FURTHER OBSERVATIONS ON INDIAN LOWER GONDWANA SPHENOPHYLLALES
P. Ie MAITHY Birbal Salmi Institute of Palaeobotany, Lucknow-226007, India
ABSTRACT
New morphological details and the possible habit of Trizygia speciosa Royle are described alongwith two new forms of Sphenophyllales, Paratrizygia rhodesii (Rigby) n. comb. and Parasphenophyllum crenulatum sp. novo from the Damuda Series of India.
INTRODUCTION has two distinct ridges, which do not alternate from one internode to the next, but are co'ntinuous. The internodal length T'RIZYGIAthe only SphenophyllalesSPECIOSA Royleknown(1839)fromis of the stem varies from 0·4-2·2 cm and the the Indian Lower Gondwana succes- width from 0·1-0·2 cm. The distance in sion. Recently, several specimens of Trizygia between two nodes show progressive increase speciosa have been collected from the Rani• in length from apical to basal end. In ganj Formation of Raniganj Coalfield. The cellulose pull of the stem, its substance study gives new information o'n the morpho• generally appears to be black or dark logy and the possible habit of plants. brown, but at some places it is light brown The same is detailed here along with the and translucent. At such points the stem record of two new forms of Sphenophyllales substance shows xylem elements with scalari• from the Lower Gondwana of India. form thickenings (Text-fig. 2A) or bordered The specimens are preserved in the form pits (Text-fig. 2B). Bordered pits are in of impressions or compressions. Observa• one row, occasionally in two rows, circular• tions. on epidermal structures and other ana• oval in shape with circular pore. They are tomical structures were possible by means of either contiguous or arranged separately. cellular pulls from compressions. For the Bordered pits have also been reported identification of Sphenophyllales the nomen• earlier by Pant 2nd Mehra, (1963), but it clature proposed by Asama (1970) has been is not definite if this is of secondary wood. followed. These elements are flanked on either side All the figured specimens in this paper by surrounding fibres. The stem surface are preserved at the Museum of Birbal shows elongate-rectangular epidermal cells, Sahni Institute of Pdaeobotany, Lucknow. which are placed end to end by their end walls. DESCRIPTION Leaves are arranged in whorls on nodes (PI. 1, fig. 1). Each whorl comprises Trizygia Royle, 1839 six leaves. The six leaves are arranged in three distinct pairs (Text-fig. 1A), i.e. Trizygia speciosa Royle, 1839 two bigger pairs and one small pair. Leaves PIs. 1, 2, figs. 1-9; Text-figs. 1-6 are attached ± at right angles to the axis by their cuneate base. Two bigger Synonymy- See Maheshwari, 1968, p. 283. pairs of leaves occupy the lateral position while the third smaller pair occupies the Two hundred speCImens of Trizygia central position in preserved condition. speciosa have been examined. All of them A leaf whorl encircles completely round are either incomplete at the apical or basal the node (Text-fig. 1B) and not one side ends, or both the ends. of the node as Feistmantel (1879, p. 166) The stems bearing the leaves are slender, believed them to be. The whorls are sprea• unbranched, articulated with slightly swollen ding horizontally. During fossilization the nodes (PI. 1, fig. 1). The internodal region lamina and the a,xis aYe so preserved tha.t ~66 MAITHY - OBSERVATiONS ON INDIAN LOWER GONDWANA SPHENOPHYLLALES 267
A o
c
TEXT-FIG. 1. - A, leaf whorl laterally preserved X l·S. B, leaf whorl dorsi-ventrally preserved X l·S. e, venation in the lateral leaf X 3. D, venation in the central small leaf X 3.
