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Two new records of Riccia L. (Ricciaceae) for the Eastern Himalaya Bryogeographical Region of India
Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx
Contents lists available at ScienceDirect
Journal of Asia-Pacific Biodiversity journal homepage: http://www.elsevier.com/locate/japb
Original Article
Two new records of Riccia L. (Ricciaceae) for the Eastern Himalaya Bryogeographical Region of India Naorem Premita Devi, S.D. Yumkham*, P.K. Singh Ethnobotany & Plant Physiology Laboratory, Centre of Advanced Studies in Life Sciences, Manipur University, Canchipur, 795 003, India
a r t i c l e i n f o
a b s t r a c t
Article history: Received 12 January 2019 Received in revised form 2 July 2019 Accepted 20 August 2019 Available online xxx
The Eastern Himalaya Bryogeographical Region is recognized as one of the richest areas for bryoflora in India. It comprises the eight North Eastern States (Arunachal Pradesh, Assam, Manipur, Meghalaya, Mizoram, Nagaland, Sikkim, and Tripura), Darjeeling, and the Hills of the West Bengal. In the present article, the occurrence of Riccia curtisii (Austin) Austin and Riccia pathankotensis Kashyap of Ricciaceae is reported for the first time for the Eastern Himalaya region from the state Manipur. Detailed descriptions and photographs of the two newly recorded species of Riccia L. are provided to facilitate easy identification. A key to species of the nine Riccia species found in Manipur is also incorporated. Ó 2019 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing Services by Elsevier. This is an open access article under the CC BY-NC-ND license (http:// creativecommons.org/licenses/by-nc-nd/4.0/).
Keywords: Riccia New record Eastern Himalaya Manipur
Introduction Riccia L. (Ricciaceae) is the largest genus among the complex thalloid liverworts with 152 species worldwide and in India, with 36 species constituting 24% of the global species diversity (Söderström et al 2016; Singh 2014). The plants possess simple linear or oblong gametophytic thalli differentiated into upper photosynthetic zone and lower storage tissue, with or without simple pores similar to other thallose members. It is unique in having fertile plants with anacrogynous condition, sporophytic sporogonia or capsules embedded singly within the thalli, absence of pseudoperianth, and cleistocarpous capsules without elaters. It differs from Ricciocarpos Corda, the other twin genus of Ricciaceae in lacking idioblastic oil cells, which are present in the latter (Crandall-Stotler et al 2009). Majority are terrestrial mesophytes which preferably grow on exposed or shaded mixed alluvial sandy soil with few aquatic and semiaquatic forms. There are eight bryogeographical regions currently recognized in India to assess the phytodiversity of liverworts and hornworts. These include the Andaman & Nicobar Islands [A & N], Central India [CI], Deccan Plateau & Eastern Ghats [DP & EG], Eastern Himalaya [EH], Gangetic Plains [GP], Punjab & West Rajasthan [P & WR],
* Corresponding author. E-mail address: rifl[email protected] (S.D. Yumkham). Peer review under responsibility of National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA).
Western Ghats [WG], and Western Himalaya [WH]. The EH comprises the eight North Eastern States (Arunachal Pradesh, Assam, Manipur, Meghalaya, Mizoram, Nagaland, Sikkim, and Tripura), Darjeeling, and the Hills of the West Bengal (Singh et al 2016). Altogether, 16 species have been recognized so far from the EH region, namely, R. attenuata Pandé, R. beyrichiana Hampe ex Lehm., R. billardierei Mont. & Nees, R. cavernosa Hoffm., R. ciliata Hoffm., R. crispatula Mitt., R. cruciata Kashyap, R. discolor Lehm. & Lindenb., R. fluitans L., R. frostii Austin, R. gangetica Ahmad, R. glauca L., R. hasskarliana Steph., R. huebeneriana Lindenb., R. sorocarpa Bisch. and R. stricta (Gottsche, Lindenb. & Nees) Perold (Singh 2014). In the present article, two unique species of Riccia, namely, R. curtisii (Austin) Austin and R. pathankotensis Kashyap, are recorded for the first time for the EH region from the North Eastern State Manipur. R. curtisii is one of the very rare species in India and is reported from GP and WH regions (Pande and Ahmad 1944; Srivastava 1964; Tewari and Pant 1983). It is taxonomically very significant because of the presence of permanently adhered spore tetrads with long spines, the only characteristic feature found in three Riccia species, the other two being R. leptothallus R.M. Schust. and R. perssonii Sultan Khan. In the remaining members of Riccia, spores are separated and no tetrads are observed at maturity. The other species R. pathankotensis is an endemic liverwort native to the Indian subcontinent which was first collected and described by Kashyap in 1916 from Pathankot of Punjab, India. Its distribution is restricted only to India, Nepal, and Pakistan. In India, it is known to occur in three regionsdWH, P & WR and WG (Dash and Saxena 2009; Gaur and Nautiyal 1981;
https://doi.org/10.1016/j.japb.2019.08.005 pISSN2287-884X eISSN2287-9544/Ó 2019 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing Services by Elsevier. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
Please cite this article as: Premita Devi N et al., Two new records of Riccia L. (Ricciaceae) for the Eastern Himalaya Bryogeographical Region of India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2019.08.005
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N Premita Devi et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx
Figure 1. Map of India showing the EH bryogeographical region and map of Manipur state showing the location of Kakching District where Riccia curtisii (Austin) Austin and Riccia pathankotensis Kashyap were recorded. EH, Eastern Himalaya.
