THE PRESENT POSITION OF INDIAN HEPATICOLOGY WITH A NOTE ON THE HEPATIC VEGETATION OF THE COUNTRY I

S. K. PANDE* AND D. c. BHARDWAJ"

INTRODUCTION DISTRIBUTION OF LIVERWORTS IN N 1936 the earlier researches in Indian hepatics were reviewed by one of the The hepatic vegetation of this subcon tinent I authors (PANDE, 19"36) at the kind has not been, so far, adequately considered suggestion of the late Professor Birbal from the distributional point of view. This Sahni, F.R.S. Since then a number of is largely due to our lack of knowledge workers have, through their investigations of the bryology of the country. The Hima­ in the field of Indian bryology, contri­ layas have received a little more attention buted to our knowledge of these plants, so in this respect. The magnificent scenery that today, after a lapse of more than a and the varied fauna and flora of these lofty decade, it seems worth while once again to mountain ranges, their perplexing geology take a census of this work and see how far and rich mineral resources have attracted we have progressed. many students of natural sciences. Some India is a vast country. It extends from of them, with their hobby of plant collect­ the tropics in the south to the temperate ing, have contributed to the advancement regions in the north, and, including Pakistan, of our knowledge of this group; but it is covers an area of about 1,581,400 sq. miles, due chiefly to the interest taken by some of with territories presenting very varied geo­ the Superintendents of the Royal Botanical graphical and ecological conditions for plant Gardens, Calcutta, and the efforts of some growth. Liverworts, as a rule, grow in shady, of the clergymen, stationed at Kurseong and moist and cool situations. Such habitats elsewhere, that our knowledge of the liver­ occur in many parts of the country (MAP 1). worts of this most interesting part of the The study of liverworts in India has not world has made some progress. received as much attention as that of some More than 50 years ago Professor V. Schiff­ other groups. Admirable work has been ner (1899), in an article based on Durel's done in phanerogams and ferns and there collection of Bhutan liverworts, pointed exist a number of excellent treatises on these out that the hepatics of this area belong plants. to purely tropical forms, most of these Hooker and Thomson, as early as 1855, corresponding to the species of the East as a result of their exhaustive studies of the Indian peninsula and the Indian archi­ angiospermic vegetation of the coun try, pelago. He further drew attention to the recognized the affinities of the angiospermic resemblances between the liverworts of flora of the Western Himalayas with that of Bhutan and those of the Indo-Malayan Western Asia and Europe, and of the Eastern region. In later years Kashyap ( 1920-21 ) Himalayas with that of China and Japan. discussed critically the distribution of liver­ Hooker ( 1904), in his sketch of the flora of worts of the Western Himalayas and the British India, again drew attention to the Punjab plain. He made extensive tours in same fact, remarking that the Himalayan the Himalayas, including Tibet, to study flora is a mixed flora. The eastern region the hepatic flora; from these he arrived at is dominated by the plants from China and a number of interesting and noteworthy Malaya, •while the western abounds in conclusions with regard to the distribution European, African and Oriental plants. of liverworts in those areas. His views were

1. Contribution from the Botany Department, Lucknow University, New Series, No.1. * Reader in Botany, University of Lucknow. ** Birbal Sahni Institute of Palaeobotany, Lucknow. PANDE & BHARDWAJ - THE PRESENT POSITION OF INDIAN HEPATICOLOGY 369

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MAP 1 later confirmed by Pande ( 1936) from his laboratory and a detailed discussion of the study of the hepatic flora of certain parts of various aspects of the flora and ecology of the Kumaon hills. Indian hepatics will be presented when During the past several years one of us these investigations have been completed. ( Pande) has made a number of tours in Meanwhile, a few preliminary remarks, based the Himalayas. and Ceylon to on the data already known and the obser­ collect material and observe the mode of vations made in the course of these travels, growth, habitat, etc., of these plants (see supplemented from a study of the specimens MAP 2). It has also been possible for us to so far examined, will be presented in this study several other collections. i.e. the rich paper. collection of the Eastern Himalayan hepatics made by Decoly and Schaul during 1897-99 PROGRESS OF RESEARCHES DURING from the neighbourhood of Kurseong (alti­ THE PERIOD 1937-1950 tude 2,000-8,000 ft.) in Darjeeling district; Pfleiderer's collection of the South Indian A. Systematics and Taxonomy - A perusal hepatics ( 1912-14), made from of the literature published during the period and its vicinity in the and under review shows marked progress in the from the Nilgiris, and Fleischer's collection study of the hepatic vegetation of India. of Ceylon liverworts. A number of friends The total of known genera has increased and and colleagues have also very kindly placed several new or otherwise interesting species at our disposal their valuable collections of have been described. Valuable informa­ liverworts from various parts of India. All tion on our hepatics has also appeared in these specimens are being studied in our some of the contemporary bryological publi- 370 THE PALAEOBOTANIST

