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On the occurrence of three species of Annularia Sternberg in the Early Permian Glossopteris flora of peninsular India
Review of Palaeobotany and Palynology 153 (2009) 394–407
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Review of Palaeobotany and Palynology j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / r ev p a l b o
On the occurrence of three species of Annularia Sternberg in the Early Permian Glossopteris flora of peninsular India Manju Banerjee ⁎, Samik Mitra, Sutapa Dutta Department of Botany, University of Calcutta, 35, Ballygunge Circular Road, Kolkata-700019, India
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Article history: Received 17 October 2007 Received in revised form 16 October 2008 Accepted 9 November 2008 Available online 14 November 2008 Keywords: In situ Annularia gondwanensis sp. nov. Annularia.sp. Annularia kurtzii Upper Lower Permian Eastern peninsular India Gondwana
a b s t r a c t Three species of articulated foliage-bearing shoots of Annularia Sternberg are described from the Barakar Formation (late Early Permian–late Artinskian) of the Saharjuri Basin, eastern peninsular India. Annularia gondwanensis sp. nov. is preserved in growth position with branches emerging from a vertical carbonized axis. The leaves, branches and axis of A. gondwanensis have preserved cuticles. Annularia sp. and Annularia kurtzii Archangelsky are preserved as impressions. The Annularia species together with diverse sphenopsids occur in a typical Gondwanan assemblage dominated by glossopterids. This late Palaeozoic Euramerian– Cathaysian genus was earlier known from the Lower Permian of various region of the Gondwana continent viz., Brazil and Argentina in South America, the Karoo Basin in South Africa and western extra-peninsular India. Now the biogeographical range of distribution of Annularia extends up to the eastern peninsular India in the Gondwana continent. The palaeoclimate, palaeoecology of the Saharjuri Basin in eastern India is discussed considering the records of diverse sterile and fertile sphenopsids and Annularia in growth position together with the upright Glossopteris plants, other glossopterid members in the upper Lower Permian. © 2008 Elsevier B.V. All rights reserved.
1. Introduction Annularia Sternberg, 1823 is a calamitalean genus with 46 species based on leaves (Table 1). The genus flourished in the warm, humid climate of the late Palaeozoic equatorial belt (Eurameria–Cathaysia) and the palaeotemperate Angaran province. Rare Annularia species also occur in the Gondwana province (Banerjee, 1991; Cleal and Thomas, 1991), in the Lower Permian of Argentina, South America, South Africa and western extra-peninsular India. (Archangelsky, 1960, 1986; Tiwari and Singh, 1981; Anderson and Anderson, 1985; Cuneo, 2000). Recently, well-preserved foliage-bearing branches of a new Annularia species attached to an erect axis, together with two other species of the genus, have been collected for the first time from upper Lower Permian strata of peninsular India. These Annularia species occur in association with diverse fertile and sterile sphenopsid shoots (viz., the bractless fertile plant Tulsidabaria Banerjee et al., 2004, the cone-bearing shoots of Rajmahaliastachys Banerjee and D' Rozario, 1999, species of Sphenophyllum, Phyllotheca and a diverse range of glossopterid leaves and seedlings (Banerjee et al.,
⁎ Corresponding author. E-mail addresses: [email protected], [email protected] (M. Banerjee). 0034-6667/$ – see front matter © 2008 Elsevier B.V. All rights reserved. doi:10.1016/j.revpalbo.2008.11.001
1991; Banerjee, 2000, 2005). In addition, several endemic plants of uncertain affinity, viz., Saharjuria, Chitraphyllum, Santhalitheca (Banerjee, 1999) and ginkgophytes occur in the associated beds. Here we describe the macromorphology of the three Annularia species and the cuticular micromorphology of the leaves and stems of A. gondwanensis sp. nov. 2. Study area and geological setting The location of study is the Saharjuri basin, Deoghar group of coal fields, Jharkhand, eastern India (Fig. 1) covering 30 km2. area between 24°05' to 24°10' N and 86°49' to 86°57 E. The geology and stratigraphic succession (Table 2) of this basin have been investigated by Niyogi (1966), Datta et al. (1983) and Pareek et al. (1988). 2.1. Materials and method The materials described in the present study are from the younger horizon of the coal bearing strata of the basin. The fossils occur in the sandstone, shale layers within coal horizon XII — Tulsidabar Top coal seam (upper Lower Permian), Barakar Formation (Fig. 2). The branches and the leafy shoots are preserved as compression on thin, finegrained carbonaceous shale layer overlying the exposed hard sandstone layer (Plate I, 1; Plate III, 1). Portions of the leafy shoots
M. Banerjee et al. / Review of Palaeobotany and Palynology 153 (2009) 394–407 Table 1 Global distribution of Annularia species Name of Annularia spp.
Phytoprovince Euramerian Cathaysian Angara Gondwana
1. A. acicularis (Dawson) White 2. A. aculeata Bell 3. A. americana Read 4. A. asteris Asteris 5. A. australis Feistmantel 6. A. batschatensis (Chachlov) Radczenko 7. A. crassiscula Halle 8. A. filiformis Jongmans et Gothan 9. A. fimbriata Walton 10. A. galioides (Lindley et Hutton) Kidston 11. A. goapingensis Xiao 12. A. gondwanensis sp. nov. 13. A. gracilescens Halle 14. A. grandifolia Schvedov 15. A. hammanskraalense Anderson & Anderson 16. A. incisa Stockmans et Mathieu 17. A. jerunakovensis Neuburg 18. A. jongmansii Walton 19. A. kurtzii Archangelsky 20. A. lanceolata Radczenko 21. A. latifolia (Dawson) Kidston 22. A. microphylla Sauveur 23. A. minima Leggewie et Schonefeld 24. A. mucronata Schenk 25. A. orientalis Kawasaki 26. A. papilioformis Kawasaki 27. A. paramucronata Schrenk 28. A. pingloensis Sze 29. A. pseudostellata H. Potonié 30. A. Punctata Zhao et Chang 31. A. radiata (Brongniart) Sternberg 32. A. raspadensis Gorelova 33. A. rigida Zhao et Liu 34. A. rigitula Xiao 35. A. sibirica Radczenko 36. A. similistellata Daber 37. A. sphenophylloides (Zenker) Gutbier 38. A. spiculosa Rasskazova 39. A. stellata (Schlotheim) Wood 40. A. subradiata Stockmans et Williere 41. A. subtenuifolia Rasskazova 42. A. shirakii Kawasaki 43. A. tajluganensis Radczenko 44. A. tenuifolia Neuburg 45. A. tridactyla Langford 46. A. vernensis (Arnold) Abbott
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with branches, erect axis and additional isolated specimens of similar leafy shoots were collected. The details of the in situ axis, leafy shoots, mode of emergence of the foliage shoots from the axis, branching pattern were examined in the field and later in the laboratory with the pieces collected. The specimens of the other two Annularia species, viz., Annularia sp. and A. kurtzii Archangelsky are impressions on gray and black shale. Macromorphological characters of the specimens were studied and photographed under a Zeiss SV II Stereo Zoom microscope and a Nikon FM 2 camera. The morphology of the leaves is described following the terminologies proposed by Dilcher (1974) and Ash et al. (1999). The coalified leaves, branches and the erect axis were macerated for cuticle extraction. Peels of coaly layers were first treated with dilute hydrofluoric acid (40%) to remove the silica particles adhered to the pieces. The isolated fragments were treated slowly with 50% to 70% nitric acid, adding KClO3 whenever necessary for oxidation. The oxidized fragments of the fossils were then treated
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with dilute (5%) potassium hydroxide solution to clear up the humic matter. Each step of chemical treatment was followed by a thorough wash with distilled water. The prepared cuticles were mounted on clear glass slides using Euperol. A few cuticular pieces were preserved in 70% alcohol for SEM study. The micromorphic details of the cuticles were studied and photographed with Zeiss Axioscope 2 and Leitz Diaplan compound microscopes, a Video Image Master Capture Kit of Leading Edge Pvt. Ltd., Australia and a Hitachi 600 scanning electron microscope. 3. Systematics Class Sphenopsida Order Equisetales Family Calamitaceae Annularia Sternberg, 1823 The genus Annularia was first described by Sternberg (1823). The macro and micro morphological characters of the genus were further elaborated by Abbott (1958) through study of additional specimens. The major distinguishing macromorphological characters of the genus are the lanceolate to obovate or spathulate shaped single-veined, dorsiventral, leaves of equal or unequal length, positioned in verticils radiating from the nodes with a narrow fused base forming a circular nodal disk. The articulated axes of all the three types of leafy shoots of the present collection show ridges and furrows which are discontinuous across nodes and the foliar characteristics are consistent with the descriptions of Abbott (1958) and Boureau (1964). The present specimens are distinctly different from other equisetalean taxa with leaf whorls at nodes recorded in the Permian strata of Gondwana. Lelstotheca (Surange and Prakash) Maheshwari (1972) has robust leaf whorls borne on a slender stem, spreading out horizontally without cup like sheath or disk at the nodes. Raniganjia Rigby (1962) is different having disk like leaf whorl; the leaves are united at the base up to more than 2/3 of their length. In Bengalia Maheshwari et al. (1989), the leaves radiate from a shallow depression with a cuneate base, suggesting attachment directly at the nodes with little or no adherent leaf sheath; the leaf lamina is flat with an indistinct median vein. Leaves of Barakaria Seward and Sahni (1920) are twice or thrice forked and the median vein dichotomizes 2–3 times. Austroannularia Rigby (1989), showing asymmetrical leaves with shorter leaves on one side forming a reniform outline of the whorl, is easily distinguishable from the present specimens. Three species of Annularia foliage shoots were collected. One of the foliage shoots, viz. A. gondwanensis sp. nov., is preserved in situ with an erect axis bearing elaborately branched leafy shoots. Annularia gondwanensis Banerjee, Mitra and Dutta sp. nov. (Plate I, 1–6; Plate II, 1–15; Plate III, 1–6)
Holotype Type slide numbers
Repository Type locality Type stratum
Etymology
Specimen number. ScTd 450, (Plate I, 2–6; Plate III, 2–3) ScTd 450 /1–5 (Plate II, 1–15; Plate III, 4–6) Sl.No.1-Macerated entire leaves on node, Sl. No. 2-Entire Leaf with mucronate tip; Sl. No.3-Lower epidermis with stomata; Sl. No 4-Upper epidermis; Sl. No 5–Trachieds showing wall thickenings SEM 1 — Impression of stomata on the coalified layer, SEM 2 — Trachieds with wall thickenings MB collection of Palaeobotany–Palynology Section, Department of Botany, University of Calcutta Tulsidabar Quary, Saharjuri Basin, Jharkhand, India Sandstone, shale, layer within coal horizon XII — Tulsidabar Top coal seam (Fig. 2), Barakar Formation, late Lower Permian (late Artinskian) The name of the species is proposed to establish the first record of Annularia from peninsular Indian Gondwana basin
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Fig. 1. Geological map of Saharjuri Basin after Datta et al., 1983; Pareek et al., 1988; Coal India — Eastern coalfields Ltd., unpublished data; showing the study area Tulsidabar quarry.
