A REVISION OF THE LOWER GONDWANA SPHENOPTERIS FROM

P. Ie MAITI1Y Dirbal Sahni Institute of Palaeobotany, Lllcknow

ABSTRACT Calcutta, and the specimens described by OCCurrence of Sphenopteris Brongn. is long Srivastava (1954), Maheshwari & Gyan known from the Lower Gondwana Formations of Prakash (1965), Kar (1968), Maithy India. Detailed morphological studies of these (1969) and Kulkarni (1970) preserved at fronds have revealed that they differ distinctly the Birbal Sahni Institute of Palaeobotanv, from Sphenopteris and represent a new morpho• Lucknow. In addition to this 300 haIid logical type. Hence, a new genus Neomariopteris has been instituted to accommodate these fronds. specimens collected from the Lower Gond• The previous records, Neomariopleris (Sphenop• wana Formations of , teris) polymorpha (Feistm.) n. comb., N. (5.) hughesi , South Karanpura Coal• (Zeil!.) n. comb. and N. (5.) lobi/olia are redefined and a new species Neomariopteris talchirensis is field, Auranga Coalfield, Daltonganj Coal• instituted. field, Ib-river Coalfield and Tattitola be s (Rajmahal Hills) have been examined. The INTRODUCTION specimens are commonly preserved in form of impressions, however, in many cases a carboniferous crust is preserved. A cellodion wan asfrondsof Indiafromshowingthe Lowerdeep cutGond•or pulls of plant substances often appears FERNlobed habit of the pinna have been brown and translucent under transmitted described under Sphenopteris Brongniart. light. Examination of pull under the micro• Three species of Sphenopteris, viz. S. poly• scope reveals various structural details. morpha Feistmantel (1876, 1881), S. hughcsi Maceration of leaf substances by usual (Feistm.) Zeiller (1902) and S. lobi/alia Schulze's method results complete dissolu• Morris (1845), are known from India. The tion of plant substance, however, in cases first nalfled two species are known from the one gets trace of cuticles. Barakar to the Raniganj Formations, where• as the third species is known only from the Raniganj Formation. DESCRIPTION In some recent collections well-preserved Neomariopteris gen. novo remains of Sphenopteris have been collected from a number of Lower Gondwana Coal• Generic Diagnosis -- Imparipinnate fronds, fields of India. The geological horizons of deeply divided, tripinnate or quadripinnate; these beds range from Barakar to Raniganj rachis winged, pinnules small to medium Formations. A critical study of these fronds size, shape variable, decurrent, attached to suggests that the forms up till now described rachis by broad base, apex variable (acute, under Sphenopteris are morphologically dis• obtuse or broadly rounded), pinnule margin tinct from typical forms of Sphenopteris entire, dentate, crenulate or undulated Brngn. and r~present entirely a new morpho• (lobed); median vein of each pinnule usually logical type. The same is discussed bere gives off simple or bifurcating nervules to• with a revision of previous records from the wards margin of pinnules (sphenopteroid Lower Gondwanas of India and other venation). countries. Genotype - N eomariopteris polymorpha (Feistm.) n. comb. MATERIAL AND METHODS • Generic Description - The shape of the complete frond is a isobilateral triangle, The observations presented in this paper because the petiole is continued as a central are based upon the examination of specimens shaft to the apex and bore its subdivisions earlier described by Feistmantel (1876, 1881, in pinnate manner (Text fig. IB). Near the 1882) and Zeiller (1902) preserved at the apical region the division of pinnule is not Museum of the Geological Survey of India, complete and one can count the number of 70 MAITBY·- REVISION OF THE LOWER GONDWANA SPHENOPTERIS FROM: INDIA 71 pinnules only by the emergence of mid• and then pass either singly or in groups veins laterally from the pinnae rachis. The to the tips or the 10bes of the pinnules." decurrent nature of the pinnules is well marked in all the species. The shape of The type species of Sphenopteris, viz. S. the pinnules and its apex is distinguishable adiantoides Schloth. (=S. elegal1s Brongniart, feature for the different species. Each in Sternberg, 1825) is a typical sphenop• pinnule has one distinct midvein. The teroid frond showing pinnules deeply lobed midvein arises at ± 450, from the point, with cueneate base. Pinnules are laterally where the lower margin of the pinnules joins free, i.e. non-decurrent, though the forms the pinnae rachis. The midvein gives lateral from the Lower Gondwana of India are nervules. The nervules passes to the margin referred to Sphenopteris because of its vena• either simple or bifurcating. Course of tion characters. However, the Lower nervules may be straight or flexuosus. Gondwana forms differ from Sphenopteris Comparison--Brongniart (1828) instituted because the pinnules are attached to the Sphenopteris to accommodate fern fronds pinnae rachis by their broad bases and are with the following characters: decurrent in nature. This feature has also " The pinnules contracted at base and been marked by previous workers. Even often attached by short stalks, usually Zeiller (1902, p. 7) while describing Sphenop• small, oval or oblong in outline and lobed teris hughesi has opined "Ces pinnules, or toothed or sometimes cut into narrow quelque peu arquees en faux, prennent acute or obtuse lobes. Midvein is straight alors une grande ressemblance de forme or flexuosus. The lateral veins depart at avec celles de certains Odontopteris du acute angle and dichotomise a few times terrain houiller, de l'Odontopteris minor

