Glossopteris flora in the Permian Weller Formation of ...

GR-01399; No of Pages 28 Gondwana Research xxx (2015) xxx–xxx

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Glossopteris flora in the Permian Weller Formation of Allan Hills, South Victoria Land, Antarctica: Implications for paleogeography, paleoclimatology, and biostratigraphic correlation Rajni Tewari a, Sankar Chatterjee b,⁎, Deepa Agnihotri a, Sundeep K. Pandita c a b c

Birbal Sahni Institute of Palaeobotany, 53 University Road, Lucknow 226007, India Museum of Texas Tech University, Lubbock, TX 79409, USA Department of Geology, University of Jammu, Jammu 180006, India

a r t i c l e

i n f o

Article history: Received 11 June 2014 Received in revised form 13 February 2015 Accepted 23 February 2015 Available online xxxx Handling Editor: M. Santosh Keywords: Glossopteris flora Permian Ice Age Allan Hills Weller Formation Antarctic Phytogeography

a b s t r a c t The Permo-Triassic Victoria Group in South Victoria Land, Antarctica, is a heterogeneous sequence of glacial tillite beds, carbonaceous and non-carbonaceous fluvial deposits, and volcaniclastic strata. The carbonaceous beds are rich in plant fossils associated with coal seams. In Antarctica, the geological record of the Late Paleozoic Ice Age is restricted to the Early Permian. After deglaciation, the Glossopteris flora thrived in polar forests in Antarctica throughout the Permian but disappeared at the end-Permian extinction. Here we describe the first comprehensive record of the Glossopteris flora from the Permian Weller Formation of Allan Hills, South Victoria Land, Antarctica. The flora is well preserved and comprises pteridophytes and gymnosperms. The pteridophytes include the sphenopsid order Equisetales and the gymnosperms comprise Glossopteridales. Equisetales are represented by branched and unbranched axes, whereas, Glossopteridales are highly diverse encompassing Gangamopteris, Glossopteris, Surangephyllum, sterile scale leaves namely Scirroma sp., Nautiyalolepis sp., Utkaliolepis indica, Scale leaf A and scale leaf of male fructification Eretmonia. The flora of the Weller Formation shows close similarity with the Late Permian assemblages of India, South Africa and Australia. Gangamopteris, an index fossil of the Early Permian formations of different Gondwana continents, had extended stratigraphic range in the Late Permian Weller Formation of Allan Hills. Antarctica played a crucial role in the dispersal of Glossopteris flora because of its central position in Gondwana. © 2015 International Association for Gondwana Research. Published by Elsevier B.V. All rights reserved.

Contents 1. 2.

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . Paleogeography and paleoclimatology of Antarctica during the Permian 2.1. Late Paleozoic Ice Age . . . . . . . . . . . . . . . . . . . 2.2. From icehouse to hothouse . . . . . . . . . . . . . . . . . 3. Geological setting of Allan Hills and sample localities . . . . . . . . 4. Age of the Weller Formation . . . . . . . . . . . . . . . . . . . 4.1. Permian–Triassic boundary . . . . . . . . . . . . . . . . . 5. Material and methods . . . . . . . . . . . . . . . . . . . . . . 6. Description of the Glossopteris flora . . . . . . . . . . . . . . . . 6.1. Order Equisetales . . . . . . . . . . . . . . . . . . . . . 6.2. Order Glossopteridales . . . . . . . . . . . . . . . . . . . 7. Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.1. Biostratigraphic correlation . . . . . . . . . . . . . . . . . 7.2. Composition of the Weller flora . . . . . . . . . . . . . . . 8. Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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⁎ Corresponding author. Tel.: +1 806 834 4590. E-mail address: [email protected] (S. Chatterjee).

http://dx.doi.org/10.1016/j.gr.2015.02.003 1342-937X/© 2015 International Association for Gondwana Research. Published by Elsevier B.V. All rights reserved.

Please cite this article as: Tewari, R., et al., Glossopteris flora in the Permian Weller Formation of Allan Hills, South Victoria Land, Antarctica: Implications for paleogeography..., Gondwana Research (2015), http://dx.doi.org/10.1016/j.gr.2015.02.003

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R. Tewari et al. / Gondwana Research xxx (2015) xxx–xxx

1. Introduction The iconic Glossopteris flora from Antarctica was discovered and collected by Robert Falcon Scott and his party near the Beardmore Glacier in the Transantarctic Mountain during their ill-fated return trip from the South Pole in January 1912 before they perished. Seward (1914) identified these fossil leaves from Scott's Terra Nova expedition as Glossopteris indica, which were well-known from the Permian coal measures of India, now located across the equator and far away to Asia. Previously, this distinctive Glossopteris flora was found in abundance on widely separated southern landmasses, namely South Africa, Australia, New Zealand, and South America. The Antarctic fossil find would have been the most crucial piece of evidence in support of Wegener's continental drift theory that Antarctica had once been part of ancient supercontinent Gondwana. The common association of the Glossopteris flora with PermoCarboniferous glacial deposits in India, Antarctica, Africa, Australia, and South America suggests that these ancient “seed-fern” trees were especially adapted to cold climates. The Glossopteris flora is diverse and incorporates several orders including Lycopodiales, Equisetales, Filicales, Glossopteridales, Cordaitales and Pinales, which all flourished in the Permian polar forests of Gondwana continents. Glossopteris was a deciduous tree with a woody trunk, reticulate-veined leaves, seedbearing fructifications, pollen-bearing organs, and roots that thrived in the swampy conditions of Gondwana during the interglacial periods of the Late Paleozoic Ice Age. Other than leaves, additional parts of Glossopteris trees such as twigs, branches, trunks, seeds and other reproductive organs, and roots have occasionally been found disarticulated, and their associations were questionable for a long time. Different taxonomic names were employed for naming different parts of the same Glossopteris tree. Gradually, over the years, the researchers have pieced together a reconstruction of the Glossopteris plant as a deciduous tree or shrub from these various parts. Glossopteris leaves differed in shape, size, apex and base, midrib development and organization of their reticulate venation (Chandra and Surange, 1979). Male and female fructifications were partially attached to the leaves (Plumstead, 1956; Surange and Chandra, 1974a, 1974b, 1975, 1978; Maheshwari, 1990). The majority of the female reproductive organs of Glossopterids, such as Scutum, Plumsteadia, Senotheca and Dictyopteridium are attached along the midrib (White, 1908; Plumstead, 1952, 1956; Surange and Chandra, 1973). The segmented roots of Glossopterids known as Vertebraria (because they resemble a vertebral column) have a specialized schizogenous architecture presumably for aeration and adaptation to swampy environments. Like modern deciduous trees, the Glossopteris leaves were shed seasonally in autumn to conserve energy over the winter. This is reflected by their dense accumulation, probably indicating accumulation of “autumnal leaf” banks at the water's edge. The leaves of Glossopteris, as described here, are abundant in the black shale deposits of the Weller Formation of Allan Hills, Victoria Land, whereas, the fossil trunks of Glossopteris trees, common in the channel sands, are marked with growth rings, reflecting the effect of strong seasonality. In the coalbearing sequences of Gondwana, Glossopteris leaves typically occur at the top of the coal seams, Vertebraria occurs at the base, and fossil tree trunks in the cross-bedded sandstones. The Glossopteris flora is widely distributed in the Permian sequences of different Gondwana basins of Antarctica. Permian megafossils of leaves, fructifications, seeds, ovules and wood are known in the form of impressions, compressions and petrifactions from various parts of East, West, and central Transantarctic Mountains, Antarctica. Seward (1914) described Glossopteris fossils from the Permian rocks of the Beacon Supergroup and later Edwards (1928) reported the occurrence of G. indica from the same horizon. Plumstead (1962) described plant assemblages from the Permian rocks of the Ross and Weddell Sea areas, which include Annularia [considered to be a possible record of Schizoneura sp. (Rigby,

1969)], Phyllotheca [included later in Paracalamites australis (Rigby, 1969)], Gangamopteris, Palaeovittaria, Glossopteris, seeds of Cordaicarpus and Stephanostoma, a microsporangium Arberiella, fructifications and Vertebraria. Schopf (1962, 1967, 1968) described Permian megafossils from Horlick Mountains and Ohio Range, and Rigby and Schopf (1969) recorded plant fossils of the same age from Central and South Victoria Land, Queen Maud, Horlick, Pensacola, Theron and Ellsworth Mountains. Rigby (1969) and Maheshwari (1972) reported sphenopsids and fossil woods from several Permian localities in Antarctica. Plumstead (1975) recorded a Late Carboniferous (? Early Permian) floral assemblage from Milorgfjella, Dronning Maud Land, Antarctica comprising lycopsids, sphenopsids, pteridosperms, wood and some glossopterid and cordaitalean taxa like Gangamopteris, Palaeovittaria, Noeggerathiopsis and Euryphyllum. McLoughlin et al. (2005) revised the identifications of plant fossils described by Plumstead (1975) and placed the species of Palaeovittaria, Noeggerathiopsis and Euryphyllum under Glossopteris and Gangamopteris, and additionally reported Glossopteris sp. cf. G. communis, Glossopteris sp. cf. G. spatulata, finely branched Vertebraria indica rootlets in situ, Phyllotheca australis, equisetalean stems and rhizomes, isolated seeds, scale leaves and fragmentary gymnosperm axes from the Middle Permian of Fossilryggen, Vestfjella, Dronning Maud Land. Subsequently, Early Permian plant fossils comprising Phyllotheca sp., Equisetum sp. and Glossopteris erehwonensis from the English Coast of eastern Ellsworth Land (Gee, 1989) and structurally well preserved Late Permian megafossils from the upper Buckley Formation of Skaar Ridge in the Beardmore Glacier region of the central Transantarctic Mountains (Smoot and Taylor, 1986; Taylor and Taylor, 1987, 1992; Pigg, 1990; Taylor et al., 1992; Pigg and Taylor, 1993; Galtier and Taylor, 1994; Zhao et al., 1995; Taylor, 1996; Klavins et al., 2001; García Massini, 2007; Cantrill and Poole, 2012; Ryberg et al., 2012a) were recorded. The floral remains from the Skaar Ridge include a moss, Merceria augustica, Glossopteris schopfii, Glossopteris skaarensis, Glossopteris stems, Gangamopteris, Plectilospermum seeds and a Choanostoma verruculosum ovule. Besides, sporadic Permian plant fossils such as Gangamopteris from Aztec Mountain (E.L. Taylor et al., 1989a), Glossopteris and Noeggerathiopsis from Kennar Valley (E.L. Taylor et al., 1989b) and Plumsteadia-type glossopterid reproductive organs from Mount Achernar near the Law Glacier, central Transantarctic Mountains (T.N. Taylor et al., 1989c) are reported. Additionally, a fossil forest of Late Permian age within the upper Buckley Formation including silicified trunks, preserved in growth position, wood specimens, megasporangiate taxa Scutum leiophyllum, Lidgettoniopsis ramulus, Arberiella sp. and isolated ovules has been recorded (Taylor et al., 1991; Ryberg et al., 2012b) from Mount Achernar. Similarly, anatomically preserved Noeggerathiopsis leaves have been reported by McLoughlin and Drinnan (1996) from the Upper Permian Bainmedart Coal Measures of eastern Antarctica. Later, Slater et al. (2012), recorded evidences of plant–animal interaction, fungal evidence (Slater et al., 2013) and peat biota (Slater et al. 2015) from the same locality. Retallack et al. (2005) reported Glossopteris browniana, from Graphite Peak, central Transantarctic Mountains, South Victoria Land. Though substantial work has been carried out on Triassic megafossils of Allan Hills (Chatterjee et al., 2013 and references cited therein), records of Permian megafossils from the area are scarce. These include two fossil woods namely, Taeniopitys scotti and Araucarioxylon allanii (Kräusel, 1962; Maheshwari, 1972), Plumsteadia ovata (Schopf, 1976), unidentified plant remains of the Glossopteris flora (Chatterjee et al., 1983) and Vertebraria australis (Retallack et al., 2005). In this paper, we report an abundant, though less diverse Glossopteris flora from the Permian Weller Formation, Allan Hills, South Victoria Land. The assemblage includes taxa of Equisetales and Glossopteridales. We describe the first records from the Allan Hills examples of sphenophytes, four species of Gangamopteris, twenty two species of Glossopteris including a new species Glossopteris taylori, Surangephyllum elongatum and scale leaf of the male fructification Eretmonia, besides four sterile scale leaves namely Scirroma sp., Nautiyalolepis sp.,