the two bigger pairs of leaves are arranged run ± straight. The angle of divergence in on the lateral sides and the smaller pair between two veins is 5°. Interconnection in the centre. The lateral pairs of leaves in between the veins are absent and the are ± double in size than the central pair. interstitial fibres too. Lateral leaves are ± asymmetrical in Epidermal structure of leaves have earlier outline, whereas the central leaves are been reported by Pant and Mehra (1963) symmetrical. Leaves of the lateral pairs and Maheshwari (see Surange, 1964). Epi• are elongate - ovate, apex obtuse, base dermal pulls of the leaves show sinuous cuneate and entire margin. The leaves of outlines in which some walls appear faint, central pair are ovate, apex obtuse, base while others are darker. In fact, the cells cuneate and entire margin. In all the arrangement as a whole indicates that leaves the inner margin are ± straight, in these pulls, the presence of two trans• while the outer margin has a strong cur• lucent layers of epidermis overlapping each vature. A progressive increase in the size other and both of them have sinuous• of leaves is well marked, in each successive walled cells, alongwith the straight-walled whorls from apical to basal end. Lateral rectangular cells of vein area. Epidermal leaves measure 0·5-2·6 cm in length while cells are 70-14-0 [J. in length and 15-40 [J. the central leaves measure 0·25-1·5 cm in in breadth. The cells of non-stomatiferous length (see Table 1). epidermis are longitudinally oriented and A single vein enters at the base of leaf tend to be in series parallel to the veins (Text-fig. Ie, D), which dichotomises into (Text-fig. 2D). The walls are seemingly less two and then into four veins. The outer sinuous. The cells of the other overlapping pair of veins divides only once before layer are irregularly ananged and their reaching to the margin. The inner pair walls are deeply sinuous (PI. 2, fig. 8; subsequently dichotomises 1, 2 or 3 times. Text-fig. 2E). The cells measure 60-80 [J. As a result of which 10-18 veinlets are at in size. The stoma.ta are haplocheilic having the apical margin of the lateral leaves and thickened lateral lamellae, measuring 18-20 [1. 10-12 veinlets in the central leaves. Veins in len~th. The sl.lbsidiary cens are like ~68 THE PALAEOBOTANIST
TEXT-FIG. 2. - A, tracheids of foliage shoot showing spiral thickenings X 500. B, tracheids of foliage shoot showing single row of bordered pits X 500. C, celts over the vein region near the basal portion of leaf X 400. D, celts of the vein cells and the non-stomatiferous surface of leaf X 400. E, celts of the stomatiferous surface X 400.
ordinary epidermal cells and the polar and ridges are continuous (PI. 1, fig. 3). subsidiary cells are not different from lateral Occasionally, the axes are branched. The ones. The basal most region of the leaf, width of the axes vary from 2-4 em. i.e. ± 2 mm from point of attachment with In some cases a distinct groove 1·5 to stem show distinct cellular structure from 2 mm broad is present in the middle (PI. 1, the laminar portion of leaf. Cells in this fig. 4). The internodal length varies from portion (PI. 2, fig. 7; Text-fig. 2C) on both 0·6 to 1·6 mm long. The nodal region of the surfaces are thick-walled, rectangular, both the halves may either fall in one level trapezoid, hexagonal in outline, margin or may be arranged at different levels (PI. 1, straight and arranged end to end. Cells are figs. 2, 6). Branch scars are preserved on 100-120 fl. long and 15-20 fL broad. Cell nodes. Branch scar is a rounded (PI. 1, wall ± 3 fl. thick. fig. 5; Text-fig. 3B) depression, measuring A large number of leafless articulated axes 1·5-2 mm in dimension. Occasionally, a are found alongwith these foliage shoots carboni sed crust of ± 4-5 mm broad, is pre• (PI. 1, figs. 2, 4; Text-fig. 3A, B). The served on both the marginal side of stem. axes are preserved in form of impressions, This carbonised crust, probably represents compressions or casts. The axes are arti• the cortical part of the axes. Usually the culated and of varying length. The grooves leafless articulated axes are found lying Pages 269-270
