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Sporoderm and Elateroderm Diversity in Some Rare and Interesting Indian Liverworts Under Sem2

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Sporoderm and Elateroderm Diversity in Some Rare and Interesting Indian Liverworts Under Sem2 J. Hauori Bot. Lab. No. 79: 129- 138 (Feb. 1996) SPORODERM AND ELATERODERM DIVERSITY IN SOME RARE AND INTERESTING INDIAN LIVERWORTS UNDER SEM2 VIRENDRA NATH1 AND A. K. ASTHANA1 ABSTRACT. Sporoderm and elateroderm pattern under SEM have been investigated in seven rare and interesting Indian liverworts: Calobryum indicum Udar et Chandra, C. dentatum Kumar et Udar (Calobryales), Schiffneria hya/ina Steph., Trichocolea tenera Udar et Singh (Jungermanniales), Fossombronia cristula Aust., F. pusi/la (L.) Dum. and Petalophyllum in ­ dicum Kash. (Metzgeriales). The diagnostic features of each taxon have also been provided. INTRODUCTION Spores in liverworts exhibit an interesting range of diversity in their sporoderm ornamentation. Distinctiveness of sporoderm sculpturing is unambiguous in hepaticae and can therefore be convincingly applied for systematic treatment of various taxa. Elaters also exhibit distinct structural patterns and ornamentations. Studies on sporoderm morphology of Indian Hepaticae and Anthocerotae under LM (Light Microscope) has already received considerable attention by Mehra and Sood ( 1969), Srivastava and Udar ( 1975), Udar ( 1976) and Gupta, and Udar ( 1986). SEM (Scanning Electron Microscope) studies have resolved some micromorphological det­ ails of sporoderm which could not be discerned using LM (Light Microscope). Utilizing the efficiency of SEM, studies on hepatic spores have been made by several workers at global level (Taylor et al. 1974; Clarke, 1979; Jovet-Ast, 1979; Geissler and Gradstein, 1982; Stienkamp and Doyle, 1984; Inoue and Hibino, 1984; Scott and Pike, 1984, 1987, 1987a, 1987b, 1988, Bischler, 1989, Brown and Lemmon, 1993; Jovet-Ast, 1993). Such studies have received attention in Indian Hepaticae by Udar and Shaheen (1983), Udar and Kumar (1983), Udar and Srivastava (1983), Udar and Srivastava, G . (1983), Udar et al. (1983), Udar and Awasthi (1983), Udar and Jain (1983), Udar and Srivastava (1984), Udar and Agarwal (1985), Srivastava (1986), Sinha et al. (1987), Asthana and Srivastava (1991), Nath and Asthana (1992), Sharma and Srivastava (1993). The present study provides SEM details of sporoderm and elateroderm of Calob­ ryum indicum, C. dentatum, Schiffneria hyalina, Trichocolea tenera, Fossombronia cristula, F. pusilla and Petalophyllum indicum. The study has shown intergeneric as well as interspecific variability in surface ornamentations both in spores and elaters, which are considered the best taxonomic tool for discrimination of genera and species. 1 Bryology Laboratory, National Botanical Research Institute, Lucknow-226 001. India. 2 NBRI Research Publication No. 436 (N.S.) . 130 J. Hattori Bot. Lab. No. 79 I 9 9 6 MATE RIALS AND METHODS Fertile plant specimens were collected from different bryologically rich localities of the Western Himalayas, Eastern Himalayas and South India. Thoroughly cleaned mature capsules were subjected to dehydration through the usual ethanol series followed by critical point drying. The dehydrated capsules were incised and non­ acetolysed spores and elaters were examined under SEM (JEOL-JSM 35c). The Voucher specimens were deposited in the Bryophyte Herbarium, National Botanical Research Institute, Lucknow (LWG). DESCRIPTIONS l. Calobryum indicum Udar et Chandra, Revue Bryol. et Lichenol. 