Kashyap 1929; Singh and Singh 2008; Srivastava 1964; Pradhan and Joshi 2009). Morphological descriptions and microphotographs of the two newly recorded species of Riccia are provided in detail. A key to species of the nine Riccia species found in Manipur is also highlighted. Material and methods On repeated field expeditions carried out from June 2016 to November 2018 across the central valley districts of Manipur, the
authors came across these two striking Riccia species, viz. R. curtisii and R. pathankotensis, growing on the moist alluvial soil deposited on the bank of Sekmai River that headed North to South in the Kakching District (Figure 1). The plants were collected from three locations namely, Kerjing, Wairi and Keirak. Specimens were carefully studied while afresh. Record-keeping, herbarium preparation, and wet preservation were carried out following the methodology given by Vanderpoorten et al (2010). The plants were identified by consulting relevant literature and publications (Austin 1869, 1879; Haynes 1920; McAllister 1916; Singh and Singh 2009).
Table 1. Comparison of Riccia curtisii and Riccia personii. Characters
R. curtisii
R. personii
Nature of thalli Thallus lobes Epidermal cell Spore diameter Band joining adhered tetrads Spine of the exine
Bright green, more spongy, dorsal surface highly lacunose Bilobed to palmate (5e71.5e2.5 mm) 80e130 60e80 mm 70e82 mm Narrow 3e6 mm long
Light green, less spongy, dorsal surface not lacunose Oblong to obovate (3e51e2 mm) 61e101 61e91 mm 88e99 mm Broad 6e9 mm long
Table 2. Comparison of Riccia pathankotensis and Riccia beyrichiana. Characters
R. pathankotensis
R. beyrichiana
Nature of thalli
Dusty green, occur singly forming complete rosette of smaller size (20 mm)
Thallus lobe Epidermal cells Number of capsules per lobe Spore diameter Spore reticulation
Linear to oblong, margin ciliated Linear to oblong More than 10, in 2e3 rows 75e98 mm Perfect, 6e8 reticulations across on distal face
Green, aggregated together forming incomplete larger rosettes (30 mm) Linear, margin entire Linear Less than 5, in 1e2 rows but never in 3 rows 80e93 mm Imperfect, 7e9 reticulations across on distal face
Please cite this article as: Premita Devi N et al., Two new records of Riccia L. (Ricciaceae) for the Eastern Himalaya Bryogeographical Region of India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2019.08.005
N Premita Devi et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx
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Figure 2. Microphotographs of Riccia curtisii (Austin) Austin.: A, habit; B, Transverse Section of sporophytic thallus; C, rhizoids; D, papillae in epidermis; E, lacunae on the dorsal surface of thallus; F, spore tetrads inside sporogonium; G, narrow band in isobilateral tetrad; H, tetrahedral tetrad showing spinous ornamentation.
Microphotographs were taken by using a phase-contrast microscope MOTIC BA210 (Motic Asia, Hong Kong). The herbarium packets and wet specimens were deposited in the Manipur University Museum of Plants (MUMP), Department of Life Sciences, Manipur University, Canchipur. For further scientific validation, the characters observed were compared with those of the allied taxa available in the literature (Tables 1 and 2). Accepted names of the species are given accordingly at http://www.theplantlist.org and http://www.tropicos.org. Systematic accounts Riccia curtisii (Austin) Austin Bull. Torrey Bot. Club 6(52): 305. 1879 (Austin 1879). Thalli bright green to yellowish green, spongy, medium-sized complete or incomplete rosettes, 10e16 mm across,
dichotomously branched, 1e2 furcate; lobes 5e7 mm long, 1.5e2.5 mm wide, bilobed to palmate, overlapping, dorsal surface highly lacunose, median groove inconspicuous, sulcus visible only at apex, margin crenate, apex truncate; ventral scales absent; rhizoids pegged and few smooth; cross section of the thallus three times as broad as high, differentiated into the upper photosynthetic zone with large air cavities and lower compact parenchymatous storage zone (Ricciella-type); epidermal cells globose to ovate, 80e130 60e80 mm, papillose. Dioecious. Female thalli larger, overcrowded above male; sporogonia few, 3e5 per thallus lobe, ventrally scattered in 1e 2 rows; spores permanently adhered in tetrads, golden brown, tetrads 70e82 mm in diameter, single spore in tetragonal tetrads 40e42 mm across, spores in isobilateral tetrads joined together by narrow band of 2.5e2.7 mm wide, exine basilar reticulations with dense slender spines, spines 3e6 mm long (Figure 2).
Please cite this article as: Premita Devi N et al., Two new records of Riccia L. (Ricciaceae) for the Eastern Himalaya Bryogeographical Region of India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2019.08.005
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Figure 3. Microphotographs of Riccia pathankotensis Kashyap.: A, habit; B, Transverse Section of sporophytic thallus; C, arrangement of sporogonia in three rows; D, spores showing perfect reticulation in both proximal and distal face.
