Proc. Indian Acad. Sci. ( Sci.), Vol. 96, No. 1, March 1986, pp. 17-24. Printed in India.

Porella platyphylla (L.) Pfeiff. in India and scanning electron microscope details of spores*

FARHAT SHAHEEN and S C SRIVASTAVA Department of Botany, University of Lucknow, Lucknow 226 007, India

MS received 3 August 1985; revised 24 February 1986

Abstraer. The occurrence of (L.) Pfeiff. in India was so far doubtful, but the present investigations have now established its definite representation in western Himalaya. The scanning electron microscope details of spores have been studied for the first time in this species showing characteristic sporoderm architecture.

Keywards, Bryophyta; Hepaticae; ; Porella platyphylla (L.) Pfeiff.; SEM spores.

1. Introduction

Kashyap (1932) described Porella platyphylla (L.) Pfeiff. from the western Himalaya but in the absence of specimens investigated by hito it was not possible to substantiate the distribution of this taxon in India (see also Chopra 1943; Parihat' 1961; Kachroo 1973). However, this species was recently reported from Kashmir Valley (Srivastava 1979). Hattori (1970, 1978) raised a doubt on the occurrence of P. platyphylla in India and described P. chinensis (St.) Hatt. from Kashmir, Narkanda and Simla instead of the former species. This possibly points towards the occurrence of both taxa is these areas with the preponderance of P. platyphylla. Both the taxa are highly distinctive in their gametophytic as well as sporophytic details which are described below. P. platyphylla differs from P. chinensis in several morphological features of both gametophyte as well as sporophyte. In P. platyphylla, the leaf-lobes and leaf-lobules are triangular-ovate with recurved apices and the underleaves are rotund-quadrate to triangular, with entire margins. The perianths are campanulate, bilabiate with generally recurved mouth. The middle and basal portion bear 3--4 plicae. The capsule usually dehisces into 5 or more irregular valves having uniformly thick capsule wall of 3-5 layers of cells. In P. chinensis, on the other hand, the leaf lobes are ovate, leaf-lobules are ligulate and the underleaves are oblong having highly crispate margins. The perianth is usually ovoid without plicae (sometimes 1-2 plicate) and the mouth is flat. The capsule dehisces typically into 4-valves. The capsule wall is not uniformly thick; it is 1-1ayered and pigmented deep brown towards apex and 2-3 layered and hyaline towards base (Shaheen and Srivastava 1985). Both the taxa further differ in the structure of capsule waU. The cells of the epidermal layer of capsule possess trigonous thickening at the angles (corners) and the cells of inner layer possess irregular thickenings on radial and end walls rarely weakly extending on tangential walls in P. platyphylla. However, in P. chinensis the epidermal cetls of the upper part of the capsule wall are elongated and have thickenings only on radial walls (in cross section).

* Contribution from the Botany Department, New Series (Bryophyta) No. 205. 17 18 Farhat Shaheen and S C Srivastava

The cells in the median and basal portions of the capsule wall are devoid of thickenings. The present study is based on several collections made from different localities in Kashmir, Himachal Pradesh and the Valley of Flowers and further supplemented by the examination of authentic specimens of P. platyphylla obtained from NICH (Japan) and Stephani Herbarium (Conservatoire et Jardin Botaniques, Geneva). The detailed critical evaluation of the above specimens revealed the possibility of occurrence of P. chinensis also in these areas in addition to P. platyphylla which is preponderant particularly in Kashmir Valley. We, however, did not succeed in collecting P. chinensis from the latter locality although it occurs in Himachal Pradesh (see Shaheen and Srivastava 1985). The become richly fertile during October-November. Both the sexes grow together in the same population and succesgfully accomplish sexual reproduction, resulting in the formation of copious sporogonia. A comparative account of Indian plants including new synonyms has been provided along with the scanning electron microscope (SEM) details of spores, hitherto unknown in this taxon.

