FLORISTICS OF THE PERMIAN AND TRIASSIC GONDWANAS OF INDIA
HARI K. MAHESHWARI Birbal Salmi Institute of Palaeobotany, Lucknow-226007
ABSTRACT Indian Peninsula The boulder bed is over• A synthesis of the floristics through the Upper lain by green laminated shales and sand• Palaeozoic and Lower Mesozoic Gondwanas of stones showing cold climate. The climate India is presented. Definite lycopsids are known gradually warmed up as is seen from the only from Middle Permian but could have been present in other ages as well.. The solitary spheno• diver~ification of the flora. The succeeding phyll SpeClOSranges from Middle Permian to basal PermIan strata contain thick coal seams Triassic. Equisetales are known by vegetative which were laid down in a warm humid shoots only, but a change in form through the time is noticed. Fertile filicinean remains known arC! climate. The early Triassic, rocks contain mostly referable to the family Asterothecaceac of undecomposed felspar which supposedly the Marattiales. The most important element of indicates arid conditions (Wadia, 1975). the period, i.e. Glossopteris-complex reprosents at The younger Triassic has thick red sandstone least two orders, viz., Ottokariales and Lidgetto• niales, as is evident from the fertile organs. This deposits which are supposed to denote complex continues into the basal Triassic and is arid desertic conditions. Robinson (1967, gradually replaced by Mosozoic 'Pteridosperm' p. 230) suggests a monsoon type of climate families Corystospermaceae and Peltaspermaceae. for the Indian Triassic. A classification Cordaltalea:l type of leJ.ves are more common in the Lower Permian; their affinity, however, is of the Permian and Triassic Gondwanas uncertain. Cycadales, Ginkgoales and Coniferales of India is shown in the table (slightly though present are meagrely represented. modified from Shah, Singh & Sastry, 1971). Plant fossils are quite abundant in the INTRODUCTION Permian and Triassic systems of the Indian Gondwana Sequence. These are of great biological interest as they comprise certain plant groups which are not represented is foundPalaeophyte-Mesophytepreserved in a thick insequenceIndia THEof mostly continental deposits known in the contemporary floras of the Northern as the Gondwana Sequence. This enormous Hemisphere. In the period under review, sequence of rocks ranges in age from the two distinct floras have been found. The Lower Permian (may be Upper Carbon,i• older, Glossopteris flora is characteristic ferous) up t.o the Lower Cretaceous (in• of the Permian though it extends into the clusive). Parallel sequences of continent.al basal Triassic. The younger, the Dicroidium deposit.s are also kno~n from the Southern flora is confined to the Triassic System only. Hemisphere cont.inents, viz., Australia, The plant groups represented in the two Africa, South America and Antarctica. It is floras are: ?Bryophyta, Lycopodophyta, supposed that the rocks of t.he Gondwana Arthrophyta, Pterophyta, Glossopterid 0• Sequence in India were deposited in slowly psida, ' Pteridospermophyta " Cycadophyta, sinking faulted troughs fed by rivers of the Ginkgophyta and Coniferophyta. In addition Gondwana country (Wadia, 1975, p. 165). there are certain plaJl,ts whose taxonomic In a few instances the deposits could have position is uncertain. been laid down in lakes. The sequence starts with a boulder bed, containing the BRYOPHYTA characteristically glaciated, striated and facetted blocks of rock embedded in a fine This group is probably represented by a silt-like matrix, suggesting a fluvio-glacial solitary capsule-like structure, Capsulites agency of transport. and deposition. The gondwanensis Saksena, obtained from a boulder bed forms a conspicuous and charac• Lower Permian shale. According to Saksena teristic datum line in the geology of the (1958), the distinct division of this structure 146 THE PALAEOBOTANIST
into foot, seta and capsule, the possibility dendron by Hislop, and later as " Stigmaria? of thE' presence of an annulus and the (Portion of the Rhizome of a Fern?)" by indication of a spore-sac extending along Bunbury (1861, pI. 12, fig. 2). The species the entire length of the capsule places it is heterosporous, its sporophylls being distri• near the Muscineae. The exact affinities buted on normal leafy shoots, without any of the specimen, however, still remain organized cone (Krausel, 1961). In the uncertain. There is a distinct possibility latter character it resembles the Lower that this structure represents a seed. Carboniferous Lepidodendropsis Lutz, the Upper Carboniferous Pinakodendron \iVeiss and probably Omphalophloios White. This LYCOPODOPHYTA could probably represent an unique line of evolution from the Lower Carboniferous Lycopsid megafossils in the Permian• to the Permian or may be that the four Triassic of India are extremely rare. The genera evolved independently of each other oldest records of the lycopsids are from the (Chaloner & Boureau, 1967, p. 510). Lower Permian Gangamopteris beds of Zewan From the Triassic of Beli, South Rewa Spur, Srinagar District, Kashmir. A frag• Gondwana Basin, another lycopsid species, mentary specimen showing crescentic ligular Lycopodites sahnii Lele (1962, pI. 1, figs. scars inside spirally disposed rhomboid leaf 1, 2, text-figs. 1, 2) is known. The frag• cushions has been tentatively identified with mentary axis bears spirally disposed linear Lepidodendron (Kapoor, 1969, pI. 2, fig. 1). delicate leaves almost at right angles to the Another specimen shows a cone-like struc• axis. The leaf-scars show a centrally situated ture probably attached to a lycopsid stem minute depression probably representing thE: (Kapoor, 1969, pI. 2, fig. 2). The presence vascular trace. The genus Lycopodites con• of a ligule in these specimens has been tinues into the Jurassic where it is repre• doubted by Surange (1971, p. 65) who opines sented by the species L. gracilis (Oldham & that "The scars described and figured as Morris) Seward & Sahni. ligules seem to correspond to the elevati?ns While the lycopsid megaplant remains are which are left on cushions when leaves dned scarce in the Permian-Triassic Gondwana without abscission like those in Sublepido• Sequence of India, a large number of dendron." A better preserved and almost microspores and megaspores, probably of complete lycopsid-like cone was found in the selaginelloid affinity, have been recorded tuffaceous shales of Liddar Valley near throughout the sequence (Bharadwaj & Pahlgam, Anantnag District, Kashmir (Sri• Tiwari, 1970; Maheshwari & Banerji, 1975). vastava & Kapoor, 1969, pI. 5, figs. 1-3, This does indicate the possibility that many text-fig. 1). The cone Lepidostrobus kash• more species of lycopsids flourished than IS mirensis is cylindrical and pedunculate. evident from the megafossil records. The The sporophylls are closely placed a~d paucity of lycopsid megaplant remains can spirally arranged. Detailed structure Of.t~IS probably be attributed to their delicate cone is not known and hence its affin1tIes nature and their suppression by an ove~• are as yet uncertain. whelming preponderance of gymnosperm1c In the peninsula the oldest lycopsid mega• megaplant remains (Maheshwari, 1974, p. 52). fossil record is from the MiddJe Permian Ironstone Shale Formation. The solitary species, Cyclodendron leslii (Seward) Krausel ARTHROHYTA is represented only by fragments of stems, which bear a number of spirally arranged, EQUISET ALES eye-shaped leaf-scars. Each leaf-scar has a more or less circular boss which perhaps The equisetalean remains, represented by indicates the position of the vascular supply. vegetative shoots only, belong to the genera: The ligular pit or the parichnos scars have Schizoneura, Phyllotheca, Lelstotheca, Bara• not been found (Kar, 1968, pI. 1, figs. 1, 2). karia and Raniganjia. In the absence of This species probably persisted into the fructifications, the relationships of the above Lower Triassic Mangali beds, from where a genera with one another or their affinities similar specimen was described as ?Lepido- within the group Equisetales are as yet MAHESHWARI - FLORISTICS OF PERMIAN AND TRIASSIC GONDWANAS 147