TABLE 1- SHOWING THE VARIATIONS IN THE SIZE OF LEAVES AND THE DISTANCE IN BETWEEN TWO INTERNODES IN DIFFERENT SPECIMENS OF TRIZYGIA SPECIOSA. (ALL MEASUREMENTS ARE IN mm UNLESS MENTIONED) L = Lateral pair of leaves, C = Central pair of leaves
SPECIMEN SIZE OF KUMBER NUMBER PLANT OF WHORLS
5'5 em 9 whorls Size of leaves in sueees- I 2 3 4 5 6 7 8 9 sive whorls L=2 L=4 L=5 L=7 L=8 L=8 L=9 L=12 L=12 C=? C=2 C=2 C=2 C=2·5 C=2'5 C=4·3 C=4 C=4 Length of internodes in 1-2 2-3 3-4 4-5 5-6 6-7 7-8 8-9 between two nodes 3 5 5 6 6 7 7'5 7
2 13 em 13 whorls Size of leaves in sueees- I 2 3 4 5 6 7 8 9 10 II 12 13 sive whorls ? L=9 L=IO L=II L=13 L=16 L=17 L=20 L=21 L=22 L=? L=24 ? Length of internodes in 1-2 2-3 3-4 4-5 5·6 6·7 7-8 8-9 9-10 10-11 11-12 between two nodes ? 7 8 9 10 II 12 13 14 IS IS
3 12 em 15 whorls Size of leaves in sueees- I 2 3 4 5 6 7 8 9 10 II 12 13 14 IS sive whorls L=5 L=6 L=8 L=10 L=12 L=13 L=14 L=? L=15 L=16 L=17 L=18 L=19 L=20 L=19 ~ C=2·5 C=3'5 C=4 C=5 C=5'5 C=' C=? C=? C=7 C=7 C=7·5 C=8 C=9 L=10 C=11 Length of internodes in 1-2 2-3 3-4 4-5 5-6 6-7 7-8 8·9 9-10 10-11 11-12 12-13 13-14 14-15 between two nodes 5 6 7 8 8'5 9 9 9 10 10 II 12 13 14
4 15 em II whorls Size of leaves in sueees- I 2 3 4 5 6 7 8 9 10 II sive whorls ? ? L=15 ? ? L=? L=23 L=' 24 L=' L=26 L=' Length of internodes in C=9 C=11 C=12 C=13 C=14 C=15 C=? between two nodes 1-2 2-3 3-4 4-5 5-6 6-7 7-8 8-9 9-10 10-11 7 7 10 11 12 14 16 18 19 ?
5 6 em 9 whorls Size of leaves in sueees- I 2 3 4 5 6 7 8 9 sive whorls ? L=7'5 L=7·5 L=IO L=12 L=I1-5 L=12 L=13 L=13 C=2 C=2 C=3 C=4'5 C=4'5 C=5'5 C=5·5 C=5·5 Length of internodes In 1·2 2-3 3-4 4-5 5-6 6-7 7-8 8-9 between two nodes 6 6 7 7 7 8 9 9
6 7'5 em 12 whorls Size of leaves in sueees- 1 2 3 4 5 6 7 8 9 10 II sive whorls L=6 L=6 L=10 L=12 L=12 L=14 L=14 L=14 L=15 L=16·5 L=18 C=2 C=2·5 C=4 C=4·5 C=4'5 C=5 C=5 C=5 C=6 C=6·5 C=7 Length of internodes in 1-2 2-3 3-4 4-5 5-6 6·7 7-8 8-9 9-10 10-11 between two nodes 5 6 7 7 8 8 7·5 8'5 9 10
7 8 em 6 whorls Size of leaves in sueees- I 2 3 4 5 sive whorls L=16 L=17·5 .L=20 L=20 L=21 Length of internodes in be- C=7 C=7'5 C=10 C=IO C=II tween two nodes 1-2 2-3 3·4 4-5 5-6 12 13 IS 16 20
.-\11 attempt has been made to place the specimens in a order to mark the parellelism in sIze of dIfferent specimens and to demonstrate the region, to which position of plant it belongs. N[AITHY - OBSERV,\.TlOl\S ON INDIAN LOWER GONDWAN,\. SPHE~ OPHYLLALES 271
A B
TEXT-FIG. 3. - A, leaflcss stem showing articulations and a distinct median groove X Nat. size. B, stem showing branch scar X Nat. sizc. obliquely in the shales, but some specimens other axis only a whorl of a fragmentary are transversely fractured showing three leaf is preserved. The anatomical structure separate regions which can only be inter• of the xylem, i.e. bordered pits from the preted as the cortex, stele and pith. The leaf-bearing shoots are identical to those of pith appears to be hollow as no cellular leafless axes. This further supports that structure is seen on examination of cellular the leafless axes and the foliage shoots of pulls from that region. Cellular pulls taken Tl'izygia are parts of the same plant. The from the stelar region show tracheids with mode of preservation indicates that the bordered pits. Pits are arranged in 1-5 leafless axes are rhizomatous structures and