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Utkaliolepis indica, Scale leaf A. The flora from the Allan Hills has been correlated with the Permian floras of other Gondwana countries, but shows close similarity with those of India, South Africa and Australia indicating shared phytogeographic province and biotic connection. 2. Paleogeography and paleoclimatology of Antarctica during the Permian During the Late Paleozoic, Antarctica was part of the supercontinent Gondwana that incorporated present-day South America, Africa, Arabia, Madagascar, India and Australia (Scotese et al., 1999). The most dramatic paleoclimatic event of Gondwana's Late Paleozoic history was the growth of vast continental glaciers. The clockwise drift of the southern Gondwana continent shifted Africa and Antarctica into polar cold regions that triggered glaciation processes (Torsvick and Cocks, 2004). At that time Gondwana continents were clustered around the South Pole that created strong latitudinal temperature gradients between the equator and the poles (Caputo and Crowell, 1985; Crowell, 1999). 2.1. Late Paleozoic Ice Age Sedimentary evidence for massive glaciation on Gondwana during the Late Paleozoic is unequivocal. Biostratigraphically-dated glacial tillites, glacial striated pavements and clasts, and ice-rafted dropstones are widespread (Caputo and Crowell, 1985). The Gondwana Ice Age has its origin in the Late Devonian that waxed and waned, but was not fully established until the Late Carboniferous to Early Permian (Scotese et al., 1999). In Antarctica, overlying the “red beds” of Taylor Group in the Beacon Supergroup of South Victoria Land is an assorted layer of tillites, debris left behind from an ancient Antarctic ice sheet. The tillites crop out in many places in the Transantarctic Mountains and tell us that approximately 300 million years ago Antarctica was ice covered much as it is today. The Metschel Tillite in Victoria Land, the Pagoda Tillite, Scott Glacier Tillite and Buckey Tillite in different sections of the Transantarctic Mountains represent the basal glacial boulder beds, indicating that the preserved record of this glacial event in Antarctica is incomplete, represented by the Early Permian. Moreover, the range of the basal glacial deposits in Antarctica is problematic. So far, no definite Late Carboniferous pollen taxa have been recorded from these beds. Most tillites contain palynomorphs such as those belonging to Parasaccites that indicate earliest Permian (Asselian) age for these beds. High-resolution paleontologic sampling requires resolving the preserved age of glaciation in Antarctica. The available evidence indicates that Antarctic glaciation, as preserved in the rock record, was largely restricted to Early Permian (Asselian) age (P1), followed by three small pulses of glacial intervals (P2 to P4) that extended to Middle Permian (~299–265 Ma) (Fielding et al., 2008; Cantrill and Poole, 2012). In the Late Permian, the post-glacial lacustrine and fluvial environments favoured the spread of the polar forest. The alternating glacial– nonglacial sequences suggest that the Late Paleozoic Ice Age was considerably more dynamic than previously thought. Most likely, early records of glacial events in Antarctica, which is widespread in western Gondwana (South America and Africa) at the end of the Late Devonian were removed by the erosive action of the much larger glacier that formed in the Early Permian (Caputo et al., 2008). The paleogeographic and plate tectonic setting of Antarctica in the united Gondwana during the Early Permian time are shown in Fig. 1 in a polar projection. At that time, Antarctica was joined with Africa on its western margin, with India on its northern margin and Australia on its eastern margin; its southern peninsular region had a coastline, facing the south polar sea. For occurrence of widespread continental glaciation, not only near-polar situation is required, but nearby open expanses of water are also needed. Both the Paleo-Pacific on the western margin of South America and Paleotethys on the northeastern margin of India and Australia provided the moisture required to ensure glaciation. In addition, warm currents along the eastern coast of Pangea

Fig. 1. A. Paleogeographic reconstruction of Gondwana in a polar projection during the Early Permian (~295 Ma) showing the extent of the Permian ice sheet by white area. B. the same reconstruction after deglaciation in Early Permian (~280 Ma) (after Moore and Scotese, 2012).

transported moist air to the South Polar Region. With the onset of Ice Age in Gondwana continents, continental glaciers pushed northwards within nearly 30° of the equator, a latitude where subtropical conditions prevailed during most of the Phanerozoic time. The expansion of glaciers during Permo-Carboniferous in Gondwana locked up water on land as ice and caused sea level to drop about 150 m than it is today. The crucial evidence of the occurrence of the Permo-Carboniferous glacial episode at several locations in South America, South Africa, Antarctica, India and Australia consists of glacially striated pavements at the base of the Gondwana basins capped by tillites and other glacigene sediments. All the evidences indicate that the Antarctic ice sheet was of continental dimensions (N2000 km radius), approximately


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comparable with the present ice sheet (Fig. 1A) and centered on east Antarctica. The orientation of striations by the moving ice suggests that the glacier moved northward from the center of accumulation in southwestern Africa and eastern Antarctica. Paleo-ice flow directions on Antarctica are towards the southeast from Victoria Land towards Ellsworth and Pensacola Mountains (Barrett and Kohn, 1975). During the warmer interglacial stages and in the outlying less frigid areas, Glossopteris and other plants, tolerant of the cool, damp climate, grew in profusion and provided the materials for thick seams of coal. At many places these remnants of past glaciations can be followed laterally with evidence of floating icebergs as revealed by laminated marine sequences with large dropstones. Fossil trees growing in polar forests document distinct seasonal growth of extreme climate in tree rings. The glacial boulder bed was followed by thick sequences of glacial lake deposits, channel sandstone, overbank mudstone, and coal seams indicating amelioration of climate and initiation of a complex fluvial system. As the ice sheet retreated, a broad, low-lying river floodplain developed, and there followed the establishment of lush stands of glossopterid forest. This deciduous swamp forest was dominated by tree-sized seed ferns — Glossopteris and Gangamopteris — along with some conifers, clubmosses and horesetails. Concentration of the carboniferous material from the abundant vegetation gave rise to several coal seams in the Late Permian deposits associated with Glossopteris fossils. The ecology of these cold-climate swamps and peatlands of Antarctica would resemble the present-day boreal taiga of Canada and Siberia.

2.2. From icehouse to hothouse During the Permian, the climate of Gondwana fluctuated from icehouse to hothouse conditions. As Pangea moved northward, Antarctica moved from the polar position (above 60°S) into paleolatitudes spanning 60° to 30°S (Torsvick and Cocks, 2004). Enhanced extensional rifting, volcano-tectonic activity during the Late Permian and widespread transgressions around the Gondwana continents caused an elevation of CO2 level in the atmosphere considerably, increasing equatorial to polar temperature gradients, which in turn, might have triggered deglaciation and concomitant rise of sea level (Stollhofen et al., 2000). As the climate became drier and more arid during the Late Permian in the northern hemisphere, coal swamps dried, too. As a result, rates of carbon burial declined and more plant debris was consumed at the Earth's surface by respiratory bacteria that released CO2 in the atmosphere leading to greenhouse warming. The process of CO2 release and temperature increase, linked in a positive feedback, caused global warming. Towards the end of the Early Permian, the ice sheet collapsed dramatically in Gondwana continents, with the development of extensive fluvial systems, which in turn opened up new environments for plants to colonize in high southern latitudes. Marine dropstones in Gondwana persisted almost till the end of Permian, when the icehouse state was abruptly replaced by the hothouse state (Fig. 2A). The Glossopteris flora was obviously an important component in the Permian polar forests that had formed coal under cool, moist conditions. This polar forest extended from within a few degrees of pole to the mid-latitudes that flourished throughout the Permian Period (Fig. 2B). The end-Permian coincides with massive Siberian Trap eruption that vented a huge amount of CO2 in the atmosphere, which caused global warming and most severe mass extinction event in geologic history (Erwin, 1994). The climate became increasingly hot and dry at the end of the Permian in Laurasia. In Gondwana, as an aftermath of the endPermian extinction, a generally warmer and less seasonal climate prevailed, when the Dicroidium flora replaced the Glossopteris flora. A slice of vanishing Permo-Triassic polar forest is preserved in the Gondwana sediments of Allan Hill, South Victoria Land of Antarctica in the form of exquisite fossils.

Fig. 2. A. Paleogeographic reconstruction of Gondwana in a polar projection during the Late Permian with complete retreat of the ice sheet (~260 Ma) and the opening of the Neotethys Ocean. B. Distribution of Glossopteris flora during the Late Permian (~260 Ma) (modified from Moore and Scotese, 2012).

3. Geological setting of Allan Hills and sample localities The Beacon Supergroup, exposed along the length of the Transantarctic Mountains, contains two major units: the older Devonian Taylor Group and the younger, Permo-Triassic Victoria Group (Collinson et al., 2006). The Victoria Group consists of Permian glacial beds (the Metschel Tillite) at the base, which are overlain successively by the Late Permian Weller Formation, and the Triassic Feather and Lashly formations. As the ice sheets retreated, the glacial beds were succeeded by dark lacustrine shale with thin bands of sandstone and limestone, deposited in a large pro-glacial basin. The Permo-Triassic sequences



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Fig. 3. A. Gondwana outcrop along the Transantarctic Mountains showing the location of Allan Hills in South Victoria Land (marked by an arrow) (after Kyle and Schopf, 1982). B. Geological map of the Allan Hills showing the plant fossil localities (after Ballance, 1977; Chatterjee et al., 1983).


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represent post-glacial flat-lying fluvial strata of shales and sandstones, intercalated with sparse coal seams (Barrett and Kohn, 1975). The plant megafossils are abundant throughout much of the Victoria Group, especially in association with the coal beds and broadly indicate the age of the sediments. The Jurassic Ferrar Group of volcanic origin caps the Victoria Group (Elliot et al., 2006). The Allan Hills comprise a nunatak of approximately 60 km2 that is part of the Transantarctic Mountains in southern Victoria Land (Fig. 3A). The excellent exposures of the Permo-Triassic Victoria Group (upper part of the Weller Coal measures, Feather Conglomerate, Lashly Formation) and units of the overlying Jurassic Ferrar Group of volcanic rocks are well developed in the Allan Hills. The plant fossils are preserved as impressions in grey and green shales exposed along the western side of the eastern arm of outcrops in the Allan Hills (Fig. 3B). Gunn and Warren (1962), Borns and Hall (1969), Barrett et al. (1971), Barrett and Kohn (1975), Ballance (1977), Chatterjee et al. (1983), Collinson et al. (1987), Kyle (1977), and Retallack (1997) have discussed the stratigraphy of the Allan Hills (Fig. 5). The Permian Weller Formation of Antarctica is exposed in a narrow belt extending along the edge of the Polar Plateau in South Victoria Land. This unit incorporates repetitive coal beds and was deposited on an alluvial plain by a braided river system (Barrett and Kohn, 1975). It disconformably overlies the glacial Metschel Tillite in most localities, indicating a rapid change from glacial to postglacial conditions and consists of conglomerate, arkosic sandstone, shale, and coal in “fining-upwards” cycle (Fig. 6). The formation is about 250 m thick and is easily recognizable from its coal-bearing horizons. It consists of three informal members: A, B and C. Exposure of the Weller Coal Measures at Allan Hills is restricted largely to Member C, which consists of 70 m of interstratified sandstone, siltstone, alternating with coal beds, which are up to 4 m thick. The plant fossils occur in Member C within the fine-grain deposits associated with both meandering and braided sandstones. The glossopterid fossil leaves, branches, and fructifications were collected from the strata directly above and beneath the coal seams, from mudstones contained within abandoned channels. This member is well exposed over a 5km2-area south of Manhaul Bay. Seven coal seams are exposed in the Allan Hills and separated by sheet-like clastic units, predominantly sandstones, subordinate siltstones and conglomerates. The carbonaceous shales and coals indicate subaquatic deposition of woody material with high water table during the peat formation. Rank of Antarctic Permian coal ranges from high-volatile C bituminous coal to anthracite, but most of the coal samples belong to low-volatile bituminous coal and semianthracite (Coates et al., 1990). Petrified logs and stumps are common in the cross-bedded, channel sandstone, indicating driftwood. The stumps are frequently shot through with zones of charcoal and the logs are universally encrusted with charcoal, indicating forest fire (Chatterjee et al., 1983). In rare cases, Glossopterid tree trunks are preserved in upright position with charcoal core indicating evidence of forest fire (Fig. 6C). Extensive Glossopteris floral assemblage was collected from the green and grey shale immediately below the second level of coal from the Member C of the Weller Formation, which is described in this paper (Fig. 5). The Weller Formation is overlain by the massive, 300-m-thick, Feather Formation consisting of pebbly conglomerate, which is interbedded with sandstone and succeeded by a light green siltstone and shale. The Feather Formation is devoid of coal but contains abundant fossil logs (Retallack, 1997). The disappearance of swamp vegetation at the beginning of the Triassic is associated with the famous “coal gap” (Retallack et al., 1996). Globally, there is no record of occurrence of coal during the 5 million years of the Early Triassic. Recovery of the peat-forming plants began in the Middle Triassic. We see the same evidence of “coal gap” in the Early Triassic Feather Formation after the endPermian extinction and subsequent recovery of coal in the Middle Triassic Lashly Formation (Fig. 4). The Lashly Formation represents the upper unit of the Victoria Group ranging in age from Middle to Late Triassic. It is 520-metre-thick and consists of a cyclic sequence of sandstone,

