Pa/ambo/GI/il/ 49 (2000) 3.13-352 0031-0174/2000/333-352 $2.00

Taxonomy and diversity of the genus Glossopteris

SHIV MOHAN SINGH*

Birbal Salmi Il15titli/e of Palaeobotany, 5] Uni\'ersiry Road, LlicknOlv 226 007, India. . Present address: TaxonolllY & Biodiversity Division, Narional Botanical Research Instirll/c, Rana Pratap Marg, LIICkl1o\v 226 001, Indic!.

(Received 08 June 1999; revised version accepted 08 June 2000)

ABSTRACT

Singh SM 2000. Taxonomy and Diversity of the genus Glossopteris Palaeobotanist49(3) : 333-352_

The speciation of Glossopteris leaves in Permian Gondwana of India is re-examined. It is based on study of thousands of specimens collected from Barakar Formation of Karanpura and Bokaro Group of Coalfields. The study of specimens of modern plants showing variation in shape and size of leaves within same species (sometimes within same plant) and survey of published literature, the author was fascinated to express the ideas about the parameters which may be helpful in speciation of the genus Glossopteris. Here. morphological characters have been critically analysed in order to find a reasonable basis for precise spe­ cific delimitations. The morphological circumscriptions have been further verified by characters of cuticle which have been taken as associated or supportive characters only. The size and shape of leaves have given secondary importance. The Diversity and the total number of species found in time and space have been tabulated.

Key·words- Glossopteris, Permian. Morphology. Cuticle. Speciation, Gondwana. India.

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INTRODUCTION is a genus of tongue-shaped fossil leaves which have a robust to flat midrib that gives off secondary veins that dichotomise Though the genus Glossopteris is the most abundant of and anastomose. The genus was first recognised by Adolphe all plants in the Permian floras, it has not been circumscribed Brongniart (1828), the father of Palaeobotany. He r~cognised in definite species within a genus. The main reason for this two species in the genus, viz., G. browniana var. allSlralisica was a complete lack of accepted criteria on which to base from Australia, G. browniana var. indica from India and G. such classification. The first attempt in this direction was made angustifolia also from India. Schimper (1869) raised var. by Arber (1905) whose system of specific circumscription has indica to the status of a species as G. indica. Dana (1849), been widely followed. In view of earlier researches on the Bunbury (1861) and Feistmantel (1876-1881, 1886, 1890) morphography and cuticular structure of leaves and on the described a number of species of Glossopteris from India nature of fructifications borne by these leaves; there is an and Australia. Zeiller (1896) reported for the first time the apparent lack of agreement between the two systems of epidermal structure of a leaf he identified as G. indica. The classification (Srivastava, 1957; Plumstead, 1958). Hence results of investigations in the genus done in the nineteenth there is a greater need for further work in this direction and century were summarised by Arber (1905). Though sporadic the evidences they obtained have to be correlated with those work continued in the early part of the twentieth century, the obtained by more critical studies on the fructification and the investigation got an impetus in the 1950s onwards when a lot cuticular structures. Though the evidences provided by the of data was generated not only on the morphography and cuticular and the fructifications are very important, their taxonomy of the leaves but also on the cuticular features and application is admittedly limited as most of the Lower attached/associated fructi fications. In spite of all this, there is Gondwana fossils occur more frequently as impressions. Hence still a controversy about the characters on which speciation in the natural thing is to device a classification based on the the genus should be based. Most workers prefelTed to separate morphographical characters. On the basis of morphography forms into different species on the basis of minor differences the retention of the taxonomic status of the present genus while others believed in maintaining a few species by merging Glossopteris seems to be most advisable and advantageous different forms of leaves into one, if most of the characters of inspite of the obvious di fficulties sometimes encountered in such leaves were the same. These two schools of speciation determination of transitional forms. It is hoped that ifdue care have been termed "splitters" and "Iumpers", respectively is taken in identifying the specimens described in the (Plumstead, 1962). Monograph of Chandra & Surange (1979), the cuticular Seward (1897, p. 317) believed that size and shape are species and the list of species given in Fig. I the most of the extremely dangerous guides in specific delimitation. Seward species will fit in one another and the number may become (1910) went to extent of suggesting that speciation based on half. In the Karanpura & Bokaro Coalfields the circumscription venation and generic delimitation on the basis of presence or of species made by studying more than one specimens and absence of a midrib is not justified. Earlier Arber (1905), a intensi ve observations have gi ven 32 species (Thesis Shi v strong supporter of maintaining a few species only, re-classified Mohan Singh, Contributions to the Early Permian flora of species distinguished by earlier workers into thirteen species Karanpura and Bokaro Coalfields, 1998). only. According to him "there existed a considerable variation in the form and shape of the leaf of the genus Glossopteris SYSTEMATICS and in the details of the nervation, even in the fronds in which there is reason to believe belonged to the same plant". He did Genus-GLOSSOPTERIS Brongniart 1828 not take into account the characters of midrib, nature of apex and angle of divergence of secondary veins from the midrib Type species-GLOSSOPTERIS BROWNIANA as characters suitable for specific delimitation. He did think Brongniart 1828 that the only constant character for speciation was the open­ Glossopteris is the most abundant fossil in the Permian ness or close-ness of the secondary veins and hence the shape floras of the Southern Hemisphere, that during the period of meshes. However, Maheshwari (1966) opined that without comprised Australia, Antarctica, Africa, South America, and knowing the whole plant. it was not possible to know whether India, and possibly also the Arabian Peninsula. Glossopteris there was such a variation in the same plant or even in the