rows, bordered, circular-oval with a oval the foliage shoots are the aerial structures. pore (PI. 2, fig. 8). When pits are in more Further record of better preserved specimen than one series, they are opposite to one will throw more light on this problem. another or arranged in groups of 2, 3 or Height of Plant - In this study an attempt 5 to 7 pits. Crossfield pits are simple and has been made to reconstruct the possible transversely oval in outline. The outer height of the leaf-bearing shoots. The task regions of the stem do not show any was difficult, since all the specimens were in• preservation of cellular structures. complete. To complete this aspect of study, Occasionally, the foliage shoots of Trizygia the size measurements of the leaves on differ• speciosa are found in close association to the ent whorls and the length of the internodes leafless articulated axis suggesting a possible have been taken from number of specimens organic oonnection. The foliage shoots are represen ting different portions of the shoots arranged ± at right angles only on one (see Table 1). Two facts came out from side of the axis. The figured specimen this study that (i) a steady mathematical (PI. 1, fig. 6; Text-fig. 4) shows a cast of increase in the size of leaves and the leafless articulated axis measuring 9 cm internodal distance is marked from apical long and 3·5 cm broad with a distinct end to basal end and (ii) the different median groove. The axis had distinct arti• shoots representing the same level of aerial culations which are continuous at nodes. shoots show equality in their sizes, i.e. the On both the lateral margins of the axes size of the leaves and the internodal distance the carbonised area of 5 mm broad are are nearly the same. Taking these facts preserved. On one side, two leaf bearing into consideration, portions of aerial shoots axes emerge from two different nodal points representing different regions were placed (see Text-fig. 4). On one axis four leaf below one after another on the basis of whorls are preserved on nodes, while on the increase in the size of the leaves. By such 272 THE P ALAEOBOT ANIST
TEXT-FIG. 4 - Leafless stem with two foliage bearing shoots arranged at the node X 1·5. placement, it was possible to get an idea preserved closely ± parallel to one another. of the height of the plant. The plant, Near the basal point, the shoots are pre• on the basis of such reconstruction, appears served fairly close suggesting that the plants to be about 20 cm in height and has were growing straight in tufts like modern 16-22 whorls of leaves (Text-fig. 5). grass. This is further supported by the size Habit - Trizygia speciosa has been re• of plant and their mode of preservation corded from many parts of Gondwanaland. that they were erect herbaceous plants Most authors have considered that the (Text-fig. 5). If the plant was a creeper leaves of Trizygia were borne on a plant then it must have shown curves in the with creeping habit. Both Seward (1941) nature of stem, which is totally absent in and Plum stead (1966) have figured Trizygia all the recorded specimens. plant as an insignificant creeper. Pant and Mehra (1963) have suggested that the shoots of Trizygia speciosa were probably strangling DISCUSSION on the ground like those of Galium of Rubiaceae, because in fossil state the leaves The Indian forms were named by Royle are spread out in single plane. However, (1839) as Trizygia speciosa due to the Rigby (1969) opined that the Gondwana characteristic arrangement of the leaves species of Sphenophyllum should not have on whorls. Subsequently McClelland (1850) been borne on a plant that differed signi• opined that the leaves agree morphologically ficantly from the plants suggested by the to Sphenophyllu111, hence it should be work of Darrah (1968). He further ex• transferred to it. This was followed by pressed that it is hard to reconcile the subsequent authors (see Maheshwari, 1968). creeping habit with such a wide distribution. However, Maheshwari (1968) preferred the In our collection there are several examples form placing under the original name Trizy• which show a number of foliage shoots gia speciosa Royle in accordance with the MAlTRY - OBSERVATIONS ON INDIA LOWER GONDWA A SPHENOPRYLLALES 273