3.1. Diagnosis Branches with leafy shoots emerge from erect, articulated axis; 6–11 small, single-veined leaves occur in whorls at nodes of the branches; leaves unequal, oblong to obovate in shape with tapering narrow base which are basally fused to form nodal disk; the bluntly obtuse apex terminate by a small mucronate tip. Two epidermal layers of the leaves have moderately thick walled cells, hypostomatic. The stomata are random in distribution. The guard cells are sunken, narrow elliptic in shape and without thickening of the walls. Subsidiary cells two, thick walled, conspicuous, reniform. Four to five epidermal cells surround the stomatal apparatus, forming a ring. Cells of non stomatiferous epidermal layer show a dark mark, commonly with circular or semicircular opening. Xylem
recovered from both erect axis and branches consist of thick walled elongated tracheids with spiral and bordered pitted wall thickenings. 3.2. Description [Plate I, 1–6; Plate II, 1–15; Plate III, 1–6] The 130 cm long and 24–28 cm wide incomplete, coalified axis of the plant is vertically preserved at right angle to the bedding plane of the hard sandstone layer within the coal horizon XII-Tulsidabar. The top coal seam is exposed at the working site (Plate I, 1, Plate III, 1). The erect axis shows nodes and internodes, ridges and furrows. The nodes are about 40 to 48 cm apart. The detailed features of nodes and internodes, ridges and furrows are not very clear in the
M. Banerjee et al. / Review of Palaeobotany and Palynology 153 (2009) 394–407 Table 2 Generalised geology and stratigraphic succession of coal seams in Saharjuri Basin, Jharkhand based on Datta et al., 1983; Pareek et al., 1988; Coal India-Eastern Coalfields Ltd. unpublished data
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Stomata are sunken. Stomatal apparatus is narrow elliptic, measuring 23.3 to 48.2 µm × 15.6 to 23.9 µm. The guard cells are without inner wall thickening, narrow, elongated, 21.4 to 34.4 µm long and 2.1 to 5.4 µm wide (Plate II, 4–6, 14; Plate III, 4–5). The stomatal opening is 8 µm in length. Two subsidiary cells are conspicuous, reniform and measure 23.7 to 48.2 µm × 7 µm (Plate II, 6; Plate III, 5). Four to five polygonal to oval-rectangular cells measuring 26.5 to 38.4 µm × 10.3 to 18.0 µm surround the stomatal apparatus forming a ring (Plate II, 4, 5; Plate III, 4, 5). Cells in the non stomatiferous layer are irregular and rectangular to polygonal in shape, moderately thick-walled (Plate II, 7–9; Plate III, 6) variable in size and measure 27.3 to 113 µm × 23.7 to 64.7 µm. Each cell shows a dark area in the middle, commonly with circular or semicircular open or collapsed hair bases. The xylem recovered from both erect axis and branches consists of thick-walled, elongated tracheids, with uni-seriate and bi-seriate bordered pits together with distinct spiral thickenings (Plate II, 10–13, 15). The bordered pits are circular to oval in shape with slightly elongated pores (Plate II, 11–13, 15). The size of the pores varies from 3.6 to 5.2 µm in diameter. The spiral thickenings are rather loosely spaced and pass through the gaps in between the bordered pits (Plate II, 11–13, 15). 3.3. Comparison
photograph as the axis is highly carbonized. A number of branches with leafy shoots emerge in radial symmetry from the relatively broad (24–28 cm) erect stem. The length and width of the axis suggest an arborescent growth habit of the plant. The vegetative shoots with elaborate articulated branch system bearing whorls of leaves are preserved on the horizontal shale layer as compressions; at least three branches bearing foliage shoots are found in organic connection to the erect axis (Plate I, 1; Plate III, 1 — marked by arrow). The primary shoots are 0.9–3.2 mm thick (Plate I, 1; Plate III, 1), branch profusely into shorter axes which bear whorls of leaves at nodes (Plate I, 2; Plate III, 2). Nodes and internodes of the branches are distinct. 6 to 9 ridges and furrows alternate in successive whorls. The number of leaves in the whorls varies from lower to higher order branches. 6 to 11 leaves occur in a whorl (Plate I, 3; Plate III, 3). The length of leaves is shorter than the length of the internodes. The leaves are 3 to 9.5 mm in length and the width varies from 1to 2.3 mm; the length-width ratio ranges between 2.3:1 and 4.8:1 (Fig. 3). The shape of the leaves in a single whorl varies from oblong to narrow oblong to oblanceolate to narrow obovate. The bases of the leaves are narrow; the bluntly obtuse apex of the leaves terminate in a small mucronate tip (Plate I, 3–4; Plate II, 1; Plate III, 2–3). The single median vein is 111 to 156 µm wide and runs straight to the apex. The margin is entire. The leaves are free up to the narrow base where they are fused to form an elliptic nodal disk of 1–1.5 mm in diameter at the point of attachment with the axis (Plate I, 2–5; Plate III, 2–3). The well-preserved coalified layers of the leaves have yielded cuticles of both upper and lower epidermal layers. The stomatiferous lower epidermal layer shows randomly arranged cells which vary in shape and size. The cells are polygonal to rectangular to slightly elongated and measure 53.2 to 136.9 µm × 20.3 to 55.0 µm. Cell walls are thick with straight or slightly oblique lateral walls (Plate II, 2–3).
The vertical axis with nodes, internodes showing longitudinal ridges and furrows is different from the upright Glossopteris plants on Vertebraria roots recorded from the associated beds of shale, sandstone layers (Banerjee et al., 1991; Banerjee, 2005) below the coal horizon XII — Tulsidabar Top coal seam (Fig. 2). The new species A. gondwanensis has been compared with the Annularia species so far known from the different phytoprovinces (Table 1) primarily on the basis of shape, number and arrangement of leaves at nodes (Abbott, 1958; Archangelsky, 1960; Boureau, 1964; Good, 1976; Barthel, 1980; Anderson and Anderson, 1985; Guanglong, 1995; Liu et al., 2000; Cuneo, 2000). The branching pattern of the articulated axes, small size of the leaves and the number of leaves in a whorl on nodes in A. gondwanensis sp. nov. are comparable to A. crassiscula Halle (1927) recorded from Lower Permian of Shanxi, China, (Cathaysia) (Boureau, 1964, page 168, figure145). However, A. crassiscula differs from A. gondwanensis in having lanceolate to narrow elliptic leaves with a sharply acute apex and in the absence of mucronate tip. Annularia pingloensis Sze and A. shirakii Kawasaki, recorded from late Lower to Upper Permian of Tibet, and South and North China (Guanglong, 1995), are different from the present specimen in the shape and size of the leaves, and in the absence of mucronate tip at the apex. The species of Annularia that have leaves with a mucronate tip, viz., A mucronata, A. latifolia, A. sphenophylloides and A. stellata (Wood, 1860; Abbott, 1958; Tiwari and Singh, 1981), differ from A. gondwanensis in the number, the shape and the arrangement of the leaves per whorl. Annularia mucronata has 10–24 leaves per verticil compared to 6–11 leaves in A. gondwanensis. The shape (spathulate) and size (4–24 mm long) of the leaves of A. mucronata are different from those in A. gondwanensis. Annularia latifolia known from Euramerica is closely comparable to A. gondwanensis in the number of leaves per verticil (6–11 in A. gondwanensis and 7–12 in A. latifolia), but differs in the variable shape and size of the leaves in a single whorl. Annularia stellata has a different in the number of leaves (13–32) per verticil which are linear to oblanceolate in shape and differ also in size (14–75 mm). Annularia sphenophylloides with 10–20 broadly spathulate leaves per verticil is different from A. gondwanensis. The oblanceolate leaves of A. hammanskraalensis (Anderson and Anderson, 1985) and the narrow elliptic leaves of A. kurtzii (Archangelsky, 1960) are different from the oblong–obovate leaves of A. gondwanensis. The other species of Annularia known from the upper Palaeozoic can be distinguished from
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Fig. 2. Comprehensive lithosuccession (not to scale) of section exposed in Chitra B mines, Tulsidabar quarry showing fossil locations.