siltstone, carbonaceous shale, and coal. Petrified logs are abundant, usually encrusted with charcoal as in the Weller Formation. Barrett and Kohn (1975) subdivided the Lashly Formation into four informal members (A through D), where the lower part is more volcaniclastic than the quartzose upper part (Members C and D). Member C is highly fossiliferous. A rich Dicroidium flora has been described recently from the Lashly Formation (Chatterjee et al., 2013) and the evidence of ancient forest fire from microscopic charcoal remains has been reported by Kumar et al. (2011). Much of the Member D is missing from Allan Hills. The Late Jurassic Ferrar Group caps the Victoria Group at the top. The basal Mawson Formation (Early Jurassic) consists of about 370-m-thick basaltic pyroclastic rocks, which are well exposed in the southern part of the lower arm of the Y-like exposure, but the overlying Ferrar Dolerite sills, dikes, and massive intrusions are relatively minor in the Allan Hills (Elliot et al., 2006). 4. Age of the Weller Formation The Weller Formation was assigned an Early Permian age (~265 Ma) based on plant megafossils (Townrow, 1967) and palynomorphs (Kyle and Schopf, 1982). Recently, Awatar et al. (2013) assigned a Late Permian age to the Formation based on palynological evidence. Additionally, the palynoflora of the Weller Formation bears similarity to the Late Permian palynoassemblages of India and South Africa (Awatar et al., 2013). The megafloral assemblage of the Weller Formation favours a Late Permian age due to its similarity with the assemblages from the Barren Measures, Raniganj, Kamthi and Bijori formations of India (Table 1). The humid climate of the eastern Gondwana (Australia, India, South Africa and Antarctica) during Lopingian was conducive to the formation of extensive peat swamps, whereas, West Gondwana (Brazil and Argentina, South America) was affected by seasonal drought during this time (Rees, 2002). Consequently, fossil floras are scarce in South America compared to Australia, India and South Africa during the Late Permian. The present study indicates that the floral assemblage of the Weller Formation was quantitatively rich, and like India, South Africa and Australia was associated with the formation of coal measures derived from the glossopterid swamp forests. 4.1. Permian–Triassic boundary The Permian–Triassic boundary records the most severe mass extinctions in Earth's history and coincides with the Siberian flood volcanism (~251 Ma). The best evidence for the Permian–Triassic boundary comes from the marine sections of Paleotethys and Neotethys oceans, such as the Meishan section of South China, Guryul Ravine in Kashmir, South Alps and Transcaucas Ali Bashi section of Iran, but the sedimentological continuity across this boundary is controversial (Erwin, 1994; Benton, 2003). This Permian–Triassic boundary is difficult to recognize in the nonmarine Gondwana sections of Antarctica and other continents, where the continental biota have coarse resolution for demarcation. Palynology provides an effective tool for cross-correlation between marine and non-marine sequences. So far, the Damodar Valley section of India seems to be a continuous section, where the Permian–Triassic boundary is generally placed between the Raniganj and Panchet formations (Fig. 5). Here, the disappearance of the Glossopteris flora constitutes the end-Permian extinction. Moreover, the palynoassemblage zone between the Permian–Triassic boundary is very distinctive in the Damodar Valley section between Raniganj (palynoassemblage zones: VI–VII) and Panchet formations (palynoassemblage zones: VIII–IX) (Tiwari and Tripathi, 1992). The Glossopteris flora in the Allan Hills is mainly restricted to Permian. Hence, most likely, the Permo-Triassic boundary in this region, if complete, would occur somewhere between the Weller Formation and the Feather Conglomerate. However, demarcation of the exact Permo-Triassic boundary in the Antarctic Gondwana sequences is highly controversial.



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Fig. 4. Generalized Permian–Triassic stratigraphic sequences of Victoria Group of South Victoria Land showing plant fossil horizons. In the Allan Hills, the oldest sequence is the upper part of the Weller Coal Measures, followed by the Feather Conglomerates, Lashly Formation, and capped by the Late Jurassic Ferrar Group (modified from Kyle, 1977; Retallack et al., 2005).


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not find any sharp negative excursions in carbon isotopes to identify the Permo-Triassic boundary at the Allan Hills. Instead, they used paleosols as a proxy for demarcating the boundary near the top of the Weller Formation, between Susanne and Dolores paleosols. In contrast, Elliot et al. (2006) placed the Weller Formation within the Permian, but showed a discontinuity between the Weller and the Feather Formations. Similar disconformable relations have been observed between Permian and Triassic sequences in different Gondwana basins in India (Fig. 5). In our discussion we have followed the Permo-Triassic boundary suggested by Elliot et al. (2006) in a broader context considering the rarity of the preservation of the boundary across the globe. Unlike the Cretaceous–Tertiary boundary, demarcated by the iridium layer and shocked quartz, no such distinctive impact layer has been found at the Permo-Triassic boundary. This is why the Permian–Triassic extinction event is difficult to recognize in lithostratigraphic sequence. However, the floral turnover in Gondwana gives some clue about this biotic crisis at the end of the Permian. The Glossopteris flora of Gondwana had formed coal under cool, moist conditions. Glossopteris and coal do not occur above the Permo-Triassic boundary, however. In their place are red beds that formed in a warm and wet interval at the beginning of the Triassic. Following the Permian–Triassic extinction, the Dicroidium flora in Antarctica and other Gondwana continents replaced the typical Glossopteris flora. Thus the replacement event of Glossopteris flora by the Dicroidium flora marks the Permo-Triassic biotic crisis in Gondwana that corroborates with our boundary in the Allan Hills (Figs. 4 and 5). 5. Material and methods

Fig. 5. Correlation of Permo-Triassic Gondwana sequences between Allan Hills, Antarctica and the Damodar Basin, India. Palynological zones of India adopted from Tiwari and Tripathi (1992).

The glossopterid fossils were collected from the Allan Hills (latitude 76°43′S, longitude 159°40′E), South Victoria Land, Antarctica (Fig. 4), during the field season 1982–1983 by one of us (SC). About 200 shale samples with well-preserved plant fossils were investigated, and representative specimens of different plant taxa were photographed. A Canon Rebel T1i camera with Canon 100 mm Macro 2.8 and Canon 28–70 mm f 3.5–4.5 lenses was used for photography. The specimens were studied with a low-power Bausch & Lomb binocular microscope and hand lens. The identifications are based on the morphological characters such as presence/absence, continuous/alternate ridges and furrows in the case of calamitalean axes and axes of uncertain affinities, and shape, nature of apex, base, margin, midrib and venation pattern in the case of leaves. The terminology of leaf shape, apex and base follows Lawrence (1955), and that of the venation pattern follows Melville (1969). In the case of Glossopteris leaves, the identification pattern of Chandra and Surange (1979) has been followed as well. The fossil material is housed at the Museum of Texas Tech University, Lubbock, Texas, USA, under accession number TTU-ATP. 6. Description of the Glossopteris flora

Barrett and Kohn (1975) placed the Permo-Triassic boundary within the Feather Conglomerate between the lower and upper Flemming Member, due to an abrupt change in the paleocurrent direction. Isbell and Cúeno (1996) regard the contact of Weller Formation and Feather Formation as a regional disconformity, which has eroded all records of Permian–Triassic boundary events in southern Victoria Land. In contrast, both Collinson et al. (2006) and Retallack et al. (2005) concluded that the Permian–Triassic boundary occurs within a relatively complete terrestrial section in Antarctica at the top of the Weller Formation. Retallack et al. (2005) studied six measured sections from the Permo-Triassic soil successions in Antarctica using carbon isotope chemostratigraphy and total organic analyses and found a shift in soil and plant types at the boundary in most sections. Latest Permian soils include coal and rooted sands, whereas, earliest Triassic soils are mainly root-filled mudstones. Unlike other sections in Antarctica, they could

The megafloral assemblage from the upper part of the Weller Formation of Allan Hills consists of pteridophytes and gymnosperms. The pteridophytes include the order Equisetales, whereas, the gymnosperms are represented by Glossopteridales. Systematic Paleobotany Pteridophytes 6.1. Order Equisetales Calamitalean axes (Fig. 7A) Well-preserved, branched and unbranched specimens are present in the collection. The main axes measure 3.6–4.8 cm in length and 0.7–1.0 cm in width. Lateral branches measure 3.4 cm long and 4–5 mm wide. The main axis of one of the specimens (Fig. 7A) measures 8.0 cm in width. All the specimens show well-preserved nodes and internodes. The distance between two nodes varies from 0.6 to



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Fig. 6. Field photographs of Allan Hills showing Permo-Triassic exposures and fossil floras. A Glossopteris leaves in the grey shales in the Upper Permian (Member C) of Weller Formation. B. Extensive Glossopteris fossil horizon in a bedding plane of grey shale in the Upper Permian (Member C) in Weller Formation. Field placement of Permo-Triassic boundary between the upper Weller Formation and the Feather Formation. C. Glossopteris tree trunk in upright position in the Weller Formation showing charcoal remains that indicate paleo forest fire. D. Coal seam (semi-anthracite) within Weller Formation. E. Dicroidium-bearing fossil horizon in the Late Triassic (Member C) of Lashly Formation.

2.5 cm. Internodes with distinct ridges and furrows. Ridges alternate or continuous, 0.6 mm–1 mm apart. Incertae sedis (Fig. 7B) Simple branched axis without nodes and internodes preserved. The main axis measures 8.0 × 0.4 cm in size and the lateral branch is 3.6 cm long and 0.2 cm broad. Both the main axis and the lateral branch are striate, striations 0.3 mm apart. Since the nodes and internodes are not preserved, it is not possible to assign the specimen to either Calamitales or Equisetales.

6.2. Order Glossopteridales Gangamopteris McCoy, 1847 Type species. Gangamopteris angustifolia McCoy, 1847 Gangamopteris angustifolia McCoy, 1847 (Fig. 7C, D) Description. Two specimens are present in the collection. Leaves are simple, linear, spatulate, apex flat, obtuse, base narrow, slightly broken on one side and elongate in one of the specimens (Fig. 7D). Leaves measure 3.3–9.8 cm in length and 2.0–2.6 cm in width, margin entire. The


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Fig. 7. A, Calamitalean axis. Specimen no. TTU-ATP 113A; B, Incertae sedis-axis. Specimen no. TTU-ATP 099A; C, Gangamopteris angustifolia. Specimen no. TTU-ATP 005; D, G. angustifolia. Specimen no. TTU-ATP 020; E, Gangamopteris clarkeana. TTU-ATP 015; F, Gangamopteris major. Specimen no. TTU-ATP 011; G, G. major. Specimen no. TTU-ATP 045.

median region of leaves is occupied by distinct, 7–8 subparallel veins 0.3 mm apart. Secondary veins arise from median veins at about 25°, arch slightly, dichotomize frequently, anastomose and form narrow, elongate meshes before meeting margin at 48°–53°. Meshes are

2–4 mm long and 0.2–0.6 mm broad near the median region, and 1.5–4 mm long and 0.2–0.5 mm broad near the margin. The vein density is 11–15 per cm near the median region and 14–32 per cm near the margin.



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Fig. 8. A, Gangamopteris walkomii. Specimen no. TTU-ATP 050; B, G. walkomii. Specimen no. TTU-ATP 111; C, G. walkomii. Specimen no. TTU-ATP 121; D, Glossopteris browniana. Specimen no. TTU-ATP 021; E, G. browniana. Specimen no. TTU-ATP 095; F, Glossopteris sp. cf. G. ampla. Specimen no. TTU-ATP 117A; G, Glossopteris arberi. Specimen no. TTU-ATP 021; H, Glossopteris bucklandensis. Specimen no. TTU-ATP 038.

Comparison. Leaves are identical in shape and venation pattern with G. angustifolia McCoy (Feistmantel, 1879, Pl. 9, Fig. 5; Maithy, 1965, Pl. 3, Figs. 9, 10) described from the Karharbari Formation of Giridih Coalfield,

Damodar Basin, West Bengal, Barakar Formation of Korba Coalfield, Son-Mahanadi Basin, Chhattisgarh (Singh et al., 2012, Pl. 2 Fig. 1), India; Early Permian of Lucky Valley Coalfield, Warwick, Queensland,


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Fig. 9. A, Glossopteris bucklandensis. Specimen no. TTU-ATP 034; B, G. bucklandensis. Specimen no. TTU-ATP 105; C, G. bucklandensis. Specimen no. TTU-ATP 018; D, Glossopteris communis. Specimen no. TTU-ATP 041; E, Glossopteris sp. cf. Glossopteris conspicua. Specimen no. TTU-ATP 132; F, Glossopteris damudica. Specimen no. TTU-ATP 117B; G, G. damudica. Specimen no. TTU-ATP 059.



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Fig. 10. A, Glossopteris longicaulis. Specimen no. TTU-ATP 096; B, Glossopteris fuchsii. Specimen no. TTU-ATP 024; C, Glossopteris sp. cf. G. gigas. Specimen no. TTU-ATP 11O; D, Glossopteris indica. Specimen no. TTU-ATP 070; E, Glossopteris sp. cf. G. gigas. Specimen no. TTU-ATP 104; F, Glossopteris karanpuraensis. Specimen no. TTU-ATP 039; G, Glossopteris longicaulis. Specimen no. TTU-ATP 060.


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Fig. 11. A, Glossopteris major. Specimen no. TTU-ATP 027; B, G. major. Specimen no. TTU-ATP 029; C, Glossopteris major. Specimen no. TTU-ATP 106A; D, Glossopteris occidentalis. Specimen no. TTU-ATP 093; E, A part of leaf in D enlarged to show venation pattern, ×3; F, Glossopteris occidentalis. Specimen no. TTU-ATP 115; G, Glossopteris pandurata. Specimen no. TTU-ATP 114; H, Glossopteris browniana A part of leaf in Fig. 12A enlarged to show venation pattern, ×3.7.

Australia (Walkom, 1922, Pl. 4, Figs. 22–23), Ross Sea Area, Antarctica (Plumstead, 1962, Pl. 3, Figs. 1, 2) and Golondrina Series, Bajo de La Leona, Laguna Polina, Santa Cruz, Argentina (Archangelsky, 1958, Fig. 50).