obscure. As far as their stems are con• shows that most of them had a similar cerned, they are all similar in being jointed epidermal pattern. In Raniganjia benga• and ribbed, the ribs continuing through lensis the guard cells are reported to possess successive nodes. In thp latter character obscure radiating striations like those seen they resemble the stems of Asterocalamites. in the modern Equisetum (Pant & Nautiyal, In that genus, however, the leaves are 1968). This character is not known either dichotomously forked many times. Such in Schizoneura gOt~dwanensis (Srivastava, forking of leaves is known only in the genus 1955) or in Phyllotheca angusta (Surange & Barakaria, and there is a report of a solitary Kulkarni, 1968). forked leaf in Phyllotheca australis (Pant & O£ the two species of the genus Schizo• Kidwai, 1968, pI. 30, fig. 6£). Further in neura, S. wardii Zeiller is restricted to the Asterocalamites the pelt ate sporangiophores Lower Permian formations only. The other are not subtencl.ed by bracts whereas species, S. gondwanensis Feistmantel ranges in P. australis the strobili have alternating from the Lower Permian to the Upper whorls of bracts and branched sporangio• Triassic but is more common in the Upper phores. The strobili of the Calamitaceae Permian and Lower Triassic. Phyllotheca have sporangiophores subtencl.ecl.by bracts makes it first definite appearance in the but branchEd sporangiophores arc not known Lower Permian by two species, viz., P. in the family except for Protocalamostachys. australis Brongniart and P. crassa Maithy. The fructification of the southern schizo• The latter species is confined to the Lower neuras is not known but ManchHrostachys, Permian whereas the former continues into the strobilus of Schizoneura manchuriensis, the Triassic. In the Middle Permian two is bracteate and has unbranched hexa• new species, viz., P. griesbachii Zeiller and sporangiate sporangiophores. Thus, while P. ampla Surange & Kulkarni (=P. angusta) the strobili of Phyllotheca australis and appear. Only the former continues into the M anchurostach.ys are somewhat similar to Upper Permian. those of the Calamitaceae, the latter family Lelstotheca robusta (Feistmantel) Mahesh• is differentiated by its foliage which lacks a wari and Barakaria dichotoma (Feistmantel) cup-like sheath and by its stems in which Seward & Sahni are characteristic Middle the grooves alternate through successive Permian species. Raniganjia bengalensis nodes. (Feistman,tel) R,igby is a typical Upper .Per• Examining the inter-relationship between mian, plant. This species is sometImes these genera. we find that Phyllotheca and found in the Lower Permian also (d. Phyllo• Schizoneura are distinguished from the other theca sahnii Saksen,a). Neocalamites foxii three genera by their leaf-sheaths which are Lele from the Upper Triassic is just an, adpressed to the stem to a certain extent. equisetalean stem impression, its foliage is Furthermore, the leaves of these two genera not known. lack the characteristic transverse striations or wrinkling. This wrinkling, which may be SPHENOPHYLLALES due to a lamellate mesophyll or could represent a transfusion tissue, was formerly The sphenophylls are probably represented supposed to be of little or no taxonomic in India by the vegetative shoots of Trizygia value (Seward & Sahni, 1920). But the speciosa Royle (1839, pI. 2, fig. 8). Trizygia association of this feature with the flat type speciosa shoots have a slender articulate axis of leaf-sheaths seems to give a certain with swollen nodes. Each node bears 3 taxonomic importance to this character unequal pairs of leaves in a whorl, successive within this assemblage. Meyen (1969) also whorls being superposed. A single vein believes that constant presence of this enters the leaf base, dichotomises a few feature may be of taxonomic importance times sending branches to the apical as well in some cases. Barakaria, however, stands as lateral margins. The species made its out by its characteristically forked leaves first appearan,ce in the Middle Permian and which superficially resemble those of some continued through Upper Permian, into the sphenophyllums, e.g. Sphenophyllum myrio• basal Triassic where it died out. phyllum Crepin. Whatever little that is Doubts have arisen as to whether this knOWnof the leaf anatomy of these genera, plant is a characteristic form of the Gond- 148 THE PALAEOBOTANIST
wanaland or is a migrant from the Northern Southern Hemisphere form still remams Hemisphere and belongs to the genus Spheno• undecided. phyllum. In fc>.ct,McClelland as early as The other fern-like forms were usually 1850 renamed the species as 'Sphaenophyllum described under various Northern Hemi• speciosa '. Later workers mostly agreed sphere genera, such as: Alethopteris, Clado• with McClelland, till Maheshwari (1968b, phlebis, Cyclopteris, Merianopteris, Pecopteris, 1974) reverted to the original name Trizygia Sphenopteris etc. Recently Maithy (1974a, after emphasizing the morphological diff('• 1974c, 1975), on the basis of a reinvestigation rences between the genera Sphenophyll1tm of sterile as well as fertile specimens, and Trizygia, like the characters of the veins has separated almost all of the Indian and the nature of the subsidiary cells. Permian ferns from the Northern Hemisphere However, on the basis of shoot morpho• fern genera. The Indian forms are noW graphy, this Southern Hemisphere form placed under the genera Dichotomopteris can be referred to the Order Spheno• Maithy, Dizeugotheca Archangelsky & Sota phyllales, if not to the genus Sphenophyllum. and Neomariopteris Maithy. Pant and Khare Its exact affinities must await the discovery (June, 1974) established ~ new gen.us of the fructifications. Damudopteris for some stenle and fertile Sphenopteris-like fronds from India. The genus is, however, nomenclaturally supe:• PTEROPHYTA fluous as it has the same type (Sphenoptens polymorpha Feistmantel, 1876) as the earlier Earliest records of fern-like plants in India published genus Neomariopteris Maithy (May, are from the Lower Carboniferous Po Series 1974). of Spiti. Ill, the Gondwana Sequence In the genus Dichotomopteris, 4-6 free unquestioned fern megaplant remains are sporangia form superficial sori, whic~ are not known in the basal Permian, the first arranged in two distinct rows, one on. elth?r side of the midvein, upon lateral vem fern-like fronds occur at the top of the endings on the underside of the pinnules. Lower Permian. Later on they increase in Annulus is absent. Spores are trilete and genera and species through the Middle and sculptured. Forms earlier described as Peco• Upper Permian and continue into the Triassic. Most forms have been attributed pteris (Alethopteris) phegopteroides (Feist• to Marattiales. There is no indication of mantel) Arber have now been transferred to the genus Dizeugotheca as they sho.w the presence of Osmundaceae