A. gondwanensis on the basis of their conspicuously different number and arrangement of leaves at the nodes, and the shape of the lamina and apex. Cuticular features are currently known for only a few Annularia species. Cuticles of A. mucronata Schenk and A. stellata (Schlotheim) Wood were studied by Abbott (1958) and Cuneo (2000). The anatomical details of a permineralised specimen of A. hoskinsii were described by Good (1976). The general features of the epidermal layers of A. gondwanensis, viz., the hypostomatic nature, the narrow elliptic stomata, sunken guard cells without wall thickenings, two reniform conspicuous, subsidiary cells on either side of stomata, the shape and the arrangement of the epidermal cells are similar to the cuticles of the above mentioned species. Hairs such as those found in the cuticles of A. mucronata, were not observed in A. gondwanensis. However, the open or collapsed circular or semicircular dark (Plate II, 8–9; Plate III, 6) areas in the middle of each cell appear to be the hair bases. Despite the similarities in the cuticular features, the macromorphological characters of A. stellata, A. mucronata are unlike those of A. gondwanensis, as mentioned above. The macromorphological features of the erect axis are not very clearly preserved in comparison with the Calamites stem. The macerated wood fragments of the erect axis as well as branches show tracheids with spiral and bordered pitted thickenings similar to
the tracheids described in the equisetaleans (Boureau, 1964; Taylor and Taylor, 1993). Annularia. sp. (Plate IV, 1–2; Plate V, 1–2)
Specimen numbers Repository Locality of collection Stratigraphic horizon
ScTd 493, ScTd 508 (Plate IV, 1–2; Plate V, 1–2) MB collection of Palaeobotany–Palynology Section, Department of Botany, University of Calcutta Tulsidabar Quary, Saharjuri basin, Jharkhand, India Shale layer within coal horizon XII — Tulsidabar Top coal seam (Fig. 2), Barakar Formation, late Lower Permian (late Artinskian)
3.4. Description The leafy shoots are articulated with long internodes. The partially preserved internodes measure 31–33 mm. The nodal disk is 4 mm in diameter and slightly elongated. The leaves 27–38 mm long and 2–5 mm wide are basally adpressed and spread out distally. The whorls at the nodes consist of 16–18 leaves, with 8–9 leaves on either side. The leaves in the middle are longer than the others. The shape of leaves varies from oblong to linear with acute apices. The mid-vein is not very prominent and measures 1/7 of width of the lamina.
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3.5. Comparison Annularia sp. resembles A papilioformis Kawasaki (1927), recorded from Lower to early Upper Permian of Shanxi, northern Cathaysian province, China (Guanglong, 1995), in the number, shape and arrangement of the leaves. The leaves of A. papilioformis are moreor-less uniform in size, however, they are basally welded in a single plane at the nodal region while the leaves in Annularia sp. are free, unequal in size and with closely adpressed bases. Annularia kurtzii Archangelsky 1960 (Plate IV, 3–6; Plate V, 3–4) Specimen numbers Repository Locality of collection Stratigraphic horizon
ScTd 491, (Plate IV, 3–4; Plate V, 3), ScTd 494, (Plate IV, 5–6; Plate V, 4), MB collection of Palaeobotany–Palynology Section, Department of Botany, University of Calcutta Tulsidabar Quary, Saharjuri basin, Jharkhand, India Shale layer within coal horizon No .XII — Tulsidabar Top coal seam (Fig. 2), Barakar Formation, late Lower Permian (late Artinskian)
3.6. Description The axis is articulated with nodes and internodes; branches emerge from the nodes. The internodes are longitudinally striated, 16–18 mm long and 1.5–2 mm wide. The leaves are narrow elliptic in
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shape with a distinct midrib, an acute apex, an entire margin and they emerge radially from the nodes. The basal portions of leaves are joined to form a nodal disk. The number of leaves per whorl varies from 9 to 10. The leaf length covers almost the internodal length, but does not exceed the internode. Leaves measure from 12 to 24 mm in length and the width varies from 1.5 to 2.5 mm. 3.7. Comparison Because of the similarity of characters, i.e., shape, number and arrangement of leaves, the present specimens are assigned to Annularia kurtzii described by Archangelsky (1960) from Lower Permian sediments of Parana Basin, South America. 4. Discussion 4.1. Biogeographical range of occurrence of Annularia The leaf genus Annularia, Asterophyllites, the stem genus Calamites and the fertile genera Calamostachys or Palaeostachya represent different parts of the plant Calamites. The plant flourished in tropical, warm, humid forests of Euramerica during late Palaeozoic together with various lepidophytes, the sphenophyte genera, Sphenophyllum, Bowmanites, Phyllotheca and other members of Euramerian flora (Meyen, 1987; Cleal and Thomas, 1991). During late Early Permian
Plate I. (see Plate I on page 400)
1. 2. 3. 4. 5. 6.
Shows in situ occurrence of upright coalified axis preserved in the thick sandstone layer; the branches emerge radially from the upright axis (the attachment points are marked by arrow) with leaves in whorl at nodes; A. gondwanensis sp. nov. (specimen photographed in the field). A portion of the branch system showing the branching pattern. A. gondwanensis sp. nov., Holotype Sp. No. ScTd 450. Enlarged view of the branch system showing unequal nature of leaves, pattern of emergence of leaves from nodes and nodal disk. A. gondwanensis sp. nov., Holotype Sp. No. ScTd 450. Portion of branches enlarged to show the details of leaf characters viz; unequal size, contracted base, mucronate apex and the mode of attachment to the axis forming nodal disk. A. gondwanensis sp. nov., Holotype Sp. No. ScTd 450. Macerated preparation of the nodal portion showing arrangement of unequal leaves with thick midvein and mucronate apex. A. gondwanensis sp. nov., Sl. No. ScTd 450/1. Enlarged view of a single leaf .A. gondwanensis sp. nov., ScTd 450.
Plate II. (see Plate II on page 401)
1. 2. 3. 4. 5. 6. 7. 8–9. 10. 11–13. 14. 15.
Macerated preparation of a leaf showing mucronate apex. A. gondwanensis. sp. nov., Sl. No. Sc Td. 450/2. Stomatiferous layer of the cuticle shows random distribution of stomata and epidermal cells of variable shape and size. A. gondwanensis sp. nov., Sl. No. ScTd 450/3. Enlarged view of portion of Fig. 2 .A. gondwanensis sp. nov. Stomatal apparatus and the surrounding epidermal cells. A. gondwanensis. sp. nov., Sl. No. ScTd 450/3. Stomatal apparatus and the surrounding epidermal cells (Marked by arrow- a) guard cell, b) subsidiary cell, c) epidermal cell). A. gondwanensis. sp. nov., Sl. No. ScTd 450/3. Stomata enlarged showing sunken nature of guard cells without wall thickening, two reniform subsidiary cells with thick walls. A. gondwanensis sp. nov., Sl. No. ScTd 450/3. Non stomatiferous upper epidermal layer showing epidermal cells of variable shape and size; each cell shows dark area of hair base. A. gondwanensis. sp. nov., Sl. No. ScTd 450/4. Non stomatiferous upper epidermal layer showing collapsed (8) or open (9) hair bases. A. gondwanensis sp. nov., Sl. No. ScTd 450/4. Wood elements showing tracheids with thickenings. A. gondwanensis sp. nov., Sl. No. ScTd 450/5. Enlarged view of wood elements with bordered pits and scalariform thickenings. A. gondwanensis sp. nov., Sl. No. ScTd 450/5. SEM photo of coalified layer of leaf on rock showing stomata and epidermal cells. A. gondwanensis . sp.nov.. Portion of Holotype Sp.No. ScTd 450. SEM photo of portion of macerated wood showing bordered pits and scalariform thickening. A. gondwanensis sp. nov., Holotype Sp.No. ScTd 450.
Plate III. (see Plate III on page 402)
1. 2. 3. 4. 5. 6.
Shows in situ occurrence of upright coalified axis preserved in the thick sandstone layer; the branches emerge radially from the upright axis (point of attachment marked by arrow) with leaves in whorls at nodes; A. gondwanensis sp. nov. (specimen photographed in the field). A portion of the branch system showing the branching pattern. A. gondwanensis sp. nov. Holotype Sp.No.ScTd 450. Portion of axis enlarged to show the alternate arrangement of ridges and furrows, details of leaf characters viz., unequal size, contracted base, mucronate tip, mode of attachment to the axis forming nodal disk. A. gondwanensis sp. nov., Holotype Sp.No.ScTd 450. Stomatiferous layer of the cuticle showing epidermal cells of variable shape and size, stomata, guard cells, reniform subsidiary cells and additional surrounding epidermal cells. A. gondwanensis sp.nov., Sl. No. ScTd 450/3. Enlarged view of stomatal layer showing epidermal cells, subsidiary cells with thickening, sunken guard cells, stomatal opening (Marked by arrow — a) guard cell, b) subsidiary cell, c) epidermal cell). A. gondwanensis sp. nov. Sl. No. ScTd 450/3. Epidermal cells of the upper cuticle; each cell with hair base, showing broad open or collapsed hair bases. A. gondwanensis sp. nov., Sl. No. ScTd 450/4.
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Plate I (caption on page 399).
M. Banerjee et al. / Review of Palaeobotany and Palynology 153 (2009) 394–407
Plate II (caption on page 399).
401
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Plate III (caption on page 399).