Specimens examined. INDIA: Manipur, District Kakching, Wairi, 766 msl, 24 30’29" N & 93 58’25" E, on shaded deposited alluvial clayed soil, 26.01.2017, Naorem Premita Devi, 003430 (MUMP); INDIA: Manipur, District Kakching, Keirak, 761 msl, 24 30’50" N & 93 57’45" E, 06.02.2017, Naorem Premita Devi, 003441 (MUMP). Habitat & Ecology. Terrestrial. It prefers to grow in isolation mostly in the shaded portion of small sinuous curves or loops formed by the clay/sand deposition. Around 2e3 thalli are seen growing in an area of 1 square foot. In some cases, it is found on exposed soil in association with hornworts such as Anthoceros, mosses, and other Riccia species. Fertile period. December to February Distribution. India (GP: Uttar Pradesh; WH: Uttarakhand; EH: Manipur), Africa, France, North America, and South America. Remarks. Antheridium and archegonium were not observed in the collected material. R. curtisii shows striking resemblance with the young developing single rosette thallus of Anthoceros L. if overlooked because of its bright green and sponginess dorsal surface. However, it can be differentiated through keen observation as the thallus of hornworts is greasy in appearance and lacks sporophytic capsules. Riccia pathankotensis Kashyap J. Bombay Nat. Hist. Soc. 24: 48e349. 1916. Thalli green to dusty green, medium-sized complete rosette, generally occur singly, 10e20 mm across, dichotomously branch,
1e2 furcate; lobes 5e10 mm long, 2 mm wide, linear to oblong, nonoverlapping, dorsal groove inconspicuous at posterior region, broad and shallow at middle, narrow and deep forming acute at anterior end, margin thick and convex, apex obtuse or truncate; ventral scales small, hyaline, not projected beyond margin; rhizoids smooth and pegged; cross section of the thallus thrice as broad as high, upper surface concavoconvex to planoconvex, nondifferentiation of the upper photosynthetic zone with large air chambers and lower parenchymatous storage zone (Euriccia-type), epidermis well-defined, subglobose. Monoecious. Sporogonia numerous, 6e10 per thallus lobe, ventrally aggregated in 2e3 rows with little vegetative tissue in between, visible on dorsal surface; spores dark brown, tetrahedral, globose, 75e98 mm in diameter, sporoderm reticulate, reticulations 6e8 across on distal face and perfect (Figure 3). Specimens examined. INDIA: Manipur, District Kakching, Wairi, 767 msl, 24 30’29" N & 93 58’25" E, on exposed deposited alluvial clayed soil, 27.11.2016, Naorem Premita Devi, 003416 (MUMP); INDIA: Manipur, District Kakching, Kerjing, 791 msl, 24 27’05" N & 94 01’23" E, 03.02.2017, Naorem Premita Devi, 003438 (MUMP). Habitat & Ecology. Terrestrial, often on exposed semishaded to shaded moist alluvial clayed soil in association with other bryophytes. Fertile period. November to January Distribution. India (WH: Himachal Pradesh, Uttarakhand; P & WR: Punjab; WG: Orissa; EH: Manipur), Nepal, and Pakistan.
Please cite this article as: Premita Devi N et al., Two new records of Riccia L. (Ricciaceae) for the Eastern Himalaya Bryogeographical Region of India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2019.08.005
N Premita Devi et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx
Key to the species of Riccia found in Manipur 1a. Internal thallus with upper photosynthetic zone containing large air chambers (Ricciellaetype), monoecious or dioecious.......................... 2 1b. Internal thallus with upper compact photosynthetic zone having slit like pores (Euricciaetype), monoecious ............................ 6 2a. Thallus nonespongy, loose semierosette, dichotomy wide, upto 3 times furcate, lobes ribbonelike, capsules in 1 row, spores golden yellow ...............R. huebeneriana 2b. Thallus spongy, complete or incomplete rosette, dichotomy narrow, 1e2 furcate, thallus lobes not ribbonelike, capsules in 1e3 rows ......................... 3 3a. Permanently adhered isobilateral spore tetrads .......................... R. curtisii 3b. Mature spores free from tetrads .............................. 4 4a. Plants dioecious, lobes very small, 1 mm wide, spores lamellate, female thalli larger, male thalli with pinkish pigmentation ......................... R. frostii 4b. Plants monoecious, lobes broad, upto 6 mm wide, spores reticulate ........................... 5 5a. Thallus cruciate, lobes 5e6 mm wide, ventral scales present, capsules in 1e2 rows, spores golden brown with complete areolate reticulation ................. R. cruciata 3b. Thallus cavernous, lobes 2e4mm wide, ventral scales absent, capsules in 1e3 rows, spores reddish brown to black with incomplete areolate reticulation ......... R. cavernosa 6a. Thallus margin thick, convex and swollen, sulcus prominent at base ..........................7 6b. Thallus margin undulate, sulcus prominent towards apex ............................. 8 7a. Dichotomy 2e4 furcate, capsules few in 1 row per lobe, spore reticulations imperfect, exine strongly granuloseepapillate.................... R. beyrichiana 7b. Dichotomy 1e2 furcate, capsules numerous in 2/3 rows per lobe, spore reticulations perfect, exine weakly granulose ..................... R. pathankotensis 8a. Dichotomy 1e2 furcate, margin purple, sulcus prominent at tip, spores golden brown to black, spore diameter (99e120) mm ...................... R. billardierei 8b. Dichotomy 2e3 furcate, margin white, sulcus prominent upto middle, spores dark brown, spore diameter (62e83) mm ........................ R. sorocarpa