33(3-4): 555-559 (1964-65). Pl. 1, figs. 1, 2. Plants strictly acrogynous, dioecious, 6- 12 mm long, gametophores erect arising from upturned tips of an interwoven tangled mass of rhizome; leaves mostly crowded at apex of axis, broader than long with entire margin; antheridia borne at apex in group forming a dome-like structure. Spores: More or less spherical, minutely papillate yellowish-brown, 2(}-25 µm in diameter, sporoderm exhibited dense round headed papillae over the entire surface, exine occasionally folded. Elaters: Yellowish-brown, bispirally thickened tapering ends, 35(}-700µm long, elateroderm exhibiting the presence of minute tubercles scattered over the surface. Specimen examined: India. Eastern Himalayas: West Bengal, Darjeeling, (Gauri­ bas 27 °N, 88.25 °E; alt. ea. 3000m), 23.5.1981, leg. S. Chandra and V. Nath, LWG 203814-B. Growing on soil. 2. Calobryum dentatum Kumar et Udar, J. Indian bot. Soc. 55: 23- 30 (1976). Pl. 1, figs . 3, 4. Plants strictly acrogynous, dioecious 8- 12 mm (- 15 mm) long, axis leafy through­ out; leaves usually longer than broad; upper two cycles including the female bracts prominently dentate; oil-bodies 5(}-70 in leaf cells; antheridia borne in the axils of two upper cycles of leaves or terminal. Spores: Yellowish brown, 18- 24 µm (- 27.5 µm) in size, sporoderm conspicuously folded and studded with minute tubercles. Elaters: Light brown, bispirally thickened, with tapering ends remarkably shorter than C. indicum up to 440 µm long, elateroderm appearing more or less smooth as compared to C. indicum except two or three (or even more) minute tubercles. Specimen examined: India. Eastern Himalayas: West Bengal, narjeeling (Gauri­ bas, 27 °N, 88.25 °N; alt. ea 3000m), 23.5.1981, leg. S. Chandra and V. Nath, LWG 203814-A. Growing on soil. 3. Schiffneria hyalina Steph. Oest. bot. Zeitschr, 44: 1 (1894). Pl. 1, figs. 5, 6. Plants dorsiventrally flattened, thalloid, 17 mm X 3 mm, laterally lobed; sex organs V. NATH & A. K. AsTHANA: Sporoderm and elateroderm diversity 131 Plate 1. Figs. I, 2: Calobryum indicum Udar et Chandra. I. Spore ( X4000); 2. Elater ( X 3000). Figs. 3, 4: Calobryum dentatum Kumar et Udar. 3. Spores in cluster ( X 2400); 4. Elater ( X 2400). Figs. 5, 6: Schijfneria hyalina Steph. 5. Spore ( X 4800); 6. Elaters with few spores ( X 540). 132 J. Hattori Bot. Lab. No. 79 I 9 9 6 borne on leafy shoots arising from ventral surface or apex: antheridium occurring singly in the axil of saccate bracts, female shoots bearing terminal perianth, oblong, smooth-walled with dented mouth; capsule wall bistratose having nodulose-slightly elongated thickenings on each alternate longitudinal wall of epidermal layer. Spores: Light yellowish brown, 12.5- 15 µm in diameter. Under SEM sporoderm exhibiting densely verrucose to pseudoreticulate pattern, proximal face marked with a thick triradiate mark. Elaters: Yellowish brown, 140-280µm long, bispirally thickened with rounded ends, under SEM elateroderm appearing more or less smooth. Specimen examined: India. Eastern Himalayas: West Bengal, Darjeeling (Sanda­ kphu-Tanglu 27.05°N, 88 °E, alt. ea 3000m), 26.4.1965, leg. S. Chandra, LWG 202450 Bry. Growing on decayed logs in association with Plagiochila sp. and mosses. 