m- and p-coumaric acid, and caffeic acid, among the moss cutin monomers. Phenolic monomers are not usually found in the cutin from vascular plants, but are important components of suberin. The presence of phenolic monomers in the moss gametophyte shows that there are clear chemical differences between the cutin of vascular plants and bryophytes. Furthermore, as already pointed out by others, this could indicate that suberin and cutin in vascular plants share a common evolutionary origin [45,46]. The bryophyte cutin could represent a primitive cutin, similar in structure and chemical composition to the lipid barrier polymer of the earliest land plants.

4. Mosses and Liverworts as Model Systems for Cuticle Function and Assembly

In recent years, the genomes of several bryophytes, such as the mosses P. patens and Sphagnum fallax, as well as the liverwort M. polymorpha have been sequenced [47,48,49]. Today, M. polymorpha and P. patens are established land plant model species with an array of available tools for genetic, developmental and molecular analyses [50,51,52]. Because the dominant phase of the bryophyte life cycle is the haploid gametophyte, the effects of transgenes can be studied already a few weeks after transformation. There are established protocols for obtaining gene knock-outs or knock-downs via RNAi, Transcription activator-like effector nucleases (TALENs), CRISPR/Cas9, homologous recombination or artificial miRNAs [53,54,55,56,57,58,59

INHALT

ABOUT THE AUTHOR

PREFACE

ACKNOWLEDGEMENTS

ABOUT THE BOOK
1 INTRODUCTION GEOLOGY, CLIMATE AND VEGETATION OF INDIA
2 METHODOLOGY

Discussion

REFERENCES

ABOUT THE AUTHOR

DR. AFROZ ALAM is currently working as an Associate Professor in the Department of Bioscience and Biotechnology, Banasthali University, Rajasthan. He obtained his M.Sc. and Ph.D. from University of Lucknow, (U.P.). He also qualified ASRB-NET (ICAR) examination. His areas of specialization are Bryophytes’ systematics and bryodiversity. He was awarded J.R.F. and S.R.F. in “All India coordinated Project on Taxonomy Capacity Building for Bryophytes” (AICOPTAX) sponsored by Ministry of Environment & Forests, New Delhi at Bryology Unit, Department of Botany, University of Lucknow during his research. Dr. Alam has collected over 3000 accessions of bryophytes in several plant collection trips to different localities of the country, including some arduous and treacherous terrains of Nilgiri and Palni Hills of Southern Peninsula, Eastern and Western Himalayas and Rajasthan.

Dr. Alam has vast experience of teaching and research in bryology. He has over 50 research publications in prestigious International and National Journals and 4 text books. Dr Alam has also attended and presented papers in a number of seminars. He is a Life member of Association of Plant Taxonomy (APT), India and Indian Bryological Society (IBS). Presently, he is working as an Associate Professor in the Department of Bioscience and Biotechnology, Banasthali University, Rajasthan. He is one of the curators of BURI herbarium. Botanical Survey of India (ENVIS-BSI) approved him as one of the experts of bryology in India.

Dedicated

To

Late Prof. Jan-Peter Frahm

‘The bryologist beyond boundaries’

PREFACE

Mosses comprise an imperative component of the mountain forest ecosystem. Mixed climatic conditions have granted India with prosperous moss diversity, yet there is no newer update regarding moss flora for India. The most recent checklist was published in the year 2005 which incorporated 1623 taxa of mosses distributed under 342 genera and 57 families. Ever since, not a single attempt was made for updating Indian moss flora. To fill this lacuna, an effort has been made to accumulate the latest status of Indian mosses in the form of present title “Moss Flora of India”, based on the all accessible and existing reports on mosses so far from the India. It provides the most up-to-date account of floristic diversity in the Indian mosses. This comprehensive listing is supplemented with present status (valid/synonym/doubtful) of each taxon based on www.theplantlist.org. Each species have been dealt with as far as achievable complete distributional details inside India. It also incorporates information about their endemism status. A modified classification scheme of Buck and Goffinet (2000) has been used for preparation of this updated checklist. This work would be useful for future studies related to Indian mosses.

The physical map has been adopted from the website of the Department of Tourism, India, while the map of India has been adopted from School Atlas (based on survey of India, 1989). Any deviation from the original map, however, has no significance, political or otherwise.

ACKNOWLEDGEMENTS

The author acknowledges the help rendered by the bryologists from various parts of India for providing research articles related to Indian bryophytes.

Thanks are due to the Prof. Aditya Shastri, Vice Chancellor, Banasthali University, Rajasthan and Prof. Vinay Sharma, Dean, Faculty of Science and Technology, Banasthali University, Rajasthan for providing basic facilities.

Prof. S. C. Srivastava, ex- Head, Department of Botany, University of Lucknow deserves special thanks for their support and encouragement during the course of present study.

Last but not least thanks are due to my family and friends for their continuous support.

I offer my genuine apologies for any inaccuracy and request suggestions from all the readers of this book to improve the publication in succeeding editions.

Afroz Alam

ABOUT THE BOOK

The immense forest cover, sufficient precipitation and vital relative humidity in India provide highly encouraging environment for the lavish prevalence of both terricolous and corticolous mosses. The mosses which occur in specialized habitats like soil surface, soil covered rocks (both moist as well as dry), shaded or exposed rocks, extremely wet rocks are considered as terrestrial while those on other larger trees as corticolous. Majority of substrate receives sufficient precipitation and humidity which provide eco-friendly conditions for the growth and development of these remarkable plants.