TEXT-FIG. 5 -- Reconstruction of Trizygia speciosa Royle. views of Krausel (1928) and Edwards (1955) dermal cells observed by Maheshwari must that the northern form identified in Glosso• be the straight-walled cells preserved only pteris flora are possibly different from at the basal portion of leaves. similar looking elements occurring in Angara A comparison of the epidermal structure with which they have been identified. of Trizygia speciosa from the known fossil Asama (1970) has reclassified the members of articulates shows that the morphological Sphenophyllales on the basis of morphology. structure of stomata in all of them is of According to him the trizygoid attachment advanced nature than Trizygia speciosa. of leaves to the stems at nodes represents The subsidiary cells in Sphenophyllum majus a distinct line of evolutionary series in described by Abbott (1958, figs. 44, 45) and Sphenophyllales. The leaves of trizygoids Remy (1962) and S. thonii described by may either show straight or curved veins. Meyen (1970) are of specialized nature, Trizygia has been restricted by him for the i.e. they form more or less regular ri'ng forms with stuight veins, while Paratrizygia around the stomata. Similar ring is also has been used for the forms with curved seen in Asterophyllites equisetiformis, Annu• veins. Following this view, the name Tri• laria mucronata and Annularia stellata (Ab• zygia is used for the Indian forms. bott, 1958). The subsidiary cells in S. Epidermal structure of Trizygia speciosa oblongifolium (Germ. & Kaulf) Unger (see has earlier been described by Pant and Meyen, 1970) are less specialized but the Mehra (1963) and Maheshwari (see Surange, stomata are arranged in linear rows. The 1964). According to Pant and Mehra (1963), subsidiary cells of Phyllotheca angusta the epidermal cells on both the laminar Surange and Kulkarni (1968) show trans• surfaces are sinuous, whereas Maheshwari verse thickenings on guard cells. observed (see Surange, 1964, p. 25) straight• The leafless axes in their gross morphology, walled cells on one surface and sinuous cells when ill preserved, show close resemblance with stomata on the other surface. The with Vertebraria indica Royle due to presence present study confirms the observations of of distinct nodes and internodes. However, pa.nt and Mehra.. The straight-wa.Ued epi- the a.xis differs from Vertebraria in its 274 THE PALAEOBOTANIST characteristic arthrophytic habit, i.e. parallel, South Karanpura Coalfield agrees morpho• continuous running ridges and grooves and logically to Sphenophyllum rhodesii Rigby the branch scars. Anatomically too, the (1966). However, the placement of these axis differs from Vertebraria. The pit aper• specimens under Sphenophyllum is wrong tures in Vertebraria are vestured, while in according to the new classification proposed these axes they are circular. Recently Pant by Asama (1970) for Sphenophyllales. The and Kidwai (1968) have described ana• specimens morphologically agree to Para• tomical details of the tracheids from the trizygia Asama (1970) due to the trizygoid leaf-bearing axis of Phyllotheca indica. The arrangement of leaves in whorl and arched pits compare in their bordered structure, but veins towards the margin of leaves. Hence differs by crossed pit pores. Surange (1955) the Australian and Indian specimens are has described the rhizome of Phyllotheca transferred to Paratrizygia. which in its organizational aspect agrees with the leafless axis of Trizygia. However, in the rhizome of Phyllotheca, the character• Parasphenophyllum Asama, 1970 istic ridges and grooves were absent. Parasphenophyllum crenulatum sp. novo Paratrizygia Asama, 1970 Pl. 2. figs. 11, 12
