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Indian -an overvIew

B. D. Sharma

Sharma BD 1992. Indian Williamsoniaceae-an overview. Palaeobolanisl 40 260·265.

The morphology, anatomy and phylogeny of the known extinct representatives of the family Williamsoniaceae from India have been overviewed. The family occurs in the Upper rocks and is represented by fronds, stems, male, female and bisexual fructifications. Though the fronds vary in shape, size and venation, all possess identical syndetocheilic stomata. The stems Bucklandia and Sahnioxylon show minor variations in struc1ure of tracheary elements and are related with cycadeoideas and homoxylous angiosperms. The male fructification Wellrichia is built on very distinct and different plan than the seed-bearing fructification- , which has been derived from Cordaianlhus. The bisexual fruclification-Amarjolia is terminal and exposed like Williamsonia, while in structure and arrangement of microsporophyll resembles . Relationship among different organs is suggested and phylogeny of Williamsoniaceae is discussed. Key-words-Williamsoniaceae, Morphology, Phylogeny, India

B. D Sharma, Deparlment of BOlany, University of jodhpur, jodhpur 342001, India. mllT ~ ~ f4\'\WHftf"ll{f11~

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FAMILY Williamsoniaceae, belonging to the class spread with two rows of appendages on which Bennettitopsida, flourished and dominated the microsynangia are produced. /seeds are vegetation during the Mesozoic Era (Harris, 1969). produced intermingled with sterile scales in a Members of this family originated sometimes during compact layer covering the . In bisexual Middle and disappeared by the end of Lower fructification baloon-shaped microsporophylls . The family includes extinct in encircle the central receptacle with a covering of which the stem surface has spirally arranged ovules and sterile scales. rhomboid leaf bases, parenchymatous pith and In India, members of this family occur at several cortex well developed with mucilage ducts; vascular Mesozoic localities, e.g., Rajmahal Hills, Bihar; zone narrow made up of collateral, conjoint and Golapilli near Elore; Madras outliers, )abalpur, endarch bundles; secondary wood mano-or )aisalmer, Kachchh, etc. (Oldham & Morris, 1963; picnox)r\ic; leaf trace is unilacunate. Leaves are Feistmantel, 1876, 1877, 1879; Seward & Sahni 1920; pinnate or partly pinnate, with parallel or reticulate Sitho!ey, 1954; Gupta, 1966; Sharma, 1967a, 1977, venation; stomata transverse to veins and syndetocheilic. Fructifications are terminal, exposed, Four Decades of Indian Palaeobotany seed-bearing and bisexual. Microsporophylls are Editors: B. S. Venk3t3chala & H. P. Singh Publisher: Birbal Sahni Institute of Palaeobotany, Lucknow SHARMA-INDIAN WILLIAMSONIACEAE-AN OVERVIEW 261

1990; Bose, 1953a, 1953b, 1968; Bose & Kasat, India. Search should also be made to find out their 1972a; Bose & Banerji, 1981, 1984). The associated stems and fructifications. Williamsonian plants described from the Indian rocks are: Pteropbyllum Brongniart Fronds-Ptilophyllum (18 sp.), (10 Frond pinnate, pinnae linear, base syrnmetrical sp.), Otozamites (7 sp.), Anomozamites C3sp.) and attached on lateral side of rachis, apex acute, obtuse Dictyozamites (7 sp.). or truncate, veins parallel with few forkings. Stems-Bucklandia (4 sp.) and Sahnioxylon (2 Epidermal and anatomical characters of Indian sp.) are not known. Oldham and Morris (1863) Male jructzfication- Weltrichia C3sp.) and Feistmantel (1877) identified a number of Seed-bearing jructzfication- Williamsonia (12 species of this from India, but Seward and sp.) Sahni (1920) merged many of these species into the Bisexual jructijication-Amarjolia (1 sp.) genus Nilssonia Brong. Sharma (l969b) also remains of Williamsoniaceae occur as transferred two species of Pterophyllum into impressions, incrustations and petrifactions. The Nilssonia. On the other hand Bose and Benerji impressions show external morphology. In (1981, 1984) rE:transferred all known Nilssonia incrustations the epidermal characters are studied species from India into Pterophyllum