The present compilation of the moss flora of India revealed the occurrence of total 1578 species of mosses which belong to 21 orders, under 66 families and 328 genera. Out of these 897 retained their valid status, while 437 species are now considered as a synonym and status of 244 species is still unresolved i.e. doubtful name. 130 taxa have been reported as endemic to India. The updated checklist of mosses of India reveals that the most diversified order is Hypnales with 28 families, followed by Dicranales (6 families); Pottiales and Bryales (4 families each). Order Timmiales, Splachnales and Rhizogoniales have 2 families each and rests of the orders are represented by single families. Whereas, in terms of family, family Pottiaceae is the most prominent having 38 genera followed by Hypnaceae with 20 genera. Genera like Fissidens (72 spp.), Brachythecium (38 spp.), Pogonatum (33 spp.), Mnium (25 spp.), Calymperes (23 spp.), Brachymnium (22 spp.), Sphagnum (21 spp.), Barbula (21 spp.), Entodon (20 spp.) and Tortula (15 spp.) are most diversified in India. This great diversity of mosses revealed the potential of India in terms of bryodiversity particularly, the mosses.

1 INTRODUCTION GEOLOGY, CLIMATE AND VEGETATION OF INDIA

India engages only 2.50% of the overall land of the planet. With a physical area of 329 million ha, it is world’s seventh leading country after Russia, Canada, USA, Brazil, Australia and China. India is one of the signatory members of Convention on Biological Diversity (CBD), and also one of the 17 megadiversity countries globally recognized with their geographic extent both on land as well as sea (www.wikipedia.org). It has diverse edaphoclimatic circumstances ranging from the icy mountain climate in the north to a humid tropical one in the south to dry hot desert in the North West (Fig. 1). This mixture of climatic conditions is also reflected in the forest and the biodiversity wealth of the country (Champion and Seth, 1968). Historically, India had over 65% of its total land under forests as early as in 1925 (at present only 20.55%; Ravindranath et al, 2014). The resultant varied climate supplemented with topography created 10 biogeographic zones (Lakshminarayana et al., 2001) namely the Trans-Himalayan, Himalayan, Indian desert, Semi arid, Western Ghats, Deccan peninsula (including the eastern Ghats), Gangetic plains, North-east India, Coasts and Islands comprising their own fragile and unique ecosystem and well defined flora ranking third in Asia and eleventh in the world harboring over 45,000 phytobinomials including lower non-vascular cryptogams, the ‘Bryophytes’. India is discernible by diversity of physical features such as peaks, uplands, plains, coastline and isles (Figs. 1 & 2). The brief descriptions of these are given below:

The Himalayas

The Himalayas considered the youngest series of folded mountains on the planet. The Himalayan Region of India is a range that extends ten states of India including Jammu & Kashmir, Himachal Pradesh, Uttarakhand, Arunachal Pradesh and Sikkim, Assam and West Bengal. It forms a curve which is about 2,400 km long. There are three similar ranges in its longitudinal extent (Ahmad, 2011). The region is liable for providing water to a hefty part of the Indian subcontinent. The region physiographically, starting from the foothills of south (Siwaliks) and extends up to Tibetan highland to the north (Trans-Himalaya). The width differs from 150 km in Arunachal Pradesh (eastern) to 400 km in Kashmir (western). The altitudinal deviations are bigger in the eastern part than in the western division. Three major geographical entities, the Himadri (greater Himalaya), Himanchal (lesser Himalaya) and the Siwaliks (outer Himalaya) extending almost continuous throughout its length, are separated by major geological fault lines. Grand rivers like the Indus, Sutlej, Kali, Kosi and Brahmaputra have cut through sharp ravines to run away into the Great Plains. Some of the utmost mountains on this planet are found in this region.

Trans-Himalayas:

This region is the Northern most area in the India extends in the states of Jammu and Kashmir and Himachal Pradesh.

The Himadri:

This is the most constant range of Himalayas (Karakoram Mountains). It holds the highest peaks of the Himalayas. The middling height of the peaks in this range is ca. 6,000 m. The folds of the Great Himalayas are irregular in nature and the center of this part is composed of granite. Because of the supercilious heights, the peaks of this range are snow-bound all through the year.

The Himanchal:

This is the southern part of the Himalayas. The elevation of peaks in this range differs from 3,700 m to 4,500 m. The Middling width of this range is ca. 50 km. This range is mainly composed of extremely compacted and distorted rocks.

The Shiwaliks Himalaya:

This is the farthest range of the southern Himalayas. It lies to the south of the Dhaula Dhar, the average height of this range is 1,500 to 2,000m and the width ranges 10 to 50 km. It includes the Jammu hills and extends to Kangra. On Uttarakhand side, it stretches from Dehra Dun to Almora and finally merge with the southern borders of Nepal. This series is made of uncombined sediments. This can be divided into four sub regions, viz. Kumaon Himalayas, Assam Himalayas, Punjab Himalayas and Nepal Himalayas.

The Indian Desert:

The western part of India has the Great Indian Desert. This is also known as the Thar Desert. It lies towards the western boundaries of the Aravali hills. It is a large (17th largest subtropical desert of the world), arid region in the northwestern part of the Indian subcontinent. In India, it covers about 320,000, of which 60% is in Rajasthan and extends into Gujarat, Punjab, and Haryana. The rainfall is very scanty (<150 mm/year), hence the vegetation of this region is meager.

The Semi Arid Region

The semi arid regions in India encompass largely of the Rann of Kutch and the semi -arid regions of Punjab and Gujarat. The Southern arid regions are in the rain shadow of the Western Ghats covering states of Maharashtra, Karnataka and Tamil Nadu.

The Peninsular India

To the south of the northern plains lies the peninsular plateau. It is triangular in outline. The relief is extremely bumpy. This is the expanse with numerous hill ranges and valleys. Aravali hills, one of the oldest ranges of the world, border it on the north-west side. It is made of the ancient rocks since it was formed from the drifted part of the Gondwana. Expansive and superficial valleys and smoothed hills are the attributes of this plateau. The Vindhyas and the Satpuras are the important ranges. The river Narmada and Tapi flow through these ranges. The upland can be largely alienated into two expanses-the Middle Uplands and the Deccan Plateau.

