On Some Plant Fossils from the Pariwar Formation, Jaisalmer Basin, Rajasthan

ON SOME PLANT FOSSILS FROM THE PARIWAR FORMATION, J AISALMER BASIN, RAJASTHAN

HARI K. MAHESHWARI Birbal Sahni Institute of Palaeobotany, Lucknow-226007 & N. P. SINGH Oil and Natural Gas Commission, Jodhpur

ABSTRACT The Pari war Formation is named after the Pari war Village (27° 14'30": 70°44'30") The paper reports on plant megafossils found in the lower part of the Pari war (Parihar) Formation and consists of a poorly exposed but exposod in the Jaisalmer Basin, Rajasthan. The stratigraphically thick sequence of sandstone plant fossils belong to the groups Filicales, Cyca• beds. The formation in the north-east dales, Bennettitales and Coniferales. The identi• part around Pariwar Village consists mainly fiable species are: ?Gleichenites sp., Phlebopteris sp., Frond type I, Taeniopteris vitlata, T. densinervis, of pebbly, current-bedded sandstones T. spatulata, Pterophyllum sp., Otozamites imbricatus, but to the south-east it is difficult to Ptilophyllum acutifolium, Elatocladus conferta, establish the stratigraphy dUE to paucity ?Elatocladus sp. and Pagiophyllum sp. In general of the outcrops. The lower fossiliferous composition the fossil assemblage is more akin to that known from the Kutch Basin of Gujarat, member consists mostly of variegated and is probably Upper Jurassic in age. (yellow, yellowish-white, brown and ochreous brown) siltstone, sandstone and shale/clay beds of considerable thickness. Das-Gupta INTRODUCTION (1974) assigned a Lower Cretaceous (Neoco• mian) age to the formation on the basis of subsurface stratigraphical correlation in this area. However, the present study of plant a numbervast Rajasthanof sedimentaryshelf basinsincludesof THEwhich the Jaisalmer Basin is the megafossils collected from the type area largest. In the north-north-west and south suggests an Upper Jurassic affinity of the of Jaisa1mer, is exposed a thick sequence Pari war Formation flora. of marine and continental sedimentaries The fossils are preserved as impressions of post-Triassic age on a truncated, pene• in ochre-yellow, argillaceous shale and in plained basement. The Mesozoic strata buff-coloured siltstone. Some of the comprising Lathi, Jaisalmer, Baisakhi, impressions are partly coloured red, prob• Pari war (formerly Parihar) and Habur ably due to leaching of iron. Preservation (formerly Abur) [ormations have till recently is usually poor but sometimes wetting the not yielded any p1ant fossils though a specimens with xylol or liquid paraffin brings mioflora of probable Lower Jurassic age out certain structures clearly. (Srivastava, 1966) is known from the Lathi Formal ion. DESCRIPTION During the 1972-73 field season, a field party of the Oil and Natural Gas Commission, Genus - Gleichenites Goppert 1836 of which the second author was a member, discovered plant megafossils in the lower Type Species - Gleichenites porsildii part of the Pari war Formation, east of the Seward 1926. Habur Village. The thick arenaceous sequence of Pari war Formation was earlier ? Gleichenites sp. described as the 'Parihar beds' by Oldham PI. 1, fig. 2 (1886) and 'Parihar Formation' by Swami• nathan, Krishnamurthy, Verma and Description - Small fragmentary speci• Chandiok (1956). men, leaf as a whole not known, pinnules 116 MAHESHWARI & SINGH - PLANT FOSSILS FROM PARI WAR FORMAT [ON 117 small, about 2 mm long, margin entire, 1968, p. 22) in having less oblique pinnae. apex: rounded, veins not preserved. Phlebopteris sp. of Roy (1968, p. 110, pI. 1, Remarks - Due to its extremely bad figs. 6-7, pI. 2, fig. 9) is known only from preservation the specimen is doubtfully a fragmentary specimen. Our specimens referred to Gleichenites. are further different from all the other phlebopterids in having a mucronate apex.