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MAP 2 - Kumaon Himalayas. Heavy lines indicate route of the journey. cations of other countries. Excellent descrip­ accurate information on various aspects of tions of some of our hepatics by eminent Indian bryology becomes available. bryologists like Verdoorn (1930,1934,1935), A. R. Chavan (1937) described a new Carl (1931), Evans (1934) and Hoffmann species of Cyathodium Kze., C. barodae ( 1935) can be found in them. Similar in­ Chavan and the same has been included by formation on some Indian species is avail­ Professor V. Schiffner (1938) in his excellent able in Fullford's monograph (1946) of the monograph on the genus, which also gives a genus Bazzania, Castle's account ( 1936) of critical account of all the other Indian Radula, and Eifrig's monograph (1936) of species, including a new species, C. acro­ the Indo-Malayan species of Taxileje2mea trichum Schffn., proposed by Schiffner for a

( Spruce ~ Schffn. specimen collected by Decoly and Schaul A critical study of Indian species of some from Kurseong (Darjeeling, Eastern Hima­ of the genera has also been taken up, and layas) in 1899. a certain amount of progress made in this R. S. Chopra ( 1938), on the basis of an direction. However, a good deal of work examination of collections made by seve­ still needs to be done before adequate and ral persons from a number of localities in PANDE & BHARDWAJ - THE PRESENT POSITION OF INDIAN HEPATICOLOGY 371

Asp. !ergussoni, from the same area. A critical study of all the three spe­ cies, and a careful comparison with SI K K\M Asp. mangaloreus St., known from Mangalore in the Western Ghats, is ,..-._-_... highly desirable. 2 - ,j --, Pande and Srivastava ( 1<942)des­ cribed a new species o( NoweUia Mitt., N. indica Pande & Srivastava, from the Western Himalayas and a year later Pande and Misra ( 1943 ) published a new Leptocoelea Evans, L. himalayensis Pande & Misra, z:r!oo from the same area. The latter is a very variable species. The corti­ colous form, collected from near v Bageshwar (3,500 ft.), shows marked '?" differences, especially in the lobule, l'=~~ from the rupicolous form gathered ~ ,~...... from the surface of moist rocks in ,, ~ the neighbourhood of Loharkhet ,C.All / C)-tuNBA"TTY (6,000ft.). In another paper, dealing Q t ~\ RANG'TONG MllE.S \ with the epiphyllous hepatics of the country, Pande and Misra ( 1943 ) described two new species of liver­ MAP 3 - Heavy lines indicate route of the journey in worts, (I) Cololejeunea himalayensis Darjeeling region of the Eastern Himalayas. Pande & Misra and (2) Leptolejeunea himalayensis Pande & Misra, from Mungpoo (4,000 ft., Darjeeling, South India, published a list of the hepatics Eastern Himalayas) and added some details containing more than a hundred species, to the description of three other liverworts, including several new records and 3 new i.e. (3) Leptocolea lancilobaSt., (4) Rectolejeunea species. In this census 25 species are assign­ aloba ( Sande. Lac.) St. and (5) Cololejeunea ed to , 20 to Kodaikanal, 18 to hispidissima (St.), Herz. The last two Shambaganur, 14 to Madras, 11 to Maha­ of these were recorded in that paper for baleshwar, 8 to Negapatam and 7 to the Nil­ the first time from India, while for the giris. In a similar list of the liverworts of first one it was the first record from South the Eastern Himalayas and Bengal, Chopra India. In another paper on this interesting ( 1938) gives about 130 species, including group of liverworts Pande and Ahmad again 3 new species and a number of new ( 1942) reported four more epiphyllous records. The majority of the species are species, i.e. (I) Taeniolejeunea parafJinis from Darjeeling while the remaining come (Schffn.) Zw., (2) Radula protensa Ldbg., from other localities, e.g. Kurseong 13, (3) Leptolejeunea schiffneri St., from Gersoppa Phallot 11. Herzog (1939), in a communi­ Falls (Mysore State, South India) and cation dealing with two bryophytic collec­ (4) Leptocolea pseudofloccosa Horikawa from tions from the Sikkim Himalayas, lists more Pedro's Peak (Ceylon). Finally, Pande than 50 species of liverworts from this and Ahmad (1943) added eight more region, including more than a dozen new species to this list. These are: (I) M icro- species proposed by the author. The paper gives a detailed taxonomic account with 2. Proskauer (Bull. Torr. Bot. Club, 1951. p. 345) suggests that the genus Aspir011lilus St. should be illustrations of all the new species and some abolished. He lists Asp. dixitianus as Anthoceros valuable information about several others. dixitiamts com b. nov., remarking that A. fergusso11i T. S. Mahabale ( 1941 ) described a new Apte & Sane may probably be a synonym of the species of A spiromitus St.. Asp. dixitianus former. He further doubts the identity of this Mahabale, from soon liverwort with A. weistii }{hanna and of the latter and after with A. QI'gillaceous St. He refers Asp. khanda­ V. V. Apte and Sane (1942) added two lensis Apte & Sane as AnthocC1'OS Ilhandalcnsis comb. more new species, Asp. khandalensis and nov. 372 THE PALAEOBOTANIST

