The genus Glossopteris from the Late Permian beds of Handapa, Orissa, India

Review of Palaeobotanyand Palynology, 75 ( 1992): 183 218 Elsevier Science Publishers B.V., Amsterdam

183

The genus Glossopteris from the Late Permian beds of Handapa, Orissa, India Shaila Chandra

and Kamal Jeet Singh

Birbal Sahni Institute c?fPalaeobotany, 53, University Road, Lucknow, 226 007, India (Received May 21, 1991; revised and accepted June 16, 1992)

ABSTRACT Chandra, S. and Singh, K.J., 1992. The genus Glossopteris from the Late Permian beds of Handapa, Orissa. India. Rev. Palaeobot. Palynol., 75: 183-218. Forty two species of the form genus GIossopterishave been reported from Hinjrida Ghati section, near Handapa village in Dhenkanal District of Orissa. This is the first report of such a large number of species of this genus from a single locality and formation from India and perhaps from the Southern Hemisphere. The huge collection of Glossopterisspecimens, with a great diversification of species, clearly indicates that the genus Glossopterisgrew as luxuriantly in the Kamthi Formation as in the equivalent Raniganj Formation, both of Late Permian age. Abundant leaves of GIossopterisin the fine sediments indicate seasonal falling and favourable conditions for plant growth. A vegetational reconstruction of the Handapa area representing Late Permian time is attempted. It is assumed that the thick forest must have been dominated by a variety of GIossopteris trees and shrubs with small patches of open area for the development of pteridophytic plants as undergrowths as shown by the composite fossil flora of the beds.

Introduction T h e genus Glossopteris c o m p r i s e s a b o u t 85% o f the total p l a n t fossils collected at the H a n d a p a locality. Q u a l i t a t i v e l y and q u a n t i t a t i v e l y the flora o f the H a n d a p a beds is one o f the richest floras recovered from I n d i a n P e r m i a n beds. T h e c o m p l e t e flora is represented by v a r i o u s groups, e.g. lycophytes, a r t h r o p h y t e s , filicales, glossopteridales, cycadales a n d ginkgoales, c o m p r i s i n g in all 18 genera a n d 64 species. L y c o p h y t e s are represented by the genus Cyclodendron; equisetales by four genera, i.e. Schizoneura, Phyllotheca, Lelstotheca a n d Raniganjia; s p h e n o p h y l l a l e s by two genera, i.e. Trizygia a n d Sphenophyllum; filicales by five genera, i.e. Dizeugotheca, Neomariopteris, Pantopteris, Damudopteris a n d Asansolia; g l o s s o p t e r i d a l e s by three genera, i.e. Glossopteris, Surangephyllum and Vertebraria; cycadales by two genera, i.e.

Correspondenceto: Dr. (Mrs) S. Chandra, Birbal Sahni Institute of Palaeobotany, 53, University Road, Lucknow, 226007, India. 0034-6667/92/$05.00

Pseudoctenis a n d Senia; a n d ginkgoales by one genus, i.e. Handapaphyllum. Besides these, eleven genera o f g l o s s o p t e r i d e a n fructifications representing seventeen species are also k n o w n from these beds. All the a f o r e s a i d genera have been r e c o r d e d or described by various workers, i.e. S u b r a m a n i a n a n d R a o (1960), K h a n (1969), S u r a n g e a n d M a h e s h w a r i (1970), S u r a n g e a n d C h a n d r a (1973a, b, c, 1974a, b, c, d), M a i t h y (1977), C h a n d r a a n d S u r a n g e (1977), C h a n d r a a n d R i g b y (1981, 1983), C h a n d r a (1984), P a n t et al. (1985), a n d C h a n d r a a n d Singh (1986, 1988, 1989). Recently, Singh a n d C h a n d r a (1987) erected seven new species o f the genus Glossopteris f r o m the same beds. S u b r a m a n ian a n d R a o (1960) r e c o r d e d four species o f this genus, n a m e l y GIossopteris indica, G. communis, G. ?angustiJolia a n d G. ?ret(fera. K h a n (1969) r e c o r d e d three m o r e species from the same beds, i . e . G , conspicua, G. browniana, a n d G. cf. damudica. Except for these brief reports, no detailed w o r k has been carried out on the genus Glossop-

,~' 1992 Elsevier Science Publishers B.V. All rights reserved

184

S. C H A N D R A A N D K J . S I N G H

teris in this area. For this study, several systematic fossil collections were made in the area. Our findings indicate the presence of many more species of Glossopteris than reported earlier. M a t e r i a l and m e t h o d s

The Glossopteris species described in this paper have been collected from the fossiliferous beds exposed in Hinjrida Ghati section (20°58'N,

84°43'E) near Handapa village in Dhenkanal, District of Orissa, India (Fig. 1). The leaves were found as impressions on pinkish yellow to pinkish brown, hard, compact, fine-grained calcareous shales. All the photographed and figured specimens are housed in the museum of the Birbal Sahni Institute of Palaeobotany, Lucknow, vide statement No. 777. Terms and methodology as applied by Chandra and Surange (1979) in identifying the species of the genus Glossopteris have been fol-

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Fig. 1. Geological map of a part of the Mahanadi Basin showing the fossil locality and the adjoining areas (after V. Ball, 1877).

THE G E N U S G L O S S O P T E R I S FROM THE LATE PERMIAN BEDS OF HANDAPA, INDIA

lowed in this paper. Detailed repetitive descriptions and comparative accounts of the well-known species reported from Handapa beds have been avoided in this paper, however, the specific features are depicted in Table I for comparative account. The species are arranged in three broad categories following Feistmantel's broad divisions (1881). For convenience sake these categories are based on visual aspects because any other parameter is not possible at present for classifying the species of Glossopteris. The specific morphographic characteristics are divided into thirteen major columns. Identification of the 34 species is generally based on these characters and matching them with the holotypes, either with actual specimens or their photographs. It is evident from Table I that identification up to the species level is not based on any particular constant characteristic but mostly on permutation and combination of characteristics. It seems that the shape of the leaf and the venation pattern in the middle part of the leaf are quite constant and reliable characteristics for specific delimitations. Epidermal characteristics, for that matter, are also not very useful as the cuticles of most gymnospermous genera from the Southern Hemisphere exhibit more or less similar features. Discussion and conclusion

Plant fossils are hardly known from the Kamthi Formation. The first palaeobotanical investigation of the Kamthi Formation was published by Bunbury (1861) reporting fossil plants from the Kamptee and Silewada localities in Nagpur area. The species recorded were Dizeugotheca phegopter-

oides (Pecopteris phegopteroides), Rhabdotaenia danaeoides ( Taeniopteris danaeoides), Noeggerathiopsis hislopii, Filicites and further certain plants of uncertain affinities, namely Phyllotheca indica, ?Vertebraria, ?Knorria, Yuccites, a fern rhizome and some stems with ridges and furrows. Apart from the above mentioned taxa, he also reported five species of Glossopteris, namely Glossopteris indica (G. browniana var. indica), G. browniana (G. browniana vat. australasica), G. musaefolia, G. leptoneura, and G. stricta. All these species of Glossopteris are found in the Handapa assemblage except for G. musaefolia.

185

Ball (1877) reported plant fossils from the Kamthi group of the Mahanadi Basin: Schizoneura

gondwanensis, Vertebraria indica, Filicites, Neomariopteris polymorpha (Sphenopteris polymorpha), and Pecopteris sp. The genus GIossopteris was represented by three species, namely G. indica, G. browniana, and G. communis. All three species are well represented in the Handapa flora. Hughes (1877) reported plant fossils from various places around Nagpur, Isapur near Chanda, Charvat, Kawarsa, and Anur near Antargaon. The fossils were later on described by Feistmantel (1880). The plant taxa were Raniganjia bengalensis

(Actinopteris bengalensis), Phyllotheca indica, Schizoneura sp., some seeds as Cycadinocarpus, and four species of Glossopteris, i.e.G, browniana, G. indica, G. musaefolia, and G. leptoneura. Except for G. musaefolia all other species of the genus are common to the Handapa flora. Oldham (1880) described plant fossils from the Kamthi area, namely Phyllotheca indica, Vertebraria indica, Pecopteris sp., Gangamopteris hughesi, Angiopteridium cf. A. maclellandi, Rhabdo-

taenia danaeoides (Macrotaeniopteris danaeoides), Macrotaeniopteris feddenii, Noeggerathiopsis hisIopii and seven species of Glossopteris i.e., G. communis, G. indica, G. browniana, G. damudica, G. stricta, G. musaefolia and G. leptoneura. Of the Glossopteris species except G. musaefolia all are recorded in the Handapa flora. Feistmantel (1881) reported the following plant fossils from the Kamthi Formation of the WardhaGodavari Valley Coalfields and from Chanda of Nagpur area: Dizeugotheca phegopteroides (Pecop-

teris phegopteroides), Rhabdotaenia danaeoides (Macrotaeniopteris danaeoides), M. feddenii, Angiopteridium cf. A. maclellandi, Gangamopteris hughesi, Noeggerathiopsis hislopii, Anthrophyopsis, Rhipidopsis densinervis and eight species of Glossopteris, i.e.G, communis, G. stricta, G. musaefolia, G. indica, G. damudica, G. angustifolia, G. leptoneura and G. browniana. All these species except G. musaefolia are commonly found in the Handapa flora. Chandra and Prasad (1981) reported a number of plant fossils from the Kamthi Formation of the Kanhargaon and Bazargaon villages located in

186

S, C H A N D R A A N D K.J. SINGH

TABLE 1 Comparative description of Glossopteris species from the Handapa area Species name

Number of specimens studied and leaf type

Shape of leaf

Apex

Base

Width

Length/ width ratio

Variation in size Length

A. N A R R O W MESH F O R M S O F GLOSSOPTERIS SPECIES

Gh;ssopteris angust(lolia

35. 11 almost complete

3.8 13 cm

0.6 2 c m

6 8:1

Lorate-oblong to narrow-oblanceolate

Obtuse

Acute-cuneate

4, 2 almost complete, small, very narrow

4.6 6.1 cm

0,9 I.I cm

6: I

Oblong

Acute

Acute-cuneate

18, 3 almost complete, medium to large, broad in middle, tapering towards both ends