or any other groups of 4 free sporangia, arranged. m tree fern in the Permian-Triassic. However, linear fashion on the pinnule margm.s. fertile fern-fronds are rarely found in the Of the 4 known species of the genus N eo• Gondwana Sequence and hence the affinities mariopteris, only N. hughesii (Feistmantel) of most of the species remain open. The oldest fern-like form from the Gond• 'Maithy and N. polymorpha (Feistmantel) Maithy are known in fertile st~.te. The wana of India is Botrychiopsis (Gondwani• sporangia are arranged in groups above dium) valida (Feistmantel) Archangelsky & lateral veinlets, a little removed from the Gamerro, which is a marker species of the margin. Each group probably comprises Lower Permian Karharbari Formation. The 4-6 sporangia in N. hztghesii and 2-9 fronds' are long and bear two rows of large sporangia in N. polymorpha. In N. poly- and more or less deeply lobed leaflets. The . morpha the sporangia are exindusiate, shortly fertile struc.ture of this species is not known stalked and annulate. The annulus is trans• and hence its pteridophytic affinities have verse and multiseriate (Pant & Khare, been quest! oned. Seward (1903) presumed 1974). Sporangial details are not known in that Gondwanidium had gymnospermous, N. hughesii. Spores arc not known in the rather than filicinean, fructifications. Similar former whereas in the latter these are forms from the Kuznetsk Basin were sup• triangular trilete, echinate, similar to those posed to have bore gymnospermous fructi• of Diehotomoptert's major (Feistmantel) fications of the type Gondwanotheca sibirica Maithy. Asterotheca santhalensis Vimal & Neuburg. The Kuznetsk forms are now Singh (1971) is incompletely known. The supposed to be different from Botrychiopsis• genera Dichotomopteris and Dizeugotheca can Gondwanidium and hence the affinity of the provisionally be referreq to the family lIIAl-IESHWARI - FLORISTICS OF PERMIAN AND TRIASSIC GONDWANAS 149
Asterother:ace<'.e of th~ M,!,rattiales. The daries dichotomize but do not anastomose. sphenopterid Neomariopteris shows some The genus Euryphyll1tl1t is also sometimes resemblance to the gleicheniaceous Oligo• included in this group. carpia. Detailed epidermal studies on the leaves In the Lower Triassic the group is pro• of the above genera have shown that certain bably represented by Sphenopteris (? := Neo• generalized epidermal types are common to mariopteris) polymorpha Feistmantel and more than one genus. According to Surange Sphenopteris spp. Two forms, viz., Pecopteris and Srivastava (1957) the various glossop• concinna Presl ,md Cyclopteris pachyrhachis terid species studied by them "fall into Gappert are restricted to the basal Triassic six groups which may provisionally be (FeistmCl.ntel,188b) but their identifications considered as of generic rank ". This is need confirm~.tion (Bose, 1974, p. 285). if we take it for granted that each glossop• In the Upper-Midcl.leTriassic we find M arat• terid genus will have only one generalized tiopsis sp. and Danaeopsis gracilis. In the epidermal type. However, during my in• former species at each vein enc1.ingis situated vestigations on the glossopterid cuticles, a sorus-like swelling. The position of the several times I noted a marked inconsistency sorus appEars to be MarattiCl.ceous (Lele, in epidermal structure of different portions 1962b). Parsorophyll1t1n indicum Lele could of the same specimen, as well as in different be another fern, characteristic of the Upper but morphographically similar specimens of Triassic. On the whole the records of the same species. Features, like the degree Triassic ferns in India arc extremely poor of cutinization of the guard cells, the surface and th,')ir rebtionships with the Permian papillae, and the thickness of the lateral types are as yet uncertain. cell walls etc. seem to be affected by preservation as well as laboratory processes. On the other hand, the epidermal structure GLOSSOPTERIDOPSIDA of Pteronilssonia gopalii Pant & Mehra, having large pinnate leaves, is like that of Thi.s is the most dom;nc>.n.tgroup of plants certain glossopterid leaves. While the im• in the Indian Permian floras 8.ppearing portance of cuticular characters in delimita• almost at the base of the Gondwana tion of species within a genus can not be Sequence, readling its zenith in the Upper denied, it is doubtful if this character can Permian and then continuing through the be utilized for delimitation of generic or Lower Triassic and becoming extinct before higher categories of taxa. Discussing the the Liassic. Glossopteris is the most im• importance of epidermal features, Seward portant member of the group and gives the and Sah'1i (1920, p. 7) have said that name Glossopteris flora. "the direct utility of this line of work in At present five leaf genera, viz., Gangamo• phylogeny is open to doubt." pteris, Glossopteris, Palaeovittaria, Rhabdo• Records of follage shoots with glossopterid taenia and Rubidgea are tentatively assigned leaves in organic conntction are rather few, to this group. The most important genus probably due to" an extraordinary efficiency Glossopteris has tongue-shaped leaves with a in abscission" (Schopf, 1967, p. 114). That definite midrib giving off secondaries whi.ch the glossopterids were deciduous is suggested dichotomize and anastomose. The anasto• not only by the occurrence of a very large moses either only connect the lateral veins number of isolated leaves but also by the in which case the secondaries can easily be presence of sharp growth rings in almost traced throughout the lamina, or change the all the Gondwana fossil wood (Maheshwari, venation to a regular reticuloid pattern 1972). Leaves or shoots have not been (Mah,eshwari, 1965a, 1966). Palaeovittaria found attached to a main stem and, there• and Rhabdotaenia also possess a midrib but fore, the various reconstructions of the ha.bit lack vein anastomoses. In the former the of the glossopteridean plants are largely secondary veins meet the leaf margin at based upon subjective extrapolation of the acute angles whereas in the latter they available circumstantial evidence. reach the margin almost at right angles. There seems little doubt, however, that the Gangamopteris and Rubidgea both lack a glossopterid leaves were borne in terminal midrib, but in the former the secondaries clusters in a tight spiral (Feistmantel, 1881a, anastomose whereas in the latter the secon- pI. 40, fig. 1, pI. 41, figs. 3, 4; Rigby, 150 THE PALAE0130TANIST