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MAP 4 - South India and Ceylon, indicating localities from which liverworts have been examined. lejellnea Sp., from Rimbick (Darjeeling, S. Ahmad ( 1942), from Lucknow, pub­ Eastern Himalayas). (2) Drepanalejeunea/ol­ lished a brief note on three new Indian licola Horikawa, (3) Aphanolejeunea sp. 1., species of Riccia. One of these, R. orien­ (4) Aphanolejeunea sp. II from ]orpokhari, talis Ahmad, a ciliate species, is based on a Darjeeling, Eastern Himalayas (the last­ specimen collected by Pande from the named species occurs also at Rimbick), Western Himalayas, in 1936; the second one, (5) Frullania sp., (6) Microlejeunea sp. II, R. gangetica Ahmad, is a monoecious species, (7) Lejeunea sp. and (8) Drepanolejeunea sp. growing during rains in the neighbourhood from Pedro's Peak (Ceylon). The total of Lucknow, often associated with R. discolor number of species of this interesting group L. & L. and R. melanospora Kash., and the of hepatics dealt with in these three papers third one, R. mangalorica Ahmad, was pro­ is 18. posed for a. plant collected by. A. R. Rao Hameed ( 1942 ), in a note on Iiverworts from Mangalore (South India). In sug­ of the Murree hills, records 24 species from gesting R. mangalorica as a new species, this region. Ahmad drew attention to the close similarity PANDE & BHARDWAJ - THE PRESENT POSITION OF INDIAN HEPATICOLOGY 373