20-35 cm

5.6 10cm

3-4: I

Elliptic

Acute

Acute-cuneate

6, 2 complete, small

7.3 9.6 cm

2.2 2.3 cm

3 4:I

Spatulateoblanceolate

Obtuse

Acute-cuncate

GhJssopteris vulgaris Pant and Gupta. 1968 (Plate IV. 2)

1, medium, strap shape, narrow. tapering toward both ends

I 1.7 c m

1.6 c m

7:I

Linear-oblong

Acute

NOt preserved

Gh~ssopteri~ tenu([blia Pant and Gupta, 1968 (Plate IV, I; Plate V, I, 2)

27, 4 complete, leaves medium to large, mostly narrow

10.8-

1.2 2.9cm

5 9:1

Oblong-oblanceolate

Acute obtusely pointed

Acute-cuneate

Gh~ssopteris varia Pant and Gupta, 1968 (Plate IV. 3)

3 complete, small, narrow

6.1 7.4cm

1.2-1.6 cm

4-5:1

Ellipticaloblanceolate

Acute or obtusely pointed

Acute-normal

GIossopteris zeilh.ri Pant and

3 small, narrow, lamina broader in apical part, narrows down to form an elongate base

4,9cm

1.5cm

4-5: I

OblanceoIate

Obtuse

Acute-cuneate

9, medium to large. broad at apical end, taper towards basal end

Most complete leaf 18,8 cm

3-4.8 cm

3 5: I

Oblanceolatc

Obtuse

Acute-cuneate

17 leaves quite big. broad, lamina abruptly narrows down towards base

24.5

9.8

2 3:1

Elliptical

13.4 cm

Acute-obtusely pointed

Acute-cuneate

27.2 cm

Brongniart, 1828 (Plate I. 1, 3; Plate II, I, 4)

Gh~ssopteris h,ptoneura Bunbury, 1861 (Plate L 2, 4; Plate I1, 2, 3)

Glossopteris communis Feistmante[, 1879 (Plate 1. 5; Plate 11, 5)

GlossoplerL~ stenoneura FeistmanteL 1881 (Plate III, 2: Plate IV, 4)

Gupta. 1968 (Plate IV. 51

Glossopteris ~patuhtta Pant and Singh, 1971 (Plate II1, I: Plate IV, 6)

Gh~ssopter&gig~ Pant and Singh 1971 (Plate 111, 3: Plate VI, 1)

16.4 cm

i87

T H E G E N U S GLOSSOPTERIS F R O M T H E LATE P E R M I A N BEDS O F H A N D A P A , I N D I A

Variation in petiole size

Midrib Type

Thickness at base

0.4 0.8 cm long

Medium to broad, flat, striated, evanescent towards apex

1.5 2 . 5 m m

Not preserved

Medium, persists up to apex

2-2.5 mm

Not preserved

Thick, flat. strong, persists up to apex

5 m m 1.2 cm

Not preserved

Thick, very thin towards apical region, ultimately dissolving into lateral veins

2 3 mm

Not preserved

Medium to thick, very broad at base. flat, striate, persists up to apex

3.5 m m

Petioles preserved in two leaves, measure 17 cm in length

Narrow to medium, fiat, striate, persists up to apex

1.5-3 mm

Not preserved

Wide at base. gradually tapering upwards, persists up to apex

2-2.5 m m

Not preserved

Medium, striate, becoming evanescent near apex

1.5 2 m m

Not preserved

Medium to broad, fiat, striate. persists up to apex

2.5-4.5 mm

Not preserved

Very wide, flat, striate, persists up to apex

0.8-1 cm

Angle of emergence of secondary veins and type of venation

Average concentration of veins per cm 2

Mesh type

Average concentration of meshes per cm 2

% age occurrence of species in flora

Emerge at 10 15<'. meeting margin at 2 5 - 4 0 . veins generally fork once or twice or sometimes thrice from midrib to margin Emerge at 12 I Y . meeting margins at 35 40 °

40-45

Elongate, oblong to linear-polygonal

33 48

10.17

Per 1/2 cm 2 16-25 at middle part of leaf

Per I / 2 c m ~ 22 24 at middle part

1.16

Emerge at 12 20', pass out to margins in graceful curves meeting at 50-65% veins run almost parallel to each other, thin, dense Emerge at 12 20 °, arch out in graceful curves to meet margin at 45-55 . venation very dense Emerge at 12-15% meeting margin at 35 40°, thin, comparatively dense E m e r g e a t 10 18, arch slightly, run straight upto margin meeting at 35 45", 2 3 meshes formed between midrib to margin, venation very dense Emerge at 10 15', arching backward. meeting margin at 35-40% venation dense Emerge at 12 15', arch slightly backwards and continue straight to meet margin at 30-35

45 60 in middle part of leaf

Elongate. narrow. oblong-polygonal, slightly broader near midrib than at margins Very long, very narrow, uniform in size throughout lamina, oblong in shape

40 45 in middle part

5.23

30 32

Narrow. elongate. oblong in shape

38 42

1.74

Per I / 2 c m 2 16 18

Few. narrow, elongate, uniform. linear-oblong to polygonal in shape

Per I / 2 c m 2 15-17

0.29

38 42

Very long, narrow. uniform in size throughout lamina, polygonal or spindle shape

40 55

7.85

Per l / 2 c m 2 18 20

Per l / 2 c m 2 20 25

0.87

Per I / 2 c m 2 20 25

0.87

Emerge at 10 17°, arching outwards, meeting margin at 50-65" in middle part, venation very dense Emerge at 12-20% arching outwards, pass out in gentle curves meeting margin at 55 70" in middle part, venation dense

30-32 near midrib, 34 37 near margin

Long, narrow, polygonal in shape, uniform in size throughout lamina 2-3 meshes formed from midrib to margin, very long, narrow near midrib, short and narrow at margins, linearpolygonal in shape Very long, narrow linear-polygonal or spindle shaped. 3 ~ . meshes between midrib and margin 5 7 very narrow, elongate meshes formed between midrib and margin, meshes uniform throughout

20 28 near midrib. 25 34 near margin

2.62

Per 2 cm 2 28-30 at midrib, 70-75 at margin

4.94

Per 1/2cm 2 22-26

Per 2 cm 2 35 40 in middle part

188 TABLE I

S. C H A N D R A A N D K.J. SINGH

(continued)

Species name

Number of specimens studied and leaf type

Variation in size Length

Glossopteris pandurata Pant and Gupta, 1971 (Plate VI, 3)

Glossopteris angusta Pant and Gupla. 1971 (Plate IV. 7)

Glossopteris gopadensis Banerji, Maheshwari and Bose, 1976 (Plate V, 3; Plate VII, I )

3, almost complete. small, broadest near apex. gradually tapering towards base 4. small to medium. broader in apical region, tapering towards base

4, small, narrow. strap shaped

Width

Length/ width ratio

Shape of leaf

Apex

Base

3.7 5.4 cm

1.3 2 cm

2 3: I

Narrow obovate oblanceolate

Round-flattened with an indication of a slight notch

Acute-cuneate

6.9 9.3 cm

2.1-2.4 cm

4 6:I

Oblanceolate

Retuse to emarginate

Not preserved

Most complete leaf 7.7 cm long

1.8 Cm

4 7:1

Narrow-oblong lorate-oblong

Acute

Acute-normal

B. M E D I U M MESH F O R M S OF GLOSSOPTERL~ SPECIES

Glossopteris strieta Bunbury,

3, 1 almost complete

Most complete leaf 12.5 cm

2.7 cm

5:1

Oblong-elliptical

Acute

Not preserved

65, 3 complete, small to large, more elongate than broad

I0 31 cm

3.3-3.4 cm

5 10:

Narrow-elliptic very narrow-elliptic

Acute-normal

Acute-normal

3. 1 complete, large, longer than broad, small to medium

22.1 cm

3.3 4.4 cm

56:

Oblong

Acute

Acute-cuneate

18.4 almost complete, medium size. broader near apical part tapering towards base

10 15cm

2-3.5 cm

46:

Elliptic-oblanceolate

Obtuse

Acute-normal acutecuneate

15, none complete. medium size. longer than broad

Most complete leaf 12.4 cm

Most complete leaf 2.9 cm

56:

Oblong

Acute

Acute-cuneate

3. 2 almost complete, small, broad

Most complete 7.0 cm

2.2-4 cm

23:

Elliptical

Acute-obtuse

Base imperl~ctly preserved

8, small to medium, not very broad

Most complete leaves 10 10.4 cm

1.6 2.3 cm in most complete leaves

56:

Narrow-ellipticlorate-oblong

Retuse

Acutc-cuneate

1861 (Plate V, 4; Plate VI, 2)

Gh~ssopteris indica Schimper, 1869 (Plate VII. 2: Plate VII 1. I)

Glossopteris arberi Srivastava 1956 (Plate VII. 3: Plate VIII. 2)

Glossopteris subtilis Pant and Gupta. 1971 (Plate VIII. 3; Plate IX. 1)

Glossopteris lanceolatus Pant and Singh, 1971 (Plate IX, 3)

Glossopteris barakarensis Kulkarni, 1971 (Plate IX, 4: Plate X. I)

Glossopteris tenuinervis Pant and Gupta. 1971 (Plate IX, 2; Plate X. 2)

189

T H E G E N U S GLOSSOPTERIS F R O M T H E LATE P E R M I A N BEDS O F H A N D A P A , I N D I A

Variation in petiole size

Midrib Type

Thickness at base

Moderately thick, flat. tapering upwards, persists up to apex Medium to thick, fiat. persistent