Paratrizygia rhodesii (Rigby) n. comb. Diagnosis - Whorls comprising of six PI. 2. fig. 10 leaves of equal size; leaves spread out at rignt angles to the stem; triangular in Synonymy: outline, petiolate base, obcordate apex, apical margin undulate and lateral margin 1966 Sphenophyllum rhodesii Rigby, p. 126, entire; a distinct median vein emerges pI. 33, figs. 30-32; text-fig. 2 at the base of leaf which dichotomises into Emended Diagnosis - Six leaves in a two, each supplying one half of the leaf whorl arranged in a trizygoid pattern, by repeated dichotomy, veins are arching two pairs bigger and one pair smaller, in the lateral portion and straight in the obcuneate shape and rounded apex, venation middle portion, anastomoses absent. dichotomous, arching towards the margin Holotype-l0/1644, Birbal Sahni Institute and straight in the middle. of Palaeobotany. Holotype - WA 23684 (Rigby, 1966). Locality - Pit no. 7, Damodar Colliery, Figured Specimen-l03/1216, Birbal Sahni East Raniganj Coalfield, West Bengal. Institute of Palaeobotany, Lucknow. Horizon - Raniganj Formation (Lower Horizon - Barakar Formation (Lower Gondwana). Gondwana). Age - Upper Permian. Age - Lower Permian. Description - Ten specimens are in collec• Description - A leaf whorl consisting of tion, out of which only 2 are nearly com• six leaves arranged around a central point. plete. The detached whorls are composed Two pairs of leaves are bigger in size than of six leaves. Leaves are of equal size and the third pair. The bigger pairs are ob• 2·5-3·5 cm long and 2·2-2·5 cm broad. cuneate in shape, broadly rounded, and Leaves are isobilateral triangular in shape. emarginate apex, contracted base and non• They are broadest at the apex. At the petiolate. Leaves are almost as long as apex, the median region of the leaves are broad, measuring 10X 8 mm. The smaller deeply notched, while the marginal ends leaves are obcuneate, broadly rounded apex, of the apex are acute. The apical margin non-emarginate, contracted base and sessile. has crenulations, while the lateral margins Leaves measure 8 X 5 mm. Venation re• are entire. A single vein emerges at the base peatedly dichotomizing, arising from a of leaf. Immediately after its entry into the single stouter vein that enters the leaf at lamina, it dichotomizes into two veins. Sub• base. All veins are curving towards the sequently, the two vei'ls dichotomize into margin except those in the middle. 30-36 veinlets. Veins in the middle portion of Comparison 0- Discussion - The specimen leaf are straight while in the lateral portion recorded from the Lower Nakari Seam of they show strong outward curvatllre, 1L\lTHY - OBSERVATioNS ON iNDiAN tOWER GONDWANA SPHENOPHYLLALES 275
Comparison- The Lower Gondwana speci• Southern hemisphere (Rigby, 1966; Wal• men is comparal;>le to Parasphenophyllum ton, 1929; Huard-Moine, 1965; Lacey & thonii (Mahr) Asama, 1970 and P. thonii var. Huard-Moine, 1966; Archangelsky, 1958). minor (Sterzel) Asama, 1970 which differ Most of the records from the different in their obovate shape. In P. spinulosum Gondwana continents have been referred to (Yabe & Oishi) Asama, 1970 the leaves have Trizygia speciosa. However, some records toothed margin. P. neofimbriattt1n (Halle) of Sphenophyllales from Wankie District, Asama, 197U differs from