Number of specimens. Two. Gangamopteris clarkeana Feistmantel, 1890 (Fig. 7E) Description. The middle part of an incomplete leaf is present in the collection. Leaf measures 4.6 × 2.6 cm in size, margin entire. Median



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Fig. 12. A, Glossopteris browniana. Specimen no. TTU-ATP 106B; B, G. browniana. Specimen no. TTU-ATP 037; C, Glossopteris spatulata. Specimen no. TTU-ATP 068; D, Glossopteris longicaulis. Specimen no. TTU-ATP 109B; E, G. spatulata. Specimen no. TTU-ATP 068; F, G. spatulata. Specimen no. TTU-ATP 137; G, Glossopteris damudica. Specimen no. TTU-ATP 030; H, Glossopteris subtilis. Specimen no. TTU-ATP 099B.

region occupied by 8 subparallel veins. Distance between two veins is 0.3 mm. Veins are evanescent towards the apex. Secondary veins arise from median veins at acute angles of about 28°, arch slightly backwards,

dichotomize frequently, anastomose and form narrow, elongate, arcuate to oblong, 2–4 mm long and 0.3–0.5 mm broad meshes near the median region and narrow elongate trapezoid, 2.0–3.0 mm long and


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0.3–0.5 mm broad meshes near the margin before meeting it at about 53°. Vein density is 14–18 per cm near the median region and 19–27 per cm near the margin. Comparison. The leaf is identical in venation pattern with G. clarkeana Feistmantel (1890, Pl. 20, Fig. 3) described from the Upper Coal Measures, Bowenfels, New South Wales, Australia. The species is additionally known from the Talchir Formation, Auranga Coalfield, Damodar Basin (Tewari and Srivastava, 2000a, Pl. 2, Fig. 5), Talchir Coalfield, Mahanadi Basin (Singh and Chandra, 2000), the Karharbari Formation, Giridih Coalfield, Damodar Basin, West Bengal (Maithy, 1965, Pl. 4, Fig. 25), the Barakar Formation, Umrer Coalfield (Tewari et al., 2012, Fig. 5C, D), Wardha Basin, Maharashtra and Korba Coalfield, Son-Mahanadi Basin, Chhattisgarh (Singh et al., 2012, Pl. 3, Fig. 3, Pl. 4, Fig. 4), India. Gangamopteris major Feistmantel, 1879 (Fig. 7F, G) Description. Basal halves of incomplete leaves measuring 5.5–5.6 cm long and 2.5–2.6 cm broad are present in the collection. Leaves are simple, base narrow, elongate, one side broken, margin entire, median region occupied by 5–11 distinct sub-parallel veins. Secondary veins arise at about 20° from median veins arch slightly backwards, dichotomize frequently, anastomose and form elongate, oblong meshes before meeting margin at 52°–65°. Meshes 5.8–8.0 mm long and 0.2–0.8 mm broad near the median region and 4.0–6.0 mm long and 0.2–0.8 mm broad near the margin. The density of secondary veins is 8–15 per cm near the median region and 13–15 per cm near the margin. Comparison. Leaves are identical in venation pattern with G. major Feistmantel (1879, Pl. 14, Fig. 3, Pl. 16, Figs. 1, 2, 2a), Maithy (1965, Pl. 1, Fig. 7) described from the Karharbari Formation, Giridih Coalfield, Damodar Basin, West Bengal, India. They also compare with the leaf recorded from the Talchir Formation, Auranga Coalfield, Damodar Basin, Bihar (Tewari and Srivastava, 2000a, Pl. 1, Fig. 2). Gangamopteris walkomii Rigby, 1967 (Fig. 8A, B, C) Description. Leaves incomplete, apical, middle and basal portions preserved separately. Approximately half of the right side of the leaf is just below the apical region broken in one specimen (Fig. 8B) and only the right side of leaf is preserved in another specimen (Fig. 8C). Leaves measure 5.2–8.5 cm long and 1.7–2.2 cm broad, simple, obovate, apex obtuse to rounded, bluntly pointed in one of the specimens (Fig. 8A), base gradually narrows to a flat, 7 mm wide short petiole, margin entire, apparently lobed in the specimen in Fig. 8A, median region occupied by 3–7 distinct sub-parallel veins 0.2–0.5 mm apart which gradually become evanescent towards the apex. Secondary veins arise at acute angles of about 18°–37° from median veins, arch backwards, dichotomize, anastomose and form narrow, elongate, arcuate and reticulate meshes before meeting margin at 52°–55°. Meshes are 3.0–7.0 mm long and 0.4–1.5 mm broad near the median region, and narrow, small trapezoid, 2–5 mm long and 0.3–0.6 mm broad near the margin. The density of secondary veins is 8–21 per cm near the median region and 15–20 per cm near the margin. Remarks. The leaves resemble in shape, nature of apex, distinct subparallel veins, the density of veins and presence of petiole with G. walkomii Rigby (1967, Pl. 25, Figs. 3, 5, 7) described from Lithgow Coal Measures of Permian age, from Duncan's Pass, Narrow Neck, near Katoomba, New South Wales. However, the leaves described here are bigger in size and petiole is wider. Number of specimens. Four. Glossopteris Brongniart, 1828 Type species. Glossopteris browniana Brongniart, 1828 Glossopteris browniana Brongniart, 1828 (Figs. 8D, E, 11H, 12A, B) Description. Complete and incomplete leaves are present in the collection. Leaves are simple, spatulate or spatulate-ovate, apex obtuse, base tapering, narrow, elongate, petiolate, 3.0–21.8 cm long and

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1.5–5.8 cm wide, margin entire, midrib distinct, strong, striate, persistent, 0.4 to 0.6 mm wide at the base, and thinning towards the apex, secondary veins arising at about 20°–40° from the midrib, arching, dichotomizing, anastomosing to meet the margin at about 44°–65°, meshes angled, deltoid, broad, polygonal, 3.0–8 mm long and 0.3–1.5 mm broad near the midrib and narrow, elongate, 1.5–7 mm long and 0.2–0.6 mm broad near the margin, the density of veins is 6–18 per cm near the midrib and 11–29 per cm near the margin. Comparison and remarks. The leaves are identical in shape, nature of apex, base, and venation pattern with G. browniana Brongniart (Feistmantel, 1890, Pl. 13, Fig. 1, Pl. 16, Figs. 3, 4, Pl. 17, Figs. 1, 3–5, 7, Pl. 20, Fig. 2) described from Australia from the Lower Coal Measures, New South Wales and Queensland, and Mersey beds (Early Permian), Tasmania, Permian Coal Measures of Greta, Gulong and Maitland (Rigby et al., 1980, Figs. 7–11) and Late Permian of New Castle beds, New South Wales and Bowen Basin, Queensland (McLoughlin, 1994a, 1994b, Pl. 1, Figs. 1–8), Australia; Bajo de La Leona, Laguna Polina, Santa Cruz, Argentina (Archangelsky, 1958, Figs. 36–38) and Lower Beaufort Subgroup (Late Permian), Mooi River District, Natal, South Africa (Lacey et al., 1975, p. 364). The leaves described by Archangelsky (1958, Figs. 36–38), though compare in venation pattern, differ in the presence of acute apex. The leaf described by Cridland (1963, Fig. 20) from Mount Glossopteris Formation, Mount Schopf, Ohio Range, Antarctica is not G. browniana. It is apparently more similar to Glossopteris damudica. The leaves also compare with those assigned to morphotype C2c and C7 (Prevec et al., 2009, Pl.4, Figs. 7, 8, Pl. 6, Fig. 5) recorded from Normandien Formation (Lopingian) of the Clouston farm locality, KwaZulu-Natal, northeastern Karoo Basin, South Africa. The species is rather well distributed in the Permian sediments of India and is recorded from Karharbari, Barakar, Barren Measures, Raniganj, Kamthi and Bijori formations by several workers (Srivastava, 1956; Saksena, 1962; Maithy, 1965; Maheshwari and Prakash, 1965; Kar, 1968; Kulkarni, 1971; Srivastava, 1979; Singh et al., 2005; Goswami et al., 2006; Singh et al., 2006a, 2006b; Tewari, 2007; Srivastava and Agnihotri, 2010) from almost all the Lower Gondwana basins namely Damodar, South Rewa, Rajmahal, Wardha, Satpura and Mahanadi. The specimens described here compare very well with those described from Early Permian (Barakar Formation) of Umrer Coalfield (Tewari et al., 2012, Fig. 5G–J), and Nand Coalfield (Singh et al., 2005, Pl. 1, Fig. 1) Wardha Basin, North and South Karanpura coalfields (Singh and Maheshwari, 2000, Pl. 2, Fig. 1), Damodar Basin, and Lower part of Kamthi Formation (Late Permian), Kamptee Coalfield (Tewari, 2007, PI. 1, Fig. 1, PI. 5, Fig. 3, PI. 6, Fig. 3, PI. 7, Fig. 7, PI. 9, Fig. 1), Wardha Basin. Number of specimens. Twenty. Glossopteris sp. cf. G. ampla (Fig. 8F) Description. Complete and incomplete leaves representing apical, middle and basal portions present in the collection. Leaves are simple, lanceolate, apex acute, base narrow, extreme basal end not preserved, 8.3–19.0 cm long and 4.5–10.2 cm broad, margin entire, midrib distinct, stout, 1–5 mm wide, persistent, thinning upwards, striate, secondary veins arise at about 31° from the midrib, arching to meet the margin after dichotomizing and anastomosing at about 50°, meshes arcuate, deltoid, angled, 5–11 mm long and 0.5–1 mm broad near the midrib, and elongate, narrow, trapezoid, 3–8 mm long and 0.4–0.8 mm broad near the margin, venation dense, density of veins 9–17 per cm near the midrib and 11–27 per cm near the margin. Remarks. Large and broad nature of leaves, nature of the midrib and venation pattern is comparable with Glossopteris ampla Dana (Feistmantel, 1890, Pl. 19, Figs. 1, 2) recorded from the New Castle beds (Upper Coal Measures) at New Castle, Australia; and Illawara (? Permian) and Mersey beds, Tasmania; Bajo de La Leona, Laguna Polina,

Fig. 13. A Glossopteris damudica. Specimen no. TTU-ATP 123; B, Glossopteris taylori sp. nov. Specimen no. TTU-ATP 067; C, Glossopteris tenuifolia. Specimen no. TTU-ATP 100; D, Glossopteris sp. A. Specimen no. TTU-ATP 098; E, Glossopteris sp. B. Specimen no. TTU-ATP 108; F, Glossopteris sp. C. Specimen no. TTU-ATP 022; G, Scirroma sp. Specimen no. TTU-ATP 006; H, Scirroma sp. Specimen no. TTU-ATP 012.


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Santa Cruz, Argentina (Archangelsky, 1958, Fig. 35) and Mount Glossopteris Formation, Mount Schopf, Ohio Range, Antarctica (Cridland, 1963, Figs. 8, 17). The leaves described by Anderson and Anderson (1985, Pl.133, Figs. 1a, b, 2) as Glossopteris cf. ampla Dana (1849) from Vryheid Formation, Middle Ecca (Early Permian) of South Africa are also comparable with the present leaves but they are much longer (350 mm) in size. The leaves described here have not been assigned to G. ampla since they are smaller in length. Number of specimens. Eight. Glossopteris arberi Srivastava, 1956 (Fig. 8G) Description. Leaves are incomplete, lorate, longer than broad, 4.6–16.0 cm long and 1.4–2.5 cm wide, extreme apical portion broken, apex apparently obtuse, base narrow, elongate, margin entire, midrib flat, striate, persistent, 0.7–4 mm wide, thinning towards the apex, secondary veins arising at about 30° from the midrib, arching, dichotomizing and anastomosing to meet the margin at about 63°, meshes arcuate near the midrib, narrow, elongate, trapezoid through the remainder of the lamina, 4–8 mm long and 0.2–0.4 mm broad near the midrib and 2–4 mm long and 0.2–0.3 mm broad near the margin, the density of veins is 10–19 per cm near the midrib and 18–31 per cm near the margin. Comparison. Leaves are identical in shape, nature of apex, base, and venation pattern with G. arberi Srivastava (Srivastava, 1956, Pl. 9, Fig. 7; Chandra and Surange, 1979, Pl. 7, Figs. 4, 7, Pl. 8, Fig. 6, Pl. 16, Figs. 4, 9, Pl. 17, Fig. 2, Pl. 40, Fig. 1) recorded from the Raniganj Formation, Raniganj Coalfield, Damodar Basin, West Bengal, the specimens described from the Barakar Formation of Umrer Coalfield (Tewari et al., 2012, Fig. 5E, F) and from the Kamthi Formation, Kamptee Coalfield (Tewari, 2007; Pl. 1, Fig. 6, Pl. 4, Figs. 3, 4), Wardha Basin, Maharashtra, India. The leaves also resemble with the morphotype C2b (Prevec et al., 2009, Pl.4, Fig. 6) recorded from the Normandien Formation (Lopingian) of the Clouston farm locality, KwaZulu-Natal, northeastern Karoo Basin, South Africa. Number of specimens. Two. Glossopteris bucklandensis McLoughlin, 1990 (Figs. 8H, 9A–C) Description. Both complete and incomplete leaves are present in the collection. Leaves are oblanceolate, apex acute, apparently mucronate in one of the specimens (Fig. 9A), base-acute cuneate, margin entire. Leaves measure 7.7–15.7 cm long and 2.5–4.5 cm broad, the midrib is thick in the basal region, elevated, flat in one of the specimens (Fig. 9B), striate, tapering upwards, persistent, 2–3 mm wide. Secondary veins arising at about 20°–37° from the midrib, arch slightly or pass straight after dichotomizing and anastomosing to meet the margin at about 37°–53°, meshes long, relatively broad near the midrib, arcuate, 4–9 mm long and 0.5–1.5 mm broad, shorter, narrower and 2–7 mm long and 0.3–0.8 mm broad near the margin. The density of veins is 8–23 per cm near the midrib and 11–32 per cm near the margin. Remarks. Leaves are identical in shape, nature of apex, base, midrib and in venation pattern to G. bucklandensis McLoughlin (1990, Pl 5, Figs. 1, 5, Pl. 6, Figs. 1, 6–9) described from the Black Alley shale, Blenheim Subgroup, Back Creek Group (Late Permian) of Bowen Basin, Queensland, Australia. Number of specimens. Four. Glossopteris communis Feistmantel, 1876 (Fig. 9D) Description. Complete leaves and different parts of apical, middle and basal portions of incomplete leaves are preserved. Shape elliptic, apex obtuse to slightly acute, base narrow, elongate, attenuate, margin entire, leaves are 5.0–19.3 cm long and 3.7–7.5 cm broad, midrib flat, striate, persistent, 2–6 mm wide, gradually thinning upwards, secondary veins arise from midrib at about 35°, arching to meet the margin at about 60° after dichotomizing and anastomosing, meshes arcuate, 6–8 mm long and 0.4–0.7 mm wide near the midrib, narrow, elongate, trapezoid, 5–7 mm long and 0.4–0.8 mm wide elsewhere. Venation dense, density of veins is 12–18 per cm near the midrib and 10–17 per cm near the margin. Comparison and remarks. Leaves are identical in venation pattern with G. communis (Feistmantel, 1879, Pl. 17, Figs. 1, 2; Feistmantel,