I 2 mm

Not preserved

Narow to broad, striate, grooved. persists up to apex

2 3 mm

Not preserved

Broad, strong, persistent, striate, attenuating rapidly in upper 1/3 part

3 4 mm

1.6 2.9 cm

Medium to very wide, stout, flat, grooved, striate, persists up to apex

2.5-5 mm

Not preserved

Very broad, fiat, tapering upwards, persists up to apex

5 mm

1.1 2 . 8 c m

Medium, flat, very gradually tapering in apical region, persists up to apex

1.5 3.5 m m

Not preserved

Narrow, fiat, persists up to apex

2 mm

Not preserved

Broad, flat, tapers rapidly above middle region, persists up to apex

2.5 4 m m

Not preserved

Medium, fiat, persists up to apex

1.5-2.5 mm

4 6 mm

Not preserved

1.5-3 mm

Angle of emergence of secondary veins and type of venation

Average concentration of veins per cm 2

Mesh type

Average concentration of meshes per cm 2

% age occurrence of species in flora

Emerge at l0 25 ° , arch backwards, meeting margin at 40 60 ° Emerge at 10 1 5 , arch backwards near midrib, run straight to margin to meet it at 30-40 ~', dense Emerge at 40-60 °, archout slightly near midrib, meeting margin at 70-85 °, dense

34 36

2 3 long, narrow, linear-polygonal meshes

45-48

0.87

30-32, in middle part

2-3 very long. narrow, linear meshes between midrib and margin

28-35 in middle part

1.16

30 35 per I/2 cm 2 in middle part

Very narrow, elongate. polygonal

Emerge at 20-35 °, arch backwardly near midrib, travel straight and horizontally, meeting margin at 80-90 °, very dense Emerge at 12 20°~ arch slightly backwards near midrib, pass out straight to margin meeting at 4 5 - 6 0 , veins anastomose 4 - 6 times upto margin Emerge at 20 25 °, arch outwards near midrib, continue upto margin meeting at 65 70 °, veins dichotomies 4 6 times Emerge at 15-20 °, arch backwards, continue straight to margin meeting at 45-60 '~

Near midrib 12-14, near margin 28 30 in middle part

Oblong polygonal, broad near midrib, very narrow, long near margin

Near midrib 30-32 near margin 45-50 in middle part

0.87

30 35 in middle part

Pentagonal to hexagonal, broadest near midrib, narrowing towards margin

30-45 near midrib, 45 60 near margin in middle part

18.90

28 30 in middle part

Oblong-polygonal, very long, uniformly narrow, subparallel to each other

30 35 in middle part

0.87

38 40 in middle part

Comparatively narrow, elongate, oblong-polygonal in shape, uniform in shape and size throughout Elongate, pentagonal or polygonal near midrib, oblongpolygonal in rest of lamina Closely spaced, in a straight line, oblong-polygonal, 6 - 7 meshes formed between midrib and margin 4 - 5 oblongpolygonal meshes~ long, narrow, uniform throughout lamina

35 45 in middle part

5.23

40 45 near midrib, 55-60 near margin

4.36

50 55 in middle part

0.87

65-70.in middle part

2.32

Emerge at 35 55 °, turn backwards with a very gentle curve, passing out straight to margin meeting at 70 85 ° Emerge at 40 60 °, curve slightly downwards, meeting margin at 70-80 ° Emerge at 20-25 °, arching backwars, continue upto margin meeting at 55 75 °

12 14 near midrib. 25-27 near margin in middle part

24 28 in middle part

30 32 in middle part

1.16

190

TABLE

S. C H A N D R A A N D K.J. SINGH

I (continued)

Species name

Number of specimens studied and leaf type

Variation in size

Length/ width ratio

Shape of leaf

Apex

Base

Acute

Not preserved

Length

Width

Largest preserved leaf 8.9 cm long, complete leaf might be 18 26cm in length 9 13cm

4 cm

2.2-3.2cm

3-5:1

Elliptic

Obtuse-round with slight depression

Acute-normal

2. leaf medium, broadest near apical portion, gradually narrowing towards base

15cm

6.6cm

2.5: I

Narrow-obovateoblanceolate

Seems to be obtuse

Not preserved

Glossopteris taeniensis Chandra and Surange, 1979 (Plate XI, 2)

4. 2 almost complete. small comparatively broad

4.6cm

1.4cm

3 4:1

Oblanceolate narrow-elliptic

Obtuse

Acute-normal

Glossopteris nimishea Chandra and Surange, 1979 (Plate X, 3; Plate XV, I)

15, 1 complete, medium to large, broader in middle part, tapering towards both ends

Most complete leaf 16.7 cm

2.6-4.7 cm

4-6:1

Narrow-elliptical

Seems to be acute

Acute-normal-acutecuneate

Glossopteris mohudaensis

4, medium, not very broad, tapering towards base

Most complete leaf 15.8 cm long

2.9 5.4 cm

3-4:1

Narrow-elliptical oblanceolate

Acute

Acute-normal-acutecuneate

13, 5 almost complete, small to medium, broad

6 I4 cm

2-4.4 cm

4:1

Oblanceolate-oblong to elliptic

Acute or obtuse

Acute-normal or acute-cuneate

20, 2 almost complete, medium to large, broad in middle part

11.415.4 cm

2.7-3.9 em

4-5:1

Oblong ellipticoblanceolate

Not preserved

Acute-normal-acute. cuneate

I incomplete, seems to be small to medium, broad, margin of lamina wavy having a wide notch at apex

3.5 cm

5 cm

42:1

Elliptical

Emarginate

Not preserved

I specimen, almost complete, medium, broad, margin of lamina slightly undulating

18.2 cm

6.8 cm

3:1

Elliptic

Obtuse

Acute-cuoeate

Glossopteris nautiyalii Pant and Singh, 1971 (Plate IX. 6)

Glossopteris obscura Pant and Singh, 1971 (Plate IX, 5)

Glossopteris oldhamii Pant and Singh, 1974 (Plate XI, I)

Chandra and Surange. 1979 (Plate XI, 4)

4, incomplete. medium size. broad in middle region, apical part very sharp, lamina widens abruptly towards base

8. only I complete. small to medium, broadest in middle part, tapering towards both ends

C. B R O A D MESH F O R M S OF GLOSSOPTERIS SPECIES

Glossopteris retifera Feistmantel, 1881 (Plate XV, 2)

Glossopteris conspicua Feistmantel, 1881 (Plate XIV. 2; Plate XV, 3)

Glossopteris sp. cf. G. divergens Feistmantel, 1881 (Plate X, 4; Plate XI, 5)

Glossopteris damudica Feistmantel, 1881 (Plate XII, I; Plate XIII, I)

191

T H E G E N U S G L O S S O P T E R I S F R O M T H E LATE P E R M I A N BEDS O F H A N D A P A , I N D I A

Variation in petiole size

Midrib

Angle of emergence of secondary veins and type of venation

Average concentration of veins per em 2

Mesh type

Average concentration of meshes percm 2

% age occurrence of species in flora

Type

Thickness at base

Not preserved

Flat longitudinally striate, persisent

Towards apex 0.25-0.5 mm

Emerge at 15-20 ~, slightly arch backwards passing straight to margin meeting at 45-55 ~

34-36

4 - 6 elongate, narrow, polygonal, uniform throughout

28 30 near midrib. 40 42 near margin

1.16

0.7-0.9 cm

Medium, flat, persistent

2 2.5 mm

34 38 in upper part, 28-32 in middle part

Broad, elongate, oblong-polygonal in shape

48-52 in upper part, 3 4 - 4 0 i n middle part

2.32

Not preserved

Flat. having longitudinal strands throughout. persists up to apex

3 mm

Emerge at 20-25", arch backwards close to midrib, run straight, meeting margin at 50 65 ~' Emerge at 45-65 ° , meeting margin at 75 90 ~'

10 11 near midrib, 16-18 near margin in middle part

19 20 near midrib, 22-24 near margin in middle part

0.58

Not preserved

Medium to broad, flat, striate, attenuating towards apex, persists up to apex Narrow, slightly grooved, striate, persists up to apex

I - 2 mm

Emerge at 20-30 °, arch slightly near midrib, meeting margin at 50 70 °

11 13 near midrib, 22-24 near margin

2-3.5 mm

In middle part emerge at 35-55 °, run straight upto margin meeting at 65 80 E, towards apex emerge and meet margin at 45 55' Emerge at 20-30". arch slightly backwards near midrib, run straight meeting margin at 65 80 °

14-15 near midrib, 19 20 near margin

Large, broad near midrib, progressively long, narrow towards margin, elongate. polygonal in shape Broad, hexagonal to polygonal, longer near midrib, smaller towards margin Broad near midrib, long narrow towards margin, mostly pentagonal or hexagonal

25-27 near midrib, 32-34 near margin

4.36

12-13 near midrib, 20 22 near margin

Very narrow, elongate, polygonal

14 16 near midrib, 30-32 near margin

1.16

1.6-2.7 cm

1.16

Not preserved

Thick, flat, striate. persists up to apex

4 7 mm

Preserved in three, 0.6 I cm

Medium to broad, flat, gradually attenuating towards apex, persistent Flat, persistent

1.5 3 m m

Emerge at 20-30 ° , run straight to margin, meeting at 45-50 °

14-17

Broad, short, almost parallel, pentagonal to hexagonal

15 20

3.78

2 3.5 mm

Emerge at 20-35 ° , run straight without curving, meeting margin at 35 45 ° Emerge at 20-30 °, meeting margin at 40-45 ° with an upward curvature. lateral veins at apex bend inwards towards midrib Emerge at 45 70 °, follow straight parallel course upto margin, meeting at 70-85 ° , in upper part veins slightly curved upwards towards margin

II 15

Very long, broad, pentagonal to hexagonal, oblongpolygonal, uniform throughout lamina Very broad near midrib, becoming narrower towards margin, 3 4 meshes formed between midrib and margin Slightly broad, short near midrib. becoming narrower towards margin, polygonal