of its spore with that of R. plana Tay., and and hills, states that these regions pointed out that it maybe a variety of the have favourable climatic and edaphic factors same. for a rich hepatic flora; but due to a long Sane ( 1942 ) described two new species of dry period, extending to about eight months A nthoceros L., A. dixiti from and in the year, only such species flourish as can A. sahyadl'ensis from Poona, recognizing its greatly tolerate xerophytic conditions and varieties, A. sahyadrensis var. purandhar can mature quickly. The hepatic flora of and A. sahyadrensis val'. poona. these places, in his opinion, conforms to Pande and Ahmad ( 1944), in a note on the "xerogeophytes" type of Gam and Riccia discolor L. & L., reduced Riccia hirna­ Verdoorn. layensis St. to a synonym of Riccia discolor D. C. Bhardwaj (1948) published an L. & L. In another communication the account of a new species of Aspiromitus St., same year Pande and Ahmad listed from Asp. mamillispora Bhardwaj, a dioecious Lucknow and its neighbourhood 14 species species with very characteristic mamillate of liverworts, representing six genera. spore, collected by Pande from Kandy In 1943 A. K. Ghose reported a species ( Ceylon) in January 1940. The paper in­ of Pallavicinia from Dacca in East Bengal. cludes a comparative table noting the diag­ Chavan and Mahabale ( 1945), in a note nostic features of all the known dioecious dealing with the distribution of hepatics in species of Aspiromitus. Gujrat, recognized three well-marked regions This species, as pointed out by Dr. Pros­ in this area. For the dry and arid region, kauer in a letter to one of the authors, is extending from Mt. Abu to Khed Bramha, apparently identical with the Andaman Vireshwar and Taranga hills, they list such specimen (co-type) of A. spinisporous St., xerophytic forms as (I) Plagiochasrna appen­ of which we had an opportunity to examine diculaturn L. & L., (2) Riccia discolor L. & L. a specimen and a traced copy of the figures and (3) Asterella (Firnbriaria) angusta St.; of the spore and elateI', kindly supplied to for the humid regions of Pavagarh and us by Dr. Proskauer. The plant from Rajpipla hills they record: (4) Conocephalurn Ceylon, however, does not fit in well with (L.) Necker, (5) Anthoceros hirnalayensis Stephani's description and figures of the Kash., (6) Cyathodium barodae Chavan and spore. The type specimen of Stephani's (7) Notothylas levieri Schffn.; and for the species comes from New Guinea and with­ southem and western plains on the coastal out reference to this it is not possible for the line, an admixture of the species from the authors to settle this issue finally. Should above two regions, e.g. Ricciadiscolor L. & L., the two be identical, A. mamillispora Bhar­ Cyathodium barodae Chavan, Riccia frostii dwaj will be reduced to a synonym of A. Aust. (noted as R. sanguinea Kash.) and spinisporous St. N otothylas levieri Schffn. Pande and Bhardwaj (1949), in a note R. L. Nirula (1945) described a new on some liverworts new to Indian flora, dealt species of Notothylas Sull., N. Chaltdhurii with three hepatics, i.e. (I) Anthoceros crispu­ Nirula, a non-columellate monoecious species lus3 (Mont.) Douin., previously known from from Nagpur. The capsules are bivalved Europe and North America and recorded by and each valve has a row of specially Pande and Ahmad in 1943 from Lucknow ; thickened cells along the margin. Nirula (2) Conocephallurn supradecompositum (Lindb.) holds Notothy/as as a reduced genus. St., a rather rare liverwort originally known Gupte ( 1945) described a yellow-spored from China and Japan and recently collected species of Notothylas from Poona, drawing by Maheshwari from Darjeeling in the Eastern attention to its close similarity with N. Himalayas; and (3) Aspiromitus harrisanus decurva Mit t. St., a Jamaican hepatic, previously known A. R. Rao ( 1947 ), in a note on the liver­ to us only through its Latin description by worts of Yercaud ( 5,200 ft.) in the Shevaroy Stephani (Sp. Hep., Vol. V, p. 965). Its hills of South India, recorded for this area: discovery near Munsyari (6,200 ft.) in the (I) Marchantia L., (2) Plagiochasma L. & L., Western Himalayas is of some interest. (3) Lejeunea Lib., (4) Plagiochila Dum., Pande, Bhardwaj and Ram Udal' ( 1949 ), (5) Fossombl'.onia Raddi, (6) Lepidozia Dum. in a note on South Indian hepatics, listed and P) Hygrolejeunea (Spr.) Schffn. P. D. Bhate (1947), in a note on a preli­ 3. According to Proskauer ( 1948 ), A. cJispulus minary survey of the liverworts of Lonavala is a synonym of A. punctatus L. 374 THE PALAEOBOTANIST ten members of the Acrogyneae; nine of khet ( 7,000 ft.) in the Western Himalayas. these, i.e. (1) Bazzania nudistipula (=Masti­ He observes tha t the species is not strictly di­ gobryum nudistipulum) St., (2) Saccogyna oecious, as has been previously recorded, but alternifolia St., (3) Plagiochila richteri St., may bear both antheridia and archegonia (4) P. wornoffi St., (5) Archilejeunea apiculi­ on the same thallus, sometimes even on the folia St., (6) Cheilolejeunea indica St., (7) Har­ same branch. Details of the development of palejeunea indica St., (8) E ulejeunea lindbergii the sexual organs and the sporophyte are St. and (9) Frullan'ia moniliata (R. Bl. Nees) also given. This follows the same course Dum. from Kudremukh (6,212 ft.), Western as in other species of the genus. The Ghats and (10) Ptychocoleus fer/ilis Trev. elaterophore is well developed, extending from Mangalore. to about one-third the length of the capsule, Pande and Ram Udar (1950), in the and there are a few short elaters attached to second paper of the same series, listed seven it. It breaks up into four pieces when the more acrogynous liverworts adding some capsule dehisces and the pieces of the ela­ valuable information about some of these: terophore, with the elaters sticking to them, (1) Radula compan'igera Mont., (2) Ptychan­ remain attached to the valves of the capsule. thus str'iatus Nees, (3) Thysananthus semire­ Bicellular endogenous gemmae are frequently pandus ( Nees) Ver. ( = Thysananthus rotun­ formed. distipulus St.), (4) Taxilejeunea tenerima St., Pande and Misra ( 1937), in a communi­ (5) Herberta pinnata St. All the liverworts cation on Sewardiella tuberifera Kash., an described in these two papers have been al­ interesting monotypic West Himalayan ready determined by Stephani. The 17 liver­ member of the Codoniaceae, have described worts enumerated in the last two papers are some details of development of the sexual out of Pfleiderer's rich collection of the South organs and the sporophyte of this liverwort Indian hepatics, running to some 124 species, and have discussed the systematic position distributed over 44 genera representing of the genus. 17 families. The spec'imens were collected Chavan ( 1937) described the life-history during the years 1911-13 from Udipi, Manga­ of Cyathodium barodae Chavan, collected lore, Bajape and Karkal, in the coastal strip from Baroda. He notes that the species is of the Western Ghats; Doddabetta and monoecious and the thallus bears only Kotagiri in the Nilgiris; Coorg (Mercara), smooth-walled rhizoids and a variable type north of the Nilgiris, and Kudremukh of ventral scales. The male receptacle is ( 6,212 ft.) in the Western Ghats. produced from a single cell and is almost Recently Schifiner and Pande (1950), sessile, and occupies a variable position. in a paper on the East Himalayan hepatics, The details of the antheridial development, listed 138 species of liverworts belonging to he states, are of the usual type found in 45 genera representing 20 families spread related forms but the septa separating the over various groups. These specimens were antheridia are persistent. The archegonial collected by Decoly and Schaul (1897-99) receptacle arises on the upper surface near from a number of localities, varying in the thallus margin and each receptacle may altitude from 2,000 to 8,000 ft. in the neigh­ produce up to seven archegonia. The in­ bourhood of Kurseong in the Darjeeling dis­ volucre is bi-lipped and develops only after trict of the Eastern Himalayas. The paper the full number of archegonia have been includes several new species and many new formed. The embryo may be of the fila­ records for the province and the country. mentous or of the octant type and the The discovery of M egaceros Campb., repre­ structure of the capsule is of the type senting the first record of the genus from characteristic of the genus. The haploid India, is of special interest. This is a very number of chromosomes is 8, each of which exhaustive treatment of the East Himalayan has a characteristic form. hepatics. P. N. Mehra (1938) has studied the B. Morphology and Cytology - Although chromosome number of three Indian genera only a few species of Indian liverworts have of the Codoniaceae, i.e. Fossombronia ht"ma­ been studied in detail, the investigations have layensis Kash., Petalophyllum indicum Kash. yielded interesting and important results. and Sewardiella tuberifera Kash. He gives Pande ( 1937) communicated a paper on 18 as the 2X number for these and states the morphology of Riccardia levieri Schfin., that the morphology of the chromosomes collected from the side of a stream in Rani- of all the three is different. A table show- PANDE & BHARDWAJ - THE PRESENT POSITION OF INDIAN HEPATICOLOGY 375