The Middle Uplands: The Middle Uplands lies to the north of the Narmada waterway. It covers the most important portion of the Malwa highland. The rivers in this region run from southwest to northeast, which points out the slant of this area. The upland also extends eastwards into the Chhotanagpur upland.

The Deccan Plateau: The Deccan plateau of Gondwanaland comprises of one of the most fragile ecosystem “the Western Ghats”, one of the ‘hot spots’ of India, hosting 30% endemic flora and fauna. The Sahyadris or Western Ghats border the plateau in the west and the Eastern Ghats offer the eastern frontier.

The Western and the Eastern Ghats:

The Western Ghats are the chief tropical evergreen region of India, spread over cardinal 220 N to 80 latitude, covering the length of 1400 km. The Western Ghats, known locally as the Sahyadri Hills, are formed by the Malabar Plains and the chain of mountains running parallel to India's western coast, about 30 to 50 km inland. The Western Ghats instigate oceanographic rains as they face the rainy winds from the west. They cover an area of about 160,000 km² and stretch for 1,600 kilometers from the country's southern tip to Gujarat in the north, broken up only by the 30 kilometers Palghat Gap.

Unlike, Western Ghats, the Eastern Ghats are broken and jagged. The plateau is loaded with various minerals. The average elevation of Eastern Ghats is about 650m (Alam et al., 2007). The Eastern Ghats, also known as Mahendra Pravata are broken and jagged range of mountains along India's eastern shore. The Eastern Ghats run from West Bengal state in the north, through Odisha and Andhra Pradesh to Tamil Nadu in the south passing some parts of Karnataka. They are wrinkled and cut through by the four major rivers of peninsular India, known as the Mahanadi, Krishna, Godavari and Kaveri.

North East India: The Brahmaputra River indicates the eastern boundary of the Himalayas. Ahead of the Dihang gorge, the Himalayas curve stridently towards south and form the Eastern hills. These hills run all the way through from the north eastern states of India. They are typically composed of sandstones. These hills are identified as Naga Hills, Manipuri Hills, Patkai Hills, and Mizo Hills.

The Gangetic Plains:

The northern Indian plains of India lie to the south of the Himalayas. They are usually flat and level. These are formed by the alluvial deposits laid down by the three rivers- the Indus, the Ganga, the Brahmaputra and their tributaries. These river plains provide fertile land for cultivation. The entire area of the northern plain is about 7 lakh sq. km. It is about 240 to 320 km broad and 2400 km long. The northern plain is divided into three sections, viz. the Ganga Plain, the Brahmaputra Plain and the Punjab Plain.

The Coastal Plains:

The Western Coastal Plains consist of a thin strip of coastal plain ca. 50 km in girth between the west coast of India and the Western Ghats hills, which starts close to the south of Tapi. They are squeezed in between the Western Ghats and the Arabian Sea. The plains initiate at Gujarat in the north and finish at Kerala in the south. It also embraces the states of Maharashtra, Goa and Karnataka. The western coastal plain has two parts. The northern part is known as Northern Circar, while, the southern part is called the Coromandel Coast.

The Eastern Coastal Plains refer to an extensive expand of Indian landmass, lying between the Eastern Ghats and the Bay of Bengal. These plains are wider and level as compared to the western coastal plains. It widens from Tamil Nadu (south) to West Bengal (north). The western coastal plains are very narrow. The eastern coastal plains are much broader. They run along the Arabian Sea on the west and along the Bay of Bengal on the east. There are number of east flowing rivers. The rivers Mahanadi, Godavari, Krishna and Kaveri drain into the Bay of Bengal.

The Islands

Two groups of Islands also form part of India. The Lakshadweep Islands are located in the Arabian Sea. Its area is 32 sq km. This cluster of islands is prosperous in terms of biodiversity. The Andaman and the Nicobar Islands lie to the southeast of the Indian mainland in the Bay of Bengal. They are larger in size and has added a number of islands. These islands also have rich biodiversity.

Prevailing factors of vegetation and flora

Land is one of the most important factors which directly and indirectly governs the vegetation and flora in nature. Varied soil types are vital for the establishment of different types of vegetation and flora.

Climatic conditions like temperature and humidity are the chief factors which govern the nature and level of flora and vegetation of an area. In general, an area with elevated temperature and elevated humidity supports evergreen forest, while an area with elevated temperature and low humidity supports xerophytic vegetation. Likewise, photoperiod (duration of light) is another factor which depends on latitude, altitude, season and duration of the day length. Precipitation also controls the nature and type of vegetation. If an area gets heavy rainfall, it is suitable for the growth of dense vegetation. Conversely, an area with insufficient rainfall is suitable for xerophytes. Ecosystem also has detrimental effects on vegetation. All the biological organisms in an area are mutually dependent on each other. These, along with their physical environment make the ecosystem. A very large ecosystem is called a biome which is identified on the basis of plant communities (www.envindia.com).

NATURAL VEGETATION

Natural vegetations are offerings of nature. They grow naturally. They follow the climatic variables. Because of a mixture of climates, a wide range of natural vegetation grows in India. Types of natural vegetation vary according to climate, soil and altitude. The naturally growing plants cover without human any interference or help is called natural vegetation. They have their characteristic flora (www.ncert.nic.in).