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Fig. 3. Leaf length and breadth ratio of A. gondwanensis sp.nov.
and Late Permian, several species of Annularia flourished in the Cathaysian province of North China, South China and the Tibetan Plateau together with Sphenophyllum and other Euramerian and endemic taxa. These assemblages are recognized as equatorial belt plant assemblage (Meyen, 1987; Cleal and Thomas, 1991; Li et al., 1995; Guanglong, 1995; Yao et al., 2000; Mei et al., 2000; Wang et al., 2000b,a). Annularia also thrived in the Angara flora (Meyen, 1987). In the Gondwana, Annularia is not as frequent in the upper Palaeozoic as in the Euramerian and Cathaysian floras. The genus occurs in the Glossopteris floral assemblage at restricted stratigraphic sequence. Unequivocal records of the genus are from the Lower Permian of the Parana Basin, Brazil, the Paganzo Basin, Argentina in South America (Archangelsky, 1960, 1986; Cuneo, 2000), the Karoo Basin in South Africa (Anderson and Anderson, 1985), western extraPeninsular India (Tiwari and Singh, 1981) and is now reported from upper Lower Permian strata of Saharjuri Basin in eastern peninsular India. The global distribution pattern of the Annularia species thus reveals the biogeographical range of occurrence up to the eastern peninsular India in the Gondwana province during the Early Permian (Fig. 4; Table 1). 4.2. Palaeoclimate and palaeogeography The postglacial Early Permian climate of Gondwanaland is known to be cool-temperate. However, palaeobotanical data from the western extra-peninsular India in the Indian subcontinent indicate a warm, humid climate since the Early Carboniferous (Pal and Chaloner,
1979; Pant et al., 1984; Pant and Srivastava, 1995) to the Late Carboniferous and Permian (Srivastava and Kapoor, 1969; Tiwari and Singh, 1981; Singh et al., 1982). Annularia species viz; A. stellata occur in Lower Permian rocks of Kumaon Himalaya (Tiwari and Singh, 1981), equivalent to the Mamal Formation (Lower Permian) of Kashmir Himalaya, India. The present record from Saharjuri Basin is very similar to the sphenopsid-rich assemblages of warm, humid climate known from the Lower Permian basins of western extra-peninsular India (Singh et al., 1982; Pant et al., 1984), South America (Archangelsky, 1960, 1986, 1990; Cuneo, 2000; Escapa and Cuneo, 2005; Escapa and Cuneo, 2006) and South Africa (Anderson and Anderson, 1985). The plants could thrive in the areas as suitable climatic condition prevailed during late Early Permian (Artinskian). Li Xingxue (1986) suggested that towards the end of the Sakmarian the climate in Gondwana became warmer. Since then the warm climate continued to the Artinskian. This is evidenced by the abundant warm water faunas. The reason was cited as the start of drift of Indian subcontinent (a part of Gondwanaland) and the southern Tibetan Plateau since Late Carboniferous towards warm subtropical areas. Archangelsky (1990) suggested ‘a latitudinal control of floral distribution in the Early Permian’ and a changed palaeogeography of the areas of South American, African basins due to improved latitudinal positions during late late Carboniferous and Early Permian. The palaeobotanical data suggest a lower latitudinal palaeogeography of the Saharjuri Basin, western extra-peninsular basins in the Indian subcontinent and the basins of South America, South Africa during Early Permian. 5. Conclusion 5.1. Palaeoecology The preserved length of 130 cm and 24–28 cm width of the erect axis suggest arborescent growth habit of the plant. The occurrence of the calamitalean genus Annularia in growth position is a rare and important addition to the Glossopteris flora. The occurrences of A. gondwanensis sp. nov., Annularia. sp, A. kurtzii, assemblage of diverse articulate fertile and sterile taxa viz.; the bractless fertile organ Tulsidabaria indica, the bracteate fertile cone bearing plant Rajmahaliastachys elongata, vegetative shoots of Sphenophyllum sp., Phyllotheca spp. suggest the occurrence of a swamp in a warm and humid environment. The arborescent A. gondwanensis and upright Glossopteris plants in the associated sediments (Fig. 2) indicate in situ deposition of the forest in the Saharjuri Basin during late Early Permian. The evidence of Annularia spp. in the Lower Permian rocks of South America in the Parana Basin of Brazil, the Paganzo basin, Argentina, in the upper Lower Permian of Patagonia, Argentina, South America, Karoo Basin of South Africa, western extra-peninsular basin, the present record from late Lower Permian of Saharjuri Basin, eastern
Plate IV. (see Plate IV on page 404)
1. 2. 3. 4. 5. 6.
Incompletely preserved leafy shoot; leaves ad pressed basally at nodes and spread out distally. Annularia. sp., Sp.No.ScTd 493. Partially preserved leafy shoot of Annularia. sp. showing leaves emerging from the node, Sp.No.ScTd 508. Leafy shoot with three whorls of leaves. A. kurtzii Archangelsky, Sp.No ScTd 491. A single leaf whorl magnified showing arrangement of leaves of A. kurtzii. Archangelsky, Sp.No. ScTd 491. Leafy shoot of A. kurtzii Archangelsky with branch system, Sp. No. ScTd 494. A portion of leafy axis magnified showing leaf arrangement of A. kurtzii Archangelsky, Sp.No. ScTd 494.
Plate V. (see Plate V on page 405)
1. 2. 3. 4.
403
Partially preserved nodal region showing the arrangement of leaves. Annularia. sp., Sp.No.ScTd 493. Partially preserved leafy shoot of Annularia. sp. showing leaves emerging from the node, Sp.No.ScTd 508. Leafy shoot with three whorls of leaves .A. kurtzii Archangelsky, Sp.No.ScTd 491. Leafy shoot of A. kurtzii Archangelsky with branch system, Sp. No. ScTd 494.
404
M. Banerjee et al. / Review of Palaeobotany and Palynology 153 (2009) 394–407
Plate IV (caption on page 403).
M. Banerjee et al. / Review of Palaeobotany and Palynology 153 (2009) 394–407
Plate V (caption on page 403).
405
406
M. Banerjee et al. / Review of Palaeobotany and Palynology 153 (2009) 394–407
Fig. 4. Global distribution of Annularia spp. ( ) including present record rocks of Gondwana continents.
during Late Palaeozoic plotted in the base map after Ziegler et al., 1997. ( ) Records from Early Permian
peninsular India plotted in the base map of Ziegler et al. (1997) reveal that a rich assemblage of sterile and fertile sphenopsids and diverse glossopterids flourished in the lowland areas of the Gondwana continent (Fig. 4) during the Early Permian. Acknowledgements The authors are grateful to Professor W. G. Chaloner FRS, Geology Department, Royal Holloway University of London for critical examination of the original manuscript and for valuable suggestions encouraging early publication of the paper. The constructive suggestions, comments by the anonymous reviewers of Review of Palaeobotany Palynology are gratefully acknowledged. Thanks are due to the Department of Science and Technology, Life Science, New Delhi, for providing financial assistance (SP/SO/A-61/98) to carry out the investigation. Grateful thanks are due to the Officers of Coal India Ltd., Eastern Coalfields Ltd., General Managers and members of Chitra Coal fields, Chitra, Jharkhand for their kind permission to undertake fieldwork also for all kinds of assistance during the collection of materials.
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Maheshwari, H.K., Singh, V.K., Bajpai, U., 1989. Bengalia raniganjensis, a new sphenopsid from Raniganj Formation (Upper Permian). Palaeobotanist 37, 169–172. Mei, M.T., Li, S.S., Hu, S.S., 2000. Carboniferous–Permian Peat forming plant in China. Acta Palaeontologica Sinica 39 (Sup.), 57–62. Meyen, S.V., 1987. Fundamentals of Palaeobotany. Chapman and Hall, London, pp. 1–432. Niyogi, D., 1966. Lower Gondwana sedimentation in Saharjuri coalfield, Bihar, India. Journal of Sedimentary Petrology 36 (4), 960–972. Pal, A.K., Chaloner, W.G., 1979. A lower carboniferous Lepidodendropsis flora in Kashmir. Nature 281 (5729), 295–297. Pant, D.D., Srivastava, P.C., 1995. Lower carboniferous plants from Wallarama spur of Punjab-Kashmir Himalaya. Paleogeographica 235, 23–49 Abt. B. Pant, D.D., Nautiyal, D.D., Srivastava, P.C., 1984. The occurrence of Cathaysian elements in Glossopteris flora of Kashmir. Phyta 4 (5), 47–52. Pareek, H.H., Bardhan, B., Chakraborty, N.C., 1988. Seam sedimentary cycles, sedimentary petrography of the bore hole core samples of Damuda (L. Permian) group and petrography of the coal seams, Saharjuri coal field, Dist. Santhal Parganas, Bihar, India. Records of the Geological Survey of India, Calcutta 118 (20), 9–32. Rigby, J.F., 1962. On a collection of plants of Permian age from Baralaba, Queensland. Proceedings of the Linnean Society of New South Wales 87 (3), 341–351. Rigby, J.F., 1989. Austroannularia gen. nov., an asymmetrical Permian sphenopsid leaf whorl from Gondwanaland. Palaeobotanist 37 (2), 189–191. Seward, A.C., Sahni, B., 1920. Indian Gondwana plants — a revision. Memoirs of the Geological Survey of India. Palaeontologia Indica 7, 1–54. Singh, G., Maithy, P.K., Bose, M.N., 1982. Upper Palaeozoic flora of Kashmir Himalaya. Palaeobotanist 30 (2), 185–232. Srivastava, J.P., Kapoor, H.M., 1969. Discovery of Lepidostrobus Brongn. from Lower Gondwana formation of Kashmir. Journal - Palaeontological Society of India 12, 44–47. Sternberg, C., 1823. Versuch einer geognostisch-botanischen Darstellung der Flora der Vorwelt. Leipzig 1 (3), 1–39. Taylor, T.N., Taylor, E.L., 1993. The Biology and Evolution Fossil Plants. Prentice Hall, Englewood Cliffs, New Jersey. Tiwari, B.S., Singh, R.Y., 1981. The Late Paleozoic fossils from Kumaon Himalaya and their stratigraphic significance. In: Khosla, S.C., Kachhara, R.P. (Eds.), Proc. 9th Indian Colloq. Micropal. Strat.. Udaipur, Udaipur Univ., Rajasthan, pp. 206–219. Wang, J., Shen, G.L., Zhang, H., Zhang, S.Q., 2000a. A preliminary study on the Carboniferous and Permian phytostratigraphy of Weibei coalfield, Shaanxi, China. Acta Palaeontologica Sinica 39 (Sup.), 96–103. Wang, J., Liu, H.Q., Shen, G.L., Wu, X.Y., 2000b. Megaflora from the carboniferous– Permian Tongwei Formation in Hejiashan, Tongwei, Gansu. Acta Palaeontologica Sinica 39 (Sup.), 76–96. Wood, H.C., 1860. Contributions to the Carboniferous flora of the United States. Acad. Nat. Philadelphia Proc. 12. Yao, Z.-Q., Liu, L.-J., Mapes, G., Rothwell, G.W., 2000. Leaf morphology and cuticular features of Sphenophyllum in the Gigantopteris flora from SouthChina. Review of Palaeobotany and Palynology 110, 67–92. Ziegler, A.M., Hulver, M.L., Rowley, D.B., 1997. Permian world topography and climate. In: Martini, I.P. (Ed.), Late glacial and post-glacial environmental changesQuaternary, Carboniferous-Permian and Proterozoic. Oxford University Press, New York, pp. 111–146.