ing the chromosome number of the various The sporophyte is rather simple. The members of the Codoniaceae is also given. elater-mother-cells belong to 3-5 generations Mehra gives 9 as the basic chromosome later than the spore-mother-cells. Like number of the family and states that, in the Kashyap, the authors assign Stephensoniella evolution of the species in this family, a place near Exormotheca, pointing out polyploidy is rather rare. that it has a comparatively simpler structure. S. Ahmad ( 1938) studied the life-history Arguments are given in support of'the view of Aitcht"soniella himalayeusis Kash., a mono­ that Stephensoniella has been derived from typic West Himalayan liverwort. He des­ forms like Exormotheca by a process of cribes the detailed structure of the gameto­ reduction. They also record certain round phyte, noting that the thallus grows by bodies or " spherules" in the fertilized egg means of an apical cell cutting off segments and the embryo. The haploid number of on four sides. The air chambers originate chromosomes is given as 9. Adventitious schizogenously and are empty. The anthe­ ventral shoots or tubers serving as vegetative ridia develop in two rows and lie embedded reproductive bodies have also been described. in the thallus behind the female receptacle, Tubers of the male plants form antheridia and the latter may be terminal, lateral or which, under unfavourable conditions, be­ may lie in the fork between the two lobes. come arrested in different stages of their Ordinarily the receptacle has one or two development, completing the process in the involucres; occasionally it has three or four. next vegetative season. The tissue of the receptacle has empty air K. S. Srinivasan (1939) described the chambers, opening to the exterior by simple development of androgynous receptacles in pores. The embryo is of the filamentous M archantia palmata Nees, from Ootacamund type. Several stages in the development of in the Nilgiris. The narrow, elongated the sporophyte are described. The classi­ antheridia-bearing lobes, which, according fication of the Marchantiales is discussed and, to Srinivasan, are ventral proliferations of in the light of his observations, the author the archegonia-bearing lobes of the purely concludes that Exormotheca can no longer female receptacles, produce in the beginning be held as the lowest type of the Compositae sexual structures of the intermediate type since A itcht"soniella breaks down the line but later give rise to typical antheridia, of demarcation between the Compositae and suggesting the homologous nature of the the Targioniaceae. The view that the two sex organs. A fungus, forming pycnidia, families should be merged into one, earlier associated with the liverwort, is also re­ proposed by Kashyap, is supported by ported. In another paper Srinivasan ( 1944 ) Ahmad. gave a detailed account of the life-history of P. N. Mehra and H. L. Mehra (1938) Marchantia palmata. The minute structure published an accoun t of the life-history of of the thallus and the differentiation of the Stephensoniella brevipedunculata Kash., an­ various tissues as well as the behaviour of other interesting monotypic genus of the the apical cell are fully described. The Marchantiales, confined to the Western details of the development of the receptacles Himalayas, i.e. Kulu, Simla, Mussoorie ( alti­ are also given. Once again he describes tUde6,000-7,000ft.). Theauthorsdescribe the androgynous receptacles of the previous various structural and developmental details type and notes that" the female receptacles of both the gametophyte and sporophyte. always end as androgynous receptacles in They ascribe the growth of the thallus to an their final stages" (SRINIVASAN, 1944, p. apical cell and state that the air chambers Ill). The details of the development of originate schizogenously and are in a row. the sex organs, the author states, are of the These are large, empty and directed obliquely usual type found among the liverworts. forward, opening by a simple pore. The The archegonium develops in two ways: antheridial receptacle is of the simplest type. "One of these is, more or less, similar to The female receptacle is tenninal and never that recorded by Humphrey (1906) in strictiy dorsal. It has a short stalk and Fossombronia longiseta and by Campbell two involucres, each of which may have 1-4 ( 1896) in Geothallus tuberosus. The second archegonia. The tissue and pores of the type shows a certain amount of similarity receptacle are of the same type as those to that seen in Pallavicinia leylii (Haupt, of the thallus, and its stalk has a shallow 1918) and Preissia quadrata, as figured by groove with a few tuberculate rhizoids. Haupt (Haupt, 1926, Figs. 34 & 35 )." 376 THE PALAEOBOTANIST