Genus - Phlebopteris Brongniart 1836 FROND TYPE-1 Type Species - Phlebopteris polypodioides Brongniart 1836. PI. 1, fig. 8; Text-fig. 1975 Cladophlebis sp. (?): Das Gupta et. al., p. 2::36,' fig. 3. Phlebopteris sp. PI. 1, figs. 6-7 Description - This lS another very common type in the present assemblage. Description - This is one of the more Frond bipinnate, main rachis winged, about common plants in this assemblage but 1·75 mm wide, pinna rachis less than 0·5 unfortunately preservation is extremely mm wide at base and gradually tapering poor. All the specimens are incomplete towards apex. Pinnae opposite, arising at and none shows the apical or basal parts an interval of about 9-10 mm and at an of the frond. Specimens pinnate, largest angle of 35°-45°. Most complete pinna specimen measuring 10 cm in length, rachis 5·5 mm long, lanceolate. Typical pinnules stout, up to 1·5 mm broad. Pinnae 5·5 to 6·5 mm long, slightly smaller in the alternate or subopposite, almost at right upper pinnae, apex narrow obtuse, pinnules angles to the rachis, straight or slightly arising at an angle of 30°-35° to the pinna falcate, adjacent pinnae confluent at base. rachis, almost straight, pointing forwards, Pinnae longer in the middle part of the leaf pinnule margin smooth, adjacent pinnules (up to 2·5 cm) and becoming smaller to• confluent at base. First pinnule arises on wards apical and basal parts; typically 2-5 the basiscopic side and sometimes is almost mm broad at the widest. Pinna apex on the rachis (Text-fig. 1). Each pinnule characteristically mucronate. Midrib pro• served with a distinct midrib which con• minent, persisting up to apex. Veins arising tinues up to apex, further secondary veins at an angle of approximately 75°-80° to not seen. the midrib, almost parallel, a few cross Remarks - Due to nonpreservation of connections observed in some specimens secondary veins, assignment of these speci• (PI. 1, fig. 6). mens to a genus is not possible. In habit Remarks - Harris (1961, p. 101, 112) these fronds resemble Pachypteris, Onychi• has remarked that the genera Phlebopteris, opsis, Scleropteris and Cladophlebis. Pachy• Matonidium and Matonia appear to pteris as instituted by Brongniart (1828), intergrade. Matonidium can be distin• is characterized by the absence of veins or guished from Phlebopteris only by its indusi um by the presence of only a single primary formed from the expanded end of the vein in the pinnules. Later st udies have placenta. In sterile condition the distinc• shown that Pachypteris is in its venation, tion between these two genera is difficult as well as in other characters, very much to make. We have provisionally referred like Thinnfeldia. Pachypteris, can further our specimens to Phlebopteris on the basis be ruled out altogether as the fronds of of their general similarity with the figured that genus are thick and coriaceous. specimens of Phlebopteris from India (see Scleropteris is an ill-defined genus, but the Bose &Sah, 1968,pI. 5,fig. 31; Sukh-Dev, 1970, midrib in that genus is usually indistinct or pI. 1, figs. 6, 7, 9) and due to occurrence of lacking. Onychiopsis has a typical venation. occasional anastomoses. Our specimens, Because of the welldefined midrib the however, differ from those of Phlebopteris specimens could belong to Cladophlebis but polypodioides Brongniart (in Sukh-Dev, 1970, the preservation being imperfect a definite p. 201) and Phlebopteris sp. (Bose & Sah, determination is not possible. 118 THE PALAEOBOTANIST