Types of Natural Vegetation

The following are the principal types of natural vegetation in India (Kishwan et al.,2009):

(1) Tropical Rain Forests
These forests grow in areas where precipitation is > 200 cm/year. They are mainly found on the slants of the Western Ghats and the north-eastern regions of Arunachal Pradesh, Meghalaya, Assam, Nagaland, the foothill areas of the Himalayas and the Andaman groups of Islands. Vegetation in such a forest has a multilayered structure. Sisthu, chaplash, bamboos, garjan, ebony, mahogany, rosewood, rubber, cinchona and sandalwood are some of the commercially important trees of these rainforests.
(2) Tropical Deciduous or Monsoon Forests
In India these forests are the most prevalent also known as Monsoon forests. They are common in those regions which acquire annual precipitation between 70 cm and 200 cm/year. The trees are deciduous which shed their leaves during summer.
On the basis of availability of water these can be divided into following two types.
(a) Moist Deciduous Forest: The moist deciduous forests need annual rainfall between 100 cm and 200 cm. Northeastern states hold these forest. However, they are also found on the eastern slopes of the Western Ghats, the foothills of the Himalayas, Jharkhand, West Orissa and Chhattisgarh.
(b) Dry Deciduous Forest: The dry deciduous forests grow in areas where the annual rainfall is between 70 cm and 100 cm. These are found in areas of the central Deccan plateau, southeast of Rajasthan, Punjab, Haryana and parts of Uttar Pradesh and Madhya Pradesh. Most of these areas are used for agriculture.
(c) Semi-deserts and Desert vegetation :These forests exist in the regions that receive less than 70 cm of annual rainfall. North-western parts of India (Rajasthan and parts of Gujarat, Punjab aand Karnataka) hold this type of vegetation. The main plant species in such a forest are xerophytes.
(d) Montane Forests : In India, Montane forests are classified into three categories; the montane wet temperate forests, Himalayan moist temperate forests and Himalayan dry temperate forests. These forests differ significantly according to altitude with varying rainfall and temperature along the slopes of the mountain.
(i) The montane wet temperate forests include shola forests. These are found between a height of 1000 and 2000 m. Evergreen broad-leaf conifers are abounding in these forests.
(ii) The Eastern Himalayan forest is an eco-region of temperate broadleaf forest. These are found in the middle elevations (1500 -3000m) of the eastern Himalayas. Gymnospermous trees of family Coniferaceae flourish in such forests.
(iii) Alpine plants grow in the alpine climate, which occurs at high altitude (>3600m) and above the tree line. Alpine plants grow mutually as a plant community. Alpine plants are limited to a single taxon. Rather, many diverse plant species live in this environment. These plants must adapt to the harsh conditions of the alpine environment, which include low temperatures, dryness, ultraviolet radiation, and a short growing season. These forests are mainly found along the southern slopes of the Himalayas and at high altitudes in southern and north-eastern India.
(e) Mangrove Forests

Mangrove forests are found in the deltas of the Ganga, the Mahanadi, the Krishna, the Godavarai and the Kaveri. They are also known as ‘Tidal Forests'; because their dense growth depends upon tidal water, which submerges the deltaic lands during high tides (www.envindia.com; www.importantindia.com).

illustration not visible in this excerpt

Figure 1: Major Physiographic Divisions of India ( Oxford, 2014)

BRYOFLORISTIC WEALTH

After angiosperms, bryophytes constitute the second most varied land plant group. There are three major bryophyte phyla: Hepatophyta the “liverworts” approx. 6000-8000 species, Anthocerophyta the “hornworts” approx. 150 species, Bryophyta the “mosses” includes about 680 genera and approx. 10000-12000 species. Bryophytes are characterized by an alternation of generations (heteromorphic) with the haploid generation (gametophyte) being the dominant phase of the life cycle. They are also referred as “Lilliputians of Plant Kingdom” due to their minuscule size. They have a dichotomous branching pattern, a primitive type a branching. They do not have any vascular supply in the midrib. They are habitually known as ‘Amphibians of the plant kingdom’ as they need water to complete the process of fertilization (Glime, 2007).

Mosses

As mentioned earlier, mosses have utmost species diversification. They habitually grow in moist and shady places as crowded greenish cluster or carpet. The plants are typically composed of simple, unistartose leaves, covering a thin axis that supports them. Plants are atracheate, as they do not have vascular tissue. At maturity, they produce sporophytic structures with massive capsules containing spores. They are usually 1–12 cm tall, however, some species like, Dawsonia are much larger, which can grow to 50 cm in length and considered as the tallest moss in the world (Schofield, 1985).

Mosses belong to phylum (division) Bryophyta, which previously also included liverworts and hornworts. Now liverworts and hornworts have separate divisions. There are roughly 10,000-12,000 taxa of moss exist under the Bryophyta (Vitt, 1984). The foremost viable use of mosses is for ornamental purposes, such as in gardens and in the florist trade. Conventional uses of mosses included as filling and as they have great capability to soak up liquids up to 20 times their weight (Sphagnum can absorbs 200 times ) they are extensively used in nursery practices (Glime, 2007).

Life Cycle

The life-cycle of moss exhibits a heteromorphic alternation of generations. It has two very distinct phases, the haploid or gametophytic and the diploid or sporophytic phases. The gametophytic phase is the dominant phase that commences with germination of haploid spore which considered as the first cell of gametophyte. The gametophyte has two growth stages: (i) Protonema stage, which is the juvenile stage represented by prostrate, green and branched thread-like structure. This is an ephemeral stage in the life cycle of a moss; and (ii) leafy stage or gametophores, which is an erect cylindrical shoot with persistent leaves and sex organs. Usually, a solitary carpet of protonema may develop several shoots, ensuing in an aggregation of moss. The main axis can be branched or unbranched. The branches always arise below the leaves. The leaves are simple, minute sessile and usually are one cell in thickness (Shaw and Goffinet, 2000). There are two mode of reproduction:

(i) Vegetative reproduction: It takes place by fragmentation, stolons, branching of protonema, special leafy shoots, gemmae and persistent apices.
(ii) Sexual reproduction: From the apices of main stem and branches of the gametophores development of sex organs in groups takes place. The plants are dioicous or monoicous. In dioicous species, male and female sex organs are borne on different plants (Glime and Knoop, 1986). In monoicous (also known as autoicous) species, both the sex organs are borne on the same plant. The female sex organs are called archegonia, which are protected by a group of modified protective leaves known as the perichaetum. The archegonia are stalked with a much elongated neck and massive venter. These are flask-shaped structure with an open neck through which the male sperm swim towards the venter. The male sex organs are termed as antheridia, these are club-shaped, narrow and elongated. The antheridial jacket is single layered. These are enclosed by modified protective leaves called the perigonium. The surrounding leaves of atheridium form a cup like cavity, allowing the sperm contained in the cavity to be splashed to adjacent stalks (female branches) by falling rain drops, for this reason this is called ‘Splash cup mechanism’ (Glime, 2007).