1882, Pl. 21, Figs. 13, 14; Feistmantel, 1890, Pl. 17, Figs. 2, 6; Chandra and Surange, 1979, Pl.1, Figs. 2, 3). They compare well with the leaves assigned to Arberia hlobanensis (Anderson and Anderson, 1985) recorded from the Vryheid Formation, Middle Ecca (Early Permian) of South Karoo Basin, South Africa. G. communis is well represented in almost all the Lower Gondwana formations of India namely, Talchir, Karharbari, Barakar, Barren Measures, Raniganj and Kamthi and has been recorded from nearly all the Indian Gondwana basins such as Damodar (Lakhanpal et al., 1976; Chandra and Tewari, 1991; Bajpai and Singh, 1994; Tewari, 1994; Srivastava and Tewari, 1996; Tewari and Srivastava, 1996, 2000a; Srivastava and Tewari, 2001), Wardha (Sundaram and Nandi, 1984; Singh et al., 2005; Tewari, 2007, 2008; Tewari et al., 2012), Godavari (Tewari and Jha, 2006), Mahanadi (Goswami et al., 2006), Satpura (Feistmantel, 1880; Crookshank, 1936; Srivastava and Agnihotri, 2010), Rajmahal (Maheshwari and Prakash, 1965) and South Rewa (Lakhanpal et al., 1976). The species is, additionally recorded from Early Permian of the extra peninsular north-east Himalayan region from Arunachal Pradesh (Tewari and Srivastava, 2000b; Maithy et al., 2006) and from northwestern Kashmir region (Singh et al., 1982; Srivastava, 2004). Other than India, the species is known from Australia from Permian sediments of Bowenfels (Feistmantel, 1890), and from South America from Early Permian Rio Bonito Formation, Parana Basin, Brazil (Iannuzzi, 2010). Number of specimens. Fourteen. Glossopteris sp. cf. Glossopteris conspicua (Figs. 9E) Description. The leaves are incomplete. Different parts of apical, middle and basal portions of the leaves are preserved. One of the leaves (not illustrated), though apparently complete, is fractured at two places and is ovate in shape with obtuse apex and narrow, elongate base. Margin entire. Leaves measure 6.3–15.2 cm in length and 2.3–4.0 cm in width. Midrib distinct, 2–8 mm wide, elevated with prominent striations, persistent, thinning towards the apex, secondary veins arising from midrib at about 45°–50°, arching slightly, dichotomizing and anastomosing to meet the margin at about 45°–55°, meshes angled, deltoid and 4–8 × 0.8–1.5 mm in size near the midrib, broad, polygonal, oblong and 3–7 × 0.5–1.3 mm in size elsewhere. The density of veins is 7–12 per cm near the midrib and 9–19 per cm near the margin. Remarks. Venation pattern of the leaves is comparable with G. conspicua Feistmantel (1881, Pl. 28A, Figs. 1, 5, 6, 8, 9), Chandra and Surange (1979, Pl. 5, Fig. 5 Pl. 7, Fig. 1, Pl. 21, Fig. 6). However, the meshes apparently are far too narrow as compared to G. conspicua. Hence, the leaves are described here as a variety of G. conspicua. Number of specimens. Four. Glossopteris damudica Feistmantel, 1881 (Figs. 9F, G, 12G, 13A) Description. The leaves are incomplete. Middle and basal parts of the leaves preserved, base narrow, extreme basal end broken, margin entire. Leaves measure 13.11–19.4 × 5.2–6.1 cm in size. Midrib distinct, elevated, stout, 1–9 mm wide, longitudinally striate, striations strong. Secondary veins arising at about 40°–50° from the midrib and after dichotomizing and anastomosing run straight to meet the margin at about 68°–85°, meshes angled, deltoid, short, broad, polygonal, 3–8 mm long, 0.5–1.5 mm broad near the midrib and elongate, narrower, 3.5–8 mm long and 0.4–0.8 mm wide elsewhere. The density of veins is 10–15 per cm near the midrib and 15–26 per cm near the margin. Comparison and remarks. Leaves compare in midrib and venation pattern with G. damudica (Feistmantel, 1881, Pl. 30A, Figs. 1, 2, Pl. 31A, Figs. 1–3, Pl. 32A, Fig. 1, Pl. 40A, Fig. 6; Plumstead, 1962, Pl. 5, Figs. 1, 7; Chandra and Surange, 1979; Pl. 4, Fig. 2, Pl. 20, Fig. 1, Pl. 36, Fig. 1). The species occurs quite frequently in India and has been recorded from the Karharbari, Barakar, Ironstone shale, Raniganj and Kamthi formations of Giridih and Raniganj coalfields, Damodar Basin, West Bengal (Feistmantel, 1881), Barakar Formation of Ramkola Coalfield, South Rewa Gondwana Basin, Madhya Pradesh (Feistmantel, 1881) and Umrer Coalfield, Wardha Basin, Maharashtra (Tewari et al., 2012), Barren Measures Succession of Jharia Coalfield, Damodar Basin, West Bengal (Srivastava and Tewari, 2001) and Late Permian/ Early Triassic



Kamthi Formation, Talcher Coalfield, Mahanadi Basin, Odisha (Singh and Chandra, 2000). The species has additionally been recorded from Bajo de La Leona, Laguna Polina, Santa Cruz, Argentina, South America (Archangelsky, 1958, Fig. 34). However, the venation pattern of this leaf differs from that of G. damudica since the secondary veins apparently meet the margin at right angle. Besides, G. damudica has been recorded from Early Permian sediments of Ross Sea Area, Antarctica (Plumstead, 1962). The leaves are comparable with the one recorded from Normandien Formation (Lopingian) of the Clouston farm locality, KwaZulu-Natal, northeastern Karoo Basin, South Africa and assigned to morphotype C6a (Prevec et al., 2009, Pl.6, Fig. 2). This particular leaf, though placed under morphotype C6a, differs from other leaves of this type (Prevec et al., 2009, Pl.5, Figs. 6, 7, Pl.6, Fig. 1) in venation pattern. The secondary veins meet the margin at right angles in these leaves, whereas, they are arched in the leaf photographed in Pl.6, Fig. 1. Number of specimens. Five. Glossopteris fuchsii Plumstead, 1962 (Fig. 10B) Description. Leaves are incomplete. Apical, middle and basal parts of leaves preserved separately. Shape lanceolate, apex rounded, base narrow, margin entire, leaves measure 15.8–18.0 × 4.8–6.8 cm in size; midrib distinct, elevated, persistent, 4 mm wide at the base, thinning towards the apex, secondary veins arising at about 38° from the midrib, slightly arching, thereafter, follow a straight course to meet the margin at about 53° after dichotomizing and anastomosing, meshes broad, polygonal 4.4–7 mm long, 0.6–1.1 mm broad near the midrib, and narrow, trapezoid, 1.8–5 mm long and 0.4–0.6 mm wide near the margin. The density of veins is 10–15 per cm near the midrib and 16–21 per cm near the margin. Comparison and remarks. Leaves are identical with G. fuchsii Plumstead (1962, Pl. 12, Figs. 1–3) described from the PermoCarboniferous of the Weddell Sea Area, Theoron Mountains, Antarctica in shape, nature of the midrib and in venation pattern. However, one of the leaves described by Plumstead (Pl. 12, Fig. 1) has an acute apex, whereas, the other (Pl. 12, Fig. 3) has an obtuse apex. Plumstead's specimens, additionally, show denser venation (28 to 38 veins per cm) near the margin. Number of specimens. Three. Glossopteris sp. cf. G. gigas (Figs. 10C, E) Description. Two incomplete leaves measuring 8.8–20 cm long and 5– 10 cm broad are present in the collection. Shape apparently ovate, apex though broken on one side, appears rounded, base broken, margin entire, midrib distinct, flat, striate, persistent, 3– mm wide at the base, thinning towards the apex, secondary veins arising at about 24°–36° from the midrib, arching slightly to meet the margin at about 39°–45° after dichotomizing and anastomosing, meshes narrow, elongate throughout, 6–15 mm long and 0.6\1.0 mm broad near the midrib, and 5–8 mm long and 0.4–0.8 mm wide near the margin. The density of veins is 9–15 per cm near the midrib and 15–20 per cm near the margin. Remarks. Leaves are identical with Glossopteris gigas (Pant and Singh (1971, Pl. 3, Figs. 10, 14; Chandra and Surange, 1979, Pl. 12, Fig. 1, Pl. 25, Fig. 1; Tewari, 2007, Pl. 2, Fig. 3, Pl. 3, Fig. 2) described from the Raniganj Formation of Raniganj Coalfield, West Bengal and Kamthi Formation of Kamptee Coalfield, Maharashtra, India in shape, large size and nature of the midrib. However, in the leaf described by Pant and Singh (1971) the veins arch outwards within 1 cm after emerging from the midrib and thereafter, travel straight to meet the margin at an angle of 70°. The veins of the leaf described here, run parallel to the midrib for about 2 to 3 cm before arching and do not follow a straight course. Therefore, the leaf, though differing in venation pattern has been described here as a variety of G. gigas on the basis of other similar characters. Number of specimens. Two. Glossopteris indica Schimper, 1869 (Fig. 10D) Description. Complete leaves and different portions of apical, middle and basal regions of incomplete leaves are preserved separately. Shape is oblanceolate, apex acute to rounded, base narrow, attenuate, margin