Conclusion With the present addition of two species from Manipur, the genus Riccia L. is represented by 18 species in the EH bryogeographical region. Among the states of EH, Manipur shows maximum species diversity with 9 species, followed by West Bengal with 8, Assam and Sikkim 6 each, Meghalaya and Mizoram 3 each, Nagaland 2, and no record so far from Arunachal Pradesh and Tripura. Geographically, above 95% of the total coverage area of North Eastern states is made of hilly tracts of ferruginous red soil, and as such, Riccia species which is suitably grown on alluvial soil is scarcely found. As Riccia show strong seasonality, the specimens studied were collected from the moist alluvial deposits of Sekmai
5
River during peak fertile season, i.e., November to March. The study provides the easternmost distributional limit of both the species in the country. The results are solely based on the preliminary collection of liverworts from the valley districts of Manipur. Perusal of literature from across the country and from the state revealed that little research is conducted so far on liverworts. It is a nondeniable fact in the field of plant sciences that liverworts are among the most neglected group of plants because of their miniature thallic body and difficult taxonomy. Neither the state is fully explored nor is critical examination of the genus conducted so far. A number of generic records and new species can be expected with further systematic exploration from this bryoflora-rich EH region. Conflict of interest The authors declare that there is no conflict of interest. Acknowledgment The present work was supported by CSIR, India bearing Grant No. 09/476/(0083)/2017-EMR-1. The authors are thankful to Dr S. K. Singh, Scientist D, Botanical Survey of India, for providing extremely useful comments and help in identification of the species. The authors also thank Laishram Premananda, Department of Geology, Assam University, Silchar, for the help in preparation of distributional map of Manipur. References Austin CF. 1869. Characters of Some New Hepaticæ (Mostly North American), Together with Notes on a Few Imperfectly Described Species. Proceedings of the Academy of Natural Sciences of Philadelphia 21:218e234. Austin CF. 1879. Notes on Hepaticology. Bulletin of the Torrey Botanical Club 6 (52): 301e306. Crandall-Stotler B, Stotler RE, Long DG. 2009. Phylogeny and Classification of the Marchantiophyta. Edinburgh Journal of Botany 66 (1):155e198. Dash PK, Saxena DK. 2009. Bryoflora of Khandadhar hill ranges, Orissa, India. Geobios 36. 713e116. Gaur RD, Nautiyal S. 1981. Thallose liverworts from Srinagar and adjacent areas. Indian Journal of Forestry 4:75e76. Haynes CC. 1920. Illustrations of Six Species of Riccia, with Original Descriptions. Bulletin of the Torrey Botanical Club 47 (7):279e287. Kashyap SR. 1929. Liverworts of the Western Himalayas and the Panjab Plain I. Lahore: The University of the Panjab. McAllister F. 1916. The Morphology of Thallocarpus Curtisii. Bulletin of the Torrey Botanical Club 43 (3):117e126. Pande SK, Ahmad S. 1944. Liverworts of Lucknow and its neighbourhood. In: Proceedings Of the 31st Indian Science Congress. p. 80. part 3. Pradhan N, Joshi SD. 2009. Liverworts and hornworts of Nepal: a synopsis. Botanica Orientalis 6:69e75. Singh DK, Singh SK. 2008. Diversity in Liverworts and Hornworts of Great Himalayan National Park, Western Himalaya, India. In: Mohamed H, Baki BB, NasrulhaqeBoyce A, et al., editors. Bryology in the New Millenium. Kaula Lumpur: University of Malaya. pp. 57e82. Singh DK, Singh SK. 2009. Hepaticae and Anthocerotae of Great Himalayan National Park and its environs (HP), India. Botanical Survey of India. pp. 398e400. Singh DK, Singh SK, Devendra S. 2016. Liverworts and Hornworts of India an annotated checklist. Kolkata: Botanical Survey of India. pp. 22e23. Singh SK. 2014. An appraisal of genus Riccia in India with a note on diversity and distribution of species. International Journal of Sustainable Water and Environmental Systems 5 (1):35e43. http://doi.org/10.5383/swes.06.01.0004. Söderström L, Hagborg A, Konrat MV, et al. 2016. World checklist of hornworts and liverworts. Phytokeys 59:1e828. http://doi.org/10.3897/phytokeys.59.6261. Srivastava KP. 1964. Bryophytes of India I. Ricciaceae. Bulletin of the National Botanical Gardens 104:1e103. Lucknow. Tewari SD, Pant G. 1983. Riccia curtisii (Aust.) James. from Kumaon Himalayas. Current Science 52 (4):164e165. Vanderpoorten A, Papp B, Gradstein R. 2010. Sampling of Bryophytes, vol. 8. Belgium: Abc taxa. pp. 331e345.