species through maceration and peel techniques using nitric without studying the epidermal and anatomical acid as etching acid (Jacob & Jacob, 1954; Bose & characters. Kasat, 1972a; Bose & Banerji, 1981, 1984; Sukh-Dev & Rajnikanth, 1988a, 1988b). The petrifactions have Otozamites Braun been studied by the usual method of cutting, Frond pinnate, pinnae small to linear to grinding and polishing techniques. At Amarjola, the triangular or round; base asymetrical and auriculate, material 'being soft and fragile needs cooking in apex acute, obtuse or round; veins diverging and Canada balsam prior to sectioning. Canada balsam is dichotomised. Epidermal cells sinuous, stomata used as mounting medium. Sometimes, the polished hypostomatic, syndetocheilic and restricted to surface is examined with a water film under stomatal bands. reflected light. Oldham and Morris (1963) and Feistmantel (1876, 1877, 1879) described a number of species of DESCRIPTION this frond genus from India. Seward and Sahni (1920) merged many of the species into the genus A systematic account of the extinct plants of Ptilophyllum. Roy (1963) described Otozamites Williamsoniaceae known from India is given as bellus from Kachchh. Bose and Banerji (1981, 1984) under: described the morphology and epidermal characters Fronds of O. imbricatus Feistmantel 1876, O. Ptilophyllum Morris Walkamotaensis Bose & Zeba-Bano 1981, 0. kachchhensis Bose & Banerji 1984. Otozamites Frond pinnate, pinnae linear to round, base occurs more frequently in the Mesozoic rocks of asymetrical, upper basal angle round, lower Kachchh than any other exposure in India. decurrent; apex acute, obtuse, truncate or round; veins parallel with forkings. Epidermal cells mostly Dictyozamites Oldham sinuous, stomata hypostomatic, transversely oriented and syndetocheilic. Rachis with a number of bundles Frond pinnate, pinnae linear, falcate or round, arranged in double U-manner. base asymetrical and auriculate, apex acute, obtuse Sharma (1967a), Bose and Kasat (1972a) and or round, venation reticulate, stomata hypostomatic, Bose and Banerji (1981, 1984) published reviews on syndetocheilic and restricted to stomatal bands. the Indian species of the genus Ptilophyllum and Rachis has vascular bundles arranged in double U­ identified 18 species. Some of these fronds are manner similar to that of Ptilophyllum (Bose & Kasat, morphologically alike but separated on epidermal 1972a). In Dictyozamites, the number of areoles characters, e.g., P. nzpanica Mittre 1956, P. indicum present in the middle of pinna is used in the Jacob & Jacob 1954, P. sahnii Gupta & Sharma 1968. identification of species. Seven species, viz., D. Minor variations in morphological and epidermal jalcatus Oldham 1963, D. indicus Feistmantel 1877, characters may be due to environmental differences D. hallei Sahni & Rao 1933, D. sahnii Gupta & and age of the frond. A re-investigation is likely to Sharma 1964, D. jeistmantelii Bose & Zaba-Bano reduce the number of species of Ptilophyllum in 1978 and D. gondwanensis Sukh-Dev & Rajnikanth 262 THE PALAEOBOTANIST

1988a are known from India. Associated stems and are also seen. Whether Sahnioxylon had leaf bases fructifications are yet to be discovered. on surface and what kind of leaf traces were present, is yet to be discovered. Anomozamites Schimper Fructifications Frond lobed or pinnate, pinnae unequal, base symmetrical, attached on lateral side of rachis, apex Weltrichia Braun obtuse to truncate, veins parallel with forkings. This male fructification was described earlier Epidermal cells sinuous. Stomata hypostomatic, under the generic name Williamsonia. Sitholey and transversely oriented and syndetocheilic. Important Bose (1953) instituted Williamsonia santalensis species of the genus known from India are: A. jissa which was later on transferred to Weltrichia (Feist.) Sharma 1969b, A. amarjolense Sharma et al. santalensis by Sitholey and Bose (1971). Sitholey 1971, and A. haburensis Bose & Banerji 1981. and Bose (1953) described a single whorl of Associated stems and fructifications remain microsporophylls surrounding a cup-shaped unknown. receptacle, while