1967, pIs. 25, 26). The records showing others bore ovules. The affinities of the lateral attachment of leaves are not con• pollen producing types are uncertain but vincing enough and may probably be due to the ovule-bearing fructifications seem to chance overlapping eeL fig. 2, in Schopf, indicate the presence of two orders, which 1967). Therefore, views postulating a presently are informally ,....amed as Otto• dimorphic arrangement of leaves on short kariales and Lidgettoniales. In Ottokariales, and long shoots need more evidence. The a multiovula1.e capitulum was born in the axes which bore glossopterid leaves have axil of an ordinary foliage leaf and closely sometimes been identified with Vertebraria adpressed to its petiole or midrib. In (Surange & Maheshwari, 1962) but mostly Lidgettoniales, the fertile leaf was scale-like these axes are without any characteristic and had 2 or more ovule-bearing capitula, exomorphic features. In anatomical details arising from its midrib region. these axes are reported to be not signi• The male fructifications, Eretmonia du Toit ficantly different from those of Vertebraria emend Surange & Maheshwari, Glossotheca (Pant & Singh, 1974). The surface striations Surange & Maheshwari and Kendostrobus found in some of these axes probably Surange & Chandra, stand apart, in their represent the growth rings in radial view. organization, from the known southern or In the smooth axes it is further possible northern hemisphere male fructifications. that preservation of a thick cortex masked In Kendostrobus, groups of 4-5 sponmgia the characteristic transverse septae of Ver• were disposed in close spirals on an axis. tebraria. The ribbed axes could represent The sporangia were exannulate and produced young vertebrarias which were probably monolete, sculptured spores. Superficially, penta- to sept agonal in cross-section. As Kendostrobus resembles the spike of Hel• Vertebraria represents only the stelar aspect minthostachys but according 1.0Surange and of an axis, leaves, even if borne on such all Chandra (1975, p 174) these two are quite axis, will be difficult to find in organic distinct in their detailed structure. Eret• connection. But Schopf (1965) insists that monia and Glossotheca possessed one and Vertebraria is a root. He of course agrees three pairs of sporangial clustcrs rEspectively, that it is allied to the glossopterids. Gould borne on a pedicel which was fused with (1975) thinks that" Vertebraria indica was the petiole of a scale-like glossopterid leaf. part of the underground system of Arau• The fructification was probably axial. carioxylon arberi (Seward) Maheshwari type Thcse fructifications show some superficial of wood and possibly represents an adapta• resemblance with the Peltaspermaceae whose tion to a semiaquatic environment". male orgaJ] Antevsia was a bipinnate Whether Vertebraria was a root or shoot, sporophyll, ultimate branches of which bore its very characteristic stelar construc• 4-12 sporangia. Eretmonia and Glossotheca tion seems to preclude its alliance with also carried their sporangia on a branch the known petrified tree trunks of the system but it Can not be regarded as Glossopteris flora. Therefore, it could have a sporophyll, and moreover each ultimate been allied only to some of the glossopterids branch carried a single sporangium only. which were large shrubs or small trees. In external characters an ind ividual sporan• Other glossopterids were large trees as is gium resembles those of the genus ArberieUa evident from the length and diameter of Pant & Nautiyal instituted for dispersed some of the petrified tree trunks found in sporangia. These two genera probably come the Gondwana sediments. under the order Lidgettoniales. Fructifications of the glossopterids are TIle ovulate glossopterid fructifications infrequent, organically attached ones being belonging to the order Ottokariales are more so. Leaves with attached fructifica• referred to the genera Dictyopteridium Feist• tions and having a cuticular crust are mantel, Ottokaria Zeiller, Gonophylloides extremely rare. In recent years, however, Maheshwari, Scutum Plumstead and Seno• well-preserved glossopterid fructifications theca Banerjee. These organs have been have been described (Sen, 1955; Maheshwari, interpreted as receptacles of various sizes 1965b; Muld1erjee, Banerjee & Sen, 1966; and shapes around which ovules are attached Banerjee, 1969; Surallge & Maheshwari, 1970 in close spirals, without bracts or ovuliferous and Surange & Chandra, 1975). Some of scales; the whole receptacle, however, is these were pollen producing members while believed to have been subtended by a scale. MAHESHWARi - FLORISTICS OF PERMIAN AND TRIASSIC GONDWANAS 151
But the stroblus-like organization for all adpressed to a dehisced fructification. of these fructifications is as yet far from Even a fructification with mature ovules proved. It is possible that in certain types will require exposure to pollen rather than the ovules/seeds were borne embedded on protection. The genus continued through the adaxial surface of a scale-like leaf the Upper Permian into the basal Triassic. (see Maheshwari, 1965, pI. 1, figs. 4, 5;' It is in the Upper Permian, where the glos• Banerjee, 1969, pI. 2, figs. 6-8; Schopf, 1970, sopterid leaves reached their acme, that the fig. 6; Rigby, 1971, p. 3; Holmes, 1974, p.l largest number of ovulate fructifications 6, fig. 1; Pant & Singh, 1974, pI. 31, figs. is known, other types being Scutum, 64, 65). Some of these fructifications are Senotheca and Gonophylloides (= Cistella, borne on a stalk which seemingly arises probably also Plumsteadia). The seeds of from the petiole of a typical glossopterid Scutum and Senotheca are Indocarpus and leaf. If that is the case then the petiole Cornuspermum respectively. Protohaploxy• can be considered as forked, and the pinus type of pollen has been found inside fructifications as foliar modifications. It the micropyle of Cornuspermum ovules is also possible that the fructifications were (Banerjee, 1969). It has been suggested borne in the axil of a leaf and the stalk of that some of the Ganophylloides type the fructification is fused w.ith the petiole of fructifications might be the seedless of the leaf to various extents giving the receptacles of Scutum (Surange & Chandra, appearance of the fructification as coming 1974, p. 12). out of the midrib. In that case the In their organization the 'strobilate' fructifications will be axial (Pant & Singh, ovulate fructifications compare with the 1974, pI. 31, figs. 64, 65). Jurassic Rajmahalia paradoxa Sahni & Rao As the glossopterid fructifications are emend Bose (1966), where, too, the seeds/ extremely rare, their evolutionary aspect is ovules are arranged in close spirals on a unknown. The oldest fructification is Otto• receptacle. Another Jurassic fructification karia bengalMsis Zeiller (1902) which is so far Carnaconites also had a similar mode of known only from the Lower Permian. This attachment of seeds (Banerjee, 1969, p. fructification arising from the midrib of a 365). Therefore, Surange and Chandra (1975, leaf, probably Glossopteris indica Schimper, p. 176) suggested that the plant group to is supposed to have been protected by a bell• which Rajmahalia paradoxa belonged could shaped veined bract which was borne on have had its roots in the Glossopteropsida a common stalk with the fructification and if RajmahaZia and Carnocot.ites of the (Surange & Chandra, 1975, p. 171, text• Pentoxyleae are related to each other, it fig. 14A-B). O. kathwaiensis Virkki (1943, is possible that the Pentoxyleae had its photo 1, text-fig. 1) is an elliptic-obovate roots in the Permian strobilate ovulate frond attached to a slender stalk. If it types. is an Ottokaria at all, then it probably re• The order Lidgettoniales comprises the presents the protective scale of the fructi• genera Denllania Surange & Chandra, fication. In the Middle Permian comes Lidgettor.ia Thomas and Partha Surange up another fructification, viz., Dictyopte• & Chandra, all known from the top of Upper ridium sporijerum Feistmantel (J 881) which Permian only. These fructifications, show is so far known only in detached state. superficial resemblance to the pteridosper• Maheshwari (1965b) observed that one mous fructifications, but are not identical surface of the fructification bore oval scars with any of the known forms. In aligned in oblique rows, while on the other Denkania about 6 seed-bearing appendages surface a regular reticuloid venation was (cupules) are attached by long pedicels on present. Surange and Chandra (1973), how• the petiolar part of a scale-like glossopterid ever, opined that the net venation belonged leaf. Each 'cupule' probably contained to a closcly adpressed subtending scale and a single seed. It seems possible that each that the fructification proper has ovules or 'cupule' was borne on the tips of ultimate seeds of Platycardia type attached all around branches of a pinnate structure which in the central receptacle. However, the speci• turn arose from the leaf-scale petiole. Thus mens studied by Maheshwari had evidently the whole structure becomes compound shed their seeds, and as such it is unlikely with a forked rachis. If this was the case that a subtending bract will remain closely then this plant could have led to the 152 THE PALAEOBOTANIST