Please cite this article as: Premita Devi N et al., Two new records of Riccia L. (Ricciaceae) for the Eastern Himalaya Bryogeographical Region of India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2019.08.005
Contents lists available at ScienceDirect
Journal of Asia-Pacific Biodiversity journal homepage: http://www.elsevier.com/locate/japb
Original Article
Two new records of Riccia L. (Ricciaceae) for the Eastern Himalaya Bryogeographical Region of India Naorem Premita Devi, S.D. Yumkham*, P.K. Singh Ethnobotany & Plant Physiology Laboratory, Centre of Advanced Studies in Life Sciences, Manipur University, Canchipur, 795 003, India
a r t i c l e i n f o
a b s t r a c t
Article history: Received 12 January 2019 Received in revised form 2 July 2019 Accepted 20 August 2019 Available online xxx
The Eastern Himalaya Bryogeographical Region is recognized as one of the richest areas for bryoflora in India. It comprises the eight North Eastern States (Arunachal Pradesh, Assam, Manipur, Meghalaya, Mizoram, Nagaland, Sikkim, and Tripura), Darjeeling, and the Hills of the West Bengal. In the present article, the occurrence of Riccia curtisii (Austin) Austin and Riccia pathankotensis Kashyap of Ricciaceae is reported for the first time for the Eastern Himalaya region from the state Manipur. Detailed descriptions and photographs of the two newly recorded species of Riccia L. are provided to facilitate easy identification. A key to species of the nine Riccia species found in Manipur is also incorporated. Ó 2019 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing Services by Elsevier. This is an open access article under the CC BY-NC-ND license (http:// creativecommons.org/licenses/by-nc-nd/4.0/).
Keywords: Riccia New record Eastern Himalaya Manipur
Introduction Riccia L. (Ricciaceae) is the largest genus among the complex thalloid liverworts with 152 species worldwide and in India, with 36 species constituting 24% of the global species diversity (Söderström et al 2016; Singh 2014). The plants possess simple linear or oblong gametophytic thalli differentiated into upper photosynthetic zone and lower storage tissue, with or without simple pores similar to other thallose members. It is unique in having fertile plants with anacrogynous condition, sporophytic sporogonia or capsules embedded singly within the thalli, absence of pseudoperianth, and cleistocarpous capsules without elaters. It differs from Ricciocarpos Corda, the other twin genus of Ricciaceae in lacking idioblastic oil cells, which are present in the latter (Crandall-Stotler et al 2009). Majority are terrestrial mesophytes which preferably grow on exposed or shaded mixed alluvial sandy soil with few aquatic and semiaquatic forms. There are eight bryogeographical regions currently recognized in India to assess the phytodiversity of liverworts and hornworts. These include the Andaman & Nicobar Islands [A & N], Central India [CI], Deccan Plateau & Eastern Ghats [DP & EG], Eastern Himalaya [EH], Gangetic Plains [GP], Punjab & West Rajasthan [P & WR],
* Corresponding author. E-mail address: rifl[email protected] (S.D. Yumkham). Peer review under responsibility of National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA).
Western Ghats [WG], and Western Himalaya [WH]. The EH comprises the eight North Eastern States (Arunachal Pradesh, Assam, Manipur, Meghalaya, Mizoram, Nagaland, Sikkim, and Tripura), Darjeeling, and the Hills of the West Bengal (Singh et al 2016). Altogether, 16 species have been recognized so far from the EH region, namely, R. attenuata Pandé, R. beyrichiana Hampe ex Lehm., R. billardierei Mont. & Nees, R. cavernosa Hoffm., R. ciliata Hoffm., R. crispatula Mitt., R. cruciata Kashyap, R. discolor Lehm. & Lindenb., R. fluitans L., R. frostii Austin, R. gangetica Ahmad, R. glauca L., R. hasskarliana Steph., R. huebeneriana Lindenb., R. sorocarpa Bisch. and R. stricta (Gottsche, Lindenb. & Nees) Perold (Singh 2014). In the present article, two unique species of Riccia, namely, R. curtisii (Austin) Austin and R. pathankotensis Kashyap, are recorded for the first time for the EH region from the North Eastern State Manipur. R. curtisii is one of the very rare species in India and is reported from GP and WH regions (Pande and Ahmad 1944; Srivastava 1964; Tewari and Pant 1983). It is taxonomically very significant because of the presence of permanently adhered spore tetrads with long spines, the only characteristic feature found in three Riccia species, the other two being R. leptothallus R.M. Schust. and R. perssonii Sultan Khan. In the remaining members of Riccia, spores are separated and no tetrads are observed at maturity. The other species R. pathankotensis is an endemic liverwort native to the Indian subcontinent which was first collected and described by Kashyap in 1916 from Pathankot of Punjab, India. Its distribution is restricted only to India, Nepal, and Pakistan. In India, it is known to occur in three regionsdWH, P & WR and WG (Dash and Saxena 2009; Gaur and Nautiyal 1981;
https://doi.org/10.1016/j.japb.2019.08.005 pISSN2287-884X eISSN2287-9544/Ó 2019 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing Services by Elsevier. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
Please cite this article as: Premita Devi N et al., Two new records of Riccia L. (Ricciaceae) for the Eastern Himalaya Bryogeographical Region of India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2019.08.005
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N Premita Devi et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx
Figure 1. Map of India showing the EH bryogeographical region and map of Manipur state showing the location of Kakching District where Riccia curtisii (Austin) Austin and Riccia pathankotensis Kashyap were recorded. EH, Eastern Himalaya.