10-15

5.81

9 10 near midrib, 15-18 near margin

0.29

25 27 near midrib, 38 42 near margin

0.29

0.9 cm long

Not preserved

Medium, persists upto apex

1.5mm

I cm long

Thick, persistent, attenuating gradually towards apex

4 mm

8 10 near midrib, 14-16 near margin

10-12 near midrib, 30 32 near margin

192 TABLE I

S. C H A N D R A A N D K.J. SINGH

(continued)

Species name

Glossopteris tortuosa Zeiller. 1902 (Plate XII, 3, Plate XIV, 3; Plate X l l l , 4)

GIossopteris gondwanensis Pant and Gupta, 1971 (Plate XII, 4; Plate XIII, 2)

Glossopteris radiata Pant and Singh, 1971 (Plate XI, 3; Plate XII, 2)

Glossopteris sastrii Pant and Singh, 1974 (Plate X l l l , 3; Plate XIV, I)

Shape of leaf

Apex

Base

Length

Width

Length/ width ratio

7, I complete, medium, narrow, contracting gradually towards base

8.8 cm

2 cm

5-6 : 1

Lorate-oblong

Obtuse widely acute

Acute-cuneate

12, 5 complete, small to medium, mostly narrow, some slightly broad in middle part

5.4-7.6 cm

0.9- h6 cm

4 7:I

Lorate-oblong

Acute-obtusely pointed

Acute-normal acutecuneate

1, incomplete, medium, distinctly broad

4.8 cm

3.6 cm

Not

Not known

Not preserved

Acute-cuneate

3 complete, small, gradually tapering towards both ends, broad in middle part

7.1 7.3cm

Narrow-oblongnarrow-elliptic

Obtuse

Obtuse-normal

Number of specimens studied and leaf type

Variation in size

preserved

1.9 2.2 c m

Maharashtra State, namely Neomariopteris hughesi, Trizygia speciosa, Schizoneura gondwanensis, Vertebraria indica, Dictyopteridium sporiferum and thirteen species of Glossopteris, i.e.G. musaefolia, G. stricta, G. leptoneura, G. surangei, G. venustus, G. raniganjensis, G. mohudaensis, G. indica, G. angustifolia, G. tenuifolia, G. lanceolatus, G. bosei, and G. arberi. Agashe et al. (1984) described Glossopteris, Phyllotheca sp., and Vertebraria indica from Chanda, Maharashtra. Subramanian and Rao (1960) and Khan (1969) recorded seven species of Glossopteris from the same Handapa rocks and erected a supposedly cycadalean genus Senia reticulata. Surange and Maheshwari (1970) described some

3 4:1

male and female fructifications referable to Glossopteridales from the same Handapa beds, namely Glossotheca utkalensis, Eretmonia utkalensis, E. hinjridaensis, Lidgettonia indica, Scutum sp., and Cistella sp. Surange and Chandra, in a series of papers (1973a, b, c, 1974a, b, c, d, 1978), described many male and female fructifications belonging to the Glossopteridales from the Handapa beds. Described were Dictyopteridium sporiferum, Denkania indica, Partha indica, P. spathulata, Scutum sahnii, S. elongatum, S. indicum, Cistella ovata, Indocarpus elongatus, Lidgettonia mucronata, Glossotheca immanis, Glossotheca orissiana, and Eretmonia ovata. Chandra and Rigby (1981, 1983) reported a

193

T H E G E N U S GLOSSOPTERIS F R O M T H E LATE P E R M I A N BEDS O F H A N D A P A . I N D I A

Variation in petiole size

Midrib Type

Thickness at base

0.7 l cm

Broad, stout, persists up to apex

2 3mm

Not preserved

Medium, Ilat, striate, persists up to apex

I 2 mm

Not preserved

Very stout, flat with prominent striations

3 mm

1.3 1.9 cm

Medium to broad, flat, striate. persistent

1.5-2 mm

Angle of emergence of secondary veins and type of venation

Average concentration of veins per cm 2

Mesh type

Average concentration of meshes per cm 2

% a g e occurrence of species in flora

Emerge at 40 45 ~, arch slightly close to midrib, follow a sinuous/zigzag course towards margin, meeting it at 65-70 ° Emerge at 20 30", continue straight upto margin meeting at 30 45 °

Per 1/2cm2 9 12

Oblong-polygonal to trapezoidal, broader near midrib than at margin

Per I / 2 c m z 10 15

2.03

12 14

16-18

3,49

Veins run horizontally in middle part. pass oat upwardly in upper region, run downwards in lower part, exhibiting thus a radiating or diverging nature Emerge at 25-30 °, arching backwards. meeting margin at 4 5 - 5 0 ' in upper region, at 60-70 ~ in lower region

13 15 near midrib, 18-20 near margin

Broad penta to hexa to polygonal, uniform throughout, arranged almost parallel to each other Uniform throughout, oblongly hexagonal to polygonal, longer than broad

35-40

0.29

20-25

0.87

large number of non-glossopterid plant fossils from Handapa area, namely Cyclodendron leslii, Phyllo-

theca indica, Lelstotheca robusta, Sphenophyllum churulianum, S. crenulatum, Trizygia speciosa, Raniganjia bengalensis, R. etheridgei, Schizoneura gondwanensis, Pseudoctenis balli, Senia reticulata, Neomariopteris polymorpha, N. hughesi, N. khanii (Maithy, 1977), Damudopteris bengalensis, Dizeuogotheca phegopteroides and erected a supposedly fern genus Pantopteris gracilis. In recent years new seed bearing glossopteridean fructifications Utkalia dichotoma and Khania dhenkanalensis were instituted (Chandra, 1984; Chandra and Singh, 1988). Some new genera and species were also erected from the same beds, namely Sphenophyllurn utkalensis, Asansolia cf. phegoptero-

5 6 near midrib, 12 13 near margin

2-3 between midrib and margin, very broad, large near midrib, becoming smaller, narrower towards margin, hexagonal to polygonal

ides (Pant et al., 1985), Surangephyllum elongatum (Chandra and Singh, 1986) and a ginkgoalean leaf Handapaphyllum indicum (Chandra and Singh, 1989). Recently Singh and Chandra (1987) studied the Handapa flora and erected seven new species of Glossopteris, namely G. kamthiensis, G. maheshwarii, G. acuminata, G. utkalensis, G. hinjridaensis, G. inaequalis, and G. dhenkanalensis. In the present study, 34 species of Glossopteris have been studied in detail for the first time from Handapa beds. The total number of Glossopteris species in the Handapa flora is 42 taking into consideration seven new species (Singh and Chandra, 1987) and a finding of G. browniana by Khan (1969). So far 87 species of Glossopteris are recog-

194

S. C H A N D R A A N D K.J. S I N G H

TABLE

II

Stratigraphic

range

of

Glossopteris

species

in different

formations

of Lower

Gondwana

in Peninsular

India

(including

the

Handapa

species)

SN.

Glossopteris

species

Talchir

Karharbari

Barakar

Barren

Raniganj

Kamthi

Panchet

Measures

1. 2. 3. 4. 5. 6. 7. 8. 9. I0. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. 51.

Glossopteris browniana B r o n g n i a r t , 1828 G. angust(folia B r o n g n i a r t , 1828 G. linearis M c C o y , 1 8 4 7 G. ampla D a n a , 1 8 4 9 G. leptoneura B u n b u r y , 1 8 6 1 G. musaefolia B u n b u r y , 1 8 6 1 G. stricta B u n b u r y , 1 8 6 1 G. indica S c h i m p e r , 1 8 6 9 G. taeniopteroides F e i s t m a n t e l , 1 8 7 8 G. communis F e i s t m a n t e l , 1879 G. decipiens F e i s t m a n t e l , 1 8 7 9 G. Iongicaulis F e i s t m a n t e l , 1879 G. intermedia F e i s t m a n t e l , 1 8 8 0 G. damudica F e i s t m a n t e l , 1 8 8 1 G. divergens F e i s t m a n t e l , 1 8 8 1 G. formosa F e i s t m a n t e l , 1881 G. stenoneura F e i s t m a n t e l , 1 8 8 1 G. conspicua F e i s t m a n t e l , 1 8 8 1 G. retifera F e i s t m a n t e l , 1881 G. orbicularis F e i s t m a n t e l , 1881 G. intermittens F e i s t m a n t e l , 1 8 8 1 G. taenioides F e i s t m a n t e l , 1 8 8 2 G. tortuosa Z e i l l e r , 1 9 0 2 G. arberi S r i v a s t a v a , 1 9 5 6 G. sahnii S r i v a s t a v a , 1 9 5 6 G. srivastavae S u r a n g e a n d M a h e s h w a r i , G.Jeistmantelii R i g b y , 1 9 6 4 b G. euryneura M a h e s h w a r i , 1965 G. emarginata M a h e s h w a r i and Prakash, G. retusa M a h e s h w a r i , 1965 G. zeilleri P a n t a n d G u p t a , 1 9 6 8 G. vulgaris P a n t a n d G u p t a , 1 9 6 8 G. tenu~folia P a n t a n d G u p t a , 1 9 6 8 G. brongniartii P a n t a n d G u p t a , 1 9 6 8 G. varia P a n t a n d G u p t a , 1 9 6 8 G. senii S r i v a s t a v a , 1 9 6 9 G. papillosa S r i v a s t a v a , 1 9 6 9 G. angusta P a n t a n d G u p t a , 1 9 7 1 G. pandurata P a n t a n d G u p t a , 1 9 7 1 G. giridihensis P a n t a n d G u p t a , 1 9 7 1 G. karanpurensis K u l k a r n i , 1 9 7 1 G. barakarensis K u l k a r n i , 1 9 7 1 G. lanceolatus P a n t a n d S i n g h , 1 9 7 1 G. rhabdotaenioides P a n t a n d S i n g h , 1 9 7 1 G. nautiyalii P a n t a n d S i n g h , 1 9 7 1 G. gondwanensis P a n t a n d G u p t a , 1 9 7 1 G. subtilis P a n t a n d G u p t a , 1 9 7 1 G. cord(lbrmis P a n t a n d S i n g h , 1 9 7 1 G. spatulata P a n t a n d S i n g h , 1 9 7 1 G. gigas P a n t a n d S i n g h , 1 9 7 1 G. nidpurensis S r i v a s t a v a , 1 9 7 1

-

+

+

+

+

+

+

-

+

+

+

+

+

+

-

+

-

+

+

+

-

-

+

-

+ -

+

-

-

-

+

-

+

+

+

-

+

+

+

+

+

+

-

-

+ +

+

+

+

-

+

+

+

+

-

+

-

+

-

+

+

-

-

+

-

+

+

-

-

+

-

+

-

-

-

+

+

+ +

+

-

+

-

-

+

+

+

+

+

+

-

+

+

+

+

+

+

+

+

+

+

+

-

+ -

-

+ -

+

-

-

+

-

+

-

-

-

-

+

+

-

-

+

+

-

-

-

+

-

-

-

+

-

+

-

-

+

-

-

1962 -

-

-

1965

+

+ +

-

-

+ +

+

-

-

+

.