Corysto~permaceae. In Partha and Lidget·· of the r2.chis has not been observed. It is tonia the seeds arc arranged on the underside believed that' these were large forms and of disc-like heads probably borne on tips only their small parts have been preserved. of the ultimate branches of a branched Such leaves with unforked rachis are in• structure as in the Pelta"permaceae. How• ",luded in the gen,us Dicroidium on the ever, the cupules of PeJtaspermaceae are ba::,is of their epidermal structure. Recent borne spirally whereas in Partha and Den• opinion is that some of the so-called kania the capitula are borne in one plane. dicroidia may in f3.ct be Lepidopteris or may A relationship between male and female even belong to a separ2,te genus (Bose, 1974, fructifications is not known. But it is p. 287). In fact, the epidernial structurE possible that Kendostrobus is related to the of Dicroidium fronds described from Nidh• Ottokariales whereas Eretmonia and Glos• puri beds by Bose an,d Srivastava (1971) otheca might be related to Lidgetton iales. does not strictly conform with that pres• The latter types could have led to the cribed for typical dicroidia (see Lele, 1962a). Triassic 'pteridospermous' plants. The separate identity of the Nidhpuri dicroid ia is furthE.I confirmed by the dis• covery, in the same beds, of two fructifi• 'PTERIDOSPERMOPHYTA' cations Nidistrobus harrisia1bus and Nidia ovalis whose epidermal structure resemblES As discussed earlier there is n,o definite most that of Dicroidium 1~idpurensis (Bose evidence of the presence of Pteridosper• & Srivasta,va, 1973, p. 78). In. case these males in the Permian Gondwanas of India two fructifications were borne on di• though certain, plants with 'cupulate' female croidia fron,c1.s,the plant would be quite fructictifications have provisiouctl1y been unlike any of the known pteridosperm referred to this group. Two families of fa.milies. N. harrisianus has spirally dis• 'Pteridosperms', viz., Peltaspermaceae and posed 'pad-shaped' pollen-bea.ring organs. Corystospermaceae, are, however, known Each 'pad' hCl.'3 a row of 7-8 poJ.len sacs on in the Triassic Goudwanas of Ind i2.. Pelta• its adaxial surfar;e. Pollen grains were spermaceae is known on,ly in sterile state probably bisaccate, non-striate. Lacey by the fronds or Lepidoptcris indica Bose (1974, p. 40, pl. 1, figs. 1-3) has recently & Srivastava (1972). The fronds arc bipin• described app2.rently similar, though com• nate, have an unforked rachis, and a few para.tively larger, 'ribbed 'object' associated pinnules are atta,ehed directly on to the with Dicroidium lancifolium in th,e Molteno rachis in between the pinnae. The pinnules flora (Middle-Upper Triassic) of Rhodesia. have a well-marked midrib and forked Nidia ovalis has spirally disposed mega• lateral veins. Rectontly Srivastava (1975) sporophylls, each bearing two pelt ate seeds• has described a microsporangiate fructifi• ovules. Thus it is more cycad-like. cation, Bosea indica which he suspects to Satsangia eampanulata Srivastava & be the pollen-producing organ of Lepi• Maheshwari (1973) is anolher fructifica• dopteris indica. Organizationally Bosea is tion-like structure. It shows oval scars quite different from Antevsia whicb is sup• arranged in oblique rows as in the genus posed to be the pollen-producing member Dictyopteridi'um. However, in Satsangia the of the genus Lepidopteris (Harris, 1937, p. scars seem to be present on the inner 35). Lepidopteris, therefore may not be surface of a bell-like organ. The organ an organ genus. probably bore ovules at the scars. Details The family Corystospermaceae is repre• 2.re, however, not known. sented in the Triassic of India by the Typical carystospermaceous pollen-bearing foliage of the genus Dicroidium Gothan and organs are also known from the Nidhpuri pollen-bearin.g organs Pter'uchus Thomas and bed. Pteruchus indicus Pant & Basu (1973) possibly Indotheca Sitholey. The fronds of is known from sporangiferous stalks, borne typical Dicroidium have a forked rachis. spirally at wide angles around a central The epidermis of Dicroidium is amphisto• axis. Ea,ch sporangiferous stalk gradually matic, subsidiary cells are 4 in number widen.s out terminating in an 'umbrella• (rarely more) but do not form a ring and like' head, on the under surface of which normally the cell surface is papillate. are arranged sporangia. Pollen is bisaccate, But in most of the Indian forms forking nOll-striate. A somewhat similar organ is MAHESHWARi- FLORISTics OF PERMiAN AND TR[ASSiC GONDWANAS 15~
11ldotheca sakeSare1~sis Sitholey (1943b, pI. basis of epidermal characters; the Bennet• 2, figs. 14-16) from the Triassic of So.1t Range. titales being an exception. The gtllUS Indotheca, however, differs from Pteruchus Pterophyllum has also been reported from in having a defin.ite lamina which bears the the Upper Triassic beds (Lele, 1956, 1962b). sporangial cluster. The species Pterophyllum sahnii is pinnately segmented, segments being closely set, at• tached to the rachis hy the whole base. CYCADOPHYTA Veins usually fork close to the emergence, rarely forking again. Th~ group Cycadophyta is poorly re• Senia reticulata Khan (1969), from the presented by folio.ge only. One of the top of Upper Permian, is a very imperfectly species known is Pseudoctenis balli (Feist• known cycad-like leaf. 7'aeniopteris spatu• mantel) S\ward. Feistmantel (1881b) des·· lata from the Parsora Formation andT. cribed the species under Anomozamites but glandu,lata from the Nidhpuri beds are also later (1886) changed to Platypterigium, suspected to have ha.d cycadean affinitiE.s. Schimper. Seward (1917) transiE,rred the species to his genus Pseudoctenis without reasoning out the necessity for such a change. 