B A

TEXT-FIG. 1 -A'-Branching pattern in Mariopteris Kidston. B. Br"llching pattern in Neornariop• leris gen. novo 72 THE PALAEOBOTANIST

Brongniart, notamment, bien que la nerva• tion, totalement differente, ne permette pas de les confondre". Dueto these morphological peculiarities the Indian forms warrant their separation from Sphenopteris. In its decurrent nature of pinnules the Lower Gondwana forms are comparable to Alethopteris Sternberg, Alloiopteris H. Potonie, and M ariopteris Zeiller. Alethop• teris Sternberg' differs from the Gondwana form in possessing a distinct midvein in the pinnules arising at right angle from pinnae rachis and the lateral nervules are also emerging oppositely from midvein at + 90°. These nervules are either simple Or- branched into two. Alloiopteris is charac• terized by very small pinnules alternately arranged, decurrent and more or less united laterally, The veins entering the pinnule branches two or three times and the veinlets TEXT-FIG. 2 - AA complete pinnae of Mariop• teris showing dimorphic nature of pinnules. pass at a steep angle to the apex where they B.A complete pinnae of Neomarioptcris. terminate usually in three more or less prominent teeth., Beside this the fertile kaf ot Alloiopterzs (Corynopterts) is known. The Lower Gondwana forms show a close appearance to frond or narrow winged agreement both in t~e attachment. of t~e giving frail appearance to frond. pinl1ules and vcnatlOns to Marwpterts. 2. Outline of the pinnae: Ovate or lanceo• late or linear or obovate. !Iowever, M ariopter.is has dimorphic pinnul~s, l.e. basal, most pmnule on the postenor 3. Angle of attachment of pinnae to the side of pinnae is distinctly larger than the rachis: Pinnae may be attached at right other pinnules and is prominently bilobed. angles to the rachis or at an angle of 45° or This pinnule also receives two distinct mid• less than 45°. veins. Beside this the branching pattern in 4. Shape of the pinnules: Oblong, lanceo• both the fronds is different. In Mariopteris late, spathulate, ovate or obovate, etc. fronds, the petiole are forked near the top 5. Apex of the pinnules: It may be either into equal branches, which has a second acute, obtuse or broadlv rounded. dichotomy producing four branches int 0 6. Margin of the pirmules: It may be two pairs (Text-fig. 1A)whereas in the present entire, serrate, crenulate or undulated. specimens the petiole is continued as a central 7. Attachment of pinnules to pinnae shaft to the apex and bore its subdivisions rachis: Pinnules are either attached at right in pinnate manner (Text-fig. 1B). angles to pinnae rachis or obliquely to rachis. 8. Shape of terminal pinnule: The termi• MORPHOLOGtCAL CHARACTERS nal pinnule may be either broadly triangular FOR SPECIFIC DELIMITATION in shape or lanceolate shape or linear shape. 9. Course of midvein and lateral veins: The fronds of Neomariopteris show super• Veins may be either straight or flexuosus. ficial resemblances to one another. But a 10. Number of nervules in one pinnule. close and careful study of the morphological details of the fronds recorded in our collec• Neomariopteris polymorpha (Feistm.) tion shows that certain characters are n. comb. variable from one species to another species and they can be well utilized for the identi• PI. 1, Figs. 1-4; Text-fig. 3A fication of species. The following characters are recognized important for specific delimita• Synonymy tion: 1876 - Sphenopteris polymorpha Feistmantel, 1. Nature of rachis: The main rachis may p. 365, pI. XVI, figs. 5-7, pI. XVII, be either broadly winged giving a robust figs. 1-3. MAITHY - REVISION OF niE LOWER GONDWANA SPHENOPTERis FROM INDIA ,~