.

.

.

+

-

-

+

-

.

+

-

.

.

.

-

-

+

+

+

+

-

+

-

+

.

.

.

.

.

.

.

.

.

.

-

+

-

+

+

-

+

.

-

+

-

-

+

+ -

-

+ +

.

.

-

-

+ +

-

.

+

-

-

-

-

+

+

+

.

.

.

.

+

-

.

.

.

.

+

+

.

.

.

.

+

+

-

-

+

+

-

-

+

-

+

+

+

+

.

.

.

.

.

.

.

.

.

-

-

+

195

T H E G E N U S G L O S S O P T E R I S F R O M T H E LATE P E R M I A N BEDS O F H A N D A P A , I N D I A

TABLE II (continued) SN. Glossopteris species

Talchir Karharbari

G. tenuinervis Pant and Gupta, 1971 G. major Pant and Singh, 1 9 7 1 G. radiata Pant and Singh, 1971 G. obscura Pant and Singh, 1971 G. recurva Pant and Singh, 1974 G. sastrii Pant and Singh, 1 9 7 4 G. maculata Pant and Singh, 1 9 7 4 G. oldhamii Pant and Singh, 1 9 7 4 G. searsolensis Pant and Singh, 1 9 7 4 G. gopadensis Banerji, Maheshwari and Bose, 1976 G. churiensis Srivastava, 197;/ G. talchirensis Chandra and Surange, 1979 G. taeniensis Chandra and Surange, 1979 G. karharbariensis Chandra and Surange, 1979 G. nimishea Chandra and Surange, 1979 G. syaldiensis Chandra and Surange, 1979 G. saksenae Chandra and Surange, 1979 G. nakkarea Chandra and Surange, 1979 G. mohudaensis Chandra and Surange, 1979 G. rewaensis Chandra and Surange, 1979 G. bosei Chandra and Surange, 1979 G. pantii Chandra and Surange, 1 9 7 9 G. raniganjensis Chandra and Surange, 1979 G. ornatus Chandra and Srivastava, 1981 G. surangei Chandra and Prasad, 1981 G. venustus Chandra and Prasad, 1981 G. shailae Bajpai, 1986 G. kamthiensis Singh and Chandra, 1 9 8 7 G. maheswarii Singh and Chandra, 1987 G. acuminata Singh and Chandra, 1987 G. utkalensis Singh and Chandra, 1987 G. hinjridaensis Singh and Chandra, 1987 G. inaequalis Singh and Chandra, 1987 G. dhenkanalensis Singh and Chandra, 1987 G. nilssonioides Pant and Pant, 1987 G. sidhiensis Pant and Pant, 1 9 8 7

52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 80. 81. 82. 83. 84. 85. 86. 87.

T O TA L

n i s e d ( T a b l e I I ) in d i f f e r e n t G o n d w a n a

.

.

.

Barakar

Barren Measures

.

+ . . . .

. . . .

. . . .

+ +

-

-

+

-

+

-

+ + + +

.

-

formations

Panchet

+ +

.

.

.

.

.

+ + + + .

-

. + + -

-

+

+ -

+ +

-

+ + + + + + + + + + -

54

47

+

. -

-

.

. 27

16

. 10

Glossopteris a r e k n o w n

+

.

+ + + + +

. T

+

.

-

. -

+ + + + + + + + -

. -

Kamthi

-

. . . .

+ -

Raniganj

-

-

-

+ + 2~

from the Triassic of India

o f p e n i n s u l a r I n d i a ( T a b l e V ) . O f t h e s e 87 s p e c i e s ,

( T a b l e I I ) , o f w h i c h 11 s p e c i e s , ( = 5 0 % ) a r e c o m -

42

m o n in t h e T r i a s s i c a n d t h e H a n d a p a

are

known

from

the

Handapa

area

(Table

III). A s is a p p a r e n t 52%)

from

o f Glossopteris

Table are

I I I , 28 s p e c i e s ( =

common

Raniganj

Formation

and

Eighteen

species

Glossopteris

present

Handapa

Kamthi

Formation,

occur

area (Table

III).

both

Handapa

in

of

both

in the Handapa

of which Kamthi

the

flora.

(excluding

species) are reported the

in

from

the the

13 s p e c i e s ( = 7 2 % ) Formation

area (Table III). Twenty

and

the

two species of

T a b l e I V s h o w s t h e r e l a t i v e p e r c e n t a g e o f Gloss-

opteris s p e c i e s in H a n d a p a angustifolia a p p e a r common From have

b e d s , G. indica a n d G.

to be the most dominant

and

species. T a b l e I I , it is e v i d e n t t h a t

the maximum

species known

number

the Kamthis

( 4 7 ) o f Glossopteris

from the Raniganj

Formation

(54)

along with some endemic species exclusively known

196

s. CHANDRA AND K.J. SINGH

TABLE

III

Comparison

of

Glossopteris s p e c i e s f r o m H a n d a p a

area with the

Glossopteris s p e c i e s f r o m t h e R a n i g a n j , K a m t h i

and Panchet

Formations SN.

1. 2. 3. 4. 5. 6. 7. 8. 9. 10. I1. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42.

Handapa

area

Raniganj

Glossopteris browniana B r o n g n i a r t , 1828 + G. angustifolia B r o n g n i a r t , 1828 + G. stricta B u n b u r y , 1861 + G. leptoneura B u n b u r y , 1861 + G. indica S c h i m p e r , 1869 + G. communis F e i s t m a n t e l , 1879 + G. retifera F e i s t m a n t e l , 1881 + G. conspicua F e i s t m a n t e l , 1881 + G. cf. G. divergens F e i s t m a n t e l , 1881 + G. damudica F e i s t m a n t e l , 1881 + G. stenoneura F e i s t m a n t e l , 1881 + G. tortuosa Z e i l l e r , 1902 + G. arberi S r i v a s t a v a , 1956 + G. vulgaris P a n t a n d G u p t a , 1968 + G. tenuifolia P a n t a n d G u p t a , 1968 + G. varia P a n t a n d G u p t a , 1968 + G. "-eilleri P a n t a n d G u p t a , 1968 G. subtilis P a n t a n d G u p t a , 197l + G. spatulata P a n t a n d S i n g h , 1971 + G. lanceolatus P a n t a n d S i n g h , 1971 + G. gondwanensis P a n t a n d G u p t a , 1971 + G. gigas P a n t a n d S i n g h , 1971 + G. radiata P a n t a n d S i n g h , 1971 + G. barakarensis K u l k a r n i , [971 G. pandurata P a n t a n d G u p t a , 1971 G. angusta P a n t a n d G u p t a , 1971 G. tenuinervis P a n t a n d G u p t a , 1971 + G. nautiyalii P a n t a n d S i n g h , 1971 + G. obscura P a n t a n d S i n g h , 1971 + G. sastrii P a n t a n d S i n g h , 1974 + G. oldhamii P a n t a n d S i n g h , 1974 + G. gopadensis B a n e r j i , M a h e s h w a r i a n d B o s e , 1976 G. taeniensis C h a n d r a a n d S u r a n g e , 1979 G. nimishea C h a n d r a a n d S u r a n g e , 1979 G. mohudaensis C h a n d r a a n d S u r a n g e , 1979 + G. kamthiensis S i n g h a n d C h a n d r a , 1987 G. maheshwarii S i n g h a n d C h a n d r a , 1987 G. acuminata S i n g h a n d C h a n d r a , 1987 G. utkalensis S i n g h a n d C h a n d r a , 1987 G. hinjridaensis S i n g h a n d C h a n d r a , 1987 G. inaequalis S i n g h a n d C h a n d r a , 1987 G. dhenkanalensis S i n g h a n d C h a n d r a , 1987 -

TOTAL

28

from the Kamthi beds. Quite a large number of Glossopteris species are common both in the Handapa and the Raniganj Formation (28 species, see Table III) in general. The form genus Glossopteris reached its greatest species diversification during the deposition of the

Formation

Kamthi +

Formation

Panchet Formation/Triassic +

+

+

+

+

+

+

+

+

+

+

+

+

+

+

-

-

+

+

-

+

-

-

+ + -

-

-

-

-

-

-

-

+

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

+

-

-

-

-

+

-

-

-

-

-

-

-

13

lI

Raniganj Formation (Late Permian) in Damodar Valley Coalfield. The large number of Glossopteris species (42) recovered from the Kamthi Formation of the Handapa area indicates that various species of Glossopteris grew luxuriantly even during Kamthi time in Mabanadi Basin (see Table V).

197

T H E G E N U S G L O S S O P T E R I S F R O M T H E LATE P E R M I A N BEDS O F H A N D A P A , I N D I A

TABLE IV Relative percentage of various GIossopteris species from Handapa area in the collection SN.

Glossopteris species

Percentage

DOMINANT

1. 2.

Glossopteris indica Schimper, 1869 G. angustifolia Brongniart, 1828

18.90 10.17

SUB-DOMINANT

3. 4. 5. 6.