2. Taxonomic description

Porella platyphylla (L.)Pfeiff., F;. Niederhessen und Mª 2:243 (1855). Syn: Madotheca platyphylla Dum, Comm. Bot. p. 111 (1822); Kashyap in, Liverw. W. Himalaya and Panjab PI. Part 2: 34(1932); Madotheca tri#onifolia St., sp. Hep. 4:293 (1910) Syn. nov.; Madotheca virens St., Sp. Hep. 4:294 (1910) Syn. nov.; Udar and Shaheen in, Geophytology 12(2): 342 (1982) Syn. noo. Plants small sized, upto 60 mm long. Stems irregularly, densely and pinnately branched, branching Frullania-Ptychanthus to Frullania-Jubula type; cortical cells in 1-4 rows, 8-17 x 8-28 pan, thick-walled, medullary cells 14-34x 11-36/~'n, thin- walled, trigonous. Leaves densely imbricate, incubous, leaf-lobes triangular-ovate, entire 1-4-1.9 x 2-1.9 mm, apex obtuse, strongly recurved, cells trigonous: Apical cells at margin 14-28 x 14-20#m, median cells 20--42 x 20-34 #m, basal cells 22-45 x 22-39 q leaf-lobules entire, rarely with 1-more dentitions at dorsal base, ovate- triangular to lanceolate, 0-6-1 x 0-2-0-3 mm, apex usuaUy incurved, base long decur- rent, entire or with few dentitions. Underleaves usually entire, rarely with 1-2 dentitions, rotund-quadrate to triangular, 0.7-1 x 0-6-0.9 mm, apex usually recurved, base broad, long decurrent with few dentitions. Dioecious, male plants with terminal androecium on very short branches (spikate), compact, bracts delicate, densely crowded and entire; antheridia singly in the axil of bracts, with spherical body anda stalk of 2- rows ofcells; wall ofantheridium single layered at apex and multilayered below. Female plants with numerous female inflorescences on short laterals either arise from main stem or from its primary branches. Bracts usually in a single pair or sometimes in 2 pairs, lower pair of bracts small and deformed, bracts and bracteoles may be dentate or entire, dentitions very small. Perianth campanulate, formed in a series on both sides of stem, middle and basal portions with 3--4 plicae, mouth usually flat, bilabiate, generally recurved, with short teeth. Calyptra 2-cell layers thick. Sporophyte with distinct foot, seta and capsule. Capsule slightly oval or spherical, dehiscing into 5 more irregular valves; cells of the epidermal layer of capsule wall with trigonous thickenings at the angles (corner), cells of the inner layer with irregular thickenings on radial and endwalls, rarely weakly extending to tangential walls. Spores spherical, 36-54 q in diameter, Scannino electron microscope details of spores 19

sometimes slightly t¡ or oblong, yellowish-brown and fmely papillose. Under SEM the spores show collapsed exine having distinct spinulose ornamentation, spinules dense, very small, broad at the base and slightly narrow and rounded at the apex. Elaters free, elongated or short and stumpy, sometimes branched, with blunt ends, spirals usually 2-3, rarely partiaUy single but always bispiral at the ends, stumpy elaters with 2 or more bands.