Various details of the sporophyte devel­ not, they could not determine. According opment have also been described. Details to them there is not much difference in the of spermatogenesis and sporogenesis are also genomic constitution of the two species. given. The haploid number of chromosomes They also report some cases of hermaphro­ for this species is given as 9 and it is stated dite thalli in R. sanguinea Kash. that the chromosome complement in the Mahabale (1942) briefly reviewed the first heterotypic division includes a pair of data available on the cytology of liverworts small chromosomes which, though not dif­ and discussed the bearing of the cytological fering in size, show a certain resemblance studies on the phylogeny of the Marchan­ to X and Y chromosomes of A1archantia tiales. He analysed the data and, from an polymorpha, in their behaviour at the ana­ examination of the published results on the phase of the first division. chromosome numbers of various liverworts, In another paper Srinivasan (1940) concludes that the basic number of chromo­ described the morphology, life-history and somes (gametophytic number, i.e. n num­ cytology of Riccia himalayensis St. The ber) for the majority of the genera of the account given is that of a monoecious Riccia Marchan tiaceae and the ] ungermanniaceae from Madras (see P AN DE & AHMAD, is 9 and for the Ricciaceae 8. According to 1943 ). Minute details of the structure him the available cytological data can be and development of the thallus, the origin interpreted in support of both the progression of air chambers and the development as well as the reduction hypotheses of evo­ of the sexual organs are given. Some lution in the Marchantiales. He postulates instances of the presence of two archegonia that the pro-Marchantialean complex had inside an archegonial chamber are also 17 chromosomes and from this the Marchan­ recorded. Various stages in the embryo­ tiaceae and the Ricciceae originated inde­ geny and development of the sporophyte pendently and polyphyletically, and the are described. Some sterile spore-mother­ ancestral complex has been divided into cells have also been reported. In spore two series - one with 8 and the other with germination, according to Srinivasan, the 9 chromosomes. He supports such an germ tube emerges through a germ pore on assumption by a parallel case found among the convex wall. Perennation by means the Rosaceae, in which the presence of two of tubers is also recorded. The haploid basic numbers has been accounted for in the number of chromosomes is stated to be 8, same manner. of which 7 are long while 1 is very small and T. S. Mahabale and P. D. Bhate ( 1945 ) round. The last one shows precession on published an account of the structure and the spindle at the anaphase. life-history of Fimbriaria angusta St: In this Mahabale and Gorji (1941) studied the paper the authors have reviewed the work chromosomes of Riccia himalayensis St. done on the genus and have given details of ( a synonym of R. discolor L. & L.), and the structure and developmen t of the thallus, record 8 as the haploid number of chromo­ sex organs and sporophyte. They state that somes and note that 7 of these are slender, the thallus grows in the usual Marchantia­ elongated and irregular in outline while the tious manner. The air chambers have no eighth is much smaller and dot-like. The assimilatorv filaments and the older thalli attachment of the 7 long ones is atelo­ contain endotrophic mycorrhiza. The devel­ mitic while that of the eighth is telomitic opment of the sex organs and embryogeny, and there is no heterochromosome in this according to them, conforms to the type species. In a subsequent paper Mahabale found in other species of the genus but the and Gorji ( 1947 ) dealt with the chromosomal embryo is of the filamentous type. They complex of two species of Riccia, R. discotor also state that the cells of the apical part of L. & L. and R. sangiunea Kash. (a synonym the capsule show radial thickenings but the of R. frostii Aust.). They state that the operculum is not so regular as in Plagio­ chromosomal complex in both the species chasma or Cryptomitrium. They record consists df 8 slender chromosomes (n num­ 9 chromosomes for this species. The authors ber), 7 of which are more or less V-shaped consider Fimbrian'a as a reduced genus and or bent and one is a small dot-shaped have adduced some evidence in support body, lying in the middle of the other 7. of this. I t is pointed out that F. augusta Whether this small dot-shaped chromosome is an ally of F. pilosum O. Kutz., F. gollani is to be identified as the sex chromosome or St. and F. mysorensis Rash. PANDE & BHARDWAJ - THE PRESENT POSITION OF INDIAN HEPATlCOLOGY 377