Sharma (1969a) on the basis of Stems-Bennettitalean stems are simple or study of more than 50 specimens, some of which are branched and possess spirally arranged rhomboid nicely preserved counter parts, suggested two leaf bases on the surface. Two stem genera, viz., whorls, i.e., the abaxial of sterile and adaxial Bucklandia Pres!. 1825 and Sahnioxylon Bose & Sah of microsporophylls. Sitholey and Bose (1971) did 1954 are known from the Indian rocks. not agree to it. Each microsporophyll has two rows Bucklandia Prest. of appendages on which parallel rows of microsynangia occur. Sharma (1969a) suggested Stem simple (B. indica), branched (B. sahnii) restorations of W. santalensis and also described a or dichotomised (B. dichotoma). Stem surface has new species W. campanulatijormis preserved as cast close or sparse leaf bases. Pith and cortex are and mold. Bose and Banerji (1984) described parenchymatous with mucilage ducts. Vascular zone another species of Weltrichia, W. harrisiana from is made up of a large number of collateral, conjoint, Kachchh. In this fructification the fertile appendages open and endarch bundles. Secondary wood is are terminal on 12-14 microsporophylls which are compact and differentiated into growth rings. The coalescent towards base forming a circular tracheids have spiral, scalariform or bordered pits on depression. Further investigations are required on radial walls. Rays 1 to many cells high, uni-to­ W. campanulatijormis and W. harrisiana as these multiseriate, homogeneous. Pits in crossfield 1-4 or are established on insufficient and poorly preserved more, circular with narrow border. Leaf trace is \ materials. unilacunate and divides into a number of bundles (5-11) in cortex. Williamsonia Carruthers Four species of Bucklandia, Viz., B. indica Seward 1917, B. sahnii Bose 1953, B. guptai Sharma Feistmantel (1876) described W. blanjordii 1967b and B. dichotoma Sharma 1970d are known from Kachchh and W microps from the Rajmahal from India. Anatomically, B. dichotoma resembles Hills (Feistmantel, 1877). Sahni (1932b) described cycadeoideas and homoxylous angiosperms in W. sewardiana from the Rajmahal Hills and having cross-shaped pit pores and scalariform suggested its restoration and showed its association pittings (Gupta, 1934). with the stem Bucklandia indica and fronds of Ptilophyllum cf. cutchense. Gupta (1943) described a Sabnioxylon Bose & Sah probable bisexual fructification- W. sahnii in which Originally the wood was described as the bracts are spread and in the basal portion of Homoxylon rajmahalense by Sahni (1932a) who receptacle 20 markings are seen, which may be of believed that it was an angiospermous wood. But fallen microsporophylls. Gupta (1958) divided Gupta (1934) related it with cycadeoidea. Hsu and williamsonian fructification into open type and close Bose (1952) made further observations on this type. In the former, bracts spread out as in W. sahnii, wood. Bose and Sah (1954) transferred the while in the latter the bracts did not open out as in Homoxylon to Sahnioxylon as the earlier name had W. guptai. This hypothesis is applicable to all the already been used for a fossil conifer wood, so they known species of Williamsonia throughout the called it Sahnioxylon rajmahalense (Sahni) Bose & world. Bose (1966a) called W. sahnii only a seed­ Sah and also described a new species-So andrewsii bearing fructification. Bose (1968) described W. Bose & Sah 1954. In the former only wood is known, harrisiana from Amarjola in the Rajmahal Hills while in the latter partly preserved ground tissues which is identical to W. guptai Sharma 1968 SHARMA-INDIAN WILLIAMSONlACEAE-AN OVERVIEW 263