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Review of Palaeobotany and Palynology j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / r ev p a l b o
On the occurrence of three species of Annularia Sternberg in the Early Permian Glossopteris flora of peninsular India Manju Banerjee ⁎, Samik Mitra, Sutapa Dutta Department of Botany, University of Calcutta, 35, Ballygunge Circular Road, Kolkata-700019, India
a r t i c l e
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Article history: Received 17 October 2007 Received in revised form 16 October 2008 Accepted 9 November 2008 Available online 14 November 2008 Keywords: In situ Annularia gondwanensis sp. nov. Annularia.sp. Annularia kurtzii Upper Lower Permian Eastern peninsular India Gondwana
a b s t r a c t Three species of articulated foliage-bearing shoots of Annularia Sternberg are described from the Barakar Formation (late Early Permian–late Artinskian) of the Saharjuri Basin, eastern peninsular India. Annularia gondwanensis sp. nov. is preserved in growth position with branches emerging from a vertical carbonized axis. The leaves, branches and axis of A. gondwanensis have preserved cuticles. Annularia sp. and Annularia kurtzii Archangelsky are preserved as impressions. The Annularia species together with diverse sphenopsids occur in a typical Gondwanan assemblage dominated by glossopterids. This late Palaeozoic Euramerian– Cathaysian genus was earlier known from the Lower Permian of various region of the Gondwana continent viz., Brazil and Argentina in South America, the Karoo Basin in South Africa and western extra-peninsular India. Now the biogeographical range of distribution of Annularia extends up to the eastern peninsular India in the Gondwana continent. The palaeoclimate, palaeoecology of the Saharjuri Basin in eastern India is discussed considering the records of diverse sterile and fertile sphenopsids and Annularia in growth position together with the upright Glossopteris plants, other glossopterid members in the upper Lower Permian. © 2008 Elsevier B.V. All rights reserved.
1. Introduction Annularia Sternberg, 1823 is a calamitalean genus with 46 species based on leaves (Table 1). The genus flourished in the warm, humid climate of the late Palaeozoic equatorial belt (Eurameria–Cathaysia) and the palaeotemperate Angaran province. Rare Annularia species also occur in the Gondwana province (Banerjee, 1991; Cleal and Thomas, 1991), in the Lower Permian of Argentina, South America, South Africa and western extra-peninsular India. (Archangelsky, 1960, 1986; Tiwari and Singh, 1981; Anderson and Anderson, 1985; Cuneo, 2000). Recently, well-preserved foliage-bearing branches of a new Annularia species attached to an erect axis, together with two other species of the genus, have been collected for the first time from upper Lower Permian strata of peninsular India. These Annularia species occur in association with diverse fertile and sterile sphenopsid shoots (viz., the bractless fertile plant Tulsidabaria Banerjee et al., 2004, the cone-bearing shoots of Rajmahaliastachys Banerjee and D' Rozario, 1999, species of Sphenophyllum, Phyllotheca and a diverse range of glossopterid leaves and seedlings (Banerjee et al.,
⁎ Corresponding author. E-mail addresses: [email protected], [email protected] (M. Banerjee). 0034-6667/$ – see front matter © 2008 Elsevier B.V. All rights reserved. doi:10.1016/j.revpalbo.2008.11.001
1991; Banerjee, 2000, 2005). In addition, several endemic plants of uncertain affinity, viz., Saharjuria, Chitraphyllum, Santhalitheca (Banerjee, 1999) and ginkgophytes occur in the associated beds. Here we describe the macromorphology of the three Annularia species and the cuticular micromorphology of the leaves and stems of A. gondwanensis sp. nov. 2. Study area and geological setting The location of study is the Saharjuri basin, Deoghar group of coal fields, Jharkhand, eastern India (Fig. 1) covering 30 km2. area between 24°05' to 24°10' N and 86°49' to 86°57 E. The geology and stratigraphic succession (Table 2) of this basin have been investigated by Niyogi (1966), Datta et al. (1983) and Pareek et al. (1988). 2.1. Materials and method The materials described in the present study are from the younger horizon of the coal bearing strata of the basin. The fossils occur in the sandstone, shale layers within coal horizon XII — Tulsidabar Top coal seam (upper Lower Permian), Barakar Formation (Fig. 2). The branches and the leafy shoots are preserved as compression on thin, finegrained carbonaceous shale layer overlying the exposed hard sandstone layer (Plate I, 1; Plate III, 1). Portions of the leafy shoots
M. Banerjee et al. / Review of Palaeobotany and Palynology 153 (2009) 394–407 Table 1 Global distribution of Annularia species Name of Annularia spp.
Phytoprovince Euramerian Cathaysian Angara Gondwana
1. A. acicularis (Dawson) White 2. A. aculeata Bell 3. A. americana Read 4. A. asteris Asteris 5. A. australis Feistmantel 6. A. batschatensis (Chachlov) Radczenko 7. A. crassiscula Halle 8. A. filiformis Jongmans et Gothan 9. A. fimbriata Walton 10. A. galioides (Lindley et Hutton) Kidston 11. A. goapingensis Xiao 12. A. gondwanensis sp. nov. 13. A. gracilescens Halle 14. A. grandifolia Schvedov 15. A. hammanskraalense Anderson & Anderson 16. A. incisa Stockmans et Mathieu 17. A. jerunakovensis Neuburg 18. A. jongmansii Walton 19. A. kurtzii Archangelsky 20. A. lanceolata Radczenko 21. A. latifolia (Dawson) Kidston 22. A. microphylla Sauveur 23. A. minima Leggewie et Schonefeld 24. A. mucronata Schenk 25. A. orientalis Kawasaki 26. A. papilioformis Kawasaki 27. A. paramucronata Schrenk 28. A. pingloensis Sze 29. A. pseudostellata H. Potonié 30. A. Punctata Zhao et Chang 31. A. radiata (Brongniart) Sternberg 32. A. raspadensis Gorelova 33. A. rigida Zhao et Liu 34. A. rigitula Xiao 35. A. sibirica Radczenko 36. A. similistellata Daber 37. A. sphenophylloides (Zenker) Gutbier 38. A. spiculosa Rasskazova 39. A. stellata (Schlotheim) Wood 40. A. subradiata Stockmans et Williere 41. A. subtenuifolia Rasskazova 42. A. shirakii Kawasaki 43. A. tajluganensis Radczenko 44. A. tenuifolia Neuburg 45. A. tridactyla Langford 46. A. vernensis (Arnold) Abbott
+ + + + − −
− − − − − −
− − − − − +
− − − − + −
− + + +
+ − − +
− − − −
− − − −
− − − − −
+ − + − −
− − − + −
− + − − +
− − + − − + + + + − − − − + − + − − − + + +
+ − − − − − − − + + + + + − + − − + + − − +
− + − − + − + − − − − − − − − − + − − − − −
− − − + _ − − _ − − − − − − − − − − − − − −
− + +
− + −
+ − −
− + −
− − − − + +
− + − − − −
+ − + + − −
− − − − − −
with branches, erect axis and additional isolated specimens of similar leafy shoots were collected. The details of the in situ axis, leafy shoots, mode of emergence of the foliage shoots from the axis, branching pattern were examined in the field and later in the laboratory with the pieces collected. The specimens of the other two Annularia species, viz., Annularia sp. and A. kurtzii Archangelsky are impressions on gray and black shale. Macromorphological characters of the specimens were studied and photographed under a Zeiss SV II Stereo Zoom microscope and a Nikon FM 2 camera. The morphology of the leaves is described following the terminologies proposed by Dilcher (1974) and Ash et al. (1999). The coalified leaves, branches and the erect axis were macerated for cuticle extraction. Peels of coaly layers were first treated with dilute hydrofluoric acid (40%) to remove the silica particles adhered to the pieces. The isolated fragments were treated slowly with 50% to 70% nitric acid, adding KClO3 whenever necessary for oxidation. The oxidized fragments of the fossils were then treated
395