1881- Sphenopteris polymorpha Feistmantel, to pinnae rachis more or less at right p. 76-77, pIs. XVA, XVIA, XVIIIA, angle. The terminal pinnule (PI. 1, Fig. 3) fig. 3. is asymmetrical, broadly triangular in shape, 1905 - Sphenopteris polymorpha Arber, bigger than the adjoining lateral pinnules. Glossopteris flora, p. 131-133, text• The pinnule margin on the superior side is fig. 31. larger drawn than the inferior side. 1966- Sphenopteris polymorpha Surange, The specimen figured in PI. 1, Fig. 2 p. 66-67, figs. 36A-B. represents the middle or the lower portion Emended Diagnosis -- Fronds large, at of frond. The rachis is broadly winged least tripinnate. Rachis broadly winged, 3 mm broad. Pinnae are alternate or secondary rachis broad, emerge alternately subopposite arranged, ± at 90°. Pinnae or sub opposite from primary rachis at an rachis is distinct, 1 mm. broad. Pinnae angle of ± 90°. Pinnae ovate shape, rachis gradually diffuses into fine nervules arranged alternately, attached to secondary as it enters the basal region of terminal rachis at an angle of 70°-90°. Lateral pinnule. Lateral pinnules oblong shape, pinnules oblong shape, shows ± equal width decurrent at base (PI. 1, Fig. 4). Lateral in the entire length, 4-10 mm. long and margin fusion of two pinnules is ± t length 2·5 mm. broad, i.e. the length to breadth of the pinnules. Apex of the pinnules ratio of the pinnule is 2: 1; lateral pinnules broadly obtuse alternately arranged standing at right angles Comparison and Discussion - The speci• or slightly obliquely to pinnae rachis, mens earlier described by Feistmantel (1876, decurrent, attached by broad base, lateral 1881) under Sphenopteris polymorpha are now fusion of two pinnules margin is ± t length transferred to the new genus because of the of the pinnules from the base, apex broadly decurrent nature of pinnules and sphenop• rounded; margin entire; the lower margin teroid venation. of the pinnule shows strong curvature to• Kulkarni (1970) described a specimen from wards the apex, whereas the upper margin the Lower Nakari Seam (Barakar) of the is straight, terminal pinnules larger than South Karanpura Coalfield under Sphenop• the adjacent pinnules, triangular in shape, teris polymol'pha Feistm. On the basis of apex rounded. Median vein distinct, her examined specimens she expressed the emerges at 45° from the point where lower opinion that both the species S. polymorplza margin of pinnule joins the pinnae rachis, Feistm. and S. hughesi Zeiller (1902) are course straight, median vein gives fine identical, and proposed to merge the latter lateral nervules which dichotomises once species with the former species. According before reaching the margin. 12-14 nervules to her the only distinction between the two are recorded in one pinnule. species is that the pinnules are unlobed in Lectotype - 5165. Geological Survey of S. polymorpha and lobed in S. huglzesi. She India, Calcutta. had observed in her specimen that the Locality - Raniganj Coalfield, Bengal. pinnules are unlobed (d. S. polymol'pha Horizon - Raniganj Formation. type) at the apical portion of frond and Specimen Figured - 35020/1386, 35021/ lobed at the posterior side (d. S. lzughesi). 1386, Birbal Sahni Institute of Palaeo• The specimen of Kulkarni (l.c.) preserved botany, Lucknow. at the Birbal Sahni Institute of Palaeo• Locality - Raniganj Coalfield, West botany has been reexamined bv me. From Bengal. the examination of frond it appears to me Horizon - Raniganj Formation. that Kulkarni (l.c.) has failed to understand Description - 25 specimens of this type the variations present in complete fern are in the collection. Beside this the type frond from apex towards base. The present specimens of Feistmantel (1876) preserved study does not support the view of Kulkarni at the Geological Survey of India have been (l.c.) for the specific identification, rather examined. The specimens figured in PI. 1, confirms with the original specific identifi• Fig. 1 show the apical portion of frond. cation proposed by Feistmantel (1881), Both the impressions are incomplete at the Zeiller (1902) and Arber (1905). The present apical portion. The pinnae near the apex study has brought to light a number of are not completely lobed. Pinnules show distinguishable morphological characters distinct decurrent nature. Lateral pinnules present in between the two species. The oblong shape with obtuse apex and attached same has been discussed on p. 76 and Table 1. THE PAL-\EOBOTANIST