TEXT-FIGS. 1-5 MAltESHWARI & SINGH - PLANT FOSSILS FROM PARiWAR FORMATION 119

Genus - Taeniopteris Bron~niart 1832 then gradually tapers to form an acute apex, margin entire except in basal reGion where Type Species - Taeniopteris vittata Brong• it is slightly wavy. Midrib strong, about niart 1832. 3·5 mm wide at base and gradually narrow• Remarks - Nathorst (1909) described the ing towards apex, fine longitudinaUy running cuticle of a specimen which he believed to striations seen. Secondary veins arising be an example of Nilssonia tenuinervis. from midrib almost at right angles, slightly As it had sinuous cell walls he created a new curved forwards almost reaching margins, genus Nilssoniopteris for it. Thomas (1913, simple or occasionally once-forked. p. 241) found, on re-examination, that thi.s Remarks - This species has been reported particular specimen resembled ~ore .Taen:• from Kutch under the name Oleandridium opteris vittata. The genus N~lssomopterzs (Taeniopteris) vittatus (Feistmantel, 1876, was therefore later abandoned (Thomas & p.15, pI. 1, figs. 1-3; pI. 2, figs. 1-5). Bancroft, 1913; Seward, 1917). H(~.rris (1932) and Florin (1933a, 1933b) recogmzed the need for separate genera for taeniop• Taeniopteris densinervis Feistmantel 1876 terid leaves with different types of cuticle. PI. 1, fig. 5; Text-fig. 4 While Harris (1932) created a new genus Taeniozamites for taeniopterid leaves with Description - Simple leaf, oval spathu• bennettitalean cuticle, Florin (1933a, 1933b) late, base and apex not preserved, incom• retained Nilssoniopteris for such leaves. plete specimens 4'6-7·9 cm long and 1·6-3·1 We think that it is prudent to retain cm broad at the widest. Midrib stout, 1·5 Nilssoniopteris for forms with bennet.titalean mm broad at base and gradually tapering cuticle as cuticle of the type specImen of towards apex, finely pitted. Veins arise the type species of the genus Taeniopteris from midrib at angles of 75°-80° and then (i.e. T. vittata) is probably not known. curve sharply forwards meeting the margins Further under prevalent botanical practices, at low angles, mostly simple, rarely once• the species vittata (with same holotype) must forked. At one or two places there is slight not be taken as type for two similar though appearance of an anastomosis but it could unrelated genera, i.e. Taeniopteris and as well be a preservation artefact, concen• Nilssoniopteris. It is suggested that vittata tration of veins 22-24 per cm. should be retained as type species of the Remarks - In general appearance and the genus Taeniopteris and that a new type angle of emergence of the veins, the figured species should be designated for the genus specimen resembles those of Taeniopteris N ilssoniopteris. densinervis from Kutch (Feistmantel, 1876, pI. 2, fig. 6). Taeniopteris vittata Brongniart 1832 PI. 2, figs. 9, 12 Taeniopt6ris spatulata McClelland 1850 PI. 2, fig. 13; Text-fig. 5 Description - Simple leaves, linear-Iance• olate in shape, complete leaves not preserved Description - Leaf simple, narrow, strap• but probably up to 14 cm long and 2 cm shaped, apex and base broken, incomplete broad at the widest. Leaves petiolate, specimen measures 3·6 cm in length and petiole base swollen, lamina gradually in• 0·6 cm in maximum width. Lamina margin creasing in width from the petiole, reaches entire. Midrib fiat, 1·0 mm broad. Veins maximum width in the middle region and arise almost at right angles to the midrib,

+-- TEXT-FIGS.1-5 - 1, outline drawing of a part of frond to show the basal pinnule, and the midveins of the pinnules X 2. 2, Otozamites imbricatus Feistmantel 1876, line c'rawing of the specimen to show pinnule base and venation X 2. 3, Ptel'ophyllum sp., line drawing showing the nature of venation X 2. 4. Taenioptel'is densinel'vis Feistmantel 1876, line drawing to show venation X 2. 5, Taenioptel'is spatulata McClelland 1850, X 2. i20 THE PALAEOBOTANisl'

are simple and meet the margin at wide on this character, the difficulty is obvious." angles. This distinction in the relative width of Remarks - In its shape and venation the pinnae of the two genera is not a satis• pattern the s:)ecimen shows some resemblance factory distinguishing criterion, in the ab• to Taeniopteris spatulata McClelland (1850, sence of cuticular features, for in some species pI. 16, fig. 1; see also Oldham & Morris, of Anomozamites the segments are not 1863, pI. 6, figs. 1-6). necessarily as long as broad [e.g. Anomo• zamites nilssonii (Phillips) Seward in, Harris, 1969, fig. 37A, DJ. Further, so far, there Genus - Pterophyllum Brongniart 1828 is no report of the presence of true Anomo• zamites in India. Anomozamites Jissus may Type Species - PterophyUum longiJolium be a Taeniopteris (Bose, 1974b, p. 190). Brongniart 1828. Anomozamites amarjolense of Sharma et al. (1971, p. 32, pI. 1, fig. 5) is a Pterophyllum leaf. A negative evidence of the absence PterophyUum S1'. of Anomozamites in India is the nonoccur• PI. 1, figs. 3, 4; Text-fig. 3 rence of WielandieUa, a fructification usually associated with Anomozamites. Therefore, DfJscription - Leaf pinnate, as a whole depending upon the external morphology linear, strap-shaped, 11 em long and 1·6 alone, we provisionally refer our specimen em wide, base and apex not preserved. to PteropltyUum. In the absence of cuticular Rachis thick, 4 mm wide in basal region, features a satisfactory specific assignment gradually narrowing towards apex, with of the present leaf is not possible. The two longiturlinally running mar[;inal ridges. leaf, however, shows some resem• Pinnae alternate, of unequal breadth, irre• blance with a leaf figured as PterophyUum gular rectangular to more or less triangular, (Anomozamites) inconstans (Braun) Zeiller sometimes falcate, attached by the whole by Du Toit (1927, p. 379, pI. 24, fig. 1), and base, basal segments smaller in size than as A nomozamites inconstans Schimper by those in the middle, segments attached Saporta (1875, pI. 9, fig. 3). laterally on the upper surface of the rachis, hence in fossil conclilion seem to slightly Genus - Otozamites Braun in Munster 1843 overlap the rachis, typical segments 5-7 mm long and 4-7 mm wide, apex occasionally notched. Veins usually simple, rarely Type Species - Otozamites (Filicites) bechei forked, parallel to the edge of the lamina, (Brongniart 1825) Harris 1961; recom• ending at distal margin, and slightly curved mended by Harris (1969, p. 10). Andrews forwards, about 8 veins per centimetre. (1970, p. 147), on the other hand, designates Remarks - The features found in this Otozamites (Otopteris) obtusus (Lindley & Hutton 1834) Brongniart 1849 as the type specimen occur individually or collectively of Otozamites. in certain cycadalean or bennettitalean leaves, viz., Ctenozamites Nathorst 1886, Anomozamites Schimper 1870 and Ptero• Otozamites imbricatus Feistmantel 1876 phyUum Brongniart 1828. The present specimen, however, can not be referred to PI. 2, figs. 10-11; Text-fig. 2 Ctenozamites as in that genus the main rachis may fork in the middle region of the leaf Description - Overall shape of the leaf and the pinnae arise on the upper side of lanceolate, with tapering base, apical part the pinna rachis. Further the veins in not known. Incomplete specimen 9·5 em Ctenozamites are much forked. Differences in length and 2·6 em in width at the between Anomozamites and PterophyUum broadest. Leaf pinnate, pinnae alternate, are not dearly marked and according to linear-Ianceolate, almost straight or slightly Harris (1969, p. 79) "The sole distinction falcate, their long axis at an angle of about from PterophyUum is in the shape of the 60°. Pinna base asymmetrical, auriculate, lamina segment which in Pterophyllum are auricle developing at acroscopic angle. typically much longer than broad. Thus Pinnae in middle region 1·5-1·7 em long, when a specimen is not satisfactorily placed veins conspicuously marked, diverging right MAHESHWARi & SiNGH·- PLANT FOSSILS FROM PARIWAR FOR,IATION 121