'GINKG()PHYTA' From the Gentric Index (Andrews, 1970, p. 165) the species balli of Felstmantel (1886) Some of the leaf types present in the seems to be the type species of the gen,us Indian Gondwana Sequence show 3. general Platypterigium Schimper ex Feistmantel resemblance to the leaves of Ginkgo. But, (1886). Under the International Code of ill the absence of fructifications their re• Botanico.1 Nomenclature, it is doubtful if lationships with one another or with the the type species of an older gf,nus can be Ginkgoales arc not definitely established. transferred to a younger genus. This species These arc, therefore, sometimes placed in which, was originally described from the the noncommital order PalaeophyJlales. The Middle Permian was later discovered in Upper Palaeozoic ginkgoalean type of leaf the basal Triassic (Lele, 1962b). The frond belongs to two genera, viz., Psygmophyllum is pinnately divided, the segments are of and Rhipiclopsis. unequal width and attached at right angles Leaves of PsygmophyUum are known only or obliquely to a slender rachis. The veins from the Permo-Carboniferous strata of are generally forked near the point of emer• the extra-peninsular Gonc1wanas of Kashmir. gence and show a few cross connections. Three species of the genus are known, all Another leaf, Pterophyll~t11t b~£1'dwanense of which hav~ simple, lobed leaves. These, (McClelland) Feistmantel was originally des• therefore, should be ta.ken out of the genus cribed as Zamia burdwanense by McClelland Psygmophyllum Saporta emend Schimper (1850). Here the veins GO not bifurcate, which is characterized by pinnate leaves. run parallel and are spaced fairly apart. Maithy (1974b) suggests the use of the The species is known from Middle to Upper generic name Ginkgophyllum S3.porta for Permic'.n strata. Because of this, the identi• the Indian leaves. These forms are more fication of the genus has been doubted. or less intermediate between Ginllgophyllum Arber (1905) suspected that the species and Ginkgophytopsis H0eg. . may represent a torn fragment of Taeniop• Ginkgophyllum (Psygmophyllum) holland~ teris-like leaves. Pant and Mehra (1963) (Seward) is known from the Lower Permi.an studied the epidermal structure of similar of Kashmir. The lami.na is divided by d leaves and described them as Pteronilssonia deep median sinus into two bilobed segments. gopalii. As there is hardly any mOlpho• G. (Psygmophyllum) haydenii (Sewa.rd), also graphical difference from the species burd• from the Permo-Carbon,iferous of Kashmir, wanense it would have been better to is a petiolate, fiabellate leaf deeply incised describe it as Pteronilssonia burdwanense. into a number of radiating, cuneate seg• Po.nt and Mehra (1963) suspect that this ments. G. (Psygmophyllum) sahnii (Ganju) species may not have cycadean but pteri• is represented by a branch, bearing spirally dospermous a.ffinities. It is, however, disposed leaves with a short petiole and doubtful if one can postulate affinities to fiat, wedge-shaped lamina divided by a higher taxa of Gymnosperms simply on the median sinus into two equal bilobed "eg- 154 'tHE pALAEOBoTANtST
ments. Each segment is again cut into. that af Cordaites (Lele & Ma.ithy, 1964). equal labes. Ganju (1943, p. 203) remarks The leaves were barne in clase spirals that P. sahnii bears a striking resemblance (Seward & Sahni, 1920, pI. 1, 'fig. 10). to. the leaves af Ginhgo, and therefare there Further Cordaitanthus like fructificatians may be same relatian between the two.. are totally absent in the Gandwana Sequence. The gmus Rhipidopsis was reparted by Maithy (1970, p. 169) is of the opinian Feistmantel (1881, 1886) from the Permian that N oeggerathiopsis cauld as well be a af the peninsula. His specimens af R. glassapterid, Hawever, presently there is ginhgoides Schmalhausen fram the Middle no. evidence to. suppart such an apinian. Permian were later put under a new The problems met with in systematics af species, R. gondwanensis by Sew,ud (1919). cardaitalean type af leaves have recently Here the lamina is divided, almast to. the been discussed by Maheshwari and Meyen base, into. 6-10 segments; the larger are (1975, p, 199). cuneate and the smaller abavate and abtuse. A large number of fossil waad with annual In R. densinervis fram the Upper Permian rings and araucariaid trachcidal pits are the lamina is deEply divided into. abcuneate knawn fram the Barakar and Raniganj segments that appear te, be irregularly farmatians (Maheshwari, 1964, 1965c, 1967; labed at the truncate margin. Sithaley and athers). Sahni and Singh (1926) (19430.,p. 188) remalked that R. densinervis thaught that such waad prabably bare may btlang to. the some genus as Psygmo• N oeggerathiopsis type af faliage. Krause], phyllum haydenii. But, Maithy (1974b) has Maithy and Maheshwari (1962) an the ather transfern:d R. densinervis to. the genus hand, suggested that some of the Lawer Platyphyllum Dawsan. He also. describes Gandwana woad with enda.reh prataxylem a samewhat similar farm as Gondwanophyton may represent canifers. Maheshwari (1972, indicum. R. densinervis is probably also. p. 37) finds it most likely that mast af present in the Lawer Triassic (Bhatt acharyya, the fassil waad belanged to. the Glcssapte• 1963). ridapsida as almast all recards of fossil The anly ather Triassic ginkgaalean type waad are fram harizans where the glassap• af leaf knawn is Baiera ~ndica Lele (1962b, terids predaminated and the canifers 0.1' pI. 3, figs. 27, 28). The leaf is deeply dis• Noeggerathiopsis were extremely rare. sected into. a number af narraw, linear, seg• The anly fructificatian which has been ments, each having 4-5 thin, parallel veins. carrelated with Noeggerathiopsis is a mega• Sitholey and Base (1974) have daubts abaut sporaphyll-like organ originally described the affinity of the specimen with the genus by Feistmantel (18810.,p. 59, pI. 28, fig. 5) Baiera due to. farmer's peculiar made af as an "inflarescence" probably belanging dissectian. to. Noeggerathiopsis hislopi. This Was later named Arberia indica by White (1908) and carrelated with Gangamopteris. Another CONIFEROPHYTA species knawn fram India is A. umbellata Surange & Lele. Similar specimens have CORDAITALES also. been figured by Ganguly (1959, fig. 14), Pant an~ Nautiyal (1966, Ottoharia-like In the Lawer Permian certain strap• head) and Maithy (1970, Dolianitia harhar• shaped, parallel-veined leaves, knawn as barensis). The megasparaphyll has a slender Noeggerathiopsis, are very camman (Feist• stalk which expands distally and bears a mantel, 1879). The genus has stray number af abruptly truncate labes ar re• representatian nat anly thraugh the Permian curved processes. It is presumed that the but in the Triassic as well (Feistmantel, labes ar pracesses bare avules as is seen in 1882). The leaves are very much similar a specimen figured by Pant and Nautiyal to. the leaves af the Northr.rn Hemisphere (1966, pI. 1, fig-. 1) as Ottokaria-like head. Cordaites and are therdare taken as repre• Similar specimens were described by Millan senting the graup Cardaitales. Hawever, (1967) under the genus Dolianitia and Were assignment of Noeggerathiopsis to. the suppased to. have affinity with Noeggera• group Cordaitales is not free fram daubt. thiopsis. But accarding to. Rigby (1972), Epidermal structure af Noeggerathiopsis Arberia (= Dolianitia) while not belanging leaves has been faund to. be distinct fram to. the same plant which bare Noeggera- MIDDLE(.)~ 0.. BARAKAR ~ "'?l KARHARBARIIRONSTONELOWERTALCHIRHIRAPURFmTOBARAKARTALCHIRTALCHIRTALCHIRBAPbedFm:J:PALlbedsFmBoulderFillFmFrowbedsFmGr..FmFmFmbeds~0WFmSh.Fm..~FmFmTIKIBARAKARFmbedBoulderBARRENDAIGAONIRONSTONEKARHARBARIBARAKARBoulderBIJORISATPURADHARMARAMTALCHIRKARHARBARIBARRENBASINBADHAURAHIMGIRBoulderKARHARBARIMOTURPACH~~ARHIFmDUBRAJPUR"p bedMEAS.FmbedbedMEAS.NnFmFmFm?ShFmFmRANIGANJFmFm BARRENKARHARBARIFm PARSORAFmNIDHPURIMEAS.FmKAMTHIFm GrMALER PANCHETI BHIMARAM Fm MANGALIYERRAPALLI Fm '" ..J(.) Cl (/) ClCl Cl ". MIDDLESUPRA-PANCHETSRANIGANJ GODAVARIRAJMAHAL~SOUTHFVALLEYMAHANADIRAJASTHANWARDHA-HILLSBASINKASHMIRVALLEYSONREWA "'p ~ :I: UPPER -D.a:(/) z:J::>4:!b Boulder~ bedl- I-- 4: MAITUR- Fm a: ~ ~ Z4: W4: DEIIWA Fm ? UP. CARB VALLEY UPPER LOWER - ~ AU~ODDAMODARbeds a: (.) ~ ;J> ::c .. tr1 (fl ::c ~ ;J>
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thioPS1:S leaves, also docs not seem to be (Bose & Maheshwari, 197+, p. 212). related to known pteiidosperm families. Little is knowJl ;)bout conifer fructifications though some forms haVe been described as CONIFERALES Araucarites cone scales or C01~ites.