TEXT-FIG. 3 l\1.\lTHY - REYISION OF THE LOWER GONDW.i\.NA SPHENOPTERIS FROM INDIA 75

TABLE 1 - SHOWING COMPARISON OF MORPHOLOGICAL CHARACTERS OF SPECIES OF NEOMARIOPTERIS RECORDED IN THE PRESENT COLLECTION

Name of spccies N. polymol'pha N. liugltesi N. lvbifolia N. lalchircnsis lIIort'wl 'g;cal cliar,7C'el'S 1. Nature of rachis P.roadly winged Xarrowly winged XClrro\\"ly wingf'd Mediumly \vinged 2. Pinnae outline Linear Linear-lance olate Lanc{'o!ate• Ovate 3. Angle of attacl1m( nt of ± 90° ± 45° ± 45° ± 70° pinnae to rachis 4. Shape of pinnules Oh'ong Lanccolnj(' 010\"at(' O\"ate 5. Apex of pinnu!es Broadly rounded A,ut" ()I,tusc Acute' 6. Margin of pinnule s Entire Cr,"nuJatc Undll late d Entire 7. ,\ngle f)f attachm{ nt of At right angle OIJlique OI,liqll(, At right apgle pinnules to rinnae rat:hls 8. Shape of te rminal Iiroadly trian?"1ar Lanceolat" Linear Lanceolate pinnule 9. Numlxr of nernIle s in 12-H 5-7 8-12 12·16 one l'innule

According to present study the specimens Neonzariopten's lmghesi (Zeiller) n. comb. of Kulkarni (l.e.) rccorded from South PI. 2, Figs. 7-11; Text-fig. 3D,C Karanpura belongs to S. hughesi Zeiller. In view of this confusion it will be of utter importance to record here the pre• Synonymy 1877 Die1?sonia d. eoneina Feistmantel, p. viously reported specimens which confirm 198, figs. 10, 11. with the diagnosis of Ncomariopteris poly• 1881 Dicl/sonia h1lghesi Feistmantc1, Palae• morpha n. comb. ont., p. 78, pI. 23A, figs. 1, 3, 12, 12a, 13. 1876 Sphenopteris polymorpha Feistmantel, p. 356, pI. 16, fig. 5-7, pI. 17. 1902 Sphenopteris (Dieksonia) hughesi Zeiller, 1881 Sphenopteris polyrnorpha Feistmantel, p. 6, figs. 1, 2, 2a, 2d. p. 76-77, pI. 15A, fig. 3, pI. 16A bis, 1905 Sphenopteris hughesi Arber, p. 133• figs. 1-6. 135, text-fig. 32. 1922 Sphenopteris polymorpha Walkom, pI. 1966 Sphenopteris h1tghesi Surange, p. 66• 5, figs. 26, 26A. 67, figs. 37A-B. 1958 Sphenopteris polymorpha Archangclsky, 1970 Sphenopteris polymorpha Kulkarni, p. fig. 25, 26. 208-211, pI. 1, text-fig. 1-4. Emended Diagnosis - Fronds large, at The following are regarded distinct from least tripinnate. Rachis extremely nanow, Neomariopteris polymorpha: 1-2 mm, secondary rachis narrow emerge alternately, subopposite or ± opposite from 1902 ;;phenopteris polymorpha Arber, p. 12, primary rachis at an angle of 45°_60°. pI. 1, figs. 4-5. Pinnae linear in shape, attached alternately 1958 Sphenopteris polymorpha Archangelsky, to the secondary rachis at an angle of 45°. fig. 29. Lateral pinnules lanceolate shape, pinnules 1961 Sphenopteris polymorpha Lele, p. 71• show maximum width at their middle por• 72, pI. 1, fig. 5. tion, 2-6 mm. long and 1-3 mm. broad, 1969 Sphenopteris polymorpha Maithy, p. the length to breadth ratio of the pinnules 266-267, pI. 1, fig. 4. is 2: ], pinnules arranged alternate, arising

TEXT-FIG. 3 - Pinnules enlarged to show shape and \'enations. A. Neomariopteris polymorpha n. comb. B-C. Neomarioplel'is hughesi n. comb. D. Neorllm'iopleris lobifGlia n. comb. E. Neo• mariopteris lalchirensis sp. novo (_\11 specimens figured X 5). 76 THE PALAEOBOTANIST