Kashyap 1929; Singh and Singh 2008; Srivastava 1964; Pradhan and Joshi 2009). Morphological descriptions and microphotographs of the two newly recorded species of Riccia are provided in detail. A key to species of the nine Riccia species found in Manipur is also highlighted. Material and methods On repeated field expeditions carried out from June 2016 to November 2018 across the central valley districts of Manipur, the
authors came across these two striking Riccia species, viz. R. curtisii and R. pathankotensis, growing on the moist alluvial soil deposited on the bank of Sekmai River that headed North to South in the Kakching District (Figure 1). The plants were collected from three locations namely, Kerjing, Wairi and Keirak. Specimens were carefully studied while afresh. Record-keeping, herbarium preparation, and wet preservation were carried out following the methodology given by Vanderpoorten et al (2010). The plants were identified by consulting relevant literature and publications (Austin 1869, 1879; Haynes 1920; McAllister 1916; Singh and Singh 2009).
Table 1. Comparison of Riccia curtisii and Riccia personii. Characters
R. curtisii
R. personii
Nature of thalli Thallus lobes Epidermal cell Spore diameter Band joining adhered tetrads Spine of the exine
Bright green, more spongy, dorsal surface highly lacunose Bilobed to palmate (5e71.5e2.5 mm) 80e130 60e80 mm 70e82 mm Narrow 3e6 mm long
Light green, less spongy, dorsal surface not lacunose Oblong to obovate (3e51e2 mm) 61e101 61e91 mm 88e99 mm Broad 6e9 mm long
Table 2. Comparison of Riccia pathankotensis and Riccia beyrichiana. Characters
R. pathankotensis
R. beyrichiana
Nature of thalli
Dusty green, occur singly forming complete rosette of smaller size (20 mm)
Thallus lobe Epidermal cells Number of capsules per lobe Spore diameter Spore reticulation
Linear to oblong, margin ciliated Linear to oblong More than 10, in 2e3 rows 75e98 mm Perfect, 6e8 reticulations across on distal face
Green, aggregated together forming incomplete larger rosettes (30 mm) Linear, margin entire Linear Less than 5, in 1e2 rows but never in 3 rows 80e93 mm Imperfect, 7e9 reticulations across on distal face
Please cite this article as: Premita Devi N et al., Two new records of Riccia L. (Ricciaceae) for the Eastern Himalaya Bryogeographical Region of India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2019.08.005
N Premita Devi et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx
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Figure 2. Microphotographs of Riccia curtisii (Austin) Austin.: A, habit; B, Transverse Section of sporophytic thallus; C, rhizoids; D, papillae in epidermis; E, lacunae on the dorsal surface of thallus; F, spore tetrads inside sporogonium; G, narrow band in isobilateral tetrad; H, tetrahedral tetrad showing spinous ornamentation.
Microphotographs were taken by using a phase-contrast microscope MOTIC BA210 (Motic Asia, Hong Kong). The herbarium packets and wet specimens were deposited in the Manipur University Museum of Plants (MUMP), Department of Life Sciences, Manipur University, Canchipur. For further scientific validation, the characters observed were compared with those of the allied taxa available in the literature (Tables 1 and 2). Accepted names of the species are given accordingly at http://www.theplantlist.org and http://www.tropicos.org. Systematic accounts Riccia curtisii (Austin) Austin Bull. Torrey Bot. Club 6(52): 305. 1879 (Austin 1879). Thalli bright green to yellowish green, spongy, medium-sized complete or incomplete rosettes, 10e16 mm across,
dichotomously branched, 1e2 furcate; lobes 5e7 mm long, 1.5e2.5 mm wide, bilobed to palmate, overlapping, dorsal surface highly lacunose, median groove inconspicuous, sulcus visible only at apex, margin crenate, apex truncate; ventral scales absent; rhizoids pegged and few smooth; cross section of the thallus three times as broad as high, differentiated into the upper photosynthetic zone with large air cavities and lower compact parenchymatous storage zone (Ricciella-type); epidermal cells globose to ovate, 80e130 60e80 mm, papillose. Dioecious. Female thalli larger, overcrowded above male; sporogonia few, 3e5 per thallus lobe, ventrally scattered in 1e 2 rows; spores permanently adhered in tetrads, golden brown, tetrads 70e82 mm in diameter, single spore in tetragonal tetrads 40e42 mm across, spores in isobilateral tetrads joined together by narrow band of 2.5e2.7 mm wide, exine basilar reticulations with dense slender spines, spines 3e6 mm long (Figure 2).
Please cite this article as: Premita Devi N et al., Two new records of Riccia L. (Ricciaceae) for the Eastern Himalaya Bryogeographical Region of India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2019.08.005
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Figure 3. Microphotographs of Riccia pathankotensis Kashyap.: A, habit; B, Transverse Section of sporophytic thallus; C, arrangement of sporogonia in three rows; D, spores showing perfect reticulation in both proximal and distal face.