P. crenulatum Rhodesia needs reconsideration in the light in its long elliptical-spathulate shape. P. of work done by Asama (1970). Spheno• shansiense Asama, 197U is distinguished by phyllum thonii Mahr, 1868 recorded by its long obovate shape with rounded apex. Lacey and Huard-Moine (1966, p. 15, pI. 1, Lacey and Huard-Moine (1966, pl. 4, fig. 1) and S. thonii var. minor Sterzel figs. 28-29; text-fig. 2) recorded a new genus reported by Walton (1929, pI. a, figs. 3, Benlightjootia mackii from Wankie beds, 4, 8) agree morphologically to each other Rhodesia. The specimens show bifid leaves but they differ considerably from the type occurring singly or in apparent clusters, specimen of the species. The Wankie speci• i.e. in a few instances two, three, four or mens are considerably smaller than the five leaves may be associated together, European ones; the leaves are not lacinate '-nd their bases directed to a common and only two basal veins are present instead point. Leaves are cordate in shape with of four. From the illustrations it can be deep apical notch; venation open, dicho• said that the Wankie forms are different tomous arising from a single vein (or from S. thonii Mahr and represent a new possibly two veins very close together) type. at the base of leaf, repeatedly forking to The forms referred to Sphenophyllum reach the margins and extremities of two oblongifolium (Germ. et Kaulf.) Unger by lobes. The vascular system of the two Walton (1929), Huard-Moine (1965) and halves of the leaf are completely separate Lacey and Huard-Moine (1966) are different from the base. This specimen shows close from the European ones. The Europea:n morphological similarity to Parasphenophyl• forms are narrow, four lobed with indenta• lum crenulatum sp. novo described here. tions on the apical margins, while the However, for the time being the Wankie Wankie specimens are broader and two lobed specimens are kept separate in want of with only shallow indentations. Hence, it specimens showing the arrangement of leaf. is felt that the Wankie specimens are Only future record will throw light on this distinct from the European ones and they problem. sho~ld be placed separately under a new speCIes. REMARKS ON THE RECORDS OF SPHENO• Thus, we find that the Gondwana speci• PHYLLALES FROM THE GLOSSOPTERIS mens earlier referred to Euramerian ones FLORA OF THE SOUTHERN HEMISPHERE differ considerably. Hence, all the forms referred to European species from the There are a number of records of Spheno• Gondwanas need careful re-examination for phyllales from the Glossopteris flora of their proper placemen 1.
REFERENCES
ABBOTT, M. L. (1958). The American species of EDWARDS, W. N. (1955). The geographical dis• Asterophyllites, A nnularia and Sphenophyllu-m. tribution of past floras. Advan. Sci., 46: Bull. Am. Palaeont., 38 (174): 289-372. 1-12. AHCHANGELSKY,S. (1958). Estudio geologico y FEISTMANTEL, O. (1879). Note on the genus palaeontologico del Bajo de la Leona (Santa Sphenophyllwn and other Eq uisetaceae with Cruz). Acta geol. lilloana, 2: 5-133. reference to Indian form Trizygia speciosa Royle ASAMA,K. (1970). Evolution and classification of (Sphenophyllwn trizygia Ung.). Rec. geol. Sttl'V. Sphenophyllales in Cathaysia land. Bull. Natn. India, 12 (3): 163-166. Sci. Mus., Tokyo, 13 (2): 291-317. HUARD-MoINE, D. (1965). Contribution a l'etude DAHHAH, E. L. (1968). A remarkable branching de la flora dite "a Glossopteris" du bassin de Sphenophyllum from the Carboniferous of Illinois. Wankie (Rhodesie du Sud) II. leD Sphenop' Palaeontographica, HOB: 99-127. sides. Ann. Unw. I.A.R.E.R.S., 2: 68-8(). 276 THE PALAIWBOTANIST