collected from the same locality. Sharma (1968) also the Mesozoic Era in India, especially the frond genus established W. amatjolense and separated it from W. Ptilophyllum which occurs in all the Upper Mesozoic guptai on the basis of different epidermal characters exposures and is represented by nearly half the total of bracts. Bose and Kasat (1969) instituted W. number of species known throughout the world. seniana in honour of Dr J. Sen (Calcutta) from ThiS frond is associated with the stem B'ucklandia ]abalpuf. It is an incrustation and preserves the and the seed-bearing fructification Williamsonia epidermal structures. (Feistmantel, 1877; Sahni, 1932b). The male Sharma (l970a, 1970b, 1970c, 1974, 1975, 1976, fructification Weltrichia is also found in close 1980) studied petrified specimens of Williamsonia association with the fronds of Ptilophyllum (Sharma, collected from Amarjola and described the anatomy 1969a) both at Sakrigalighat and Dhokuti in the of and receptacle, structure of seed, Rajmahal Hills. The association of other fronds­ ontogeny and development of fruit. Sharma (1977) Otozamites, Pterophyllum, Anomozamites and published an illustrated review as Indian Dictyozamites with allied stems and fructifications Williamsonias. Sharma (l982a) interprets the are yet to be discovered. Similarly, the phylogeny of morphology of interseminal scales and derives reticulate venation of Dictyozamites needs Williamsonia from Cordaianthus as a result of investigation. modification and condensation of cone axis. Bose Sahnioxylon though resembles in the structure and Banerji (1984) described three new species of of tracheary elements with the stem Bucklandia, Williamsonia, viz., W. kakadbhitensis, W. specially B. dichotoma, its affinities remain doubtful trambuensis and W. sukhpurensis from Kachchh. All for want of complete and better preserved material these are incrustations and fertile structures remain to study the origin and nature of leaf traces and unknown. Identification is based mainly on structure of pith and cortex. Associated leaves and epidermal characters of bracts. In W. kakadbhitensis fertile parts of Sahnioxylon are yet to be discovered. and W. sukhpurensis the bracts are densely hairy The stem genus Bucklandia also needs further while trichomes/ramenta bases are rare in W. investigations as wide variations occur in the trambuensis. morphology and distribution of leaf bases on stem surfaces. Amarjolia Bose et al. In all the species of Weltrichia known from This bisexual fructification was originally India the microsporophylls originate from the brim described as Cycadeoidea dactylota by Bose of a cup-like receptacle. The abaxial whorl of sterile (1966b). Sitholey and Bose (1971) amplified the bracts is present in W. santalensis, while in others it description. The fructification is terminal and is yet to be seen. W. companulatiJormis and W. exposed, and the microsporophylls differ in harrisiana are based on the study of 1-3 incomplete structure from Cycadeoidea so the new genus specimens. Amatjolia was proposed by Bose, Banerji and Pal Though the seed-bearing Williamsonia is (1984) and called it A. dactylota. The bracts are hairy represented by several species (12 sp_), the basic like in W. sewardiana Sahni 1932b. There are nearly structure, i.e., presence of a compact layer of sterile 20 baloon-shaped microsporophylls, surrounding and fertile scales surrounding a receptacle, and the the central conical receptacle which bears a compact linear, simple, curved bracts which protect the fertile layer of seminiferous and interseminal scales. Each parts are identical. Despite the study of all internal microsporophyll bears appendages on which rows of details including anatomy, ontogeny and structure of microsynangia are produced. Our present seeds/ovules, and fruit development, the phylogeny knowledge on Amatjolia is based only on two qf the fructification remains doubtful. Though specimens and more collection is required for Sharma (l982a) derives Williamsonia from further investigations. Cordaianthus, the intermediate presumptions are In addition to the above described hypothetical and need proof. Further investigations fructifications, a number of incomplete (Sharma, of the petrified fructifications are likely to provide 1982b) or poorly preserved (Sharma, 1990) solution to the problem. bennettitalean organs and scales (Cycadolepis) are also known from the Mesozoic rocks of India (Bose REFERENCES & Banerji, 1984). Bose MN 1953a. Ptilophyllum amarjolense sp.nov. from (he DISCUSSION Rajmahal Hills. Proc. natn. Ins! Sci. India 19 : 605-612. Bose MN 1953b. Bucklandia sahnii sp. nov. from (he Rajmahal Hills. Palaeobotanist 2 : 41-50. The extinct representatives of the family Bose MN 1966a. Fossil plan( remains from (he Rajmahal Series in Williamsoniaceae dominated the vegetation during (he Upper Gondwana of fndia In Symposium on Floristics 264 THE PAlAEOBOTANIST