from the point of attachment, forking 1-2 apparently in two rows in dorsi ventral times, about 11-13 veins in the middle of shoots. A faintly marked midrib occasion• each pinna, mostly veins ending in lateral ally seen in some leaves. Leaf apex obtuse, margins, a few reaching the acute apex. base twisted, contracted and ?decurrent. Remarks - The specimen is morpho• Remarks - So far Elatocladus conferta was logically ident ical with the ones reported supposed to be a dorsi ventral type and any by Feistmantel (1876, p. 48, pI. 8, fig. 1) specimen showing dimorphism was placed and by Bose (1974 p. 100, pI. 46, figs. 1,2) in E. heterophylla. Specimens in the present from Kutch. The cuticular structure of this assemblage clearly show the occurrence of species was described by Roy (1965, p. 215, dimorphism in E. conferta also. In fact pI. 1, figs. 3-5). Otozamites imbricatus was there is not much difference bet ween E. so far not known from any area other than conferta and E. heterophylla, except that Kutch. in the latter species the leaves are narrowly linear and sharply acute (Halle, 1913, p. 84). It is possible that both species refer Genus - Ptilophyllum Morris in Grant 1840 to the same plant. E. australis Frenguelli (1944, p. 543, pI. 1, figs. 1-3) is also probably Type Species - Ptilophyllum acutifolium conspecific. Morris in Grant 1840.

? Elatocladus sp. Ptilophyllum acutifolium Morris in Grant 1840 PI. 2, figs. 16a, 17 PI. 1, fig. 1 Description - Fragmentary specimen with Description - Leaves pinnate, apical and dorsiventral symmetry. Leaves lanceolate, basal parts not preserved, rachis stout, almost straight, spirally arranged, disti• about .2 mm broad. Pinnae alternate, chous, with indications of a number of more obliquely attached by whole base, central or less parallel veins. axis of pinnae forming an angle of 75°-80° Remarks - The specimen shows a certain to the rachis, linear-lanceolate, almost resemblance to a figure of Elatocladus dick• straight or slightly falcate, apex broadly soniana (Heer) Seward (1926, p. 108, pI. acute, base apparently decurrent. 10, fig. 91). Remarks - The specimens compare well with the figured specimens of Ptilophyllum acutifolium (see Oldham & Morris, 1863, Genus - Pagiophyllum Heer 1881 pI. 20, figs. 1-2; Feistmantel, 1877, pI. 2, fig. 2; Bose & Kasat, 1972, pI. 1, figs. 1-6, Type Species - Pagiophyllum czrctnzcum pI. 14, figs. 115-116). (Saporata) Heer 1881.