obliquely from the rachis, attached by their the preservation is not very good but under broad base, pinnules near the base shows the binocular two layers of epidermis are lateral fusion, apex acute, margins crenulate, marked. Stomatiferous and non-stomati• both the lateral margins show equal curva• ferous surfaces. The stomatiferous surface ture, terminal pinnule lanceolate, longer is thick (PI. 2, Fig. 10). Epidermal cells are than broad, apex acute, margin crenulate; trapezoidal or polygonal in outline. Inter• a gradual increase in size of the pinnules is vening cell walls thick, the stomata at present from apex to base of the pinnae; certain places is well marked. They are venation thin, median vein arises from the haplocheilic with thickened lateral lamellae. point where the basal margin of pinnule The subsidiary cells are like ordinary epi• join, the pinnae rachis, median vein gives dermal cells, and the polar subsidiary cells fine lateral nervules, which show rare are not different from the lateral ones dichotomy before reaching to the margin. (PI. 2. Fig. 11, Text-fig. 4). Non-stomati• 5-7 nervules are recorded in one pinnae. ferous surface is thin (PI. 2, Fig. 9). Cells Lectotype - 5207 Geological Survey of are polygonal in outline. Intervening cell India, Calcutta. wall is thin. Locality - lharia Coalfield, Bihar. Comparison and Discussion - Feistmantel Horizon - Raniganj Formltion. (1881) described these fronds under Dicksonia Specimen Figured- 35022/1069, 35023/ concina Heer. The same was transferred by 1227, Birbal Sahni Institute of Palaeo• Zeiller (1902) to a new species of Spheon• botany, Lucknow. pteris, S. hughesi. Since the form shows Locality - Churulia pit, East Raniganj decurrent nature of pinnules, it is, therefore, Coalfield, West BenaI. proposed to transfer them under the new Horizon - Barakar Formation. genus. Kulkarni (1970) considered this Description - 100 hand specimens of this species similar to N eomariopteris (Spheno• type were examined. Beside this the type pteris) polymorpha. The present morpho• specimens described by Feistmantel (1881) logical study of fronds shows that both the and Zeiner (1902) were also examined. species are different. N. polymorpha is a The :.pecimen is a tripinnate frond. Pri• robust form, because of strong wide rachis mary rachis thin measuring 1 mm. broad. with bigger oblong pinnules, attached to Secondary rachises emerge at an angle of rachis at ± 900, pinnule apex is obtuse with 600 from primary rachis. They are arrang• entire margins. Contrary to this N. hughesi ed alternate or subopposite. Secondary is a weak form with a delicate narrow rachis. rachis is also narrow (1 mm.) like primary The form has small linear pinnules, attached rachis. Pinnae linear shape, varying size, to rachis obliquely, ± 450, pinnule apex is smaller towards apex and bigger towards acute with crenulate margins. Thus the proximal side. Pinnae attached to rachis two forms are morphologically different, alternately at an angle of 450• Pinnae at hence it is proposed here to keep N. hughesi the very extreme apex of the frond is un• as distinct from N. polymorpha. lobed. However, the subsequent pinnae It will not be out of place to list those show graduallobing re'Sulting in the formation previous records which agree with the diag• of pinnules. Pinnules show a variation in nosis of N. hughesi. They are as follows: their shape. Pinnules near the apical portion of frond small linear lanceolate shape with 1881 Dicksonia hughesi Feistmantel, p. 78, pI. 23A, figs. 1, 3, 12, 12a, 13. slight crenulation of pinnule (PI. 2, Fig. 7). 1882 D. hughesi Feistmantel, p. 28, pI. 12, The pinnules towards the posterior side are lancealate-spathulate shape, comparatively figs. 3, 3a, 3b. 1902 Sphenoptt-ris (Dicksonites) hughesi bigger in size and the margins of the pinnules Zeiller, p. 6, pI. 4, figs. 1, 2, 2a-2d. show pronounced crenulation (PI. 2, Fig. 8, 1965 S. hughesi Maheshwari & Gyan Text-fig. 4D). Veins fade out and arise at the point where lower margins of the pinnule Prakash, p. 120, pI. 1, fig. 10. join the pinnae rachis (PI. 2, Figs. 7, 8). 1970 S. polymorpha Kulkarni, p. 208-211, pI. 1, figs. 1-5, text-figs. 1-4. I t gives lateral nervules which dichotomise The following are recorded as distinct rarely before rcaching the margin of the from N eomariopteris hughesi: pinnules. Cuticle - Epidermal structure is seen in 1968 Sphenopteris hughesi Kar, p. 245, tra'1slucent pulls of pinnules. Although pI. 1, figs. 3, 4. MAITHY - REVISION OF THE LOWER GONDWANA SPHENOPTERIS FROM INDIA 77