Abeywickrama ( 1945) described the struc­ sued by Pande and Bhardwaj. I n this con­ ture and life-history of R. crispatula Mitt., nection, Pande and Bhardwaj ( 1949) com­ gi ving various details of the sex organs and municated a paper on the morphology of sporophyte. According to him the species A nthoceros L. (Phaeoceros ), a yellow-spored is monoecious and not dioecious, as described Indian species, and Bhardwaj (1949) in­ by Stephani. He notes that favourable con­ vestigated the life-history of a black-spored ditions for vegetative growth retard the species, A n/hoceros crispulus (Mon q Douin. formation of the sex organs and the size of Various details of the structure of the thal­ the sporangium depends upon food supply. lus and sex organs and of the development The species closely resembles R. billardieri of the sporophyte are described. Bhardwaj M. & N. from Java. has investigated the life-history of two other Mahabale and Deshpande (1947) ha ve Indian species of Anthoceros (L.) Prosk., described the ]ife-history of a common coJlected by Pande. One of the specimens Indian species of Plagiochasma, P. arti­ was collected from near Munsyari (Garhwal) culatum Kash., giving the details of the in 1936 and the other from near Mane­ structure and development of the thallus, bhanjan, Darjeeling, Eastern Himalayas, in receptacles, sex organs and sporophyte. 1939. The internal structure of the thallus, they A critical study of the various Indian state, is of the normal type. Its charac­ species of Riccia is being pursued by J~am teristic articulated appearance is due to Udal' and, in a recent paper, he ( 1950) has the intermittent development of the thal­ communicated the results of his investiga­ lus lobes. The sessile, often forked, male tions of three common monsoon species of receptacles and their development, as well Lucknow. as the development of the antheridia, are P. N. Mehra ( 1950) has described some of the same type as in other species of the details of the embryogeny of Sewardiella genus; but the female receptacle, though tuberi/era Kash. terminal at first, becomes dorsal secondarily A monograph on the Indian species of due to the development of a pseudo-branch Asterella Pal. B. is under preparation by from below the growing point. It bears Pande and Sultan Ahmad. ordinarily four archegonia, one at each corner, and has air chambers, opening to the REMARKS ON THE HEPATIC outside through barrel-shaped pores. Cases VEGETATION OF INDIA of bisexual receptacles are also recorded. The gametophytic number of chromosomes As we have already stated, in the present is gi ven as 8 and the embryo is sta ted to be state of our knowledge of Indian liverworts, of the filamentous type. The authors regard it is not possible for us to say much about Plagiochasma articulaturn as a transitional the various aspects of the flora and form between a type like Reboulia Raddi ecology of these plants; but an analysis of and P. appendiculatum L. & L. Attention is the data, gathered in the course of our tours drawn to the similarity between P. arti­ (MAPS 2, 3 & 4) as well as the study of culatum and M assalongoa Nees. A scheme the coJlections from various parts of the suggesting the derivation of the genera of country, so far carried out by. ourselves and the Operculatae by a process of reduction by others, suggest certain noteworthy fea­ from the level of M archantia is gi ven. tures of the vegetation which are presented Plagiochasma is assigned the last position here. in this table. In the end the authors have About 550 species of liverworts, distribut­ discussed the importance of the chromosome ed as shown in Map 1 and Tal:)le I, are number in the phylogeny of the genus and known from India. The figures are, how­ suggest that either polyploidy or deletion of ever, only approximate, because so long as a chromosome from the ancestral complex a critical study of our hepatics is incomplete, seems to have taken place in the evolution a certain amount of duplication of some of the species of Plagiochasma. species is unavoidable. The Anthocerotaceae and the Ricciaceae In the Himalayas, South India and Cey­ have been under investigation at the Uni­ lon there are vast areas rich in hepatics, versity of Lucknow for some time past. A the greatest number of species occurring in critical study of the morphology of some territories with abundan t rain and high Indian genera of the former is being pur- humidity ( TABLE I). The Vindhya and the 378 THE PALAEOBOTANIST

TABLE I

LOCALlTY ALTliUDE R,\INFALL RELATIVE HV;.110ITY No. OF DURING SPECIES OF ]UNE-Sf:PTEMBER LIVERWORTS

I- Western Himalayas Simla 7.000'·8,000' 63" 36 Mussoorie 0,000'·8,000' 8060" ~Iorning89:0' 65 Evening 88 /0 II - The Eastern Himalayas Nepal ( Kathmandu) 5,000'·7,000' Over 100" 38 Darjeeling 7,000'·8,000' 126-42" j\Iorning 05% 1I0 Evening 90% Kurseong 3,500'·0,000' 150" 138 Khasi Hills 4,000' -5,000' 200"·300" 65

III - Plains or the Uttar Pradesh Lucknow 303' 40' II

IV - Madhya Bharat and Madhya Pradesh

Pachmarbi 3J 5i8' 70-6l" l\Iorning 85% 15 Evening 78% Mount Abu 5,645' 75' 8 Lonavala I,QOO' 170-08" 25

V- South India j\[ahabaleshwar 4,720' Over 250'" 40 Kudremukh 0,212' Over 200" 91 l\Iangalore Sea level 134" 10 Baogalore 3,248' 3408" Morning 86% 10 Evening 62% Madras Sea level Over 100" 20 Nilgiris 8,760' Over 200" 44 Palni Hills 8,841 OVf.r 200'" 20

VI - Ceylon Kandy 1,800' 80'9" ]8

TABLE II A

R£GION TOTAL No. ENDEMIC SPF, Sf'P. COMMON TO SF?, CO~MONTO SP?, COMMON TO SP?, CO~I~ON ~ r--~ r-----'-----, GENERA ~ EAST INDIES COMMON Genera Spp. TO South Ceylon China Japan TO \V. HIMALAY,\S India Gen~a Sp~ A U5TR,\L[A E. Himalayas 83 333 40 00 23 23 40 12 26 II

B REGION ,--A-,TOT.-\L No. ENDEMIC SPP. SPP. SPP. SP? COMMON TO SPP. COMMON SPP. GENERA CO~n.ION COMMON COMMON ~ E,\ST INDIES ~ TO COMMON Genera Spp. TO TO TO TO E, HI~('\LAY,\S'A'. HUIALAYAS CEYLON Genera Spp. China Japan E, AFRICA South India 64 225 60 30 23 20 53 7 23 17

C SPP. REGION TOTAL No. ENDEMIC SPP. SPP. COMMON TO SPP. COMMON TO SPP. ~ GENERA CO;\DIQN CO,\IMON ~ EUROPE ~ToEAST COMMOl'l TO TO SOUTH Genera Spp. E. HIMALAYAS INDII\ Genera Spp. Cbina Japan INDIE:S W. Himalayas 53 170 40 30 20 31 12 ] 3