G. G. G. G.

tenuifolia Pant and Gupta, 1968 conspicua Feistmantel, 1881 subtilis Pant and Gupta, 1971 communis Feistmantel, 1879

7.85 5.81 5.23 5.23

G. G. G. G. G. G. G. G. G. G. G. G. G. G. G. G. G. G.

gigas Pant and Singh, 1971 lanceolatus Pant and Singh, 1971 nimishea Chandra and Surange, 1979 retfera Feistmantel, 1881 gondwanensis Pant and Gupta, 1971 spatulata Pant and Singh, 1971 inaequalis Singh and Chandra, 1987 tenuinervis Pant and Gupta, 1971 obscura Pant and Singh, 1971 tortuosa zeiller, 1902 stenoneura Feistmantel, 1881 hinjridaensis Singh and Chandra, 1987 leptoneura Bunbury, 1861 angusta Pant and Gupta, 1971 nautiyalii Pant and Singh, 1971 gopadensis Banerji, Maheshwari and Bose, 1976 taeniensis Chandra and Surange, 1979 mohudaensis Chandra and Surange, 1979

4.94 4.36 4.36 3.78 3.49 2.62 2.62 2.32 2.32 2.03 1.74 1.45 1.16 1.16 1.16 1.16 I. 16 1.16

RARE

7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24.

EXTREMELY RARE

25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41.

G. G. G. G. G. G. G. G. G. G. G. G. G. G. G. G. G.

stricta Bunbury, 1861 arberi Srivastava, 1956 varia Pant and G u p t a , 1968 zeilleri Pant and Gupta, 1968 barakarensis Kulkarni, 1971 pandurata Pant and Gupta, 1971 sastrii Pant and Singh, 1974 utkalensis Singh and Chandra, 1987 oldhamii Pant and Singh, 1974 maheshwarii Singh and Chandra, 1987 cf. G. divergens Feistmantel, 1881 damudica FeistmanteL 1881 vulgaris Pant and Gupta, 1968 radiata Pant and Singh, 1971 kamthiensis Singh and Chandra, 1987 acuminata Singh and Chandra, 1987 dhenkanalensis Singh and Chandra, 1987

0.87 0.87 0.87 0.87 0.87 0.87 0.87 0.87 0.58 0.58 0.29 0.29 0.29 0.29 0.29 0.29 0.29

Total number of specimens studied = 344. Dominant = >/10% (more than 34 specimens); Sub-dominant - 5-10% (17-34 specimens): R a r e = 1 5% (4 16 specimens); Extremely r a r e = 0 1% (1-3 specimens).

198

s. CHANDRA AND K.J. SINGH

TABLE V Lower G o n d w a n a Formations of India Permian standard scale

Lithostratigraphic units Damodar valley

Satpura basin

Son valley

Rajmahal region

Pali Fm.

Raniganj Fm.

Tatarian

Raniganj Fm.

Bijori Fm.

Kazanian

Kulti Fm.

M o t u r Fm.

Artinskian Upper Lower

Barakar Fm. Karharbari Fm.

Barakar Fm. Karharbari Fm.

Barakar Fm. Karharbari Fm.

Sakmarian

Talchir Fm.

Talchir Fro.

Talchir Fm.

It is apparent that in the Handapa beds Glossopteris species occur in greater numbers than ever reported from any other Gondwana locality. The Handapa beds thus represent a single locality in which such a prolific number of species of one particular genus has grown. The question arises whether it is possible in nature for such a large number of species to grow in a single forest. Is it possible that the number of species is far less and that we are in fact dealing with variants and not the natural species? The basic leaf structure with its venation pattern of Glossopteris is remarkably conservative whereas the fructifications are highly variable indicating that we are indeed dealing with more genera of leaves of the same genus Glossopteris. A plausible explanation could be that the form genus Glossopteris collectively represents a much higher taxon comprising several families and orders. This can be further explained by the fact that there is a variety of fructifications assignable to GIossopteris reported from all over the Gondwana continent. From time to time several authors suggested to raise the status of Glossopteris to a gymnosperm class with separate orders and families. At present it is not possible tO classify Glossopteris leaves on the basis of their variable fructifications as they are generally found in association and not in actual attachment with the parent plant. The present state of our knowledge clearly indicates that the assorted leaves of Glossopteris represent a class equivalent to angiosperm leaves and not just a form genus. However, a precise classification is not yet possible and we will have

WardhaGodavari valley

Mahanadi valley

Kamthi Fm.

Kamthi Fm.

Kulti Fm.

Kulti Fm.

Barakar Fro.

Barakar Fm.

Barakar Fm. Karharbari Fm.

Talchir Fm.

Talchir Fm.

Talchir Fro.

to wait for more data to take any logistic step. Till then we will have to depend upon this very artificial classification system. The Handapa flora, in general, resembles the Raniganj flora of equivalent age deposited in separate basins during the same time period. So far, no typical Triassic forms have been reported by any of the previous workers. However, Subramanian and Rao (1960) reported typical Raniganj forms from the Handapa beds but assigned a Panchet age equivalent to the Mahadevas. Khan (1969), while describing the new cycadalean plant Senia reticulata from the same locality, assigned it to the Raniganj rocks of the Talchir Coalfield. Surange and Maheshwari (1970) in a description of some new fructifications have put their specimens under "?Kamthi" but did not discuss it further. However, later Surange and Chandra (1973a, b, c, 1974a, b, c, d) always assigned these beds to a Raniganj-age. Chandra and Rigby (1981) on the basis of the similarity in the lithology of the rocks and the flora with that described by Bunbury (1861) appropriately placed these beds on the Kamthi Formation equivalent to the Raniganj Formation age. According to the latest Lexicon of Stratigraphy' published by the Geological Survey of India (1977) the Late Permian beds of the Mahanadi Basin are referred to as Kamthi. Previously the same beds of the Mahanadi Basin were referred to as Hingir Formation but later this name was considered as obsolete. The noncarbonaceous Kamthis of the Wardha-Godavari graben are treated as equivalent in age to the

THE GENUS G L O S S O P T E R I S FROM THE LATE PERMIAN BEDS OF HANDAPA, INDIA

PLATE I

(for description see p. 203)

199

200

S. C H A N D R A A N D K.J. S I N G H

PLATE

II

2

3 (for description see p. 203)

4

5

THE GENUS GLOSSOPTERIS FROM THE LATE PERMIAN BEDS OF HANDAPA, INDIA

PLATE

III

(for description see p. 203)

201

202

S. C H A N D R A A N D K.J. S I N G H

P L A T E IV

4 6

2

3

I

5

7

T H E G E N U S G L O S S O P T E R I S F R O M T H E LATE P E R M I A N BEDS O F H A N D A P A . I N D I A

203

PLATE I (see p. 199)

1. GIossopteris angust![olia Brongniart. Specimen showing a complete leaf. 36662. x 1. 2. GIossopteris leptoneura Bunbury. Specimen showing almost a complete, narrow-linear leaf with long base and very thin secondary veins. 36664. x 1.

3. Glossopteris angust([~)lia Brongniart. Specimen showing a complete, small and narrow leaf enlarged to show the details of venation. 36661. x 2.

4. Glossopteris leptoneura Bunbury. Almost a complete leaf enlarged to show the sharp apical portion and very dense venation pattern. 36665. x 2.

5. Glossopteris communis Feistmantel. Specimen showing almost a complete leaf with very dense venation. 36668. x 3/4.

PLATE II (see p. 200)

1. Glossopteris angusti[olia Brongniart. Almost a complete leaf showing the details of venation pattern. 36662. x I. 2. Glossopteris leptoneura Bunbury. Almost a complete leaf enlarged to show the sharp apical portion and very dense venation pattern. 36665. x 2.

3. GIossopteris leptoneura Bunbury. Specimen showing almost a complete, narrow-linear leaf with long base and very thin secondary veins. 36664. x 1.

4. GIossopteris angusti/blia Brongniart. Almost a complete, very small and narrow leaf enlarged to show the details of venation pattern. 36661. x 2.

5. GIossopteris communis Feistmantel. Specimen showing almost a complete leaf with very dense venation. 36668. x 3/4.

PLATE III (see p. 201)

1. GIossopteris spatulata Pant and Singh. Specimen showing almost a complete spatulate leaf with thick midrib and very thin secondary veins. 36687. x 1.

2. GIossopteris stenoneura Feistmantel. Specimen showing almost a complete leaf, midrib ultimately dissolving into lateral veins in the apical portion. 36675. x 1.

3. GIossopteris gi~as Pant and Singh. Specimen showing thick and prominent midrib, uniform fine secondary veins forming very dense venation pattern. 36693. x 3/4.

PLATE IV

1. GIossopteris tenui[olia Pant and Gupta. Specimen showing a complete leaf with persistant midrib and very fine lateral veins forming dense venation pattern. 36680. x I.

2. GIossopteris vul~aris Pant and Gupta. Specimen showing a complete, strap shape, linear-oblong leaf with prominent midrib and dense venation pattern. 36679. x 1.

3. GIossopteris varia Pant and Gupta. Specimen showing almost a complete lens-shaped leaf with thin secondary veins and narrowoblong meshes of uniform size. 36682. x 1.

4. GIossopteris stenoneura Feistmantel. Specimen showing almost a complete leaf, midrib ultimately dissolving into lateral veins in the apical portion and dense venation pattern. 36675. x 1.

5. Glossopteris zeilleri Pant and Gupta. Specimen showing a small complete leaf having long base, evanescent midrib, very long and narrow meshes and dense venation pattern. 36683. x 2.

6. Glossopteris .~patulata Pant and Singh. Specimen showing almost a complete spatulate leaf with thick midrib and very thin secondary veins. 36687. x 1.

7. GIossopteris angusta Pant and Gupta. Almost a complete oblanceolate leaf showing bending of the lateral veins towards the midrib in the emarginate apical region and very dense venation pattern. 36698. x 1.

204

S. C H A N D R A A N D K.J. S [ N G H

PLATE V

1. GIossopterZ~" tenu(l~dia Pant and Gupta. Specimen showing a complete leaf and very fine secondary veins, forming dense venation pattern. 36680. x 1.