Neomariopteris lobi/olia (Morris) n. comb. near the base. Th" lateral fusion of pinnules is about 1/5th length of the pinnule from PI. 2, Figs. 12-13; Text-fig. 3D base. Median vein of pinnule arises from the point where the lower margin of pinnules Synonymy joins the narrow pinnae rachis. Median 1845 Sphenopteris lobi/olia Morris in Streze- vein sinuate (PI. 2, Fig. 13) supplying simple lecki'" ew South Wales, p. 246, or bifurcating branches to margin. Bifur• pI. vii, figs. 3, 3a. cating branches are rare. 1847 Sphenopteris lobi/olia McCoy, p. 149. Comparison and Discussion - Due to the 1905 Sphenopteris lobi/olia Arber, p. 135-138, decurrent nature of pinnules the fronds pI. V, fig. 2, 3. earlier described under Sphenopteris lobi/olia 1954 Sphenopteris lobi/olia Srivastava, p. Morris are now transferred under the new 70-71, pI. 1, figs. 1-3. genus N eomariopteris. 1966 Sphenopteris lobi/oila Surange, p. 68• N eomariopteris polymorpha differs from 69, fig. 38A, B. N. lobi/olia in having broad and strong 1966 Sphenopteris lobi/olia Rigby, p. 128• rachis, oblong pinnules with entire margin 129, pI. 33, fig. 32. attached ± right angles to the pinnae rachis. (For other Synonymy see Arber, 1905) The present form shows a close comparison Emended Diagnosis - Frond" probably tri• to N. hughesi by the narrow rachis and pinnate, triangular in contour. Rachis ex• oblique attachment of pinnules. However, tremely narrow 1-2 mm. broad, winged. N. hughesi differs by lanceolate shape of Pinnae linear-Ianceolate shape, attached pinnules with acute apex, attached to the alternate to subopposite to the rachis at rachis by their broad base; whereas the an angle of 45°. Lateral pinnules obovate pinnules in N. lobi/alia are obovate with shape, pinnules show maximum width at obtuse apex and slightly contracted their middle portion, 3-8 mm. long and base. 2-4 mm. broad, pinnules arranged alternate, standing obliquely to the rachis, attached N eomariopteris falchirensis sp. novo by their contracted base, pinnules at the base are decurrent in nature; apex obtuse PI. 1, Figs. 5-6; Text-fig. 3E with slight lobed margin, both the margins show equal curvature, terminal pinnule Synonymy linear in shape, median vein of the pinnule 1876 Sphenopteris polymorpha (partim) sinuate, supplying simple or bifurcating Feistm., p. 356. branches to the margins. 8-12 nervules 1881 Cyathea d. tchihatcheffi Feistmantel, are found in one pinnule. p. 75-77, pI. 16A, figs. 1-2 & 4. Holotype - Morris 13,530 Mus. Geoi. Soc. 1969 Sphenopteris polymorph a Maithy, p. London. 266-267, pI. 1, fig. 4. . Locality - New South Wales, Australia. Diagnosis - Fronds tripinnate. Rachrs Horizon - Permo-Carboniferous. broad 3 mm. wide, winged. Secondary Specimen Figured - 35024/1384, Birbal rachises broad emerge alternately or sub• Sahni Institute of Palaeobotany, Lucknow. opposite from primary rachis at a wide Locality - Mahavir Colliery, East Rani• angle, i.e. 70°. Pinnae lanceolate-ovate ganj Coalfield. shape, arranged alternately, attached to Horizon - Ranigani Formation. the secondary rachis ± right angles. Lateral Description - The figured specimen is a pinnules 6-8 mm. long and 3-4 mm. broad, tripinnate frond. The extreme apical por• ovate shape, decurrent, standing ± at right tion is not preserved. Pinnae are arranged angles to the pinnae rachis, attached by alternately towards the basal portion and their broad base. Lateral fusion is about show a subopposite tendency towards t length of the pinnule from the base; apex apical part. The pinnae near the terminal acute, margin entire, lower margin of the portion are unlobed. The lobing increases pinnule shows strong curvature from base as we pass to the posterior side of frond towards apex, whereas the upper margin is (PI. 2, Fig. 12). Pinnules are attached ± 90° (straight) from pinnae rachis. Termi• obliquely (30° angle) to the pinnae rachis. nal pinnule lanceolate with acute apex. Pinnules obovate shape and closely placed. Venation distinct. Median vein somewhat Pinnules show lateral fusion of margins sinuous, dichotomises once before reaching 78 THE PALAEOBOTANIST