Fertilization takes place with the aid of water. Water confers the swimming movement of male gametes from the antheridium towards the archegonium to complete the process of fertilization. The male gametes of mosses are biflagellate structures and exhibits chemotactic movement. After fertilization, the juvenile sporophyte pushes its way out of the archegonial venter. After six months for the sporophyte becomes mature. The sporophyte is usually differentiated into foot, seta and capsule. The elongated seat raises the capsule much above the gametophyte to facilitate the dispersal off spores. The capsule is covered by a cap like structure called the operculum. The capsule and operculum both are enclosed by an additional structure called calyptra which is the remnants of the archegonial venter hence haploid in nature. It normally falls off from full-grown capsule. The capsule usually has a peristome which helps in the dispersal of spores. Within the capsule, sporogenous mass undergo reduction division and form haploid spores. Most mosses depend on the wind for their dispersal.

It has recently been found that few insects can affect fertilization process of moss because they attract towards the specific moss emitted odour. For example, fire moss, releases unusual and composite volatile organic odour. Female plants give off added compounds than male plants. Springtails prefer usually the female plants of this taxa, and increase moss fertilization, suggesting a odour-mediated relationship equivalent to the plant-pollinator relationship found in many seed plants. Similarly, Splachnum sphaericum increases insect pollination at higher rate by magnetizing flies to its sporangia with a intense odour of carrion. Bright red coloured inflated collars below each spore filled capsule serve as a strong visual hint for pollinators. Flies fascinated to the moss transmit its spores to fresh droppings of cattle, which is the conducive habitat of this taxa (Glime, 2007).

Habitat

They are better adapted to terrestrial habitats than other groups of bryophytes. Since moss gametophytes have no water conducting tissue through the plant or water holding systems to avoid tissue water from evaporation, mosses need a moist environment in which they grow, and availability of fluid water for reproduction. Since mosses are autotrophic (except saprophytic genus Buxbaumia) they need sufficient sunlight to conduct photosynthesis. Selection of substrate differs according to the specis. They grow on moist soil, rocks, damp walls, old buildings and on the stem of trees in tropical forest. Moss species growing on or in the shade big trees are often species specific such as preferring reduced leaf tree like pinus to broadleaf trees like eucalyptus. The remarkable point about the mosses is their mode of nutrition, they always grow on trees as epiphytes, they never adopt the parasitic mode on the phorophyte.

In general, mosses can grow in a variety of habitats. For example, Barbula comosa, Weisia endultula, Brachymenium walkeri grow on dry faces of cliffs of gneiss and granites; Bryum giganteum, Hypophila comosa and Barbula indica on pegmatites, lime and black loam; Fissidens lutescens, F. walkeri, Trematodon ceylonesis and Bryum wiohlii on banks of stream in shady places; Bryum ramosum and B. doliolum on dry banks; Leucoloma walkeri, Fissidens aomalous and Bryum apalodictyoides on dead wood and decaying tree trunks: Leucoloma renauldii, Tortella hyalinoblasta and Macromitrium sulcatum on large trees forming felts (in dense jungles) etc. are found growing in various parts of the country. Several mosses such as Polytrichum are reported to grow on rocks in extremely xerophytic conditions, sometimes in association with lichens. These help in succession of the vegetation (xerosere). Likewise, Porella platyphylloidea growing on rocks can withstand without water for several months. It shows extreme drought resistance.

One of the ecological significance of bryophytes is to avoid erosion by holding the soil particles trough carpet form of growth and retaining the greater amount of water. They also slow down the rapid runoff water and melted snow. These also provide a soft bed to the seeds and moist conditions favourable to their growth. Thus, yet insignificant, bryophytes play a quiet but fairly vital role in the nature (Proctor, 1984).

DISTRIBUTION OF MOSSES IN INDIA

India is one of the 17 mega biodiversity countries in the world and one of the hot spots of biodiversity.. The large area and a variety of phytoclimatic conditions contribute to the great diversity of the Indian flora (Singh, 1997; 2001; Alam et al., 2007). In case of bryophytes and especially mosses, rich diversity is found in different regions of India. According to a rough estimate about 2000 species of mosses, 816 species of liverworts and 34 species of hornworts are occurring in India. The plants are distributed in Eastern and Western Himalayas, South India, Rajasthan, Gujarat, Punjab, Central India, Andaman and Nicobar Islands (Fig. 2). According to Lal (2005) about 2480 taxa of bryophytes (including intraspecific taxa) are reported from India (including island groups, and Sikkim), comprising about 722 taxa of liverworts in 128 genera and 52 families, 36 taxa in 6 genera and 2 families of hornworts and about 1623 taxa in 342 genera and 57 families of mosses.

Several bryologists of India have assessed the moss flora of different bryological regions from time to time, such as Gangulee (1969-80) published “Mosses of Eastern India and Adjacent regions” in eight fascicles which included 990 species. Chopra (1975) dealt with nearly 2,000 species belonging to 329 genera under 56 families. He listed many species as “likely to occur” and also nomen nuda. Lal (2005) published “A checklist of Indian mosses” and listed 1623 taxa of mosses from India. After that, no valid attempt has been made to provide a complete wealth of mosses in India with their current status.

This compilation work is an attempt to provide current status of the moss flora of India which includes mosses of Western Himalayas, Eastern Himalayas, South India, Central India, Gangetic plains, Punjab, Rajasthan, Gujarat Andaman and Nicobar Islands (Figure 1 & 2).