KRAUSEL, R. (1928). In Krausel, R. & Range, P. PLUMSTEAD, E. P. (1966). The story of South Beitrage zur kenntnis der Karruformation De• Africa's Coal. Optima: 186-202. utsch-siidwest Afrikas. Beitr. geol. Erforsch. REMY, W. (1962). Sphenophyllum majus Brongn., dt. Schutzgeb, 20: 1-54. Sphenophyllum saarensis n. sp. and Spheno• LACEY, W. S. & HUARD-MoINE, D. (1966). Karoo phyllum orbicularis n. sp. aus dem Karbon des floras of Rhodesia and Malawi. Part 2. The Saargebietes. Monatsber. Dt. Akad. Wiss., 4 Glossopteris flora in Wankie District of Southern (3-4): 235-246. Rhodesia. Symposium on Floristics &> Strati• RIGBY, J. F. (1966). The Lower Gondwana floras graphy on Gondwanaland. Birbal Sahni Insti• of the Perth and Collie basins, Western Aus• tute of Palaeobotany, Lucknow: 13-28. tralia. Palaeontographica, 118B (4-6): 113• MAHESHWARI,H. K. (1968). Studies in the Glos• 152. sopteris flora of India-38. Remarks on Trizy• RIGBY, J. F. (1969). The Lower Gondwana Scene. gia speciosa Royle with reference to the genus Boletim Paranaense de GeoC'ieneias, 27: 1-13. Sphenophyllum Koenig. Palaeobotanist, 16 (3): ROYLE, J. F. (1839). Illustrations of tho botany 283-287. and other branches of Natural History of the MAHESHWARI,H. K. (1974). Palaeozoic Lycopsida Himalayan mountains, etc. London (1833• and Sphenopsida, in: Surange, K. R. et al. (Eds): 39). Aspects &> Appraisal of Indian Palaeobotany. SEWARD, A. C. (1941). Plant life through the ages. Birbal Sahni Institute of Palaeobotany, Cambridge Univ. Press: 607. Lucknow: 51-61. SURANGE,K. R. (1955). Studies in the Glossopteris MCCLELLAND, J. (1850). Report of the Geological flora of India-2. Equisetales from the Rani• Survey of India for the Session 1848-49. Calcutta. ganj Coalfield. Palaeobotanist, 4: 83-88. MEYEN, S. V. (1970). Epidermisuntersuchungen SURANGE, K. R. (1964). Indian fossil pterido• an Permischen Landpflanzen des Angaragebie• phytes. Botanical Monographs, 4. C.S.I.R., tetetes. Paliiont. Abh., 3B (3-4): 523-552. New Delhi. • PANT, D. D. & MEHRA, B. (1963). On the epider• SURANGE,K. R. & KULKARNI, S. (1968). On two mal structure of Sphenophyllum speciosum new species of Phyllotheca from the South Karan• (Royle) Zeiller. Palaeontographiea, 112B (1-3): pura Coalfield, Bihar. Palaeobotanist, 16 (1): 51-57. 95-100. PANT, D. D. & KIOWAI, P. F. (1968). On the WALTON, J. (1929). The fossil flora of the Karoo structure of stems and leaves of Phyllotheca System in the Wankie District, Southern indica Bunbury and its affinities. Palaeonto• Rhodesia. Bull. geol. Surv. S. Rhodesia, 15: graphica, 121B (4-6): 102-121. 62-75.
EXPLANATION OF PLATES
PLATE 1 PLATE 2
Trizygia speeiosa Royle Trizygia speciosa Royle 7. Epidermal structure from the basal-most 1. A nearly complete specimen with 14 whorls of region of the leaves. x 100. leaves. X Nat. size; No. 15/1644. 8. Epidermal structure of stomatiferous surface. 2. Leafless axes showing articulated stem with x 100. distinct nodes and internodes. x N at. size ; No. 97/ 9. Trachiedal structure in the cellular pulls taken 1547. from the surface of leafless axes. X 400. 3. A portion of leafless axes enlarged to show the continuity of ridges and grooves. x 3. Paratrizygia rhodesii n. comb. 4. Leafless axes with a median groove, indicating 10. A single whorl of six leaves: note the arching region of pith. x Nat. size; No. 98/1547. veins towards the margin of leaves. x 3; No. 103/ 5. Leafless axes showing branch scar (B). x Nat. 1216. size; No. 6/1644. 6. Leafless axes with two shoots of Trizygia Parasphenophyllum erenulatum sp. novo speciosa. The shoots are arranged at right angle 11. A single whorl of six leaves. xl; No. 10/ from the leafless axes at two adjacent nodes. X Nat. 1644. size; No. 90/1547. 12. Two leaves enlarged to show venation. X 3. MAtTHY - OBSERVAnONS ON tNDIAN LOWER GONDWANA SPHENOPHYLLALES 277
rLA'l'E t 278 THE PALAEOBOTANISf
PLATE 2