to the margin. 12-16 nervu1cs are recorded However, N. talchirensis differs by the in one pinnule. ovate shape of pinnules with acute apex Lectotype - 5169, Geological Survey of and lanceolate shape of terminal pinnules. India, Calcutta (PI. 16A, Fig. 4). N. hughesi n. comb. and N. lobi/olia n. Locality - Talchir Coalfield. comb. differs in having narrow rachis, Horizon - Barakar Formation. pinnules attached obliquely to the rachis, Figured Specimen - 33119/838, Birbal and in the different shape of the pinnule. Sahni Institute of Palaeobotany, Lucknow. Locality - Junction of Lobjee and Sada• baha rivers, Daltonganj Coalfield. REMARKS ON SPHENOPTERIS ALATA Horizon - Barakar Formation. (BRONGNIART) STERNBERG AND OTHER LOWER GONDWANA SPHENOPTERIS Description - The present description is based upon the specimens No. 5166 (Feistm., 1881, PI. 16A, Fig. 1),5167 (PI. 16A, Fig. 27) The species discussed here are based only and 5169 (PI. 16A, Fig. 4) preserved at the on the basis of literature. No referable Geological Survey of India, Calcutta. The specimens are available in our collections. best preserved specimen is 5169 which is a A good deal of nomenclatural confusion tripinnate frond, whereas 5166 shows the exists about the species Sphenopteris alata apical portion of frond and 5167 the middle (Brngn.) Sternberg, because many of the portion of frond. authors who have mentioned this fossil have The specimens figured here have earlier fallen into error in some form or the been described by Maithy (1969) under other. The main cause of this is due to Sphenopteris polymorpha from the Barakar the fact that Brongniart described and beds of Daltonganj Coalfield. The portion figured in 1828, two ferns with the specific of the frond figured here shows that the title alata: (1) Pecopteris alata (p. 361, pI. pinnae rachises are arranged alternately cxxvii) from New South Wales and at a distance of 0·5-0'8 em. Pinnae 0·5 em. (2) Sphenopteris alata (p. 180, pI. xlviii, to 3 em. long. Pinnae lanceolate, pointed fig. 4) from Germany. Sternberg (1820-38, towards apex. Pinnule small, 3-4 mm. Pt. ii, pp. 59 & 131) transferred the form long and 1-2 mm. broad, ovate shape, acute described under P. alata to Sphenopteris. apex, attached to pinnae rachis by their Therefore, in his work two different forms se ,sile wide base. Pinnules are decurrent in are described, one under (1) Sphenopteris nature. Margin of thp- pinTJules is entire. alata (Brngn.) and the other under (2) Sphe• Secondary veins emerge at acute angle, nopteris alata Brongniart. However, the dichotomous and flexuosus. German type (2) is now known as Sphenop• Comparison and Discussion - These speci• teris grandini (Goepp.) Schimper (1869, mens were first described by Feistmantel Vol. 1, p. 404). Thus the only Sphenop• (1876) under Sphenopteris polymorpha. Later teris with the specific title alata is the Feistmantel (1881, p. 75, 76) opined that Australian form earlier described under these specimens arc different from Sphenop• Pecopteris alata Brngn. Very little is known teris polymorpha and show a resemblance to about this form. The only specimens re• Cyathea tchihatcheffi. A reexamination of the corded are of Brongniart (1828) obtained specimens of Feistmantel (l.c.) and Maithy from the coal mines on the Hawkesbury (1969) shows that they conform with the river near port Jackson and the other speci• generic diagnosis of the new genus proposed men reported by McCoy (1847) from New here. The decurrent nature of the pinnules Castle, New South \-Vales. Dutoit (1932) at the base has also been marked by reported a specimen from a locality north• Feistmantel (Feistm. l.c. p. 76). Hence the west of Bergville, Natal (Lower Beaufort forms described under Cyathea d. tchihat• beds). All these specimens show decurrent cheffi and S. polymorpha Maithy (l.c.) from nature of pinnules and conform with the Daltonganj Coalfield are now transferred diagnosis of N eomariopteris. Since no refer- . under a new species of Neomariopteris, N. able material is in our collection, therefore, talchirensis. it is proposed to keep it under Sphenopteris The present form compares closely with till fresh material is collected or the type N eomariopteris poly morpha (Feistm.) n. comb. forms are examined. due to its winged rachis, and the pinnules The same holds true also for Sphenop• being attached ± 90° to pinnae rachis. teris hastata McCoy (1847), S. jlexuosa MAUHY - REVISION OF THE LOWER GONDWANA SPHENOPTERIS FROM INDIA 79

McCoy (l.c.), S. plumosa McCoy (l.c.) and kind permission to examine the type speci• S. germanus McCoy (l.c.) which are based mens. Thanks arc due to Mr. M. V. A. upon fragmentary specimens. Arber (1905, Sastry, Mr. S. C. Sah, Mr. Gopal Singh and p. 135) has considered them synonymous Mr. D. K. Roy of the Palaeobotany Sec• to S. lobi/alia Morris. tion of Geological Survey of India, Calcutta, for extending all the facilities to examine ACKNOWLEDGEMENT the type specimens. I am also thankful to Miss Indra Kumari Janveja for the Sincere thanks are due to Dr. G. C. preparation". of cuticles and drawings and Chatterji, the then Director-General, Geo• to Mr. N. C. Saxena for typing the manu• logical Survey of India, Calcutta, for the script ot present paper.