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entire. Leaves measure 6.7–23.3 × 4.1–7.0 cm in size, midrib strong, distinct, elevated, persistent, 2–4 mm wide, wider at the base, thinning towards the apex, secondary veins arise at about 30°–35° from the midrib, slightly arch, form gentle curves to meet the margin at about 55°–60° after dichotomizing and anastomosing. Meshes short, broad, arcuate, 4–9 mm long and 0.5–1.1 mm broad near the midrib, and long, narrow, 3–8 mm long and 0.3–0.8 mm wide elsewhere. The density of veins is 8–20 per cm near the midrib and 13–27 per cm near the margin. Comparison and remarks. Leaves are identical with G. indica Schimper (Chandra and Surange, 1979, Pl. 5, Fig. 1, Pl. 10, Fig. 4, Pl. 15, Fig. 11, Pl. 28, Fig. 1, Pl. 29, Fig. 1; Tewari and Srivastava, 2000a, Pl. 1, Fig. 4) in shape, nature of the midrib and in venation pattern. G. indica is of a common occurrence in India and is recorded throughout the Permian Period (Chandra and Surange, 1979; Bajpai and Singh, 1994; Tewari and Srivastava, 2000a; Goswami et al., 2006; Tewari and Jha, 2006; Tewari, 2007, 2008; Srivastava and Agnihotri, 2010) from various Lower Gondwana basins. Additionally, it is known from the PermoCarboniferous of Weddell Sea Area, Whichaway Nunataks (Plumstead, 1962); Early Permian (Rio Bonito Formation) of Itarare Group, Parana Basin, Rio Grande do Sul State, Brazil (Lundquist, 1919; Iannuzzi, 2010); Vereeniging (Middle Ecca Formation), Transvaal, South Africa (Plumstead, 1952); Mount Glossopteris Formation, Mount Schopf, Ohio Range, Antarctica (Cridland, 1963) and Newcastle (Rigby et al., 1980), Australia, and Late Permian of Mooi River District of Natal, South Africa (Lacey et al., 1975) and Bajo de La Leona, Laguna Polina, Santa Cruz, Argentina (Archangelsky, 1958). Number of specimens. Eleven. Glossopteris karanpuraensis Kulkarni, 1971 (Fig. 10F) Description. An incomplete leaf is present in the collection. Apical half of the leaf preserved, apex acute. Leaf measures 13.0 × 4.0 cm in size, margin entire, midrib thick, persistent, 3 mm broad in the middle region, secondary veins arise at about 52° from the midrib, gently arch to meet the margin at about 56° after dichotomizing and anastomosing, meshes broad, polygonal throughout, deltoid, 5–7 mm long and 0.7–0.9 mm broad near the midrib and narrower, 2–5 mm long and 0.6–0.7 mm broad near the margin. The density of veins is 12–20 per cm near the midrib and 21–26 per cm near the margin. Comparison and remarks. The leaf is identical in overall shape, nature of the midrib and venation pattern with G. karanpuraensis Kulkarni (Chandra and Surange, 1979, Pl. 2, Fig. 5, Pl. 22, Figs. 1, 2, Pl. 30, Fig. 1). The species is of sporadic occurrence and till date is not recorded out of India. This is the first record of G. karanpuraensis from Antarctica. In India, it is reported from the Early Permian Barakar Formation of South Karanpura Coalfield (Kulkarni, 1971) and Raniganj Coalfield (Maheshwari and Tewari, 1992), Damodar Basin, West Bengal and Ib River Coalfield (Goswami et al., 2006), Mahanadi Basin, Odisha. The leaf compares in venation pattern with morphotype C5 (Prevec et al., 2009, Pl. 5, Fig. 5) recorded from Normandien Formation (Lopingian) of the Clouston farm locality, KwaZulu-Natal, northeastern Karoo Basin, South Africa. Glossopteris longicaulis Feistmantel, 1879 (Figs. 10A, G, 12D) Description. Incomplete specimens are present in the collection, mainly basal half of the leaves preserved, base obtuse normal with long petiole. Only one of the leaves is complete apparently slightly retuse with two notches on either side of the leaf margin in the apical region in one of the specimens (12D). Leaves measure 9.8–24.3 cm long and 2.0–6.8 cm broad, margin entire, midrib broad distinct, flat to elevated, striate, occupying almost the entire width of basalmost part and petiole where 1–4 mm wide, secondary veins thin, numerous, arise at about 20°–40° from the midrib, arch to meet the margin at 40°–48° after dichotomizing and anastomosing, meshes arcuate, 3–9 mm long and 0.3–1.0 mm broad near the midrib and narrower, 2–8 mm long and 0.3–0.8 mm broad near the margin. The density of veins is 6–20 per cm near the midrib and 10–25 per cm near the margin. Comparison and remarks. The leaf is identical in the nature of the midrib, in possessing a long petiole and in the venation pattern to Glossopteris


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longicaulis Feistmantel (Chandra and Surange, 1979, Pl. 1, Fig. 4, Pl. 15, Fig. 13). The species is known from India from the Karharbari Formation, Giridih Coalfield (Chandra and Surange, 1979), Damodar Basin, West Bengal, Barakar Formation, Talcher Coalfield (Goswami et al., 2006), Mahanadi Basin, Odisha and Umrer Coalfield (Tewari et al., 2012) and Kamthi Formation, Wardha Valley Coalfield (Tewari, 2008) Wardha Basin, Maharashtra Wardha Basin, Maharashtra. Additionally, the species is also recorded from the Early Permian Rio Bonito Formation of Morro do Papaléo flora, Parana Basin, Brazil (Iannuzzi, 2010). Number of specimens. Six. Glossopteris major Pant and Singh, 1971 (Fig. 11A–C) Description. Both complete and incomplete specimens are present in the collection. Leaves are large, longer than broad, measure 9.0–12.6 cm long and 3.0–5.0 cm broad, oblanceolate in shape, apex obtuse, base acute normal, narrow, margin lobed, midrib distinct, elevated, striate, persistent, 2–3 mm wide in the basal and middle regions, gradually thinning upwards, secondary veins arise at about 35°–40° from the midrib, arch to meet the margin at 40°–50° after dichotomizing and anastomosing, meshes arcuate, 4–5 mm long and 0.4–0.8 mm broad near the midrib, long and narrow throughout the lamina, and 3–6 mm long and 0.2–0.7 mm broad near the margin. The density of veins is 12–19 per cm near the midrib and 15–25 per cm near the margin. Comparison and remarks. The leaf is identical in shape, nature of apex, base, midrib, venation pattern and lobed margin with G. major Pant and Singh (1971, Pl. 1, Fig. 1, Text-fig. 1A), Chandra and Surange, 1979, Pl. 45, Fig. 2, Text-fig. 42A) described from the Raniganj Formation, Raniganj Coalfield, Damodar Basin, West Bengal, India. Additionally, the species is recorded from the Barakar Formation and lower part of the Kamthi Formation of Ib River Coalfield, Mahanadi Basin, Odisha (Goswami et al., 2006). Other than India and Allan Hills, Antarctica, G. major is not recorded. Number of specimens. Three. Glossopteris occidentalis (White, 1908) Tybusch and Iannuzzi, 2010 (Fig. 11D–F) Description. Leaves are incomplete, large, measure 13.5–16.4 cm long and 6.3–7.0 cm broad, apex acute, basal portion not preserved, margin entire to lobed, midrib distinct, flat, striate, persistent, 1–5 mm wide in basal and middle regions, gradually thinning upwards, secondary veins arise at about 20°–30° from the midrib, bend sharply, then follow a straight and parallel course at nearly right angles to the midrib and meet the margin at 50°–70° after dichotomizing and anastomosing, meshes oblong, polygonal, short, 5–8 mm long and 0.3–0.8 mm broad near the midrib, linear, trapezoid and narrow in between the midrib and margin, and narrower and shorter, 0.6–1.1 mm long and 0.3–0.6 mm broad near the margin. The density of veins is 16–23 per cm near the midrib and 22–30 per cm near the margin. Comparison and remarks. The leaves are identical in nature of apex, midrib and venation pattern with G. occidentalis (White) Tybusch and Iannuzzi (2010, Figs. 4A–D, 5A–F, 6 A, B) described from the Early Permian Rio Bonito Formation, Itararé Group, Rio do Rasto Road, near Lauro Müller, Parana Basin, Santa Catarina, southern Brazil. Based on the occurrence in the Morro do Papaléo outcrop in Mariana Pimentel, Rio Grande do Sul State (Tybusch and Iannuzzi, 2010), southernmost Brazil the species now extends to the top of the Itararé Group. Therefore, in the Parana Basin, it ranges from the Middle Sakmarian to Early Artinskian in age. In Argentina, the species is recognized from the Arroyo Totoral and La Colina formations in the Paganzo Basin and in the lower member of the El Imperial Formation of the San Rafael Basin which are of Asselian– Artinskian age (Archangelsky, 1996). A doubtful record of the species is from the “Productive Series” of the Zambezi Basin, Mozambique which is tentatively assigned an Artinskian age (Bernardes-de-Oliveira and Pons, 1975). Apart from Brazil and Argentina, and a doubtful record from Mozambique this is the first record of G. occidentalis from the Weller Formation, Allan Hills, Antarctica. The leaves compare well in shape and venation pattern with morphotype C6a described from Normandien Formation (Lopingian) of the Clouston farm locality, KwaZulu-Natal,

northeastern Karoo Basin, South Africa (Prevec et al., 2009, Pl. 5, Figs. 6, 7, Pl. 6, Fig. 1). Number of specimens. Three. Glossopteris pandurata Pant and Gupta, 1971 (Fig. 11G) Description. A single specimen is present in the collection. Leaf is complete, small, widest near apex, measures 4.8 × 1.6 cm in size, spatulate in shape, apex broad, flat, base narrow, elongate, attenuate, extreme basal end flat, margin entire, midrib distinct, flat, striate, persistent, 2 mm wide in the basal region, gradually thinning towards the apex, secondary veins arise at about 25° from the midrib, arch to meet the margin at 63° after dichotomizing and anastomosing, meshes arcuate, 4–5 mm long and 0.3–0.6 mm broad near the midrib, long, narrow, trapezoid throughout the lamina, 3–5 mm long and 0.4–0.6 mm broad near the margin. The density of veins is 11–16 per cm near the midrib and 15–22 per cm near the margin. Comparison and remarks. The leaf is identical in shape, nature of apex, midrib and venation pattern with G. pandurata Pant and Gupta (1971, Pl. 21, Fig. 39, Text-fig. 2A), Chandra and Surange (1979, Pl. 8, Fig. 2, Pl. 17, Fig. 6, Pl. 43, Fig. 6) described from the Karharbari Formation, Giridih Coalfield, Bihar, India. Besides, the species has also been recorded from the Barakar Formation, Raniganj Coalfield, Damodar Basin, West Bengal (Maheshwari and Tewari, 1992) and lower part of Kamthi Formation, Talcher Coalfield, Mahanadi Basin, Odisha (Goswami et al., 2006). Other than India, the species is not recorded. This is the first record of the species from the Allan Hills, Antarctica. Glossopteris spatulata Pant and Singh, 1971(Fig. 12C, E, F) Description. Both incomplete and complete leaves are present in the collection. Leaves measure 9.8–18.9 cm long and 2.0–5.0 cm broad, shape spatulate, apex obtuse, base narrow, elongate, margin entire, midrib distinct, elevated, striate, persistent, 1–3 mm wide, gradually thinning upwards, secondary veins arising at about 28°–40° from the midrib, arching to meet the margin at angles between 40° and 70° after dichotomizing and anastomosing, meshes arcuate, 4.0–16 mm long and 0.3–0.7 mm broad near the midrib and narrow, elongate, trapezoid throughout the lamina, 3–6 mm long and 0.2–0.5 mm broad near the margin. Venation dense, density of veins is 8–20 per cm near the midrib and 15–22 per cm near the margin. Comparison and remarks. The leaf is identical in shape, nature of apex, midrib and in venation pattern with G. spatulata Pant and Singh (1971, Pl. 10, Fig. 60), Chandra and Surange (1979, Pl. 8, Fig. 1, Pl. 12, Fig. 5, Pl. 17, Fig. 7, Pl. 18, Fig. 1, Pl. 27, Fig. 1). G. spatulata is well distributed in almost all the Lower Gondwana horizons of India. It is recorded from the Talchir Formation, Auranga Coalfield, Bihar (Tewari and Srivastava, 2000a), Barakar Formation, Ib River Coalfield, Odisha (Singh et al., 2006a), Umrer Coalfield, Maharashtra (Tewari et al., 2012), Barren Measures Succession (Srivastava and Tewari, 2001) and Raniganj Formation (Tewari, 1994), Jharia Coalfield, Bihar, Raniganj Coalfield, West Bengal (Pant and Singh, 1971), lower part of Kamthi Formation, Ib River and Talcher coalfields, Odisha (Singh et al., 2006a, 2006b) and Kamthi Formation, Kamptee Coalfield, Maharashtra (Tewari, 2007). Additionally, the species is reported from the Early Permian Bhareli Formation of extra peninsular Himalayan region, Arunachal Pradesh (Tewari and Srivastava, 2000b), India. Except for the new record in the Allan hills, Antarctica, the species, till date, is known only from India. Number of specimens. Five. Glossopteris subtilis Pant and Gupta, 1971 (Fig. 12H) Description. Leaves incomplete. Middle parts of leaves measuring 8.1–8.3 cm long and 3.4–4.0 cm broad preserved, apex broken, apparently obtuse, margin entire, midrib thin, striate, 1–2 mm wide, gradually thinning upwards, secondary veins arise at about 44° from midrib, travel without curving to meet the margin at about 54° after dichotomizing and anastomosing, meshes elongate throughout the lamina, 5–7 mm long and 0.7–0.8 mm broad near the midrib, 2–4 mm long and 0.3–0.5 mm broad near the margin. The density of veins is 12–15 per cm near the midrib and 18–24 per cm near the margin.