Genus - Elatocladus Halle 1913 Pagiophyllum sp. PI. 2, fig. 16b Type Species - Elatocladus heterophylla Halle 1913. Description - Branched conifer twigs, largest shoot fragment 7·5 cm long. Leaves spirally borne, crowded, in the basal part Elatocladus conferta (Oldham & Morris 1863) look like obliquely inserted spines, arising Halle 1913 from a ?rhomboidal leaf base cushion at PI. 2, figs. 14-15 an angle of 40°-50° at base and spreading out in the top part. Leaves much longer Description - Branched coni fer shoots, than broad, more or less triangular to linear, branches showing a radial and/or a dorsi• sometimes falcate, keeled, apex acute, marg'in ventral symmetry. Leaves linear, linear• entire. lanceolate or oblong in shape, rarely falcate, Remarks - The specimens show some usually obliquely inserted, spirally arranged, resemblance to Pagiophyllum divaricatum adpressed in radial shoots and spread out (Bunbury) Sahni (1928) in having smaller, 122 THE PALAEOBOTA rist

spine-like leaves in the basal portion. How• flora is more akin to that known from ever, in the upper half our specimens show the Kutch region of Gujarat. Following longer and more spreading leaves as in P. plants are common between the two: Taeni• marwarensis Bose & Sukh-Dev (1972), but opteris vittata, T. densinervis, Otozamites this species is primarily based on cuticular imbricatus, Ptilophyllum acut1jolium, Pagio• features. phyllum sp., and Elatocladus conferta. Taeniopteris vittata is characteristic of the DISCUSSION Inferior Oolite of England and is also recorded from the Jurassic of Soviet Union, The megafloral assemblage from the Arctic regions, Poland, Japan, China and Jaisalmer District, Rajasthan c.escribed in Australia etc. the previous pages comprises species belong• The present megaflora seems to fit in the ing to the groups Filicales, Cycadales, Pagiophyllum - BrachYPhyllum Assemblage Bennettitales, and Coniferales. Filicales are sub-zone of the Ptilophyllum Assemblage represented by Gleicheniaceae (?Gleichenites zone. This sub-zone is assigned on Upper sp.), Matoniaceae (Phlebopteris sp.) and an Jurassic age by Sah, Singh and Sastry (1971, unidentifiable fern. The Bennettitales are table 3). not very common and are represented by Ptilophyllum and Otozamites. The Cycadales are represented by Taeniopteris. Arauca• ACKNOWLEDGEMENTS riaceae are represented by Pagiaphyllum while Elatocladus probably represents the To Shri S.N. Talukdar, Director of Geo!ogy, Podocarpaceae. Oil & Natural Gas Commission, Dehradun, The total absence of the Nilssoniales and thanks are due for permission to work out Dictyozamites, and rarity of Pterophyllum this material at Lucknow and to publish show that the beds studied are younger it. Dr. K. R. Surange, Director, Birbal than Lower Jurassic in age. On the other Sahni Institute of Palaeobotany, Lucknow hand, the total absence of Weichselia kindly granted laboratory and library reticulata and Onychiopsis paradoxus shows facilities for this work at the Institute. that the beds are older than the Lower Thanks are also due to Dr M. N. Bose, for Cretaceous. In general composition the critically going through the manuscript.

REFERENCES

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EXPLANATION OF PLATES

PLATE 1 distinct petiole, robust midrib and wavy lower margin of the lamina, X nat. size. 1. Ptilophyllum acutifoliunl Morris 1840, X nat. 10. Otozamites imbricatus Feistmantel1876 emend. Size. Roy 1965, X nat. size. 2. ?Gleichenites sp., X 4. 11. Otozamites imbricatus, a portion of specimen 3. Pterophyllum sp., X nat. size. in figure 3 enlarged to show pinnule base and 4. Pterophyllum sp., counterpart of specimen in venation, X 4. figure 3, X ca 1.2. 12. Taeniopteris vittata, note the acute apex in 5. Taenioptens densinervis Feistmantel 1876, left hand specimen, X nat. size. X nat. size. 13. Taeniopteris spatulata McClelland 1850, X 6. Phlebopteris sp., two pinnules enlarged to show nat. size. the distinct midrib and faint secondary veins, X 4. 14. Elatocladus conferta (Oldham & Morris 1863) 7. Phlebopteris sp., X nat. size. Halle 1913, X nat. size. 8. Frond type 1, X nat. size. 15. Elatocladus coniferta, X nat. size. 16. a:? Elatocladus b: Pagiophyllum sp., sp., PLATE 2 X nat. size. 17. ? Elatocladus sp., note several parallel veins 9. Taeniopteris vittata Brongniart 1832, note the in each leaf, X 4.