/" " PLATE 1

I. Glossopteris karanpurael/sis Kulkarni 1971, Specimen no. BSIP­ 2-5. CUlicle of Glossopteris karanpurael/sis, probably of the slomatiferous 38836-13 (1/4578 A I). Barakar Formation, shales associated with surface. showing barely discernible laleral cell walls. and spindle­ Naditoli Seam. Sirka Colliery. South Karanpura Coalfield, Bihar. x shaped Slomala. Specimen no. BSIP-38836-B ( 1/4578 AI). 2 x 200. nat. size. 3·5 x 400. --J I en ..._v 'i:I r» t- en ;l> 0 -3 c:; 0 tr1 --J N » Z VJ --J SINGH- TAXONOMY AND DIVERSITY OF THE GENUS GLOSSOPTERIS l37

same species of fossi Ileaves. He pointed out that in a character, leaf may be assigned to Gangalllopteris. Besides Pant and co­ the delimitation of its range is difficult and so the variation workers, H0eg and Bose (1960). Surange and Maheshwari shall necessarily be an arbitrary one. (1962). Saksena (1963), Rigby (1966). Srivastava (1969. Before 1956 the speciation of Glossopteris was based 197/). Banerjee (1971), Chandra and Surange (1~77a,b). on morphography, i.e .. external features of the leaf. Zeiller Rigby et al., (1980), Chandra and Snvastava (1981) and (1896) and Sahni (1923) described the epidermal and cuticular Maheshwari and Tewari (1992) have reported on the cuticles structures of the leaves of G. indica Schimper and G. of Glossopteris leaves. angustifolia Brongniart, respectively. Srivastava (1957) made The first Glossopteris fructification was described and an effort to delimit species on the basis of cuticular characters. ill ustrated by Feistmantel (1881). Though the lectotypes for He described features of the cuticle in 16 species of the taxon Dict)'opteridiwlI sporiferulll (Geological Survey of Glossopteris, 6 species of Gallgalllopteris and one species of India, Calcutta, Museum Specimen 5210 figured by Banerjee Palaeovittaria. Surange and Sri vastava (1957) classified these 1973) does provide some evidence of probable attachment to 23 species in six groups on the basis of totality of cuticular a Glossopteris leaf. Feistmantel thought it was a fern plllnule. features. They thought that these groups could be of generic Zei lIer (1902) reported Ottokaria (Feistll/{/ntelia) bel1galensis rank. However, later workers have not followed this line of now shown to be an ovaliferous capitulum with a long stalk thinking. It may be mentioned here that H0eg and Bose (1960) and subtended by a Glossopteris leaf (Banerjee, 1978). found that morphographically similar leaves may have different White (1908) established the genus Arberia for broadly cuticulartypes. Recently Maheshwari and Tewari (1992) have incised, coriaceous or striate and thick nerved scale leaves found that morphographically different leaves may have similar whose distant recurvate and truncate lobes appear to owe their looking cuticles. These two observations need to be examined abrupt or even slightly ragged terminations to the detachment in detail, particularly in the light of Birbal Sahni's reluctance of some sort of bodies. Presumably reproductive in nature. to accept features of the cuticle as very good indicators of His specimen came from Gangal/lopteris bed (Joaguim Branco specific variation. Horizon, Rio Bonito Formation, Guata subgroup, Tubaro Pant (1958) and Pant and Gupta (1968-1971) have Group Rigby 1972a), northeast of Minas (now Lauro Muller). described the cuticular features of a number of species of Santa Catarina, Brazil and were intimately associated with Glossopteris and Gangalllopteris. According to these authors. Sal/laJ'Opsis seeds and Gangamopteris obovata Carruthers one of the major characters which should be taken into account leaves. while differentiating the genera Glossopteris and