REFERENCES

ARBER, E. A. J. (1902). The Clarke collection South Rewa Gondwana Basin, India. Ibid. 10 of fossil plants from New South Wales. Q. (1 & 2) :69-83. J. geol. Soc. 58: 1-26. MAIIESHWARI, H. Ie & GYAN PRAKASH (1965). Idem (1905). Glossopteris flora, London. Studies in the Glossopteris flora of India. ARCHANGELSKY,S. (1958). Estudis Geologico Y 21. Plant megafossils from the Lower Gondwana Paleontologico del Bajo de la Leona (Santa exposure along Bansloi river in Rajmahal Cruz). De Acta Geol. Lillona T II: 5-133. hills, Bihar. Ibid. 13 (2): 115-128. BRONGNIART,A. (1828). Prodrome dune Histoire MAITHY, P. Ie (1969). Palaeobotany and strati• des Vegetaux fossiles. Paris. graphy of the coal bearing beds of the Dalton• DUTOIT, A. L. (1932). Some fossil plants from ganj Coalfield, Bihar. Ibid. 17 (3): 265-274. the Karoo System of South Africa. Anu. S. McCoy, F. (1847). On the fossil botany and Africa Mus. 28 (4): 369-393. zoology of the rocks associated with coals of FEISTMANTEL, O. (1876). On some fossil plants Australia. Ann. Mag. Nat. Hist. 20: 145, from the Damuda Series in the Raniganj coal• 226-298. field, collected by Mr J. Wood Mason. J. MORRIS, J. (1845). In Count Strezelecki's "Physi• Asiatic Soc. Bengal. XLV (2): 329. cal description of New South Wales and Van Idem (1881). The fossil flora of Gondwana Sys• Diemen's land", etc. London. tem. II. The flora of Damuda-Panchet Divi• SCHIMPER, W. P. (1869). Traite de Paleontologie sion. Palaeont. indica Ser. 12, 3 (2): 1• Vegetal, 1-3. 77. SRIVASTAVA,P. N. (1954). Studies in the Glossop• Idem (1882). The fossil flora of Gondwana teris flora of India. 1. Some new fossil plants System. The fossil flora of South Rewa from the Lower Gondwanas of Raniganj Coal• Gondwana Basin. Ibid. 4 (2). field. India. Palaeobotanist. 3: 70-78. KAR, R. Ie (1968). Studies in the Glossopteris STERNBERG, C. G. (1820-28). Versuch einer geog• flora of India. 36. Plant fossils from Barren nostioch botanischen Darstellung der Flora del' Measures succession of Jharia Coalfield, Bihar, Vorwelt. Leipzig. India. Palaeobotanist. 16 (3): 243-248. SURANGE, Ie R. (1966). Indian fossil Pterido• KULKARNI, S. (1970). Studies in the Glossopteris phytes. Delhi. flora of India. 40. Sphenopteris polymorpha VvALKOM(1922). Palaeozoic floras of Queensland. Feistm. (1881) emend from the Barakar Stage I. The flora of the lower and upper Bowen of South Karanpura Coalfield, Bihar, India. Series. Qld. geol. Surv. Pub I. 270: 1-64. Ibid. 18 (2): 208-211. ZEILLER, R. (1902). Observations sur quelques LELE, Ie M. (1961). Studies in the Indian Middle plantes fossiles des Lower Gonclwanas. Palae• Gondwana Flora. 2. Plant fossils from the onto indica N.S. 2 (1): 1-140.

EXPLANATION OF PLATES

PLATE 1 4. Pinnules enlarged to show venation and decurrent nature of pinnules, X 5. Neomariopteris polymorpha n. comb.

1. Apical portion of frond, X 1'5 (specimen no. 35020/1396, Raniganj Coalfield, Raniganj). Neomariopteris talchirensis sp. novo 2. Portion from the middle region of frond, X 1'5 (specimen no. 35021/1396, Raniganj Coal• 5. A portion of frond, X 1 (specimen no. 33119/ field, Raniganj). 838, Daltonganj Coalfield, Barakar). 3. A pinnae enlarged to show the terminal 6. The same enlarged to show venation and pinnule and lateral pinnules, X 3. shape of pinnules. X 5. 80 THE PALAEOBOTANIST

PLATE 2 10. Epidermal structure of stomatiferous surface, X 100. Neomariopteris hughesi n. comb. 11. A stomata and subsidiary cells enlarged (S = stomata). X 500. 7. A portion of frond showing the arrangement of pinnae and the decurrent nature of pinnules, x 5. Neomariopteris lobifolia n. comb. (specimen no. 35022/1069. Raniganj Coalfield). 8. Median portion of frond is enlarged to show 12. A portion of a frond showing arrangement pinnules and venation, x 5 (specimen no. 35023/ of pinnae, X 1'5 (specimen no. 35024/1384, Rani• 1227. Raniganj Coalfield, Barakar). ganj Coalfield. Raniganj). 9. Epidermal structure of non-stomatiferous 13. Portions of pinnae enlarged to show the surface, X 250. nature of pinnules, X 5. MAITRY - PLATE 1 THE PALAEOBOTANIST, VOL. 21

/ M.-\1THY .- PLATE 2 THE PALAEOBOTANIST, VOL. '21