The oldest conifer-like foliCl.ge known from the Indian Gondwana Sequence is the CONCLUDING REMARKS Lower Permian ?Paranoclad·us indica Surange & Lr1c (1957, pI. 1, fig. 13, 15). Till l".te there Was no evidence of plant The shoo~s 3.re irregularly bmnched, branches life in the basal boulder bed horizon of the being covered by som~what spreading, Gondwana Sequence although Sahni (1939) spirally disposed homomorphic leaves. opined that the rise of the Glossopteris A somewh.at youngrr species is Buriadia flora overlapped with the later phases of sewardii Salmi. The vegetative shoot shows the Palaeozo:c glaci PalaeoviUaria. The Glossopteris-complex is Permia.n floras. In the Upper part of best known from the Upper Permian. strata. the Lower Triassic the 'pteridospermous' Besides leal morphography and wood families Corystospermaceae and Peltasper• anatomy we have data about epidermal maceae a.ppear and continue at least up to features ?nd the fructifications. The infor• Upper Triassic. mation available so far doC'snot favour the inclusion of this complex in tht Pterido• sptlmophyta. The Glossopteris-complex is ACKNOWLEDGEMENTS therefore referred to an independent class - the Glo3sopteridopsida. This class Thanks are due to Drs. M. N. Bose, Shaila comprises at least two orders, viz., the Chandra and K. R Surange for fruitful Ottob.riales and the Lidgettonialcs. c'iscussions. To Miss Asha Bhardwaj thanks The b?.s?.lTriassic flora is extremely poor are due for assis1ance in translating some but not much different from the Upper Russian papers. REFERENCES ARBER, E. A. N. (1905). Catalogue of the fossil FEISTMANTEL, OTTOKAR (1876). 0:1 some fossil plants of Ihe Glossopteris Flora in the Depart• plants from the Damuela Series of Ra ..iganj ment of Geology, British Museum (Natural Coalfield collected by Mr. J. 'Vooel-Mason. History). London. J. asiat. Soc. Beng. 45 (2): 329-382. BANERJEE, MANJU (1969). Senotheca muruli• IDEM (1879). The fossil flora of the Talchir-Kar• dihensis, a new glossopteridoa 1 fructification harbari beds. Mem. geol. Surv. India, Palaeont. from India associated with Glossopteris tafnio• indica, ser. XII, 3 (1): 1-49. pteroides Feistmantel: pp. 359-368 in, Santapall, IDEM (1881a). The fossil flora of the Gondwana H. et al. (Editors) - J. Sen Nlemorial Volume, System-II. Tho flora of the Damuela-Panchet Bot. Soc. Bengal, Calcutta. divisions. Ibid. ser. XII, 3 (2, 3): 1-149. BHARADWA],D. C. & TIWARI, R. S. (1970). Lower IDEM (1881 b). Palaeontological notes from the Gondwana megaspores - a monograph. I'lIlae• Hazaribagh and Loharclagga districts. Rec. geol. ontagraphica. 129B: (1-3): 1-70. Surv. India. 14 (3): 241-263. BIIATTACHARYYA,A. K. (1963). The assemblage IDEM (1882). The fossil flora of the Gondwana of plant megafossils from the Lower Gond wana System-III. The fossil flora of the South Rewa rocks of the western part of Auranga valley Gond wana Basin. Ibid. ser. XII, 4 (1) : Coalfield, Palamau District, Bihar. Q. II geol. 1-52. Min. mclall. Soc. India. 35 (2): 123-128. IDEM (1886). The fossil flora of the GOl'dwana BOSE, M. N. (1966). A revision of Rajmahalia. System-IV. The fossil flora of some of the Palaeobolanisl. 14: 85-88. coalfielels in western Benga!. Ibid. ser. XII, IDEM (1974). Triassic floras: pp. 285-293 in, 4 (2): 1-65. Surange, K. R. et a.l. (Ec\itors)- A speels and GANGULY,SUBINAY(1959). Palaeontological study Appraisal of Indian Palaeobotany, Birbal Sah ~i of Lower Gondwana rocks including the coal Institute of Palaeobotany, Lucknow. SEams around Chirimiri of Surguja District, BOSE, M. . & MAHESHWARI,H. K. (1974). Meso• Maelhya Pradesh. Q. II geol. Min. metall. zoic conifers: pp. 212-223 in, Surange, K. R. Soc. India. 31 (3): 155-166. et al. (Editors)- Aspects a.nd Appraisal of GANJU, P. N. (1943). On a new species of Psygmo• Indian Palaeobotany, Birbal Salmi Institute of phyllum (P. sahnii sp. nov.) from the Lower Palaeobota'1Y, Lucknow. Gondwana beds of Kashmir. J. Indian bot. BOSE, M. N. & SRIVASTAVA,S. C. (1971). The Soc. 22: 201-207. genus Dicroidium from the Ttiassic of Nidpur, GOULD, R. E. (1975). A preliminary report on Madhya Pradesh, Inc\ia. Palaeobotanisl. 19 (1): petrifiod axes of Vertebraria from thfJ PC!mia'1 41-51. of Eastern Australia: pp. 109-115 in, Campbell, IDEM (1972). Lepidopteris indica sp. no\'. from K. S. W. (Ed.) - Gondwana Geology, paps 3rd the Lower Triassic of Nidpur, Madhya Pradesh. Gonaw. Symp. Canberra 1973, Australian natn. J. palaeont. Soc. India. 15: 64-68. Univ. Press, Canberra. IDEM (1973). Some micro- and megastrobili from HARRIS, 1'. M. (1937). The fossil flora of Scoresby the Lower Triassic of Goparl River Valley, Sound, East Greenland. Part 5. Stratigraphic id pur. Geophytology. 3 (1): 69-80. relations of the plant beels. Meddr GrGlossopteridales from the Upper PeImian CHALONER, W. G. & BOUREAU, ED. (1967). Lyco• of N.S.W. Proc. Linn. Soc. N.S. W. 98 (3): phyta: pp. 435-802 in, Boureau, Ed. (Ed.)• 131-141. TraUe de Paleobotanique, Masson et Cie, KAPOOR, H. M. (1969). Additional records of paris. fossils and fossiliferou$ banc\s from the Lower 158 THE PALAEOBOTANIST Gondwanas of Zewan Spur, Kashmir. J. IDEM (1968b). Studies in the Glossopteris Flora of palaeont. Soc. India. 13: 24-28. India-38. Remarks on Trizygia speciosa Royle KAR, R. K. (1968). Studies in the Glossopteris with reference to the genus Sphenophyllum flora of India-36. Plant fossils from Barren Koenig. Ibid. 16 (3): 283-287. Measures Succession of Jharia Coalfield, Bihar, IDEM (1972). Permian wood from Antarctica and India. Palaeobotanist. 16 (3): 243-248. revision of some Lower Gond wana wood taxa. KHAN, A. M. (1969). Senia reticulata, a new plant Palaeontographica. 138B: 1-43. fossil from the Raniganj rocks of the Talchir IDEM (1974). Palaeozoic Lycopsida and Spheno• Coalfiele(, Orissa, India: pp. 335-337 in, psida: pp. 51-61 in, Surange, K. R. et al. Santapau, H. et al. (Editors)- J. Sen Memorial (Editors) - Aspects and Appraisal of Indian Volume, Bot. Soc. Beng. Calcutta. Palaeobotany, Birbal Sahni Institute of Palaeo• KRAUSEL, R. (1961). Lycopodiopsi; derbyi Renault botany, Lucknow. une( einige andre Lycopodiales an den Gondwana• MAHESHWARI,H. K. & BANERJI, JAYASRI (1975). Schichten. Palaeontographica. 109B: 62-92. Lower Triassic palynomorphs from the Maitur KRAusEL, R., MAITHY, P. K. & MAHESHWARI, Forma ion, West Bengal, India. Palaeonto• H. K. (1962). Gymnosperrpous woods with graphica. 152B: 149-190. primary structures from Gondwana rocks - A MAHESHWARI, H. K. & MEYEN, S. V. (1975). review. Palaeobotanist. 