Specimens examined. INDIA: Manipur, District Kakching, Wairi, 766 msl, 24 30’29" N & 93 58’25" E, on shaded deposited alluvial clayed soil, 26.01.2017, Naorem Premita Devi, 003430 (MUMP); INDIA: Manipur, District Kakching, Keirak, 761 msl, 24 30’50" N & 93 57’45" E, 06.02.2017, Naorem Premita Devi, 003441 (MUMP). Habitat & Ecology. Terrestrial. It prefers to grow in isolation mostly in the shaded portion of small sinuous curves or loops formed by the clay/sand deposition. Around 2e3 thalli are seen growing in an area of 1 square foot. In some cases, it is found on exposed soil in association with hornworts such as Anthoceros, mosses, and other Riccia species. Fertile period. December to February Distribution. India (GP: Uttar Pradesh; WH: Uttarakhand; EH: Manipur), Africa, France, North America, and South America. Remarks. Antheridium and archegonium were not observed in the collected material. R. curtisii shows striking resemblance with the young developing single rosette thallus of Anthoceros L. if overlooked because of its bright green and sponginess dorsal surface. However, it can be differentiated through keen observation as the thallus of hornworts is greasy in appearance and lacks sporophytic capsules. Riccia pathankotensis Kashyap J. Bombay Nat. Hist. Soc. 24: 48e349. 1916. Thalli green to dusty green, medium-sized complete rosette, generally occur singly, 10e20 mm across, dichotomously branch,
1e2 furcate; lobes 5e10 mm long, 2 mm wide, linear to oblong, nonoverlapping, dorsal groove inconspicuous at posterior region, broad and shallow at middle, narrow and deep forming acute at anterior end, margin thick and convex, apex obtuse or truncate; ventral scales small, hyaline, not projected beyond margin; rhizoids smooth and pegged; cross section of the thallus thrice as broad as high, upper surface concavoconvex to planoconvex, nondifferentiation of the upper photosynthetic zone with large air chambers and lower parenchymatous storage zone (Euriccia-type), epidermis well-defined, subglobose. Monoecious. Sporogonia numerous, 6e10 per thallus lobe, ventrally aggregated in 2e3 rows with little vegetative tissue in between, visible on dorsal surface; spores dark brown, tetrahedral, globose, 75e98 mm in diameter, sporoderm reticulate, reticulations 6e8 across on distal face and perfect (Figure 3). Specimens examined. INDIA: Manipur, District Kakching, Wairi, 767 msl, 24 30’29" N & 93 58’25" E, on exposed deposited alluvial clayed soil, 27.11.2016, Naorem Premita Devi, 003416 (MUMP); INDIA: Manipur, District Kakching, Kerjing, 791 msl, 24 27’05" N & 94 01’23" E, 03.02.2017, Naorem Premita Devi, 003438 (MUMP). Habitat & Ecology. Terrestrial, often on exposed semishaded to shaded moist alluvial clayed soil in association with other bryophytes. Fertile period. November to January Distribution. India (WH: Himachal Pradesh, Uttarakhand; P & WR: Punjab; WG: Orissa; EH: Manipur), Nepal, and Pakistan.
Please cite this article as: Premita Devi N et al., Two new records of Riccia L. (Ricciaceae) for the Eastern Himalaya Bryogeographical Region of India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2019.08.005
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Key to the species of Riccia found in Manipur 1a. Internal thallus with upper photosynthetic zone containing large air chambers (Ricciellaetype), monoecious or dioecious.......................... 2 1b. Internal thallus with upper compact photosynthetic zone having slit like pores (Euricciaetype), monoecious ............................ 6 2a. Thallus nonespongy, loose semierosette, dichotomy wide, upto 3 times furcate, lobes ribbonelike, capsules in 1 row, spores golden yellow ...............R. huebeneriana 2b. Thallus spongy, complete or incomplete rosette, dichotomy narrow, 1e2 furcate, thallus lobes not ribbonelike, capsules in 1e3 rows ......................... 3 3a. Permanently adhered isobilateral spore tetrads .......................... R. curtisii 3b. Mature spores free from tetrads .............................. 4 4a. Plants dioecious, lobes very small, 1 mm wide, spores lamellate, female thalli larger, male thalli with pinkish pigmentation ......................... R. frostii 4b. Plants monoecious, lobes broad, upto 6 mm wide, spores reticulate ........................... 5 5a. Thallus cruciate, lobes 5e6 mm wide, ventral scales present, capsules in 1e2 rows, spores golden brown with complete areolate reticulation ................. R. cruciata 3b. Thallus cavernous, lobes 2e4mm wide, ventral scales absent, capsules in 1e3 rows, spores reddish brown to black with incomplete areolate reticulation ......... R. cavernosa 6a. Thallus margin thick, convex and swollen, sulcus prominent at base ..........................7 6b. Thallus margin undulate, sulcus prominent towards apex ............................. 8 7a. Dichotomy 2e4 furcate, capsules few in 1 row per lobe, spore reticulations imperfect, exine strongly granuloseepapillate.................... R. beyrichiana 7b. Dichotomy 1e2 furcate, capsules numerous in 2/3 rows per lobe, spore reticulations perfect, exine weakly granulose ..................... R. pathankotensis 8a. Dichotomy 1e2 furcate, margin purple, sulcus prominent at tip, spores golden brown to black, spore diameter (99e120) mm ...................... R. billardierei 8b. Dichotomy 2e3 furcate, margin white, sulcus prominent upto middle, spores dark brown, spore diameter (62e83) mm ........................ R. sorocarpa