The assets of this work include almost all previous work and reports from these regions of India, like, Dixon, 1909; Dabhade, 1969; Chopra, 1975; Vohra, 1970; Gangulee, 1969-1980; Chopra and Kumar, 1981; Tewari and Pant, 1994; Deora and Chaudhary, 1996; Dabhade, 1998; Madhusoodanan and Nair, 2004; Lal, 2005; Saxena and Gangwar, 2005; Nair and Madhusoodanan, 2006; Saxena et al., 2006; Madhusoodanan et al., 2007; Manju et al., 2007; Saxena et al, 2007; Nath et al, 2007; Daniels and Daniel., 2007; Kuamar and Krishnamurthy, 2007; Nath et al, 2008; Aziz and Vohra, 2008; Saxena and Arfeen, 2009; Saxena et al, 2010; Singh et al., 2010; Daniels, 2010; Dandotiya et al., 2011; Manju et al., 2011; Alam et al, 2012; Verma et al., 2011; Alam, 2013a,b; Daniels and Kariyappa, 2013; Asthana and Sahu, 2013, Rajesh et al.; 2013; Schwarz, 2013; Schwarz and Frahm, 2014; Rajesh and Manju, 2014; Alam et al., 2014). This work aims to be all-inclusive for species of mosses. It contains a total 757 moss species along with their upper hierarchy. The list has three categories for the species names viz. Valid name, synonym and doubtful name. These categories are based on available literature from reliable resources like ‘The Plant List’ (2010) which was prepared in collaboration of the Royal Botanic Gardens, Kew and Missouri Botanical Garden by combining multiple checklist datasets held by these institutions and other collaborators. This present list also follows this and provides the valid name, synonyms and also provides doubtful name for which the contributing data sources did not contain sufficient evidence to decide whether they were accepted or synonyms.

Therefore, this assemblage provides comprehensive, reorganized and an updated account on the moss flora of India.

2 METHODOLOGY

This study is fundamentally based on all previous and recent reports regarding moss flora of India. The orders are arranged alphabetically for feasibility. All moss species included in the list were checked against the TROPICOS database (at the Missouri Botanical Garden). A modified classification scheme of Buck and Goffinet (2000) has been used for preparation of this updated checklist.

THE CHECKLIST OF MOSSES

A. ORDER: TAKAKIALES

Family: Takakiaceae S. Hatt. & Inoue.

1. Takakia S. Hatt. & Inoue.

1. Takakia ceratophylla (Mitt.) Grolle; Present status: Valid name

Distribution in India: Eastern Himalayas

B. ORDER: ANDREAEALES LIMPR.

Family: Andreaeaceae Dum.

2. Andreaea Hedw.

2.Andreaea commutata Müll. Hal.; Present status: Synonym of Andreaea rupestris var. fauriei (Besch.) Takaki

Distribution in India: Eastern Himalayas (Endemic to India)

3. Andreaea densifolia Mitt.; Present status: Valid name

Distribution in India: Eastern Himalayas (Endemic to India)

4. Andreaea indica Mitt.; Present status: Doubtful

Distribution in India: Eastern Himalayas (Endemic to India)

5. Andreaea rigida Wilson in Mitten; Present status: Valid name

Distribution in India: Eastern Himalayas (Endemic to India)

6. Andreaea kashyapii Dixon & Vohra & Wadhwa Endemic to India; Present status: Synonym of Didymodon subandreaeoides (Kindb.) R.H. Zander

Distribution in India: Endemic to India to Western Himalyas.

7. Andreaea rupestris Hedw.; Present status: Valid name

Distribution in India: Western Himalayas and Eastern Himalayas

C. ORDER:ARCHIDIALES LIMPR.

Family: Archidiaceae Schimp.

3. Archidium Brid.

8. Archidium birmannicum Mitt. ex Dixon; Present status: Valid name

Distribution in India: Gangetic plains and South India

9. Archidium indicum Müll. Hal.; Present status: Doubtful name

Distribution in India: Western Himalayas and Central India.

10.Archidium microthecium Dixon & P. de la Varde; Present status: Doubtful name

Distribution in India: South India.

11. Archidium octosporum Dixon & P. de la Varde; Present status: Synonym of Archidium ohioense Schimp. ex Müll. Hal.

Distribution in India: South India

D. ORDER:BRYALES LIMPR.

Family: Bryaceae Schwägr.

4. Anomobryum Schimp.

12. Anomobryum astorense (Broth.) Broth. ; Present status: Valid name

Distribution in India: Western Himalayas

13. Anomobryum auratum (Mitt.) A. Jaeger.; Present status: Valid name

Distribution in India: Western Himalayas, Eastern Himalayas, South India

14. Anomobryum brachymenioides Dixon & P. de la; Present status: Synonym of Bryum brachymenioides (Dixon & P. de la Varde) Ochi

Distribution in India: South India

15. Anomobryum cymbifolium (Lindb.) Broth.; Present status: Valid name

Distribution in India: Western Himalayas, Eastern Himalayas, South India

16. Anomobryum filiforme (Griff.) A. Jaeger ; Present status: Valid name

Distribution in India: Western Himalayas, Eastern Himalayas, South India

17. Anomobryum filiforme var. concinnatum (Spruce) Loesk. (Asthana & Sahu, 2013); Present status: Valid name

Distribution in India: Western Himalayas, Eastern Himalayas, South India

18. A. kashmirense (Broth.) Broth.; Present status: Valid name

Distribution in India: Western Himalayas

19. Anomobryum latifolium Cardot & P. de la Varde; Present status: Synonym of Bryum auratum Mitt.

Distribution in India: South India

20.Anomobryum marginatum Dixon & Badhw.; Present status: Synonym of Bryum blandum subsp. handelii (Broth.) Ochi