and stratigraphy oj Gondwanaland: 143·154. Birbal Sahni mahal Hills with notes on the distribution of the genus. Proc. Institute of Palaeobotany, Lucknow. natn. Inst. Sci. India 17: 7·13- Bose MN 1966b. A petrified bennettitalean flower from the Jacob K & Jacob C 1954. Cuticular studies of Indian Ptilophyllum Rajmahal Hills, India. Curro Sci. 35 : 569· 570. frond from Cutch and Jabalpur. Mem. geol Surv. IndIa Bose MN 1968. A new species of Williamsonia from the Rajmahal Palaeont. indica 33. Hills, India. Bot. j. Linn. Soc. Lond 61 : 121·127. Mittre V 1956. Studies on the fossil flora of Nipania, Rajmahal Bose MN & Banerji J 1981. Cycadophytic leaves from · Hills-. Palaeobotanist 5 : 95·99. Lower Cretaceous rocks of India. Palaeobotanist 28-29 : 218· Morris J 1840. Appendix in Grant, C. W. "Memoir to illustrate a 300. ReoloRical map of cutch" Trans. geol. Soc. ser. 2, 5 : 289 Bose MN & Banerji J 1984. The fossil floras of Kachchh·l. 329. Mesozoic megafossils. Palaeobotanist 33 : 1·189. Oldham T & Morris J 1863. Fossil flora of Rajmahal Series in the Bose MN, Banerji J & Pal PK 1984. Amarjolia dactylota (Bose) Rajmahal Hills. Fossil flora of Gondwana System. Mem. comb. nov.: a bennettitalean bisexual flower from the geol. Sum India Palaeont indica ser. 2, 1 (1) : 1·40. Rajmahal Hills, India. Palaeobotanist 32 : 217-229. Presl CB 1825. In Sternberg C Von Versuch einer geognostisch Bose MN & Kasat ML 1969. Fossil flora of the Jabalpur 5eries·4. botanischen Darstellungder Florader Vorwelt. III. Leipzig. Williamsonia seniana sp. nov. In Santapau H et al. Roy SK 1963. Studies on the Mesozoic flora of Kutch and Kathia· (editors)-j. Sen Mem. Vol. : 305·309. Botanical Soc. Bengal, war·2. Otozamites bellus sp. nov. Proc. 50th Indian Sci. Calcutta. Congr. 3 Abst. : 397. Bose MN & Kasat ML 1972a. The genus Ptilophyllum in India. Sahni B 1932a. Homoxylon rajmahalense gen. et sp. nov., a fossil Palaeobotanist 19 : 115·145. angiospermous wood devoid of vessels from the Rajmahal Bose . MN & Kasat ML 1972b. On a petrified specimen of Hills. Mem. geol. Surv. India Palaeont. indica n. ser. 20 : 1· Dictyozamites from the Rajmahal Hills, India. Palaeobotanist 19 19 : 248·250. Sahni B 1932b. A petrified Williamsonia (W. sewardiana sp. nov.) Bose MN & Sah SCD 1954. On Sahntoxylon rajmabalense: a new from the Rajmahal Hills. Mem. geol. Sum India Palaeont. name for the Homoxylon rajmahalense Sahni and a new sp. of indica. n.S. 20 (3) : 1·19. Sahntoxylon from Amarapara in the RaJmahal Hills. Palaeo­ Sahni B & Rao AR 1933. On some Jurassic plants from the Raj· botanist 3 : 1·8. mahal Hills. j. Asiatic Soc. Bengal. n.s. 27: 183·208. Bose MN & zaba·Bano 1978. The genus Dictyozamites Oldham Seward AC 1917. Fossil plants·III. Cambridge. from India. Palaeobotanist 25 : 79-99. Seward AC & Sahni B 1920. Indian Gondwana plants. A revision. Bose MN & zaba·Bano 1981. On a new species of Otozamites from Mem. geol. Sum India Palaeont. indica, n.ser., 7 : 1·41. Kachchh, western India. Palaeobotanist 27 : 227- 231. Sharma BD 1967a. Investigations on the Jurassic flora of Raj· Feistmantel 0 1876. Jurassic (oolite) flora of Kutch. Fossil flora of mahal Hills, India 3. A review on the genus Ptilophyllum Gondwana System. Mem. geol. Surv. India Palaeont indica 2 Morris with description of rwo new species from Amarjola in (1). the Rajmahal Hills. Palaeontographica 1208 : 139·150. FeistmantelO 1877. Jurassic (Uassic) flora of the Rajmahal Group Sharma BD 1967b. Investigations on the Jurassic flora of Raj­ in the Rajmahal Hills. Mem. geol. Surv. India Palaeont. mahal Hills, India 4. On a new species of Indian Bucklandia, indica 1 (2). B. guptai with remarks on B. sahnii of Bose. Ameghiniana Feistmantel 0 1879. Upper Gondwana flora of the outlier on the 5 : 35-46. Madras coast. Fossil flora of Gondwana System. Mem. geol. Sharma BD 1968. Investigations on the Jurassic flora of the Surv. India Palaeont. indica ser. 2, 1 (4): 191· 233. Rajmahal Hills 5. Epidermal studies in the bracts of rwo new Gupta KM 1934. On the wood anatomy and theoretical signifi· species of Williamsonia, W. guptai and W. amarjolense. Acta cance of homoxylous angiosperms. j. Indian bot. Soc. 13 : bot. Hung. 14 : 373·383. 