with dilute (5%) potassium hydroxide solution to clear up the humic matter. Each step of chemical treatment was followed by a thorough wash with distilled water. The prepared cuticles were mounted on clear glass slides using Euperol. A few cuticular pieces were preserved in 70% alcohol for SEM study. The micromorphic details of the cuticles were studied and photographed with Zeiss Axioscope 2 and Leitz Diaplan compound microscopes, a Video Image Master Capture Kit of Leading Edge Pvt. Ltd., Australia and a Hitachi 600 scanning electron microscope. 3. Systematics Class Sphenopsida Order Equisetales Family Calamitaceae Annularia Sternberg, 1823 The genus Annularia was first described by Sternberg (1823). The macro and micro morphological characters of the genus were further elaborated by Abbott (1958) through study of additional specimens. The major distinguishing macromorphological characters of the genus are the lanceolate to obovate or spathulate shaped single-veined, dorsiventral, leaves of equal or unequal length, positioned in verticils radiating from the nodes with a narrow fused base forming a circular nodal disk. The articulated axes of all the three types of leafy shoots of the present collection show ridges and furrows which are discontinuous across nodes and the foliar characteristics are consistent with the descriptions of Abbott (1958) and Boureau (1964). The present specimens are distinctly different from other equisetalean taxa with leaf whorls at nodes recorded in the Permian strata of Gondwana. Lelstotheca (Surange and Prakash) Maheshwari (1972) has robust leaf whorls borne on a slender stem, spreading out horizontally without cup like sheath or disk at the nodes. Raniganjia Rigby (1962) is different having disk like leaf whorl; the leaves are united at the base up to more than 2/3 of their length. In Bengalia Maheshwari et al. (1989), the leaves radiate from a shallow depression with a cuneate base, suggesting attachment directly at the nodes with little or no adherent leaf sheath; the leaf lamina is flat with an indistinct median vein. Leaves of Barakaria Seward and Sahni (1920) are twice or thrice forked and the median vein dichotomizes 2–3 times. Austroannularia Rigby (1989), showing asymmetrical leaves with shorter leaves on one side forming a reniform outline of the whorl, is easily distinguishable from the present specimens. Three species of Annularia foliage shoots were collected. One of the foliage shoots, viz. A. gondwanensis sp. nov., is preserved in situ with an erect axis bearing elaborately branched leafy shoots. Annularia gondwanensis Banerjee, Mitra and Dutta sp. nov. (Plate I, 1–6; Plate II, 1–15; Plate III, 1–6)
Holotype Type slide numbers
Repository Type locality Type stratum
Etymology
Specimen number. ScTd 450, (Plate I, 2–6; Plate III, 2–3) ScTd 450 /1–5 (Plate II, 1–15; Plate III, 4–6) Sl.No.1-Macerated entire leaves on node, Sl. No. 2-Entire Leaf with mucronate tip; Sl. No.3-Lower epidermis with stomata; Sl. No 4-Upper epidermis; Sl. No 5–Trachieds showing wall thickenings SEM 1 — Impression of stomata on the coalified layer, SEM 2 — Trachieds with wall thickenings MB collection of Palaeobotany–Palynology Section, Department of Botany, University of Calcutta Tulsidabar Quary, Saharjuri Basin, Jharkhand, India Sandstone, shale, layer within coal horizon XII — Tulsidabar Top coal seam (Fig. 2), Barakar Formation, late Lower Permian (late Artinskian) The name of the species is proposed to establish the first record of Annularia from peninsular Indian Gondwana basin
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Fig. 1. Geological map of Saharjuri Basin after Datta et al., 1983; Pareek et al., 1988; Coal India — Eastern coalfields Ltd., unpublished data; showing the study area Tulsidabar quarry.
3.1. Diagnosis Branches with leafy shoots emerge from erect, articulated axis; 6–11 small, single-veined leaves occur in whorls at nodes of the branches; leaves unequal, oblong to obovate in shape with tapering narrow base which are basally fused to form nodal disk; the bluntly obtuse apex terminate by a small mucronate tip. Two epidermal layers of the leaves have moderately thick walled cells, hypostomatic. The stomata are random in distribution. The guard cells are sunken, narrow elliptic in shape and without thickening of the walls. Subsidiary cells two, thick walled, conspicuous, reniform. Four to five epidermal cells surround the stomatal apparatus, forming a ring. Cells of non stomatiferous epidermal layer show a dark mark, commonly with circular or semicircular opening. Xylem
recovered from both erect axis and branches consist of thick walled elongated tracheids with spiral and bordered pitted wall thickenings. 3.2. Description [Plate I, 1–6; Plate II, 1–15; Plate III, 1–6] The 130 cm long and 24–28 cm wide incomplete, coalified axis of the plant is vertically preserved at right angle to the bedding plane of the hard sandstone layer within the coal horizon XII-Tulsidabar. The top coal seam is exposed at the working site (Plate I, 1, Plate III, 1). The erect axis shows nodes and internodes, ridges and furrows. The nodes are about 40 to 48 cm apart. The detailed features of nodes and internodes, ridges and furrows are not very clear in the
M. Banerjee et al. / Review of Palaeobotany and Palynology 153 (2009) 394–407 Table 2 Generalised geology and stratigraphic succession of coal seams in Saharjuri Basin, Jharkhand based on Datta et al., 1983; Pareek et al., 1988; Coal India-Eastern Coalfields Ltd. unpublished data
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Stomata are sunken. Stomatal apparatus is narrow elliptic, measuring 23.3 to 48.2 µm × 15.6 to 23.9 µm. The guard cells are without inner wall thickening, narrow, elongated, 21.4 to 34.4 µm long and 2.1 to 5.4 µm wide (Plate II, 4–6, 14; Plate III, 4–5). The stomatal opening is 8 µm in length. Two subsidiary cells are conspicuous, reniform and measure 23.7 to 48.2 µm × 7 µm (Plate II, 6; Plate III, 5). Four to five polygonal to oval-rectangular cells measuring 26.5 to 38.4 µm × 10.3 to 18.0 µm surround the stomatal apparatus forming a ring (Plate II, 4, 5; Plate III, 4, 5). Cells in the non stomatiferous layer are irregular and rectangular to polygonal in shape, moderately thick-walled (Plate II, 7–9; Plate III, 6) variable in size and measure 27.3 to 113 µm × 23.7 to 64.7 µm. Each cell shows a dark area in the middle, commonly with circular or semicircular open or collapsed hair bases. The xylem recovered from both erect axis and branches consists of thick-walled, elongated tracheids, with uni-seriate and bi-seriate bordered pits together with distinct spiral thickenings (Plate II, 10–13, 15). The bordered pits are circular to oval in shape with slightly elongated pores (Plate II, 11–13, 15). The size of the pores varies from 3.6 to 5.2 µm in diameter. The spiral thickenings are rather loosely spaced and pass through the gaps in between the bordered pits (Plate II, 11–13, 15). 3.3. Comparison
photograph as the axis is highly carbonized. A number of branches with leafy shoots emerge in radial symmetry from the relatively broad (24–28 cm) erect stem. The length and width of the axis suggest an arborescent growth habit of the plant. The vegetative shoots with elaborate articulated branch system bearing whorls of leaves are preserved on the horizontal shale layer as compressions; at least three branches bearing foliage shoots are found in organic connection to the erect axis (Plate I, 1; Plate III, 1 — marked by arrow). The primary shoots are 0.9–3.2 mm thick (Plate I, 1; Plate III, 1), branch profusely into shorter axes which bear whorls of leaves at nodes (Plate I, 2; Plate III, 2). Nodes and internodes of the branches are distinct. 6 to 9 ridges and furrows alternate in successive whorls. The number of leaves in the whorls varies from lower to higher order branches. 6 to 11 leaves occur in a whorl (Plate I, 3; Plate III, 3). The length of leaves is shorter than the length of the internodes. The leaves are 3 to 9.5 mm in length and the width varies from 1to 2.3 mm; the length-width ratio ranges between 2.3:1 and 4.8:1 (Fig. 3). The shape of the leaves in a single whorl varies from oblong to narrow oblong to oblanceolate to narrow obovate. The bases of the leaves are narrow; the bluntly obtuse apex of the leaves terminate in a small mucronate tip (Plate I, 3–4; Plate II, 1; Plate III, 2–3). The single median vein is 111 to 156 µm wide and runs straight to the apex. The margin is entire. The leaves are free up to the narrow base where they are fused to form an elliptic nodal disk of 1–1.5 mm in diameter at the point of attachment with the axis (Plate I, 2–5; Plate III, 2–3). The well-preserved coalified layers of the leaves have yielded cuticles of both upper and lower epidermal layers. The stomatiferous lower epidermal layer shows randomly arranged cells which vary in shape and size. The cells are polygonal to rectangular to slightly elongated and measure 53.2 to 136.9 µm × 20.3 to 55.0 µm. Cell walls are thick with straight or slightly oblique lateral walls (Plate II, 2–3).