Distribution in India: Western Himalayas

21. Anomobryum nitidum (Mitt.) A. Jaeger.; Present status: Valid name

Distribution in India: Western Himalayas and Eastern Himalayas

22. Anomobryum pellucidum Dixon & Badhw.; Present status: Synonym of Bryum himalayanopellucidum Ochi

Distribution in India: Western Himalayas

23. Anomobryum schmidii (Müll. Hal.) A. Jaeger; Present status: Valid name

Distribution in India: South India

24. Anomobryum subnitidum Cardot & P. de la Varde; Present status: Doubtful

Distribution in India: South India

5. Brachymenium Schwägr.

25. Brachymenium acuminatum Harv.; Present status: Valid name

Distribution in India: Eastern Himalayas and South India

26. Brachymenium alpinum Ochi; Present status: Valid name

Distribution in India: Eastern Himalayas

27. Brachymenium bryoides Hook. & Schwägr.; Present status: Valid name

Distribution in India: Western Himalayas, Eastern Himalayas, South India

28. Brachymenium clavariaeforme (Müll. Hal.) A. Jaeger; Present status: Valid name

Distribution in India: South India

29. Brachymenium cristatum (Müll. Hal.) A. Jaeger; Present status: Valid name

Distribution in India: South India

30. Brachymenium extenuatum (Mitt.) A. Jaeger; Present status: Valid name

Distribution in India: South India

31. Brachymenium exile (Dozy & Molk.) Bosch & Sande Lac.; Present status: Valid name

Distribution in India: Western Himalayas, Eastern Himalayas, South India

32. Brachymenium fischeri Cardot & Dixon; Present status: Doubtful

Distribution in India: South India

33. Brachymenium flaccidisetum (Müll. Hal.) A. Jaeger; Present status: Valid name

Distribution in India: South India

34. Brachymenium indicum (Dozy & Molk.) Bosch & Sande Lac.; Present status: Valid name

Distribution in India: Gangetic plains

35. Brachymenium lanceolatum Hook. & Wilson; Present status: Valid name

Distribution in India: Central India

36. Brachymenium leptostomoides (Müll. Hal.) A. Jaeger; Present status: Doubtful

Distribution in India: South India

37. Brachymenium longicolle Thér.; Present status: Synonym of Brachymenium leptophyllum (Bruch & Schimp. ex Müll. Hal.) Bruch & Schimp. ex A. Jaeger

Distribution in India: Eastern Himalayas

38. Brachymenium longidens Renauld & Cardot; Presnt status: Doubtful name

Distribution in India: Eastern Himalayas

39. Brachymenium microstomum Harv.; Present status: Synonym of Pseudopohlia bulbifera R.S. Williams

Distribution in India: Eastern Himalayas and Central India

40. Brachymenium nepalense Hook.; Present status: Valid name

Distribution in India: Western Himalayas, Eastern Himalayas, South India

41. Brachymenium ochianum Gangulee; Present status: Synonym of Brachymenium capitulatum (Mitt.) Paris

Distribution in India: Eastern Himalayas

42. Brachymenium ptychothecium (Besch.) Ochi; Present status: Valid name

Distribution in India: Eastern Himalayas

43. Brachymenium rugosum (Müll. Hal.) A. Jaeger; Present status: Valid name

Distribution in India: South India

44. Brachymenium sikkimense Renauld & Cardot; Present status: Doubtful name

Distribution in India: Eastern Himalayas

45. Brachymenium velutinum (Müll. Hal.) A. Jaeger; Present status: Valid name

Distribution in India: South India

46. Brachymenium walkeri Broth.; Present status: Doubtful name

Distribution in India: Eastern Himalayas and south India

6. Bryum Hedw.

47. Bryum allionii Broth.; Present status: Synonym of Bryum mildeanum Jur.

Distribution in India: Western Himalayas, Eastern Himalayas, South India

48. Bryum alpinum Huds. & With. ; Present status: Valid name

Distribution in India: Western Himalayas, Eastern Himalayas, South India

49. Bryum alpinum var. mildeanum (Jur.) Podp.; Present status: Synonym of Bryum mildeanum Jur.

Distribution in India: Western Himalayas, Eastern Himalayas, South India

50. Bryum apiculata Schwägr.; Present status: Synonym of Bryum mildeanum Jur.

Distribution in India: Western Himalayas, Eastern Himalayas, South India and Gangatic Plains

51. Bryum ambiguum Duby; Present status: Valid name

Distribution in India: South India

52. Bryum andrei Cardot & P. de la Varde; Present status: Synonym of Bryum paradoxum Schwägr

Distribution in India: South India

53. Bryum argenteum Hedw.; Present status: Valid name

Distribution in India: Western Himalayas, Eastern Himalayas, South India, Central India and Rajasthan

54. Bryum argenteum var. lanthanum (P. Beauv.) Hampe; Present status: Synonym of Bryum argenteum Hedw.

Distribution in India: Western Himalayas, Eastern Himalayas, South India and Central India

55. Bryum. atrovirens Brid.; Present status: Valid name

Distribution in India: Western Himalayas and, Eastern Himalayas

56. Bryum badhwarii Ochi ; Present status: Synonym of Bryum kashmirense Broth.

Distribution in India: Western Himalayas and Eastern Himalayas

57. Bryum bessonii Renauld & Cardot; Present status: Valid name

Distribution in India: South India

58. Bryum bicolor Dicks.; Present status: Synonym of Bryum dichotomum Hedw.

Distribution in India: Western Himalayas, Eastern Himalayas and Rajasthan

59. Bryum billardieri Schwägr. ; Present status: Valid name

Distribution in India: South India

60. Bryum bornholmense Wink. & R. Ruthe; Present status: Valid name

Distribution in India: Rajasthan

61. Bryum bryoides (R. Br.) Ångström ; Present status: Synonym of Bryum arcticum (R. Br.) Bruch & Schimp.

Distribution in India: Western Himalayas

62. Bryum caespiticium Hedw.; Present status: Valid name

Distribution in India: Western Himalayas and Eastern Himalayas

63. Bryum capillare Hedw.; Present status: Synonym of Ptychostomum capillare (Hedw.) D. T. Holyoak & N. Pedersen

Distribution in India: Western Himalayas, Eastern Himalayas and, South India

64. Bryum cellular Hook.; Present status: Valid name

Distribution in India: Western Himalayas, Eastern Himalayas and Nicobar Island.