Comparison and remarks. The leaf is identical in venation pattern with G. subtilis Pant and Gupta (Chandra and Surange, 1979, Pl. 3, Fig. 7, Pl. 8, Figs. 5, 7, Pl. 14, Fig. 3, Pl. 17, Fig. 16, Pl. 21, Fig. 2, Pl. 22, Figs. 3, 12, Pl. 37, Fig. 2). G. subtilis is recorded from India from the Barakar Formation, Manuguru area, Godavari Graben (Tewari and Jha, 2006), Telangana, Ib River Coalfield, Mahanadi Basin, Odisha (Singh et al., 2006a), Barren Measures Succession, Jharia Coalfield, Bihar (Srivastava and Tewari, 2001) and Raniganj Formation, Raniganj Coalfield, Damodar Basin, West Bengal (Pant and Gupta, 1971), Lower Kamthi Formation, Talcher Coalfield, Mahanadi Basin, Odisha (Singh et al., 2006b) and lower part of Kamthi Formation, Kamptee Coalfield (Tewari, 2007) and Wardha Valley Coalfield, Wardha Basin (Tewari, 2008), Maharashtra. Additionally, the species is reported from the Early Permian Bhareli Formation of extra peninsular Himalayan region, Arunachal Pradesh (Tewari and Srivastava, 2000b). This is the first record of G. subtilis from the Allan Hills, Antarctica. The leaf compares in venation pattern with morphotype C6b recorded from Normandien Formation (Lopingian) of the Clouston farm locality, KwaZulu-Natal, northeastern Karoo Basin, South Africa (Prevec et al., 2009, Pl. 6, Fig. 3). Number of specimens. Two. Glossopteris taylori sp. nov. (Fig. 13B) Diagnosis. Leaf obovate, apex broad, obtuse, slightly rounded, base narrow, margin entire, midrib flat, striate, evanescent in apical region, margin entire, lateral veins arise at acute angles from midrib, arch slightly near the midrib, thereafter, run straight to meet margin, meshes elongate, trapezoid throughout, slightly broader near the midrib. Description. Part and counterpart of a single leaf are present in the collection. The leaf is almost complete except for the extreme basal end, which is slightly broken. The leaf is obovate in shape with a broad, obtuse, slightly rounded, almost flat apex, narrow base, entire margin and measures 15.7 cm long and 7.3 cm broad, widest a few centimteres below the apex, midrib distinct, flat, striate, thin, 4 mm wide in basal and middle regions narrowing upwards, almost invisible in the apical portion. Secondary veins arise at about 30° from midrib, arching slightly, continue straight to meet the margin at about 60° after dichotomizing and anastomosing, meshes elongate and broad throughout, slightly broader near the midrib, about 5–9 mm long and 0.7–0.8 mm broad and 5–6 mm long and 0.5–0.8 mm broad near the margin. The density of veins is 15–18 per cm near the midrib and 18–24 per cm near the margin. Holotype. Specimen no. TTU-ATP 067, Museum of Texas Tech University, Lubbock, Texas, USA. Locality and Horizon. Allan Hills, central Transantarctic Mountains, South Victoria Land, Antarctica, Weller Formation. Etymology. After Thomas N. Taylor for his major contribution to Antarctic plant fossils. Comparison. G. taylori is different from all the known species of Glossopteris. It shows a slight resemblance with G. pandurata in its shape (Pant and Gupta, 1971, Pl. 21, Fig. 39, Text-fig. 2A; Chandra and Surange, 1979, Pl. 8, Fig. 2, Pl. 17, Fig. 6, Pl. 43, Fig. 6). However, G. pandurata is a small leaf, with a notched apex, a persistent midrib and medium, narrow meshes. The venation is dense like G. communis (Chandra and Surange, 1979, Pl. 1, Figs. 2, 3, Pl. 15, Fig. 1; Feistmantel, 1879, Pl. 21, Figs. 13, 14). However, shape of G. communis is different and it has a persistent midrib. Glossopteris tenuifolia Pant and Gupta, 1968 (Fig. 13C) Description. A complete leaf is present in the collection. Shape linear, lorate, apex acute, base attenuate, margin entire. Leaf measures 11.6 cm long and 1.9 cm broad, midrib thin, persistent, 1 mm wide at the base, gradually tapering upwards, secondary veins arising at about 25° from the midrib, arching to meet the margin at 40° after dichotomizing and anastomosing, meshes arcuate, 3–6 mm long and 0.4–0.5 mm broad near the midrib, long, narrow, trapezoid throughout the lamina, 3–5 mm long and 0.3–0.4 mm broad near the margin. The density of veins is 21–25 per cm near the midrib and 23–31 per cm near the margin.

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Comparison and remarks. Leaf is identical in shape, apex, base and venation pattern with G. tenuifolia (Pant and Gupta, 1968, Pl. 21, Fig. 15; Chandra and Surange, 1979, Pl. 6, Fig. 1, Pl. 15, Fig. 10, Pl. 17, Fig. 10, Pl. 42, Figs. 1, 6). The species is rather common in Indian lower Gondwana horizons and has been recorded from the Talchir Formation, Jharia Coalfield, Bihar (Tewari and Srivastava, 1996), Barakar Formation, Auranga Coalfield, Bihar (Tewari and Srivastava, 2000a), Ib River and Talchir coalfields, Odisha (Goswami et al., 2006), Umrer Coalfield, Maharashtra (Tewari et al., 2012), Raniganj Formation, Raniganj Coalfield, West Bengal (Pant and Gupta, 1968; Chandra and Surange, 1979), Jharia Coalfield, Bihar (Tewari, 1994), lower part of Kamthi Formation, Talcher Coalfield, Odisha (Goswami et al., 2006), Kamthi Formation, Kamptee Coalfield (Tewari, 2007) and Wardha Valley Coalfield (Tewari, 2008), Maharashtra. The species is additionally known from extra peninsular Himalayan region of India, from the Bhareli Formation of Arunachal Pradesh (Tewari and Srivastava, 2000b). Other than India, this is the first record of G. tenuifolia from the Allan Hills, Antarctica. The leaves from Estcourt Formation (Late Permian), North East Karoo Basin, South Africa, assigned to Lidgettonia africana by Anderson and Anderson (1985, Pl.114, Figs. 1–9) compare well with the leaf described here in overall shape and venation pattern. The leaf is also comparable in shape, apex and venation pattern with leaves assigned to morphotype C2a recorded from Normandien Formation (Lopingian) of the Clouston farm locality, KwaZulu-Natal, northeastern Karoo Basin, South Africa (Prevec et al., 2009, Pl. 4, Figs. 3, 4, 5). Glossopteris sp. A (Fig. 13D) Description. Middle and basal parts of an incomplete leaf measuring 29.0 × 6.4 cm in size, preserved. A characteristic feature of the leaf is that the lamina is inflated in the basal region just a few centimetres above the base, which is broken. Leaf margin entire, midrib stout, elevated, striate, 3.0 mm wide, gradually thinning upwards. Secondary veins arising at about 27° from the midrib, arching, dichotomizing and anastomosing to meet the margin at about 52°, meshes elongate, narrow, trapezoid throughout, arcuate, 8–13 mm long and 0.6–0.8 mm broad near the midrib and 7–9 mm long and 0.5–1.1 mm broad near the margin. The density of veins is 17–21 per cm near the midrib and 18–22 per cm near the margin. Remarks. The leaf is characteristically inflated in the basal region. This feature is not observed in the known leaves of the genus Glossopteris. Since the leaf is incomplete, it is described here as Glossopteris sp. A. Glossopteris sp. B (Fig. 13E) Description. Leaf almost complete, measures 15.5 × 4.0 cm in size. Shape apparently lanceolate, apex broken, base narrow, petiolate, margin entire. In the basal part, the margin characteristically meets the petiole at about 37° on the left side and at about 39° on the right side. Midrib distinct, flat, striate, persistent, 2.0 mm wide, thinning upwards. Secondary veins arising at about 35° from the midrib, arching slightly to meet the margin at about 57° after dichotomizing and anastomosing, meshes broad, polygonal throughout, angled, deltoid, 5–7 mm long and 0.7–1.0 mm broad near the midrib and 3–5 mm long and 0.5–0.8 mm broad near the margin. The density of veins is 11–13 per cm near the midrib and 18–21 per cm near the margin. Remarks. The leaf is comparable with G. browniana Brongniart in having a similar venation pattern. However, it differs in its characteristic base. The base of the leaf is comparable with Glossopteris cf. leptoneura described by Lacey et al. (1975) from the Upper Permian of Mooi River District of Natal, South Africa, but this leaf is narrower. Since the leaf is incomplete it is described here as Glossopteris sp. B. Glossopteris sp. C (Fig. 13F) Description. A complete ovate leaf with obtuse apex, narrow, petiolate base and entire margin is present in the collection. Leaf is 3.6 cm long and 1.4 cm wide, midrib distinct, elevated, persistent, 9 mm wide, thinning towards the apex, secondary veins arising at about 47° from midrib, arching, dichotomizing once or twice, anastomosing rarely and meet the margin at about 70°, meshes arcuate near the midrib,


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Fig. 14. A, Scale leaf A. Specimen no. TTU-ATP 014; B, Nautiyalolepis sp. Specimen no. TTU-ATP 023; C, Utkaliolepis indica. Specimen no. TTU-ATP 054; D, Scale leaf of Eretmonia. Specimen no. TTU-ATP 017; E, Counterpart of specimen in Fig. D; F, Surangephyllum elongatum. Specimen no. TTU-ATP 099A.



narrow, elongate elsewhere, 2–3 mm long and 0.4–0.6 mm broad near the midrib and 2.4–3.4 mm long and 0.3–0.6 mm broad near the margin, density of veins 23–31per cm near the midrib and 25–35 per cm near the margin. Remarks. The leaf, though complete shows scratches on its surface due to which the venation is not very clear. Hence it is described here as Glossopteris sp. C. Genus — Scirroma Surange and Chandra, 1974b Type species — Scirroma angusta Surange and Chandra, 1974b Scirroma sp. (Fig. 13G, H) Description. Leaves complete, shape rhomboid, apex obtuse, base broad, margin entire, measure 0.7–1.2 × 0.6–0.7 cm in size, median region occupied by 4–5 parallel veins, lateral veins arise from median veins at about 30°–35°, dichotomize and anastomose to form narrow, oblong meshes, veins thick near the margin. Comparison and remarks. The leaves are comparable with the scale leaf Scirroma angusta described by Chandra and Surange (1977, Text fig. 1D) from the Raniganj Formation of Raniganj Coalfield, Damodar Basin, West Bengal in rhomboid shape and venation pattern. However, Chandra and Surange (1977) described the cuticular features of S. angusta which are not preserved in present leaves. Therefore, these are described here as Scirroma sp. Genus — Nautiyalolepis Tiwari et al., 2009 Type species — Nautiyalolepis lanceolata Tiwari et al., 2009 Nautiyalolepis sp. (Fig. 14B) Description. Leaf complete, measures 3.8 cm long and 1.6 cm broad, ovate in shape, apex acuminate, base long, narrow, curved and 0.4 mm in width. Veins spread from the base and run almost straight to meet the margin. Dichotomization and anastomosis are not visible. Comparison and remarks. The present leaf resembles Nautiyalolepis lanceolata described by Tiwari et al. (2009, Pl. 1, Fig. 1) from the Kamthi Formation of Handappa area, Mahanadi Basin, Odisha in having long, strong base but N. lanceolata differs with present leaf in lanceolate shape. Genus — Utkaliolepis Tiwari et al., 2009 Type species — Utkaliolepis indica Tiwari et al., 2009 Utkaliolepis indica (Fig. 14C) Description. Leaf complete, asymmetrical, measures 1.1 × 0.7 cm in size, rhomboid-triangular in shape, apex acuminate, base broad. 3–4 veins arise from the base, run parallel in the median region and reach the apex. Lateral veins also arise from the base, divert at about 25°– 30°, dichotomize and anastomose to form narrow, polygonal, small meshes. Comparison and remarks. Scale leaf is identical with Utkaliolepis indica described by Tiwari et al. (2009, Pl. 2, Fig. 2) from the Kamthi Formation of Handappa Area, Mahanadi Basin, Odisha and, Srivastava and Agnihotri (2012, Pl. 1, Fig. 5) from the Barakar Formation of Pench Valley Coalfield, Satpura Basin, Madhya Pradesh in asymmetrical rhomboid-triangular shape and venation pattern. Genus — Eretmonia (du Toit, 1932) Lacey et al., 1975 Type species — Eretmonia natalensis du Toit, 1932 Scale leaf of Eretmonia (Fig. 14D, E) Description. Part and counterpart of the specimen are present in the collection. Scale leaf complete with a short petiole, lamina fan shaped, 4.7 cm long and 4.8 cm broad at its widest part (apical), margin entire. The width of the petiole is 7 mm. 7–8 veins emerge from the base of petiole, run straight for a short distance and then dichotomize twice or thrice before meeting the margin. The density of veins in apical region is 15–20 veins/cm. Sporangia not preserved Remarks. The scale leaf is comparable with those of Eretmonia natalensis described from the Beaufort Series (Late Permian) of Karoo Basin (du Toit, 1932, Pl.40, Fig. 11), from Mooi River District, Natal (Lacey et al., 1975, NM 1221a) and from the Late Permian of New and Old Wapadsberg Pass, Eastern Cape Province (Prevec et al., 2010, Fig. 8D), South Africa in having a petiole and similar venation

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pattern of lamina. However, the lamina of this leaf is ovate to triangular or even pentagonal and the petiole is long, whereas, the herein described leaf has a fan shaped lamina, and short, wide robust petiole. Additionally, all the leaves from South Africa have sporangia preserved. Since the sporangia are not preserved in the present leaf, it is described here as a scale leaf of Eretmonia. Scale leaf A (Fig. 14A) Description. Scale leaf measures 2.4 × 1.3 cm in size, shape obovate, apex obtuse, base narrow, median region occupied by 0.9 mm thick single vein, which runs straight up to 2/3 part of the lamina. Secondary veins arise from the median vein at about 20° and after dichotomization and anastomosis form narrow, polygonal meshes near the margin. Comparison and remarks. The leaf is very characteristic due to the presence of a thick and solid median vein, which is not present in other known scale leaves. Genus — Surangephyllum Chandra and Singh, 1986 Type species. Surangephyllum elongatum (Lacey et al., 1975) Chandra and Singh, 1986 Surangephyllum elongatum (Lacey et al., 1975) Chandra and Singh, 1986 (Fig. 15 F) Description. A single leaf is present in the collection. Leaf simple, oblanceolate, petiolate, measures 17.3 cm in length and 5.4 cm in width, apex broken, base sagittate to hastate, margin entire to undulate. Leaf unicostate, lamina constricted above basal lobes, petiole 2.4 cm long. Midrib distinct, flat, striate, persistent, 3–4 mm wide, thinning towards the apex. Secondary veins arising from midrib at about 40°, arch slightly, dichotomize, anastomose and meet the margin at about 68°, meshes angled, deltoid, short, broad, 3–4 × 1.2–1.6 mm in size near the midrib, broad, elongate, polygonal, 7–8 × 0.5–0.7 mm in size near the margin. The density of veins is 7–10 per cm near the midrib and 11–21 per cm near the margin. Comparison and remarks. Leaf resembles in shape, nature of base and venation pattern with Surangephyllum elongatum (Chandra and Singh, 1986, Pl.1, Figs. 1–4) described from the Kamthi Formation exposed in the Hinjrida Ghati section near Handappa Village, Mahanadi Basin, Odisha. However, marginal veins arising from midrib on inner edge of each basal lobe are not visible in the herein described leaf. The leaf is also reported from Karharbari biozone of Talchir Coalfield (Singh et al., 2006a, 2006b) and Barakar Formation, Ib River Coalfield (Goswami and Singh 2010), Mahanadi Basin, Odisha. 7. Discussion 7.1. Biostratigraphic correlation The Glossopteris flora of Allan Hills has similarities to the floral assemblages recorded from both the Early and Late Permian of different Gondwanan countries. Accordingly, it shows resemblance with the

Fig. 15. Bar diagram showing number of species of different taxa of Glossopteridales in Weller Formation of Allan Hills.