2. GIossopteris tenu(folia Pant and Gupta. Specimen showing a complete leaf having acute apex, persistent midrib and fine secondary nervation. 36681. x 1.

3. Glossopteris gopadensis Banerji, Maheshwari and Bose. Specimen showing almost a complete leaf (although its lamina is broken all along its margin) having wide base, thick midrib and dense venation pattern. 36705. x 1.5.

4. GIossopteris stricta Bunbury. Specimen showing a complete, narrow-elliptical leaf with almost horizontal dense venation pattern. 36663. x 1. PLATE Vl

1. Glossopteris gigas Pant and Singh. Specimen showing thick and prominent midrib, uniform fine secondary veins forming very dense venation pattern. 36693. x 3/4.

2. GIossopteris stricta Bunbury. Specimen showing a complete, narrow, elliptical leaf with almost horizontally running secondary veins forming broad meshes near the midrib and very narrow at the margin. 36663. x 1.

3. GIossapteris pandurata Pant and Gupta. A complete narrow-obovate leaf showing round or flattened apex with long base and very thin secondary veins. 36696. x 1

205

T H E G E N U S G L O S S O P T E R I S F R O M T H E LATE P E R M I A N BEDS O F H A N D A P A . I N D I A

P L A T E VI

2

3

206

.

S. C H A N D R A

A N D K.J. S I N G H

PLATE VII

i'

,

i

"

]

~ Jl"'

1

2

3

1. Glossopteris gopadensis Banerji, Maheshwari and Bose. Specimen showing almost a complete leaf having wide base, thick midrib and very dense venation. 36705. x 2. 2. Glossopteris indica Schimper. A complete petiolate leaf showing details of venation pattern. 36667. × 3/4. 3. Glossopteris arberi Srivastava. Specimen showing thick midrib and uniform venation pattern. 36678. x 1.

207

T H E G E N U S G L O S S O P T E R I S F R O M T H E LATE P E R M I A N BEDS O F H A N D A P A . I N D I A

PLATE VIII i

•

....

~,

3

1. Glossopteris indica Schimper. Specimen showing a complete leaf having a petiole. 36667. x 3/4. 2. Glossopteris arberi Srivastava. Specimen showing almost a complete leaf having a thick midrib and uniform venation pattern. 36678. x I. 3. Glossopteris subtilis Pant and Gupta. Specimen showing a complete oblanceolate leaf having petiole and compact venation. 36684. x 1.

208

S C H A N D R A A N D K.J. S I N G H

PLATE IX

5

3

2 (for description see p. 213)

4

6

THE GENUS G L O S S O P T E R I S FROM THE LATE PERMIAN BEDS OF HANDAPA+ INDIA

PLATE X

( f o r d e s c r i p t i o n see p. 2 1 3 )

209

210

s. CHANDRA AND K,J. SINGH

PLATE XI

/\

1

2 (for d e s c r i p t i o n see p. 213)

3

5

THE GENUS GLOSSOPTERIS FROM THE LATE PERMIAN BEDS OF HANDAPA, INDIA

PLATE XII

(for description see p. 213)

211

212

S. C H A N D R A A N D K.J. S I N G H

PLATE XIII

2

4

I 1

3

THE GENUSGLOSSOPTERIS FROMTHE LATEPERMIANBEDSOF HANDAPA,INDIA

213

PLATE IX (see p. 208)

I. Glossopteris subtilis Pant and Gupta. Specimen showing an elliptical leaf with very fine secondary veins and dense venation pattern. 36685. x 1. 2. Glossopteris tenuinervis Pant and Gupta. Specimen showing a narrow-oblong leaf having thick midrib and dense venation pattern. 36697. × 1. 3. GIossopteris lanceolatus Pant and Singh. Specimen showing a lanceolate leaf having thin secondary veins, running almost horizontally to the margins and forming very narrow meshes. 36688. x 1. 4. GIossopteris barakarensis Kulkarni. Almost a complete elliptic leaf showing many small and uniform meshes arranged in a line from midrib to the margin. 36695. x 1. 5. Glossopteris obscura Pant and Singh. Specimen showing a complete petiolate elliptic leaf having thin lateral veins. 36701 x 1. 6. GIossopteris nautiyalii Pant and Singh. Specimen showing very sharp apical portion of the leaf and the dense venation pattern. 36700. x 1.

PLATE X (see p. 209)

I. GIossopteris barakarensis Kulkarni. Specimen showing almost a complete elliptic leaf having a thick midrib. 36695. x 1.5. 2. GIossopteris tenuinervis Pant and Gupta. Specimen showing an oblong leaf having thick midrib and dense venation pattern. 36699. x 1.5. 3. GIossopteris nimishea Chandra and Surange. Specimen showing almost a complete narrow-elliptic leaf having long petiole and the secondary veins running almost horizontally to the margins. 36709. x 1. 4. GIossopteris cf. G. divergens Feistmantei. Specimen showing the half preserved upper portion of the leaf with lateral veins diverging upwardly. 36673. x 2. PLATE XI (see p. 210)

1. GIossopteris oldhamii Pant and Singh. Almost a complete narrow-obovate leaf showing lateral veins running horizontally to the margins. 36704. x 1. 2. Glossopteris taeniensis Chandra and Surange. Almost a complete oblanceolate and small leaf having retuse apex and open venation pattern. 36707. x 1. 3. Glossopteris radiata Pant and Singh. Basal portion of the leaf showing thick midrib and secondary veins radiating in the upward, horizontal and in the downward directions from the midrib. 36694. × I. 4. Glossopteris mohudaensis Chandra and Surange. Specimen showing almost a complete leaf with thick midrib and dense venation pattern. 36711. × 1. 5. GIossopteris cf. G. divergens Feistmantel. Specimen showing the half preserved upper portion of the leaf having emarginate apex and the lateral veins diverge upwardly in the apical region with broad and elongate meshes. 36673. × 1. PLATE XII (see p. 211)

1. Glossopteris damudica Feistmantel. Specimen showing almost a complete elliptic leaf having small petiole and almost horizontally running lateral veins. 36674. x I. 2. Glossopteris radiata Pant and Singh. Basal portion of the leaf showing thick midrib and secondary veins radiating in the upward, horizontal and in the downward directions from both the sides of the midrib. 36694. x 2. 3. Glossopteris tortuosa Zeiller. Specimen showing almost a complete leaf having prominent midrib and zigzag type of venation. 36676. x I. 4. Glossopteris gondwanensis Pant and Gupta. Specimen showing 3/4 preserved leaf having sharp apical lamina and open venation pattern. 3669[. x 1.

PLATE XIII

1. Glossopter& damudica Feistmantel. Specimen showing almost a complete elliptic leaf having small petiole and almost horizontally running lateral veins. 36674. × I. 2. Glossopteris gondwanensis Pant and Gupta. Specimen showing emergence of the secondary veins at acute angles and open venation pattern. 36691. x 1. 3. Glossopteris sastrii Pant and Singh. Specimen showing a complete, oblong, petiolate leaf having open venation with broad and large polygonal meshes near the midrib. 36703. x 1. 4. Glossopteris tortuosa Zeiller. Half preserved specimen showing a thick midrib, wide base, trapezoidal meshes and sinuous/zigzag type of venation pattern. 36677. × 2.

214

S. C H A N D R A A N D K.J. S [ N G H

PLATE XIV

I. Glossopteris sastrii Pant and Singh. Specimen showing a complete oblong leaf having petiole, open venation with broad and large polygonal meshes near the midrib. 36703. x 2. 2. Glossopteris conspicua Feistmantel. Specimen showing thick midrib and long and broad meshes. 36672. x 1. 3. Glossopteris tortuosa Zeiller. Half preserved specimen showing a thick midrib, wide base and the lateral veins running horizontally to the margins. 36677. × 2.

215

T H E G E N U S G L O S S O P T E R I S F R O M I-HE LATE P E R M I A N BEDS O F H A N D A P A , I N D l A

PLATE XV

,~\'iJ~ Yt~/

........... i?

2

3 I. Glossopteris nimishea Chandra and Surange. Specimen showing a complete narrow-elliptic leaf having long petiole and the secondary veins running almost horizontally to the margins. 36709. x 1.

2. Glossopteris retit~'ra Feistmantel. Specimen showing a complete leaf and open venation pattern. 36670. x 1. 3. Glossopteris conspicua Feistmantel. An incomplete specimen showing long and broad meshes. 36671. x 1.

216

S. CHANDRA AND K.J. SINGH

f_

I ~

J

"---_ 1

t 9

c~r..,,

10

Fig. 2. Reconstruction of vegetation at H a n d a p a during Late Permian time depicting the following plants: 1 = Glossopteris; 2 = Neomariopteris: 3 = Schizoneura; 4 = Phyllotheeu; 5 = Di=eugotheca, 6 = Raniganl'ia 7 = Trizygia; 8 = Sphenophyllum; 9 = W o o d e n log; 10 = Damudopteris; 11 = Pseudoctenis.

T H E G E N U S GLOSSOPTERIS F R O M T H E LATE P E R M I A N BEDS O F H A N D A P A , I N D I A

Raniganj Formation of the Damodar Valley which is a major coal-bearing horizon. Most of the vegetation of the Kamthi-time forests is similar to the Raniganj forests. Unlike the Raniganj forests the Kamthi forests were perhaps growing on uplands. The climate was warm and humid and thus very suitable for luxuriant plant growth. The red bed facies of the ferruginous sandstones indicate seasonal variability of dry spells. This could be one of the reasons for the lack of coal in the Kamthis inspite of enormous vegetal matter. The plant fossil assemblage at Handapa indicates a dense forest type vegetation dominated by arborescent trees bearing Glossopteris foliage with few other plants belonging to lycopodiales, articulates, sphenophyllales, filicales and some other gymnosperms. It is widely believed that Glossopteris plants were deciduous and this is confirmed by the deposition of several hundred leaves separated vertically by thin layers of sediments representing annual leaf fall. The deposition site of the vegetal matter was not very far from the original growing site which is shown by the fact that most of the large sized Glossopteris leaves are preserved quite complete without much distortion. Also the deposition took place in quiet and still water, perhaps in a lake, because of the very fine grained nature of the sediment and the common occurrence of leaves in seasonal layers. The leaves are preserved in a haphazard manner in the sediment and no definite direction is indicated by the tips of the leaves. Some of the species were huge trees, while others were shrubs of varying sizes. There must have been some small open patches where small pteridophytic plants could grow. A vegetational reconstruction during Late Permian time around Handapa area has been attempted on the basis of the plant fossils (Fig. 2).