2.1 Specimens examined G. 21534, Madotheca trigonoJblia St., Loc.: Liddar Valley (alt. ca 3300 m), Kashmir, Leg.: J. F. Duthie and Brothms 13325, July 29, 1893; G. 16803, Madotheca trigonifolia St., Herb. Levier No. 2891, Loc.: Alliabad, Pirpunjal (alt. ca 3100 m), Kashmir, Leg. Inayat Khan, Aug. 9, 1901; G. 21536, Madotheca trigonifolia St., Loc.: Gulmarg (alt. ca 2700m), Kashmir Leg.: J. F. Duthie and Brothms 14281, July 1, 1893; G. 21539, Madotheca virens St., Herb. Munchen, Loc.: Narkanda, Leg. and det. ign.; G. 21543, Madotheca virens St. Herb. Munchen; Loc.: India Orient., Leg. and det. ing.; NICH 225703, Herb. K. starcs flora Bava¡ (Madotheca) Porella platyphylla (L.) Dum., Loc.: Oberpflaz, Kr. Amberg., Leg.: K. Starcs, April 1, 1946; NICH 225692, Hepaticae Scandinavicae Ex. Herb. K. A. Th. Seth, Porella platyphylla (L.) Loc.: Procincia, Leg.: Klthe Seth, May 18, 1977; NICH H-225704, Cryptogame Exsiccatae editae a Museo Hist. Natur. Vindobon› 3792, Porella platyphylla (L.) Lindenb. Loc.: Osterrich, Osttirol (alt. ca 800 m), Leg.: E. Bauer, Aug., 1910, Det.: V. Schiffner; NICH 225691, Muscinees communiquee's par l'Herbier Biossier, (Madotheca) Porella platyphylla (L.) Dum. var. communis Nees, Loc.: Suisse. Leg.: Dr H. Bernet, 1885: LWU: 6843/68, Bryophytes from western Himalaya, Porella platyphylla (L.) Pfeiff., Loc.: Kokarnag, Kashmir, Leg.: R. Udar and party, 1968, Det.: R. Udar, S. C. Srivastava and F. Shaheen; LWU: 6848 Bryophytes from western Himalaya, Poretla ptatyphytla (L.) Pfeiff, Loc.: Pehlgam, Leg. Y. K. Sharma, Det.: Ro Udar, S. C. Srivastava and F. Shaheen; LWU. Nos. 206/76, 210/76, 213/76, 215/76, 216/76, Bryophytes from western Himalayas, Porella platyphylla (L.) Pfeiff., Loc.: Yusmarg (on way to Neelganga), Kashmir, Leg.: S. C. Srivastava, Oct. 13, 1976, Det.: R. Udar, S. C. Srivastava and F. Shaheen; LWU. Nos. 214/76, 229/76, Bryophytes from western Himalaya, Porella platyphylla (L.) Pfeiff., Loc.: on way to Khilanmarg, Kashmir, Leg.: S. C. Srivastava, Oct. 28, 1976, Det.: R. Udar, S. C. Srivastava and F. Shaheen; LWU. 4089/80, Bryophytes from western Himalaya, Porella platyphylla (L.) Pfeiff, Loc. VaUey of Flowers, Leg.: S. C. Srivastava, D. Kumar and D. K. Singh, May 21, 1980, Det.: R. Udar, S. C. Srivastava and F. Shaheen.

3. Discussion

Porella platyphylla forros a natural complex with P. decurrens (St.) Hatt. and P. gracillima Mitt. but significantly differs from them in overall morphology ofleaves and underleaves (see Hattori 1969, 1976; Schuster 1953, 1980). Herzog (1938) and Hattori (1979) recorded P. platyphylla from Hindu Kush--an area which is adjacent to Kashmir and it seems that the plants prefer high altitudes for favourable growth as it also occurs in the Valley of Flowers and Himachal Pradesh. 20 Farhat Shaheen and S C Srivastava

3

2

4

7 8

9 .2 mm

3-6 ,! mm

6 | 1,2,7,8 3. mm Figures Ÿ Porella ptatyphylla (L.) Pfeiff. 1. A portion of plant with mate inflorescences (ventral view). 2. A portion of plant showing attachment of leaf-lobules and underleaves. 3. Cross-section ofstem (a portion). 4. Marginal leafcells. 5. Median leafcells. 6, Basal leaf cells. 7 and 8. Underleaves. 9. Antheridium. Scannin9 electron microscope details of spores 21

~?, t4.- / ~' ...... -,? / 13

X ! :ii.i! 16

~'~ 24 25 27

~~ 14 .2mm

18- 27 I mm

18 2~~ I0-13,15-17

Figures 10-27. Porellaplatyphylla (L,) Pfeiff, 10, A portion of plant with t'emale inflores- cences and dehisced capsuie~ (ventral vie,u). 11 and 12. Fema]e bracts. 13. Female bracteole. 14. Perianth cells (at mouth). 15. Cross-section of perianth (apical portion). 16. Cross- section of perianth (median portion). 17. Cross-section of perianth (basal portion). 18. Cells of outer layer of capsule wall. 19. Cells of inner layer of capsule wall. 20-22. Spores. 23--27, Elaters. 22 Farhat Shaheen and S C Srivastava

Figures 28-29.Porella platyphylla (L.) Pfeiff. 28.SEM of Spore. x 1305"3.29.Aportionofthe same. x 2107-3.