65. Bryum. coronatum Schwägr.; Present status: Valid name

Distribution in India: Western Himalayas, Eastern Himalayas, South India, Central India, Rajasthan, Gangetic plains and Andaman Island.

66. Bryum doliolum Duby, Present status: Synonym of Bryum coronatum var. doliolum (Duby) A. Jaeger

Distribution in India: South India

67. Bryum euryphyllum Dixon & P. de la Varde; Present status: Valid name

Distribution in India: South India

68. Bryum flaccum Wilson ex Mitt.; Present status: Doubtful name

Distribution in India: Eastern Himalayas

69. Bryum formosum Mitt. ; Present status: Synonym of Bryum wightii Mitt.

Distribution in India: South India

70. Bryum ghatense Broth. & Dixon; Present status: Synonym of Bryum mildeanum Jur.

Distribution in India: South India

71. Bryum ghatense var. satarense Broth. & Dixon; Present status: Synonym of Bryum mildeanum Jur.

Distribution in India: South India

72. Bryum heterophyllum Warnst.; Present status: Synonym of Bryum dichotomum Hedw.

Distribution in India: South India

73. Bryum klinggraeffii Schimp.; Present status: Valid name

Distribution in India: Rajasthan and Gangetic plains

74. Bryum lamprostegum Müll. Hal.; Present status: Doubtful

Distribution in India: South India

75. Bryum medianum Mitt. ; Present status: Synonym of Bryum neelgheriense var. neelghreniense

Distribution in India: Western Himalayas, Eastern Himalayas, South India

76. Bryum montagneanum Müll. Hal.; Present status: Synonym of Brachymenium pendulum Mont.

Distribution in India: South India

77. Bryum muehlenbeckii Bruch & Schimp.; Present status: Valid name

Distribution in India: Western Himalayas, Eastern Himalayas, South India (Endemic to India)

78. Bryum nitens Harv Present status: Synonym of of Gemmabryum apiculatum (Schwägr.) J.R. Spence & H.P. Ramsay

Distribution in India: South India

79. Bryum pachycladum Cardot ex P. de la Varde; Present status: Doubtful

Distribution in India: South India

80. Bryum paradoxum Schwägr.; Present status: Valid name

Distribution in India: Western Himalayas, Eastern Himalayas, South India

81. Bryum paradoxum var. reflexifolium (Ochi) Ochi; Present status: Synonym of Bryum reflexifolium (Ochi) Ochi

Distribution in India: Eastern Himalayas

82. Bryum plumosum Dozy & Molk.; Present status: Synonym of Gemmabryum apiculatum (Schwägr.) J.R. Spence & H.P. Ramsay

Distribution in India: Western Himalayas, Eastern Himalayas, South India and Gangetic Plains

83. Bryum porphyroneuron subsp. erythropus M. Fleisch.; Present status: Synonym of Bryum mildeanum Jur.

Distribution in India: Western Himalayas, Eastern Himalayas, South India

84. Bryum pseudotriquetrum (Hedw.) P. Gaertn., B. Mey. & Scherb.; Present status: Valid name

Distribution in India: Western Himalayas, Eastern Himalayas

85. Bryum ramosum (Harv.) Mitt.; Present status: Synonym of Bryum billarderi Schwägr

Distribution in India: South India

86. Bryum ramosum subsp. zollingeri (Duby) M. Fleisch.; Present status: Doubtful name

Distribution in India: South India

87. Bryum. recurvatum Broth.; Present status: Synonym of Bryum recurvulum Mitt.

Distribution in India: South India

88. Bryum retusifolium Cardot & P. de la Varde; Present status: Valid name

Distribution in India: South India

89. Bryum rubens Mitt.; Present status: Valid name

Distribution in India: Rajasthan

90. Bryum salakense Cardot; Present status: Valid name

Distribution in India: India Orientalis

91. Bryum tuberosum Mohamed & Damanhuri; Present Status: Synonym of Rosulabryum tuberosum (Mohamed & Damanhuri) J.R. Spence

Distribution in India: South India

92. Bryum turbinatum (Hedw.) Turner ; Present status: Valid name

Distribution in India: Western Himalayas

93. Bryum. uliginosum (Brid.) Bruch & Schimp.; Present status: Valid name

Distribution in India: Western Himalayas, Eastern Himalayas

94. Bryum vellei Cardot & P. de la Varde; Present Status: Synonym of Rosulabryum billarderi (Schwägr.) J.R. Spence

Distribution in India: South India

95. Bryum vellei var. robustum Dixon & P. de la Varde; Present Status: Synonym of Rosulabryum billarderi (Schwägr.) J.R. Spence

Distribution in India: South India

96. Bryum weigelii Spreng.; Present status: Valid name

Distribution in India: Western Himalayas

97. Bryum wightii Mitt.; Present status: Valid name

Distribution in India: South India

7. Mielichhobryum J. P. Srivast.

98. Mielichhobryum sahayadrense J.P. Srivast. ; Present status: Doubtful name

Distribution in India: Western Himalayas and South India

8. Mniobryum Limpr.

99. Mniobryum delicatulum (Hedw.) Dixon ; Present status: Synonym of Pohlia melanodon (Brid.) A.J. Shaw

Distribution in India: Western Himalayas

[...]

0 Thoughts to “Pogonatum Classification Essay

Leave a comment

L'indirizzo email non verrà pubblicato. I campi obbligatori sono contrassegnati *