10: 97-107. Cladostrobus and the systematics of cordaitalean LACEY, \'V. S. (1974). Somo new African Gondwana lea ves. Lethaia. 8: 103-123. plants: pp. 34-41, in, Bose, M. N. (Edltor)• MAITHY, P. K. (1970). Studies in the Glossopteris Symposium on Morphological and Stratigraphical Flora of India-39. On some new plant fossils Palaeobotany, Birbal Sahni Institute of Palaeo• from the Karharbari beds, Girinih Coalfield, botany, Spec. Publ. no. 2. India. Palaeobotanist. 18 (2): 167-172. LELE, K. M. (1956). Plant fossils from Parsora in IDEM (1974a). A revision of the Lower Gond• the South Rewa Gondwana Basin, India. wana Sphenopteris from India. Ibid. 21 (1): Palaeobotanist. 4: 23 -34. 70-80. IDEM (1962a). Studies in the Indian Middle IDEM (1974b). Studies in the Glossopteris Flora Gond wana Flora-I. On Dicroidium from the of India-41. Gondwanophyton gen. novo with South Rewa Gondwana Basin. Ibid. 10: 48-68. a revision of allied plant fossils from the IDEM (1962b). Studies in the Indian Middle Lower Gondwana of India. Ibid. 21 (3): Gondwana Flora-2. Plant fossils from the 298-304. South Rewa Gondwana Basin. Ibid. 10: 69-83. IDEM (1974c). Dichotomopteris, a new type of fern LELE, K. M. & KARIM, REHANA (1971). Studies frond from the Lower Gondwana of India. Ibid. in the Talchir Flora of India-6. Palynology of 21 (3): 365-367. the Takhir Boulder Bed in Jayanti Coalfield, IDEM (1975). Some contribution to the knowledge Bihar. Ibid. 19 (1): 52-69. of Indian Lower Gondwana ferns. Ibid. 22 LELE, K. M. & MAITHY, P. K. (1964). Studies in (1): 29-38. the Glossopteris Flora of India-IS. Revision MCCLELLAND, J. (1850). Report of the Geological of the epidermal stlucture of Noeggerathiopsis Survey of India for the season 1848-49. J. C. Feistmantel. Ibid. 12 (1): 7-17. Sherriff, Military Orphan Pless, Calcutta. MAHESHWARI,H. K. (1964). Studies in the Glos• MEYEN, S. V. (1969). The Angara member of sopteris Flora of India-16. Dadoxylon jamu• Gondwana genu5 Barakaria and its systematical dhiense, a now species of fossil wood from the position. Argumenta palaeobot. 3 (1-4): 1-14. Raniganj Stage of Jharia Coalfield, Bihar. MILLAN, J. H. (1967). Novas fructificacoes na Ibid. 12 (3): 267-269. flora Glossopteris do Gonduana Inferior do IDEM 0965a). Studies in the Glossopteris Flora Brasil. Notas prelim. Estud. Div. Geol. Miner. of India-22. On some species of the genus Bras~l, no. 140: 1-19. Glossopteris from the Raniganj Stage of the MUKHERJEE, S., BANERJEE, M. & SEN, J. (1966). Raniganj Coalfield, Bengal. Ibid. 13 (2) : Further glossopteridean fructifications from 129-143. India. Palaeontographica. 117 (B): 99-113. IDEM (1965b). Studies in the Glossopteris Flora PANT, D. D. & BAsu, NUPUR (1973). Pteruchus of India-23. On two fructifications from the indicus sp. novo from the Triassic of Nidpur, Raniganj Stage of the Raniganj Coalfield, India. Ibid. 144B: 11-24. Bengal. Ibid. 13 (2): 144-147. PANT, D. D. & BHATNAGAR,SUMAN(1975). A new IDEM (1965c). Studies in the Glossopteris Flora kind of foliage shoots Searsolia oppositifolia gen. of India-24. On two new species of fossil wood et sp. novo from Lower Gondwanas of Rani• from the Raniganj Stage of Raniganj Coalfield, ganj Coalfield, India. Ibid. 152B: 191-199. Bengal. Ibid. 13 (2): 148-152. PANT, D. D. & KHARE, P. K. (1974). Damudopteris IDEM (1966). Studies in the Glossopteris Flora of gen. nov.- a new genus of ferns from the Lower India-31. Some remarks on the genus Glosso• Gondwanas of the Raniganj Coalfield, India. pteris Stern b. Ibid. 14: 36-45. Proc. R. Soc. Land. 186B: 121-135. IDEM (1967). Studies in the Glossopteris Flora of PANT, D. D. & KmwAI, PARVEEN F. (1968). On India-28. On some fossil woods from the the sit ucture of stems and leaves of Phyllotheca Raniganj Stage of Raniganj Coalfield, Bengal. indica Bun bury and its affinities. Palaeonto• Ibid. 15 (3): 243-257. graphica. 12tB: 102-121. IDEM (1968a). Studies in the Glossopteris Flora PANT, D. D. & MEHRA, BHARATI (1963). On a of India-34. On a record of Phyllotheca australis cycadophyte leaf, Pteronilssonia gopalii gen. et Brongn. from }haria Coalfield, Bihar. Ibid. sp. nov., from the Lower Gondwanas of India. 16 (2): 167-169. Ibid. 113B: 126-134. MAHESHWARI - FLORISTICS OF PERMIAN AND TRIASSIC GONDWANAS 159 PANT, D. D. & NAUTIYAL, D. D. (t966). On two wanas: pp. 306-326 in, International Gondwana peculiar fossils of Karharbari Stage, India: Symposium, A ligarh 1970, Ann. Geo!. Dept., pp. 98-101 in, Symposium on the Floristics and Aligarh M. Upiv. 5 & 6 (Special issue). Stratigraphy of Gondwanaland, Birbal Sahni SITHOLEY, R. V. (1943a). On Psygnwphyllum Institute of Palaeobotany, Lueknow. haydeni Seward. J. Indian bot. Soc. 22: 183• IDEM (1967). On the structure of Buriadia hetero• 190. phylla (Feistmantel) Seward & Sahni and its IDEM (1943b). Plant rcmains from the Triassic fructifications. Phil. Trans. R. Soc. Lond. ser. of the Salt Range in the Punjab. Proc. natn. B, 252 (774): 27-48. A cad. Sci. India. 13 (5): 300-327. IDEM (1968). On the stt u<:ture of Raniganiia SITHOLEY, R. V. & BOSE, M. N. (1974). Mesozoic bengalensis (Feistmnntel) Rigby with a discuss• Ginkgoales: pp. 210-211 in, Surange, K. R. et al. ion of its affinities. Palaeattlographica. 121B: (Editors)- Aspects and Appraisal of Indian 52-64. Palaeobotany, Birbal Sahni Institute of Palaeo• PANT, D. D. & SINGH, R. S. (1974). On the stem botal' y, Lucknow. and attachment of Glossopt~ris and Ganga• SRIVASTAVA,J. P. & KAPOOR, H. M. (1969). Dis• mopteris leaves. Part-II. structural features. covery of Lepidostrobus Brongniart from Lower Ibid. 147B: 42-73. Gondwana Formation of Kashmir. J. palaeo!!t. RIGBY, J. F. (967). On Gangamopteri, walkomii Soc. India. 12: 44-47. sp. novo Rec. Aust. Mus. 27 (8): 175-182. Sr VIMAL. K. P. & SINGH. PRATAP (1971). A new 1.C.- Final Report, Commissao de estudos das species of fern from the Lower Gondwana, minas de Carvao de Pedra do Brazil, Rio de San thai Pargana, Bihar. ]1 palaeont. Soc. Janeiro. India. 16: 89-91. WADIA, D. N. (1975). Geology of India. 4th VIRKKI, C. (1943). In Sahni, B. & Sit holey. R. V. Edition. T2.ta McGraw-Hill Publishing Co. (Editors)- Palaeobot. India I V. Progress report New Delhi. for 1942 - Salt Range. J. Indian bot. Soc. 22: ZEILLER. R. (1902). Observations sur quel(lues 171-172. plantes fossiles des Lower Gonclwanas. Mem. \"HITE. DAVID (1908). Fossil flora of the Coal geol. Surv. India, palaeont. indica. n.S. 2: 1• Measures of Brazil: pp. 339-617 in, White, 40. -- 4/033