The plants of P. platyphylla mostly become pink under JJK reaction (see Hattori 1978). However, the specimens collected from Kashmir and the Valley of Flowers rail to respond to this treatment. Other plants of P. platyphylla located in NICH (225703, 225692, H-225704, 225691) investigated during the present work also show negative Scanning electron microscope details of spores 23 reaction. P. platyphylla is considered to be interesting showing variable chemical pattern in different seasons of growth (Asakawa et al 1980). The JJg reaction may possibly be influenced by the changing chemical components of the species, hence does not appear to be taxonomically so important. Hattori (1970) treated M. trigonifolia St. and L. virens St. as synonyms of P. chinensis but a study of the same specimens and some other specimens of the former two species located in Herb. G. shows that the plants of M. trigonifolia collected from Kashmir (G, 16803, 21534, 21538, 21536) and those ofM. virens from Narkanda (G. 21539) and India orient? (G. 21543) and the plants collected by us from Kashmir, Himachal Pradesh and Valley of Flowers are clearly referable to P. platyphylla on the basis of their morphological characters (figures 1-29). The plants of M. trigonifolia (G. 16803) were fertile with male plants showing bifurcated antheridial branches and the female ones showing campanulate perianths. The specimens collected from Kashmir are also richly fertile and dimorphic. The male infl0rescences are usually terminal on short lateral branches and may be branched (figure 1). The female plants have campanulate perianths (figure 10). The SEM observations of spores of P. platyphylla revealed distinct sporoderm ornamentation of the spinules type. The spinules form small protuberances and are very dense (figures 28, 29). In another Indian species, P. perrottetiana (Mont.) Trev., the spinules are widely spaced, rod shaped and topped with a small rounded head (Udar and Shaheen 1983). The spores under LM, however, do not show conspicuous ornamentation (figures 20-22). The elaters are usually with 2-3 spirals, rarely becoming unispiral in some portions, but are always bispirate at the ends (figures 23-27).

Acknowledgements

The authors are grateful to late Professor Raro Udar for evaluating the Indian specimens of Porella platyphylla and also critically going through the MS. Thanks are also due to Dr Patricia Geissler, Director, Conservatoire et Jardin Botaniques, Geneva and Dr S Hattori of Hattori Botanical Laboratory, Nichinan, Japan for the loan of Type/authentic specimens, to the University Grants Commission for financial assistance and to Mr V K Lal of Indian Institute of Technology, New Delhi for SEM facilities.

References Asakawa Y, Tokinaga N, Toyota N, Takemoto T, Hatto¡ S, Mizutani M and Suire C 1980 Chemosystematics of Bryophyta IV The distribution of terpenoids and aeromatic compounds in Hepaticae and Anthocerotae;J. Hattori Bot. Lab. 47 153--164 Chopra R S 1943 A Census of Indian Hepatics; J. lndian Bot. Soc. 22 237-259 Hatto¡ S 1969 Studies of the Asiatic species of the Porella (Hepaticae)II; J. Hattori Bot. Lab. 32 319-359 Hatto¡ S 1970 Studiesof the Asiaticspecies of the genus Porella (Hepaticae)III; J. Hattori Bot. Lab. 33 41-87 Hattori S 1976 Studies of the Asiatic species of the genus Porella (Hepaticae)VI; J. Hattori Bot. Lab. 40 121-138 Hattori S 1978 A note on Asian Porella platyphylla; Misc. Bryol. Lichenol. 8 1-2 24 Farhat Shaheen and S C Srivastava

Herzog T 1938 Bryophyta in Haeckel und Troll, Botanische Ergebniss der Deutschen Hindukush Expedition 1935; Feddes Repert. Beih. 108 3-12 Kachroo P 1973 Hepaticae of India--a taxonomic survey and census III Plagiochilaceae through Pleuroziaceae; Kashmir J. Sci. 1 141-161 Kashyap S R 1932 Liverworths of the western Himalayas and the Panjab Plains II (The University of Punjab, Lahore) Parihar N S 1961 Ah annotated revised census of Indian Hepatics; Unir. Allahabad Stud. Bot. (Sect.) 1-56 Schuster R M 1953 Boreal Hepaticae. A mannual of the liverworts of Minnesota and adjacent regions; Ara. Midl. Nat. 49 257-684 Schuster R M 1980 The Hepaticae and Anthocertotae ofNorth America IV (New York, London: Columbia Unir. Press) Shaheen F and Srivastava S C 1985 Porella chinensis (St.) Hatt. from India with remarkable capsule wall morphology; J. lndian Bot. $oc. (in press). Srivastava S C 1979 Hepaticae of Kashmir Valley; Nova Hedwigia Z. Kryptogamenkd. 63 333-338 Udar R and Shaheen F 1983 Morphotaxonomy of Porella perrottetiana (Mont.) Trev. from south India; J. lndian Bot. Soc. 62 319-325