Conclusion With the present addition of two species from Manipur, the genus Riccia L. is represented by 18 species in the EH bryogeographical region. Among the states of EH, Manipur shows maximum species diversity with 9 species, followed by West Bengal with 8, Assam and Sikkim 6 each, Meghalaya and Mizoram 3 each, Nagaland 2, and no record so far from Arunachal Pradesh and Tripura. Geographically, above 95% of the total coverage area of North Eastern states is made of hilly tracts of ferruginous red soil, and as such, Riccia species which is suitably grown on alluvial soil is scarcely found. As Riccia show strong seasonality, the specimens studied were collected from the moist alluvial deposits of Sekmai
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River during peak fertile season, i.e., November to March. The study provides the easternmost distributional limit of both the species in the country. The results are solely based on the preliminary collection of liverworts from the valley districts of Manipur. Perusal of literature from across the country and from the state revealed that little research is conducted so far on liverworts. It is a nondeniable fact in the field of plant sciences that liverworts are among the most neglected group of plants because of their miniature thallic body and difficult taxonomy. Neither the state is fully explored nor is critical examination of the genus conducted so far. A number of generic records and new species can be expected with further systematic exploration from this bryoflora-rich EH region. Conflict of interest The authors declare that there is no conflict of interest. Acknowledgment The present work was supported by CSIR, India bearing Grant No. 09/476/(0083)/2017-EMR-1. The authors are thankful to Dr S. K. Singh, Scientist D, Botanical Survey of India, for providing extremely useful comments and help in identification of the species. The authors also thank Laishram Premananda, Department of Geology, Assam University, Silchar, for the help in preparation of distributional map of Manipur. References Austin CF. 1869. Characters of Some New Hepaticæ (Mostly North American), Together with Notes on a Few Imperfectly Described Species. Proceedings of the Academy of Natural Sciences of Philadelphia 21:218e234. Austin CF. 1879. Notes on Hepaticology. Bulletin of the Torrey Botanical Club 6 (52): 301e306. Crandall-Stotler B, Stotler RE, Long DG. 2009. Phylogeny and Classification of the Marchantiophyta. Edinburgh Journal of Botany 66 (1):155e198. Dash PK, Saxena DK. 2009. Bryoflora of Khandadhar hill ranges, Orissa, India. Geobios 36. 713e116. Gaur RD, Nautiyal S. 1981. Thallose liverworts from Srinagar and adjacent areas. Indian Journal of Forestry 4:75e76. Haynes CC. 1920. Illustrations of Six Species of Riccia, with Original Descriptions. Bulletin of the Torrey Botanical Club 47 (7):279e287. Kashyap SR. 1929. Liverworts of the Western Himalayas and the Panjab Plain I. Lahore: The University of the Panjab. McAllister F. 1916. The Morphology of Thallocarpus Curtisii. Bulletin of the Torrey Botanical Club 43 (3):117e126. Pande SK, Ahmad S. 1944. Liverworts of Lucknow and its neighbourhood. In: Proceedings Of the 31st Indian Science Congress. p. 80. part 3. Pradhan N, Joshi SD. 2009. Liverworts and hornworts of Nepal: a synopsis. Botanica Orientalis 6:69e75. Singh DK, Singh SK. 2008. Diversity in Liverworts and Hornworts of Great Himalayan National Park, Western Himalaya, India. In: Mohamed H, Baki BB, NasrulhaqeBoyce A, et al., editors. Bryology in the New Millenium. Kaula Lumpur: University of Malaya. pp. 57e82. Singh DK, Singh SK. 2009. Hepaticae and Anthocerotae of Great Himalayan National Park and its environs (HP), India. Botanical Survey of India. pp. 398e400. Singh DK, Singh SK, Devendra S. 2016. Liverworts and Hornworts of India an annotated checklist. Kolkata: Botanical Survey of India. pp. 22e23. Singh SK. 2014. An appraisal of genus Riccia in India with a note on diversity and distribution of species. International Journal of Sustainable Water and Environmental Systems 5 (1):35e43. http://doi.org/10.5383/swes.06.01.0004. Söderström L, Hagborg A, Konrat MV, et al. 2016. World checklist of hornworts and liverworts. Phytokeys 59:1e828. http://doi.org/10.3897/phytokeys.59.6261. Srivastava KP. 1964. Bryophytes of India I. Ricciaceae. Bulletin of the National Botanical Gardens 104:1e103. Lucknow. Tewari SD, Pant G. 1983. Riccia curtisii (Aust.) James. from Kumaon Himalayas. Current Science 52 (4):164e165. Vanderpoorten A, Papp B, Gradstein R. 2010. Sampling of Bryophytes, vol. 8. Belgium: Abc taxa. pp. 331e345.
Please cite this article as: Premita Devi N et al., Two new records of Riccia L. (Ricciaceae) for the Eastern Himalaya Bryogeographical Region of India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2019.08.005