The vertical axis with nodes, internodes showing longitudinal ridges and furrows is different from the upright Glossopteris plants on Vertebraria roots recorded from the associated beds of shale, sandstone layers (Banerjee et al., 1991; Banerjee, 2005) below the coal horizon XII — Tulsidabar Top coal seam (Fig. 2). The new species A. gondwanensis has been compared with the Annularia species so far known from the different phytoprovinces (Table 1) primarily on the basis of shape, number and arrangement of leaves at nodes (Abbott, 1958; Archangelsky, 1960; Boureau, 1964; Good, 1976; Barthel, 1980; Anderson and Anderson, 1985; Guanglong, 1995; Liu et al., 2000; Cuneo, 2000). The branching pattern of the articulated axes, small size of the leaves and the number of leaves in a whorl on nodes in A. gondwanensis sp. nov. are comparable to A. crassiscula Halle (1927) recorded from Lower Permian of Shanxi, China, (Cathaysia) (Boureau, 1964, page 168, figure145). However, A. crassiscula differs from A. gondwanensis in having lanceolate to narrow elliptic leaves with a sharply acute apex and in the absence of mucronate tip. Annularia pingloensis Sze and A. shirakii Kawasaki, recorded from late Lower to Upper Permian of Tibet, and South and North China (Guanglong, 1995), are different from the present specimen in the shape and size of the leaves, and in the absence of mucronate tip at the apex. The species of Annularia that have leaves with a mucronate tip, viz., A mucronata, A. latifolia, A. sphenophylloides and A. stellata (Wood, 1860; Abbott, 1958; Tiwari and Singh, 1981), differ from A. gondwanensis in the number, the shape and the arrangement of the leaves per whorl. Annularia mucronata has 10–24 leaves per verticil compared to 6–11 leaves in A. gondwanensis. The shape (spathulate) and size (4–24 mm long) of the leaves of A. mucronata are different from those in A. gondwanensis. Annularia latifolia known from Euramerica is closely comparable to A. gondwanensis in the number of leaves per verticil (6–11 in A. gondwanensis and 7–12 in A. latifolia), but differs in the variable shape and size of the leaves in a single whorl. Annularia stellata has a different in the number of leaves (13–32) per verticil which are linear to oblanceolate in shape and differ also in size (14–75 mm). Annularia sphenophylloides with 10–20 broadly spathulate leaves per verticil is different from A. gondwanensis. The oblanceolate leaves of A. hammanskraalensis (Anderson and Anderson, 1985) and the narrow elliptic leaves of A. kurtzii (Archangelsky, 1960) are different from the oblong–obovate leaves of A. gondwanensis. The other species of Annularia known from the upper Palaeozoic can be distinguished from
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Fig. 2. Comprehensive lithosuccession (not to scale) of section exposed in Chitra B mines, Tulsidabar quarry showing fossil locations.
A. gondwanensis on the basis of their conspicuously different number and arrangement of leaves at the nodes, and the shape of the lamina and apex. Cuticular features are currently known for only a few Annularia species. Cuticles of A. mucronata Schenk and A. stellata (Schlotheim) Wood were studied by Abbott (1958) and Cuneo (2000). The anatomical details of a permineralised specimen of A. hoskinsii were described by Good (1976). The general features of the epidermal layers of A. gondwanensis, viz., the hypostomatic nature, the narrow elliptic stomata, sunken guard cells without wall thickenings, two reniform conspicuous, subsidiary cells on either side of stomata, the shape and the arrangement of the epidermal cells are similar to the cuticles of the above mentioned species. Hairs such as those found in the cuticles of A. mucronata, were not observed in A. gondwanensis. However, the open or collapsed circular or semicircular dark (Plate II, 8–9; Plate III, 6) areas in the middle of each cell appear to be the hair bases. Despite the similarities in the cuticular features, the macromorphological characters of A. stellata, A. mucronata are unlike those of A. gondwanensis, as mentioned above. The macromorphological features of the erect axis are not very clearly preserved in comparison with the Calamites stem. The macerated wood fragments of the erect axis as well as branches show tracheids with spiral and bordered pitted thickenings similar to
the tracheids described in the equisetaleans (Boureau, 1964; Taylor and Taylor, 1993). Annularia. sp. (Plate IV, 1–2; Plate V, 1–2)
Specimen numbers Repository Locality of collection Stratigraphic horizon
ScTd 493, ScTd 508 (Plate IV, 1–2; Plate V, 1–2) MB collection of Palaeobotany–Palynology Section, Department of Botany, University of Calcutta Tulsidabar Quary, Saharjuri basin, Jharkhand, India Shale layer within coal horizon XII — Tulsidabar Top coal seam (Fig. 2), Barakar Formation, late Lower Permian (late Artinskian)
3.4. Description The leafy shoots are articulated with long internodes. The partially preserved internodes measure 31–33 mm. The nodal disk is 4 mm in diameter and slightly elongated. The leaves 27–38 mm long and 2–5 mm wide are basally adpressed and spread out distally. The whorls at the nodes consist of 16–18 leaves, with 8–9 leaves on either side. The leaves in the middle are longer than the others. The shape of leaves varies from oblong to linear with acute apices. The mid-vein is not very prominent and measures 1/7 of width of the lamina.
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3.5. Comparison Annularia sp. resembles A papilioformis Kawasaki (1927), recorded from Lower to early Upper Permian of Shanxi, northern Cathaysian province, China (Guanglong, 1995), in the number, shape and arrangement of the leaves. The leaves of A. papilioformis are moreor-less uniform in size, however, they are basally welded in a single plane at the nodal region while the leaves in Annularia sp. are free, unequal in size and with closely adpressed bases. Annularia kurtzii Archangelsky 1960 (Plate IV, 3–6; Plate V, 3–4) Specimen numbers Repository Locality of collection Stratigraphic horizon
ScTd 491, (Plate IV, 3–4; Plate V, 3), ScTd 494, (Plate IV, 5–6; Plate V, 4), MB collection of Palaeobotany–Palynology Section, Department of Botany, University of Calcutta Tulsidabar Quary, Saharjuri basin, Jharkhand, India Shale layer within coal horizon No .XII — Tulsidabar Top coal seam (Fig. 2), Barakar Formation, late Lower Permian (late Artinskian)
3.6. Description The axis is articulated with nodes and internodes; branches emerge from the nodes. The internodes are longitudinally striated, 16–18 mm long and 1.5–2 mm wide. The leaves are narrow elliptic in
399
shape with a distinct midrib, an acute apex, an entire margin and they emerge radially from the nodes. The basal portions of leaves are joined to form a nodal disk. The number of leaves per whorl varies from 9 to 10. The leaf length covers almost the internodal length, but does not exceed the internode. Leaves measure from 12 to 24 mm in length and the width varies from 1.5 to 2.5 mm. 3.7. Comparison Because of the similarity of characters, i.e., shape, number and arrangement of leaves, the present specimens are assigned to Annularia kurtzii described by Archangelsky (1960) from Lower Permian sediments of Parana Basin, South America. 4. Discussion 4.1. Biogeographical range of occurrence of Annularia The leaf genus Annularia, Asterophyllites, the stem genus Calamites and the fertile genera Calamostachys or Palaeostachya represent different parts of the plant Calamites. The plant flourished in tropical, warm, humid forests of Euramerica during late Palaeozoic together with various lepidophytes, the sphenophyte genera, Sphenophyllum, Bowmanites, Phyllotheca and other members of Euramerian flora (Meyen, 1987; Cleal and Thomas, 1991). During late Early Permian
Plate I. (see Plate I on page 400)
1. 2. 3. 4. 5. 6.
Shows in situ occurrence of upright coalified axis preserved in the thick sandstone layer; the branches emerge radially from the upright axis (the attachment points are marked by arrow) with leaves in whorl at nodes; A. gondwanensis sp. nov. (specimen photographed in the field). A portion of the branch system showing the branching pattern. A. gondwanensis sp. nov., Holotype Sp. No. ScTd 450. Enlarged view of the branch system showing unequal nature of leaves, pattern of emergence of leaves from nodes and nodal disk. A. gondwanensis sp. nov., Holotype Sp. No. ScTd 450. Portion of branches enlarged to show the details of leaf characters viz; unequal size, contracted base, mucronate apex and the mode of attachment to the axis forming nodal disk. A. gondwanensis sp. nov., Holotype Sp. No. ScTd 450. Macerated preparation of the nodal portion showing arrangement of unequal leaves with thick midvein and mucronate apex. A. gondwanensis sp. nov., Sl. No. ScTd 450/1. Enlarged view of a single leaf .A. gondwanensis sp. nov., ScTd 450.
Plate II. (see Plate II on page 401)
1. 2. 3. 4. 5. 6. 7. 8–9. 10. 11–13. 14. 15.
Macerated preparation of a leaf showing mucronate apex. A. gondwanensis. sp. nov., Sl. No. Sc Td. 450/2. Stomatiferous layer of the cuticle shows random distribution of stomata and epidermal cells of variable shape and size. A. gondwanensis sp. nov., Sl. No. ScTd 450/3. Enlarged view of portion of Fig. 2 .A. gondwanensis sp. nov. Stomatal apparatus and the surrounding epidermal cells. A. gondwanensis. sp. nov., Sl. No. ScTd 450/3. Stomatal apparatus and the surrounding epidermal cells (Marked by arrow- a) guard cell, b) subsidiary cell, c) epidermal cell). A. gondwanensis. sp. nov., Sl. No. ScTd 450/3. Stomata enlarged showing sunken nature of guard cells without wall thickening, two reniform subsidiary cells with thick walls. A. gondwanensis sp. nov., Sl. No. ScTd 450/3. Non stomatiferous upper epidermal layer showing epidermal cells of variable shape and size; each cell shows dark area of hair base. A. gondwanensis. sp. nov., Sl. No. ScTd 450/4. Non stomatiferous upper epidermal layer showing collapsed (8) or open (9) hair bases. A. gondwanensis sp. nov., Sl. No. ScTd 450/4. Wood elements showing tracheids with thickenings. A. gondwanensis sp. nov., Sl. No. ScTd 450/5. Enlarged view of wood elements with bordered pits and scalariform thickenings. A. gondwanensis sp. nov., Sl. No. ScTd 450/5. SEM photo of coalified layer of leaf on rock showing stomata and epidermal cells. A. gondwanensis . sp.nov.. Portion of Holotype Sp.No. ScTd 450. SEM photo of portion of macerated wood showing bordered pits and scalariform thickening. A. gondwanensis sp. nov., Holotype Sp.No. ScTd 450.