Acknowledgements The authors are thankful to Mr. John F. Rigby, Geological Survey of Queensland, Brisbane, Australia, for useful discussions and valuable suggestions. They are also grateful to Drs H.J. Schweitzer and J. Galtier for acting as referees and for suggesting many alterations to improve the final manu-

217

script. Thanks are also due to Mrs. Sunita Khanna, Mr. Avinesh Kumar Srivastava and Mr. D.K. Tiwari for helping us to produce this manuscript and other works on computer.

References Agashe, S.N., Gowda, P.R.N., Suresh, F.C. and Geetha, K.R., 1984. Recent advances in the palaeobotanical studies on Lower Gondwana Strata of Chandrapur District, Maharashtra. In: A.K. Sharma, G.C. Mitra and M. Banerjee (Editors), Proc. Symp. Evolutionary Botany and Biostratigraphy, Calcutta, 1979. A.K. Ghosh Commem. Vol. Curt. Trends Life Sci., 10:369 382. Ball, V., 1877. On the geology of the Mahanadi Basin and its vicinity. Rec. Geol. Surv. India, 10(4): 167-185. Banerji, J., Maheshwari, H.K. and Bose, M.N., 1976. Some plant fossils from the Gopad River Section near Nidpur, Sidhi District, Madhya Pradesh. Palaeobotanist, 23(1): 59-71. Brongniart, A., 1828. Histoire des V~g6taux fossiles ou Recherches botaniques et geologiques sur les V~g~taux renferm6s dans les diverses couches du Globe. Dufour & d'Ocagne, Paris and Amsterdam, pp. 1-136. Bunbury, C.J.F., 1861. Notes on a collection of fossil plants from Nagpur, Central India. Q. J. Geol. Soc. London, 17: 325-346. Chandra, S., 1984. Utkalia diehotoma gen. et sp. nov.--A fossil frutification from the Kamthi Formation of Orissa, India. Palaeobotanist, 31(3): 208-212. Chandra S. and Prasad, M.N.V., 1981. Fossil plants from the Kamthi Formation of Maharashtra and their biostratigraphic significance. Palaeobotanist, 28/29:99 121. Chandra S. and Rigby, J.F., 1981. Lycopsid, sphenopsid and cycadaceous remains from the Lower Gondwana of Handappa, Orissa. Geophytology, 11(2): 214-219. Chandra S. and Rigby, J.F., 1983. The Filicales from the Lower Gondwanas of Handappa, Orissa. Palaeobotanist, 31(2): 143-147. Chandra S. and Singh, K.J., 1986. Surangephyllum gen. nov. from the Kamthi Formation of Handapa, Orissa. I.S.G. Bull., 1:15 18. Chandra S. and Singh, K.J., 1988. A new seed-bearing plant organ from the Kamthi Formation of Orissa, India. Curr. Sci., 57(18): 996 998. Chandra S. and Singh, K.J., 1989. Handapaphyllum--a new leaf type from the Upper Permian of Orissa, India. Palaeobotanist, 37(2): 143-146. Chandra S. and Surange, K.R., 1977. Fertile bracts and scales of Glossopteris fructifications from the Lower Gondwana of India. Palaeobotanist, 24(3): 195-201. Chandra S. and Surange, K.R., 1979. Revision of the Indian species of GIossopteris. Birbal Sahni Inst. Palaeobot., Monogr., 2, 301 pp. Feistmantel, O., 1879. The fossil flora of the Lower Gondwanas, 1. The flora of the Talchir Karharbari beds. Mem. Geol. Surv. India, Palaeontol. Indica Ser. 12, 3(1): 1-48. Feistmantek O., 1880. The fossil flora of the Gondwana System.

218 The flora of the Damuda and Panchet divisions. Mem. Geol. Surv. India, Palaeontol. Indica Set. 12, 3(2): I 77. Feistmantel, O., 1881. The fossil flora of the Gondwana System. The flora of the Damuda and Panchet divisions. Mem. Geol. Surv. India, Palaeontol. lndica Ser. 12, 3(3): 78-149. Hughes, T.W.H., 1877. The Wardha Valley Coalfield. Mem. Geol. Surv. India, 13(I): 1-154. Khan, A.M., 1969. Senia reticulata, a new plant fossil from the Raniganj rocks of the Talchir Coalfield, Orissa, India. In: H. Santapau, A.K. Ghosh, S.K. Roy, S. Chanda and S.K. Chaudhuri (Editors), J. Sen Memorial Volume. Bot. Soc. Bengal, Calcutta, pp. 335-338. Kulkarni, S., 1971. Glossopteris and Gangamopteris species from South Karanpura Coalfield. Palaeobotanist, 18(3): 297 304. Maithy, P.K., 1977. Three new fern fronds from the GIossopteris flora of India. Palaeobotanist, 24(2): 96-101. Oldham, R.D., 1880. Fossil plants from Kamthi Formation. Palaeontol. lndica Ser. 12, 3: 19. Oldham, R.D., 1893, A Manual of the Geology of India. Calcutta, 2nd ed., pp. 1-543. Pant, D.D. and Gupta, K.L., 1968. Cuticular structure of some Indian Lower Gondwana species of Glossopteris Brongniart, Part 1. Palaeontographica B, 124(1/3): 45 81. Pant, D.D. and Gupta, K.L., 1971. Cuticular structure of some Indian Lower Gondwana species of GIossopteris Brongniart, Part 2. Palaeontographica B, 132(1/4): 130-152. Pant, D.D. and Singh, K.B., 1971. Cuticular structure of some Indian Lower Gondwana species of Gh~ssopteris Brongniart, Part 3. Palaeontographica B, 135(1/2): 1-40. Pant, D.D. and Singh, R.S., 1974. On the stem and attachment of G/ossopteris and Gangamopteris leaves, Part II structural features. Palaeontographica B, 147(1/3): 42 73. Pant, D.D., Srivastava, P.C. and Das, P.K., 1985. Some new pteridophytic remains from the Lower Gondwana rocks of Hinjrida Ghati, Orissa. Curr. Sci., 54(2): 90 92. Sastry, M.V.A., Acharyya, S.K., Shah, S,C., Sutsangi, P.P., Ghosh. S.C., Raha, P.K., Singh. G. and Ghosh, R.N. (Editors). 1977. Stratigraphic Lexicon of Gondwana Formations of India. Geol. Surv. India, Misc. Publ., 36, pp. 1-170. Schimper, W.P,, 1869. Trait6 de Pal~ontologie V6g&al ou la Flore du Monde Primitif dans ses Rapports avec les Formations Geologiques et la Flore du Monde Actual, 1. Balliere, Paris, 740 pp.

S. C H A N D R A A N D K.J. S I N G H

Singh, K.J. and Chandra, S., 1987. Some new species of GIossopteris from the Kamthi Formation of Handapa, Orissa. Geophytology, 17(I): 39 55. Srivastava, P.N., 1956. Studies in the Glossopteris flora of India, 4. Glossopteris, Gangamopteris and Paloeovittaria from the Raniganj Coalfield. Palaeobotanist, 5(1): 1-45. Subramanian, K.S. and Rao. C.N., 1960. GIossopteris from the Mahadevas of Hinjrida, Talchir Coalfield, Orissa. Proc. 47th Indian Sci. Congr., Bombay, Part 3 (Abstr.): p. 278. Surange, K.R. and Chandra, S., 1973a. Dictyopteridium spori,[~,rum Feistmantel-female cone from the Lower Gondwana of India. Palaeobotanist, 20(1): 127-136. Surange, K.R. and Chandra, S., 1973b. Denkania imtica gen. et sp. nov. A glossopteridean fructification from the Lower Gondwana of India. Palaeobotanist, 20(2): 264-268. Surange, K.R. and Chandra, S., 1973c. Partha A new type of female fructification from the Lower Gondwana of India. Palaeobotanist, 20(3): 356-360. Surange, K.R. and Chandra, S., 1974a. Fructifications of Glossopteridae from India. Palaeobotanist, 21 ( 1): 1-17. Surange, K.R. and Chandra, S., 1974b. Lidgettonia mucronata sp. nov. A female fructification from the Lower Gondwana of India. Palaeobotanist, 21 ( 1): 121 - 126. Surange, K.R. and Chandra, S., 1974c. Further observations on Glossotheca Surange and Maheshwari: A male fructification of glossopteridales. Palaeobotanist, 21(2): 248-254. Surangc, K.R. and Chandra, S., 1974d. Some male fructifications of glossopteridales. Palaeobotanist, 21(2): 255 266. Surange, K.R. and Chandra, S., 1978. Morphology and affinities of G/ossopteris. Palaeobotanist, 25: 509-524. Surange, K.R. and Maheshwari, H.K., 1962. Studies in the Glossopteris flora of India, 11. Some observations on I~2,rtebraria from the Lower Gondwanas of India, Palaeobotanist, 9(1/2): 61-67. Surange, K.R. and Maheshwari, H.K., 1970. Some male and female fructifications of glossopteridales from India. Palaeontographica B, 129(4/6): 178 192. White, Mary E, 1978. Reproductive structures of the Glossopteridales in the plant fossil collection of the Australian Museum. Rec. Aust. Mus., 31(12): 473 505. Zeiller, R., 1902. Observations sur quelques plantes fossiles des Lower Gondwanas. Mem. Geol. Surv. India, Palaeontol. Indica, 2(1): 1 40.