4. Trichocolea tenera Udar et Singh, Geophytology 7: 69 (1977). Pl. 2, figs. 1- 4. Plants in tufts, usually 60-90 mm long, regularly bi pinnate, stem delicate, main stem 281 - 346 µm wide, 8-12 cells across diameter, cortex single layered; leaves in three rows, underleaves smaller, about half as wide as lateral leaves, lamina of underleaves on main stem bearing rhizoidal disc bearing tufts of rhizoids, androecial branches similar to vegetative branches bearing 1 or rarely 2 antheridia in the axil of each bract; female plant bearing terminal coelocaule, oblong, wider at mouth with paraphyllia. Spores: Yellowish-brown, 15-17.5 µm in diameter, sporoderm densely vermiculate with ridged exine. Sometimes these ridges form a reticulum enclosing irregularly shaped lumina. Proximal face bearing a prominent elevated triradiate mark. Elaters: Brown, 13 7. 5- 192. 5 µm long, bis pi rally thickened with prominent taper­ ing ends. Elateroderm more or less smooth in nature. Specimen examined: India. Eastern Himalayas: West Bengal, Darjeeling (Senchal lake, 27 °N, 88.25 °E, alt. ea 3000m), 18.5.1981, leg. S. Chandra and V. Nath, LWG 203723. Growing on moist humus on road side. 5. Petalophyllum indicum Kash., J. Indian bot. Soc. 7: 14 (1928). Pl. 2, figs . 5, 6. Plants dioecious, simple or furcate, 12 mm X 7 mm, basal portion cylindrical and wingless with fanning apex having parallel, erect leaf appendages forming lamellae on the dorsal surface; wings or lamellae multi-layered at base gradually becoming single layered above, wavy along the margin; antheridia in groups behind the apex, protected by scattered scales; sporogonia 1- 4, usually one in each perianth; capsule wall usually 3-layered; inner layers with thick annular or sometimes semiannular bands. Spores: Dark brown, spherical, 40 µm in diameter, sporoderm exhibiting membra­ nous elevated lamellae forming a perfect reticulate pattern, reticulations penta­ hexagonal in shape, 8- 10 µm, usually 3- 4 reticulations across the diameter, marginal wings possessing fine striations. Elaters: Bi-trispiral, lightly coloured, spirals very distinct, brown, 280-400 µm long, 8- 10 µm broad, sometimes branched, elateroderm exhibiting prominent striations except on spiral band (Pl. 2, fig . 6). V . NATH & A . K. ASTHANA: Sporoderm and elateroderm diversity 133 Plate 2. Figs. 1-4: Trichocolea tenera Udar et Singh. 1. Group of spores ( x 3600); 2. Bunch of elaters and spores ( X 720); 3. Spore ( X 4800); 4. Elater with few spores ( X 1000). Figs. 5, 6: Petalophyllum indicum Kash. 5. Spore ( X 2000); 6. Elater (a portion X4000). Specimen examined: India. Western Himalaya: Uttar Pradesh, Chamoli (30.4°N, 79.5 °E; alt. ea. 2500m), March 1978, leg. S.S. Bir, LWG Bir/78. Growing on soil. 6. Fossombronia cristula Aust., Acad. Philad. p. 288 (1866). Pl. 3, figs. 1- 3. Plants monoecious, leaves flat, somewhat wavy at the margin, each lobe with mucilage papilla at apex, unistratose except at base. Spores: Yellowish brown to dark brown, 43- 58 µm in diameter, sporoderm exhibiting a perfectly reticulate pattern formed by coalition of elevated lamellae on 134 J. Hattori Bot. Lab. No. 79 I 9 9 6 distal face, usually 5- 7 luminae (meshes) across the diameter, perispore well developed at periphery, with crenulate margin. Proximal face entirely devoid of lamellae and possessing minute papillae. Elaters: Inadequately developed pale yellow, 24- 115 µm long having annular­ bispiral thickenings, elateroderm appearing more or less smooth, without any ornamen­ tation. Specimen examined: India. South India: Kerala, Wayanad (Periyar, 10. l 0 N, 76.4°E; alt. ea. 600m), 9.8.1984, leg. S. Chandra and V. Nath, LWG 204069.
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