Plate III. (see Plate III on page 402)
1. 2. 3. 4. 5. 6.
Shows in situ occurrence of upright coalified axis preserved in the thick sandstone layer; the branches emerge radially from the upright axis (point of attachment marked by arrow) with leaves in whorls at nodes; A. gondwanensis sp. nov. (specimen photographed in the field). A portion of the branch system showing the branching pattern. A. gondwanensis sp. nov. Holotype Sp.No.ScTd 450. Portion of axis enlarged to show the alternate arrangement of ridges and furrows, details of leaf characters viz., unequal size, contracted base, mucronate tip, mode of attachment to the axis forming nodal disk. A. gondwanensis sp. nov., Holotype Sp.No.ScTd 450. Stomatiferous layer of the cuticle showing epidermal cells of variable shape and size, stomata, guard cells, reniform subsidiary cells and additional surrounding epidermal cells. A. gondwanensis sp.nov., Sl. No. ScTd 450/3. Enlarged view of stomatal layer showing epidermal cells, subsidiary cells with thickening, sunken guard cells, stomatal opening (Marked by arrow — a) guard cell, b) subsidiary cell, c) epidermal cell). A. gondwanensis sp. nov. Sl. No. ScTd 450/3. Epidermal cells of the upper cuticle; each cell with hair base, showing broad open or collapsed hair bases. A. gondwanensis sp. nov., Sl. No. ScTd 450/4.
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Plate I (caption on page 399).
M. Banerjee et al. / Review of Palaeobotany and Palynology 153 (2009) 394–407
Plate II (caption on page 399).
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Plate III (caption on page 399).
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Fig. 3. Leaf length and breadth ratio of A. gondwanensis sp.nov.
and Late Permian, several species of Annularia flourished in the Cathaysian province of North China, South China and the Tibetan Plateau together with Sphenophyllum and other Euramerian and endemic taxa. These assemblages are recognized as equatorial belt plant assemblage (Meyen, 1987; Cleal and Thomas, 1991; Li et al., 1995; Guanglong, 1995; Yao et al., 2000; Mei et al., 2000; Wang et al., 2000b,a). Annularia also thrived in the Angara flora (Meyen, 1987). In the Gondwana, Annularia is not as frequent in the upper Palaeozoic as in the Euramerian and Cathaysian floras. The genus occurs in the Glossopteris floral assemblage at restricted stratigraphic sequence. Unequivocal records of the genus are from the Lower Permian of the Parana Basin, Brazil, the Paganzo Basin, Argentina in South America (Archangelsky, 1960, 1986; Cuneo, 2000), the Karoo Basin in South Africa (Anderson and Anderson, 1985), western extraPeninsular India (Tiwari and Singh, 1981) and is now reported from upper Lower Permian strata of Saharjuri Basin in eastern peninsular India. The global distribution pattern of the Annularia species thus reveals the biogeographical range of occurrence up to the eastern peninsular India in the Gondwana province during the Early Permian (Fig. 4; Table 1). 4.2. Palaeoclimate and palaeogeography The postglacial Early Permian climate of Gondwanaland is known to be cool-temperate. However, palaeobotanical data from the western extra-peninsular India in the Indian subcontinent indicate a warm, humid climate since the Early Carboniferous (Pal and Chaloner,
1979; Pant et al., 1984; Pant and Srivastava, 1995) to the Late Carboniferous and Permian (Srivastava and Kapoor, 1969; Tiwari and Singh, 1981; Singh et al., 1982). Annularia species viz; A. stellata occur in Lower Permian rocks of Kumaon Himalaya (Tiwari and Singh, 1981), equivalent to the Mamal Formation (Lower Permian) of Kashmir Himalaya, India. The present record from Saharjuri Basin is very similar to the sphenopsid-rich assemblages of warm, humid climate known from the Lower Permian basins of western extra-peninsular India (Singh et al., 1982; Pant et al., 1984), South America (Archangelsky, 1960, 1986, 1990; Cuneo, 2000; Escapa and Cuneo, 2005; Escapa and Cuneo, 2006) and South Africa (Anderson and Anderson, 1985). The plants could thrive in the areas as suitable climatic condition prevailed during late Early Permian (Artinskian). Li Xingxue (1986) suggested that towards the end of the Sakmarian the climate in Gondwana became warmer. Since then the warm climate continued to the Artinskian. This is evidenced by the abundant warm water faunas. The reason was cited as the start of drift of Indian subcontinent (a part of Gondwanaland) and the southern Tibetan Plateau since Late Carboniferous towards warm subtropical areas. Archangelsky (1990) suggested ‘a latitudinal control of floral distribution in the Early Permian’ and a changed palaeogeography of the areas of South American, African basins due to improved latitudinal positions during late late Carboniferous and Early Permian. The palaeobotanical data suggest a lower latitudinal palaeogeography of the Saharjuri Basin, western extra-peninsular basins in the Indian subcontinent and the basins of South America, South Africa during Early Permian. 5. Conclusion 5.1. Palaeoecology The preserved length of 130 cm and 24–28 cm width of the erect axis suggest arborescent growth habit of the plant. The occurrence of the calamitalean genus Annularia in growth position is a rare and important addition to the Glossopteris flora. The occurrences of A. gondwanensis sp. nov., Annularia. sp, A. kurtzii, assemblage of diverse articulate fertile and sterile taxa viz.; the bractless fertile organ Tulsidabaria indica, the bracteate fertile cone bearing plant Rajmahaliastachys elongata, vegetative shoots of Sphenophyllum sp., Phyllotheca spp. suggest the occurrence of a swamp in a warm and humid environment. The arborescent A. gondwanensis and upright Glossopteris plants in the associated sediments (Fig. 2) indicate in situ deposition of the forest in the Saharjuri Basin during late Early Permian. The evidence of Annularia spp. in the Lower Permian rocks of South America in the Parana Basin of Brazil, the Paganzo basin, Argentina, in the upper Lower Permian of Patagonia, Argentina, South America, Karoo Basin of South Africa, western extra-peninsular basin, the present record from late Lower Permian of Saharjuri Basin, eastern
Plate IV. (see Plate IV on page 404)
1. 2. 3. 4. 5. 6.
Incompletely preserved leafy shoot; leaves ad pressed basally at nodes and spread out distally. Annularia. sp., Sp.No.ScTd 493. Partially preserved leafy shoot of Annularia. sp. showing leaves emerging from the node, Sp.No.ScTd 508. Leafy shoot with three whorls of leaves. A. kurtzii Archangelsky, Sp.No ScTd 491. A single leaf whorl magnified showing arrangement of leaves of A. kurtzii. Archangelsky, Sp.No. ScTd 491. Leafy shoot of A. kurtzii Archangelsky with branch system, Sp. No. ScTd 494. A portion of leafy axis magnified showing leaf arrangement of A. kurtzii Archangelsky, Sp.No. ScTd 494.
Plate V. (see Plate V on page 405)
1. 2. 3. 4.
403
Partially preserved nodal region showing the arrangement of leaves. Annularia. sp., Sp.No.ScTd 493. Partially preserved leafy shoot of Annularia. sp. showing leaves emerging from the node, Sp.No.ScTd 508. Leafy shoot with three whorls of leaves .A. kurtzii Archangelsky, Sp.No.ScTd 491. Leafy shoot of A. kurtzii Archangelsky with branch system, Sp. No. ScTd 494.
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Plate IV (caption on page 403).
M. Banerjee et al. / Review of Palaeobotany and Palynology 153 (2009) 394–407
Plate V (caption on page 403).
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Fig. 4. Global distribution of Annularia spp. ( ) including present record rocks of Gondwana continents.
during Late Palaeozoic plotted in the base map after Ziegler et al., 1997. ( ) Records from Early Permian
peninsular India plotted in the base map of Ziegler et al. (1997) reveal that a rich assemblage of sterile and fertile sphenopsids and diverse glossopterids flourished in the lowland areas of the Gondwana continent (Fig. 4) during the Early Permian. Acknowledgements The authors are grateful to Professor W. G. Chaloner FRS, Geology Department, Royal Holloway University of London for critical examination of the original manuscript and for valuable suggestions encouraging early publication of the paper. The constructive suggestions, comments by the anonymous reviewers of Review of Palaeobotany Palynology are gratefully acknowledged. Thanks are due to the Department of Science and Technology, Life Science, New Delhi, for providing financial assistance (SP/SO/A-61/98) to carry out the investigation. Grateful thanks are due to the Officers of Coal India Ltd., Eastern Coalfields Ltd., General Managers and members of Chitra Coal fields, Chitra, Jharkhand for their kind permission to undertake fieldwork also for all kinds of assistance during the collection of materials.
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