Satpura mountains, the Pachmarhi plateau, vailing weather conditions are still less con­ , Parasnath and Mt. Abu genial, so that here only a few hepatics grow are comparatively less favourable for the in favourable localities. growth of these plants. In the plains, over The Eastern Himalayas represent, appa­ the greater part of the country, the pre- rently, the richest liverwort territory in India. PANDE & BHARDWAJ - THE PRESENT POSITION OF INDIAN HEPATICOLOGY 379

The number of species (330) reported from flora of this region and that of the East here is almost one and a half times that Indies and Japan (TABLE II C). reported from South India (225 species), The hepatic flora of the Western Himalayas nearly twice the number recorded for the is further characterized by the presence of Western Himalayas ( 170 species) and three a few endemic and monotypic genera, i.e. times of those found in Ceylon (110 (1) Aitchisoniella Kash., (2) Stephensoniella species ). Kash., (3) Sauchia Kash. and (4) Sewardiella In the Western Himalayas the most Kash. These genera have a very restricted luxuriant hepatic vegetation occurs between distribution. the altitudes of 6,000 and 8,000 ft. (see also KASHYAP, 1920 & 1921 ), but in the Eastern ACKNOWLEDGEMENTS Himalayas the richest region lies at a com­ paratively lower altitude. This contribution is a humble token of the In the outer Himalayas the number of homage of the authors to their illustrious species of liverworts, as well as the frequency guru, the late Professor Birbal Sahni, F.R.S., of the individuals of a species, at a particular who initiated research in Bryology in the altitude, decreases as one proceeds from University of Lucknow and keenly followed Darjeeling in the east to Kashmir in the its progress. The authors wish to convey west (MAP 1 and TABLE I). This is in their grateful thanks to Mrs. Savitri Sahni, conformity with the view expressed by President, Governing Body of the Palaeo­ Kashyap ( 1921) and by Pande ( 1936 ). botanical Society, and to the Editorial Com­ There are more common species between mittee of The Palaeobotanist for the oppor­ the Eastern Himalayas and South India and tunity given to them to contribute this Ceylon than between any of these regions article to the Sahni Memorial Volume. and the Western Himalayas (TABLE II). They are also grateful to those friends and The hepatic flora of the Eastern Himalayas colleagues who have placed their valuable shows affinities with the hepatic flora of the collections at the disposal of the authors. Malayan region. It also includes some To the Department of Meteorology, Poona, species in common with the flora of China, their thanks are due for the meteorological Japan and Australia (TABLE II A). The data included in this paper. hepatic flora of South India, like that of the lVlr. Sultan Ahmad Khan (Research Assis­ Eastern Himalayas, is characterized by the tant to Pande) has been helpful in many presence of Malayan species. There is a ways, especially in the collection of material lesser element of Chinese species in this flora from South India during his tour there in but it includes several species common to 1950. The work of Mr. Khan was financed East Africa ( TABLE II B ). The flora of the by the Government of Uttar Pradesh to Western Himalayas shows affinities with the whom and to the Scientific Research Com­ flora of Central Asia and Europe. There mittee, D.P., grateful acknowledgement is is a lesser element of common species in the made.

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ABEYWICI

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S,~NE,P. V. ( 1942). Observat.ions on some species SRINIVASAN, K. S. (1939). The developmental of Anthocel'os Lind. from Poona and neighbour­ morphology of androgynous receptacles 111 ing hills of the \Vestern Ghats. JOUl'n. en/v. Jial'chaillia palmata :\ees. Proc. Ind. Acad. Bam. 10: 5. Sn lOB (1); 88 . SCHIFFNER, V (1899). Beitrage zur Lebermoos- Idem (1940). On the morphology, Life-history flora 'von Bhootan. IVien. and Cytology of Nlccia himalaye'lsis St. IOUl'l1. Idem (1938). Monographie del' Gattung Cya- Madras l.luie. 12: 59·80 thodiurn. Annales Jj'yologicL 11: 10-30; 12: Idem (1944) The developmental rporphology 121-142 & 131-140 and Cytology of :1fw'chanlia palmata 1\"et:s. SCHIFFNER, V. & PANDE, S. K. (1950). On the joul'll. Jlar/ras Univ. 15 (2): 101-133. East Himalayan Hepatics. 1. Proc. 37th Ind \'ERDOORN, F (1930) Die Frllllaniacea del' Indo­ Sci. Cong.. 41 maleslscilen lnseln . ..Juna"', BnologlCl. Supp!. 1: STEPHANI, ro. ( 1900) Species Hepaticarllm. I. 1-187. Idem ( 1906). Ibid. II Idem (1934). Studien iiber Asiatische ]ubuleat:. Idem ( 19069). Ibid III. ,~unateslJryologlci. Supp!. 4: 1-231. Idem (1909-12). Ibid IV. Idem (1935). Hepaticae Seleetae d criticae Idem ( 1912-17). Ibid V. \'11 1'1 VIII. Annall'> Bryo/ngici. Supp1. 8: Idem ( 1917-24). Ibid. Vr. 150