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R. Tewari et al. / Gondwana Research xxx (2015) xxx–xxx Table 1 Distribution of Glossopteris floral elements of Weller Formation, Allan Hills in different Lower Gondwana basins of peninsular and Himalayan regions of India.



Early Permian assemblages of New South Wales (Feistmantel, 1890), Tasmania (Feistmantel, 1890), Argentina (Archangelsky, 1958, 1996), Brazil (Lundquist, 1919; Tybusch and Iannuzzi, 2008; Iannuzzi, 2010), South Africa (Plumstead, 1952; Anderson and Anderson, 1985), Damodar Basin, West Bengal (Feistmantel, 1879; Maithy, 1965; Kulkarni, 1971; Lakhanpal et al., 1976; Srivastava, 1979; Chandra and Tewari, 1991; Bajpai and Singh, 1994; Tewari and Srivastava, 1996; Srivastava and Tewari, 1996; Singh and Maheshwari, 2000; Tewari and Srivastava, 2000a, 2000b), Mahanadi Basin, Odisha (Chandra and Singh, 1992 and references cited therein, Goswami et al., 2006; Singh et al., 2006a; Singh et al., 2012), Wardha Basin, Maharashtra (Sundaram and Nandi, 1984; Singh et al., 2005; Tewari et al., 2012), Godavari Graben, Telangana (Tewari and Jha, 2006), Satpura Basin, Madhya Pradesh, (Srivastava and Agnihotri, 2012), Arunachal Pradesh (Tewari and Srivastava, 2000b; Maithy et al., 2006) and Kashmir region (Singh et al., 1982; Srivastava, 2004), India (Table 1) and other regions of Antarctica such as Milorgfjella, Dronning Maud Land (Plumstead, 1975), Mount Schopf (Mount Glossopteris Formation), Ohio Range, (Cridland, 1963) and Ross Sea Area (Plumstead, 1962), Antarctica. Besides, the flora shows similarity with the Late Permian assemblages of South Africa (Etheridge, 1902; Seward, 1908; Lacey et al., 1975; Anderson and Anderson, 1985; Prevec et al., 2009; Prevec et al., 2010), Australia (Du Toit, 1932; McLoughlin, 1990, 1994a, 1994b) and different Lower Gondwana basins of India, namely, Damodar, West Bengal (Feistmantel, 1881; Kar, 1968; Srivastava and Tewari, 2001; Tewari, 1994), Godavari, Telangana (Tewari and Jha, 2006), Rajmahal, Bihar (Maheshwari and Prakash, 1965), Wardha, Maharashtra, (Tewari, 2007, 2008), Mahanadi, Odisha (Singh and Chandra, 2000; Singh et al., 2006b; Goswami et al., 2006) and Satpura, Madhya Pradesh (Srivastava and Agnihotri, 2010) (Table 1). 7.2. Composition of the Weller flora Collective fossil records from the Weller Formation of Allan Hills, South Victoria Land, Antarctica (Kräusel, 1962; Maheshwari, 1972; Schopf, 1976; Chatterjee et al., 1983; Retallack and Krull, 1999; Retallack et al., 2005) and this study demonstrate that a Glossopteris flora comprising Equisetales and Glossopteridales existed in the area. The Equisetales include branched and unbranched axes. The Glossopteridales are represented by four species of Gangamopteris namely, G. angustifolia, G. clarkeana, G. major and G. walkomii, twentytwo species of Glossopteris- G. browniana, G. sp. cf. ampla, G. arberi, G. bucklandensis, G. communis, G. sp. cf. G. conspicua, G. damudica, G. fuchsii, G. sp. cf. G. gigas, G. indica, G. karanpuraensis, G. longicaulis, G. major, G. occidentalis, G. pandurata, G. spathulata, G. subtilis, G. taylori sp. nov., G. tenuifolia, Glossopteris sp. A, Glossopteris sp. B, Glossopteris sp. C; Surangephyllum elongatum, sterile scale leaves namely Scirroma sp., Nautiyalolepis sp., Utkaliolepis indica, Scale leaf A and scale leaf of male fructification Eretmonia;, fructification — Plumsteadia ovata and root Vertebraria australis (previous studies) (Fig. 15). Additionally, two fossil woods, Taeniopitys scotti and Araucarioxylon allanii of uncertain affinities are also recorded by earlier workers. The genera Gangamopteris, Glossopteris, Surangephyllum and scale leaf of male fructification Eretmonia, are recorded for the first time from the Weller Formation of the Allan Hills. 8. Conclusion Antarctica offers unique potential for understanding the paleobiogeography and paleoclimatology of early history of Gondwana and dispersal of plant fossils during the deglaciation of the Late Paleozoic Ice Age. The most diverse Antarctic plant assemblages come from the Permian Weller Formation of Allan Hills. The plant fossil assemblages of the Weller Formation, comprising the orders Equisetales and Glossopteridales, have morphological similarities to those of Early and Late Permian of other Gondwana continents. However, they show

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strong similarities to the fossil assemblages of Late Permian of almost all the Lower Gondwana basins of India, namely Damodar, Mahanadi, Wardha-Godavari and Satpura; Karoo Basin of South Africa and to a lesser extent with Bowen Basin of Queensland, Australia. Besides equisetalean axes, three species of Gangamopteris, thirteen species of Glossopteris, Surangephyllum elongatum and scale leaf Utkaliolepis indica are common with the plant fossils of Late Permian formations of India (Table 1); ten species of Glossopteris of South Africa are shared in the flora of Weller Formation, and only two species of Glossopteris namely, G. browniana and G. bucklandensis of Allan Hills are similar to those of Australia. The similarity of Weller flora of Allan Hills with those of India, South Africa and to some extent with Australia corroborates the fact that these Gondwana continents were part of a single phytogeographic province during the Late Permian. Because of Antarctica's central position in the Gondwana assembly, it probably played a vital role in the dispersal of Glossopteris flora. Acknowledgements We thank Nick Hotton, Hal Borns, Bryan Small, and George Jacobson for assistance in the field in the Allan Hills of Antarctica and the National Science Foundation for logistic support, Director, Birbal Sahni Institute of Palaeobotany (BSIP), Lucknow for providing necessary facilities to carry out this research work and Chris Scotese for the paleogeographic maps. We also thank Bill Mueller of the Museum of Texas Tech University and Pradeep Mohan, Technical Officer-B, Birbal Sahni Institute of Palaeobotany, Lucknow, India, for helping in the photography of specimens. One of us (RT) is thankful to Gary Edson, Director (2009) of the Museum of Texas Tech University for granting permission to study the Antarctic plant fossils and for providing hospitality during a short visit at Lubbock. David Dilcher and Bill Mueller have critically appraised the manuscript and offered valuable suggestions. We thank M. Santosh and T. Horscroft for inviting us to prepare this focus review article and three anonymous reviewers for helpful suggestions and insights that strongly improved the manuscript. References Anderson, J.M., Anderson, H.M., 1985. Palaeoflora of Southern Africa. Prodromous of South African megafloras. Devonian to Lower Cretaceous. A.A. Balkema Publishers, Rotterdam, Netherlands, pp. 1–423. Archangelsky, S., 1958. Estudio Paleontologico del Bajo de la Leona, Santa Cruz. Acta geologica lilloana 2, 5–133. Archangelsky, S., 1996. Aspects of Gondwana Palaeobotany: gymnosperms of the Palaeozoic–Mesozoic transition. Review of Palaeobotany and Palynology 90, 287–302. Awatar, R., Tewari, R., Agnihotri, D., Chatterjee, S., Pillai, S.S.K., Meena, K.L., 2013. Late Permian and Triassic palynomorphs from the Allan Hills, central Transantarctic Mountains, South Victoria Land, Antarctica. Current Science 106, 988–996. Bajpai, U., Singh, K.J., 1994. Indian Gondwana. Annotated Synopses. Permian Megaplants (III) vol. II. Birbal Sahni Institute of Palaeobotany, Lucknow. Ballance, P.F., 1977. The Beacon supergroup in the Allan Hills, Central Victoria Land, Antarctica. New Zealand Journal of Geology and Geophysics 20, 1003–1016. Barrett, P.J., Kohn, B.P., 1975. Changing sediment and transport directions from Devonian to Triassic in the Beacon Supergroup of South Victoria Land, Antarctica. In: Campbell, K.S.W. (Ed.), Gondwana Geology. Australian National University Press, Canberra, pp. 15–35. Barrett, P.J., Kohn, B.P., Askin, R., McPherson, J.G., 1971. Preliminary report on Beacon Supergroup studies between Hatherton and Mackay Glaciers, Antarctica. New Zealand Journal of Geology and Geophysics 14, 605–614. Benton, M.J., 2003. When Life Nearly Died. Thames and Hudson, London. Bernardes-de-Oliveira, M.E.C., Pons, D., 1975. Taphoflora of Karroo in the Zambezi Basin (Tete Region, Mozambique). Boletim do Instituto de Geociências 6, 33–53. Borns Jr., H.W., Hall, B.A., 1969. Mawson ‘tillite’ in Antarctica: preliminary report of a volcanic deposit of Jurassic age. Science 166, 870–872. Brongniart, A., 1828. Histoire des vegetaux fossiles ou recherches botaniques sur les vegetaux renfermes dans les diverses couches du globe, Paris 1, pp. 1–136. Cantrill, D.J., Poole, I., 2012. The Vegetation of Antarctica Through Geologic Time. Cambridge University Press, Cambridge, U. K. Caputo, M.V., Crowell, J.C., 1985. Migration of glacial centers across Gondwana during the Paleozoic era. Geological Society of America Bulletin 96, 1020–1036. Caputo, M.V., Melo, J.H.G., Streel, M., Isbell, J.L., 2008. Late Devonian and Early Carboniferous glacial records of South America. In: C. R., Fielding, C.R., Frank, T.D., Isbell, J.L. (Eds.), Resolving the Late Paleozoic Ice Age in Time and Space. Geological Society of America Special Paper 441, pp. 161–173.


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Zhao, L., Taylor, T.N., Taylor, E.L., 1995. Cupulate glossopterid seeds from the Permian Buckley Formation, central Transantarctic Mountains. Antarctic Journal of the United States 30, 54–55. Rajni Tewari is Scientist ‘F’ at Birbal Sahni Institute of Palaeobotany, Lucknow India. Her main research interests include Palaeozoic plant megafossils, Gondwana megaspores, cuticular studies including dispersed angiospermous cuticles from Tertiary of northeast India, palaeoclimate and biostratigraphy. Her current research focuses on Gondwana plant fossils from India, Antarctica and Brazil, their comparative study, evolution and palaeoclimate.

Sankar Chatterjee is Paul Whitfield Horn Professor of Geosciences and Curator of Paleontology at Texas Tech University. He led several expeditions to Antarctica, China, India, and American Southwest in search of dinosaurs and early birds. His research focuses on the origin of life, macroevolution, plate tectonics, mass extinction, and animal flight.

Deepa Agnihotri is Scientist ‘B’ at Birbal Sahni Institute of Palaeobotany, Lucknow India. She obtained her M.Sc. (2005) from Kanpur University and Ph.D. (2011) in Botany from the University of Lucknow, Lucknow. Her research interests include Gondwana plant mega- and microfossils of peninsular and extra peninsular regions of India.

Sundeep K. Pandita is a professor at the Department of Geology, University of Jammu, India. He has been working on the stratigraphy and sedimentology of the Cenozoic successions in Northwest Himalaya. His current research focuses on palaeoenvironment, biostratigraphy and tectonics in Kashmir Himalaya.


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