71·101. Sharma BD 1969a. Further observations on Williamsonia santa· Gupta KM 1943. A new species of Williamsonia (W. sahnii) from lensis Sitholey & Bose with description of a new species. the Rajmahal Hills. j. Indian bot. Soc. 22 : 191-199. Palaeontographica 1258: 93·103. Gupta KM 1958. Williamsonia fruCtifications from the Jurassic Sharma BD 1969b. On some fossil cycadean fronds from India. of Rajmahal Hills-their preservation and plan of construc­ Bull. bot. Sum India 11 : 115·119. tions. j. palaeont. Soc. India 3 : 230· 232. Sharma BD 1970a. On the structure of Williamsonia cr. W. scottica Gupta KM 1971. Investigations on the Jurassic flora of Rajmahal from the Middle Jurassic of Rajmahal Hills, India. Ann. Bot. Hills, India·9. On the struCture and affinities of Sewardioxylon 34 : 289· 296. sahnii gen. et sp. nov. Gupta.-a petrified cycadean wood Sharma BD 1970b. On the vascular organization in the receptacle from the Rajmahal Hills, India. Palaeontographica 1318 : of seed bearing williamsonias from the Middle Jurassic of 160·166. Rajmahal Hills, India. Ann. Bot. 34 : 1063-1070. Gupta KM & Sharma BD 1964. Investigations on the Jurassic flora Sharma BD 197Oc. On the struCture of the seed of Williamsonia of Rajmahal Hills, India l. On the bennettitalean genus collected from Amarjola in the Rajmahal Hills, India. Ann. Dictyozamites with the description of a new species D. Bot. 34: 1071·1077. sahnii. j. palaeontol. Soc. India 5-9: 21·28. Sharma BD 1970d. Bucklandia dichotoma sp. nov. from the Gupta KM & Sharma BD 1968. Investigations on Jurassic flora Middle Jurassic of Rajmahal Hills, India. Ameghiniana 6 : of Rajmahal Hills·2. On a new species of Ptilophyllum-P. 303·308. sahni('from Amarjola in Amarpara region. j. palaeont. Soc. Sharma BD 1974. Ovule ontogeny in Williamsonia Carr. Palaeo· India 10-11 : 1·7. nlographica 1488 : 137·143. Harris TM 1969. The Yorkshire Jurassic Flora III. Bennettitales. Shanna BD 1975. Further observations on Williamsonia sewar· Brit. Mus. (Nat. Hist.) Lond 675: 1·186. diana Sahni collected from Amarjola in the RajmahaJ Hills, Hsu J & Bose MN 1952. Further observations on Homoxylon India. Casop. Mineral. Geol. 20 : 31· 37. rajmahalense Sahni. j. Indian bot. Soc. 21. Sharma BD 1976. Fruit development in Williamsonia Carr. pcob K 1951. Dictyozamites bapjorensis sp. nov. from the Raj· (Bennettitales). Geobios 9 : 503·507. SHARMA-INDIAN WlLLIAMSONIACEAE-AN OVERVIEW 265

Sharma BD 1977 Indian wi II iamsonias-an ill uSlrated review. Sitholey RV 1954. "The Mesozoic and Tertiary floras of lndia­ ACla palaeobol. 18: 19- 29. a review_ Palaeobolanisl 3 : 55-69- Sharma BD 1980. Micropyle in Williamsonia Carr. (Bennellilales) Silholey RV & Bose MN 1953. Williamsonia samalensis sp. nov., Ann 801. 45 : 91-95. a male fructification from the Rajmahal Series with remarks Sharma BD 1982a. Morphology of imerseminal scales in William­ on the structure of Ontheanlhus polyandra Ganju. Palaeo­ sonia Carr. (Bennellitales). Indian I Earlh Sci 9: 1·5 bOlaniSI 2 : 29· 39. Sharma BD 1982b. A petrified picullar bennettitalean fructi· Sitholey RV & Bose MN 1971 Wellrichia sanlalensis (Sitholey fication from Amarjola in the Rajmahal Hills, India. ACla & Bose) and other bennettitalean male fructifications from palaeobol. 12 : 165·170. India. Palaeontograpbica 1318: 151·159. Sharma BD 1990. Morphology of bennellilalean fruClificalions. Sukh·Dev & Rajnikamh A 1988a. "The Gangapur Formation: fossil Palaeobolanisl 38 : 163-170. flora and stratigraphy. Geopbylology 18: 1·27. Sharma BD, Surana AC & Singh AP 1971. Jurassic plants from Sukh-Dev & Rajnikamh A 1988b. The Shivaganga Formation: Amarjola in the Rajmahal Hills, India. I palaeom. Soc. fossil flora and stratigraphy. Geopbylology 18 : 186· 205. India 16: 27·34