On the basis of association a relationship between Arberia Gangamopteris and also circumscribing a species is the midrib. and Gangamopteris was presumed. Rigby 1972b) interpreted According to Surange and Srivastava (1957), leaves of Arberia as a fructification that bore large numbers of naked Gangamopteris indica Srivastava and Grl/lgamopteris ovules on pinnate branchlets arranged laterally along a forked cyclopterioides Feistmantel are easily confused with racheis. Glossopteris because their median veins are more prominent, However, the restoration of a mature specimen by Rigby and forms like Glossopteris longicaulis Feistmantel and (1972b, text fig. 2) and some of specimens illustrated by Appert Glossopteris decipiellS Feistmantel, where a midrib is seen (1977, pI. 36, fig. 2) Chandra & Srivastava (1981, pI. I, fig. only in the lower portion of the frond. can be mistaken for 1) and Anderson and Anderson (1985, pI. 103, fig. I) show Gangamopteris. Maheshwari (1966) emphasised the fact that that the branching is pleiochasial. though, vertically running strands are found on the midrib of Plumstead (1952, 1956a, b, 1958) described and Glossopteris leaves yet, they never anastomose and in this illustrated a large number of fructifications in organic sense are different from the median veins of the genus connection with leaves of Gangamopteris, Glossopteris and Gangall10pteris which, though sometimes simulate a midrib, Palaeovittaria. Pant (1982) recognised 4 groups of Gondwana yet show definite anastomoses. Pant and Singh (1968) arrived Gymnosperms, viz., Glossopteridales, Noeggerathiopsidales, at the concl usion that if the median region is iII-defi ned and if Coniferopsida and incertae sedis. He remarked that the unique the cuticle of this region shows stomatiferous areas (meshes) leaf attached fructifications of Glossopteridales may suggest bounded by non-stomatiferous areas (veins) like those of the some affinity with the Pteridosperms. lamina, and if it is otherwise not clearly differentiated from From time to time various diversified structures have been the cuticle of the lamina (in thickness or cell characters), the described as fructification of Glossopteris (Arber, 1905;

./ PLATE 2 "

Glossopteris ('OllllllUllis Feistmanlel 1876. Specimen no. BSIP-38855 2-5 Cuticle or Glossopteris ('OIlIlIlUlli.\ Feistmantel, Specimen no. [lSIP­ (8/4754). Barakar Formation, Dakara Colliery. North Karanpura Coal­ 38855 (8/4754). x 100 field, Bihar. x nal. size. w w 00 SINGH- TAXONOMY AND DIVERSITY OF THE GENUS GLOSSOPTERIS 339

Walkom. 1928; Dutoit, 1927; Sen. 1954. 1955a, b. 1956; White. to find a reasonable basis for precise specific delimitations. 1962). The first attached fructification was figured by Zeiller Morphographical circumscriptions have been further verified (1902, pI. 4, fig. 9) under the name Ollokaria bcngalclIsis which by characters of the cuticle wherever available. Thus for is attached to a Glossopleris indica type of leaf. Later on a specific circulllscription, the characters of the cuticle h~lvebeen large number of attached fructifications have been described taken as associated or supportive characters only. (Plumstead, 1952, 1956, 1958; Sen, 1955a, b; Rigby, 1962; Maheshwari, 1966; Pant, 1982; Anderson & Anderson, 1985). Morphographical characters Some important records of attached fructification with genus Glossopteris are G. browllialla, G. da/lllldica, G. The morphographical characters which are considered decipiens, G. indica. G. jamollei, G. IOllgicaulis, G. reti{era important from the point of view of specific circumscription (=elongata), G. stricta, G. tort/lOS(/ var. vaolallse. G. are: angusli{olia. (a) Shape of the leaf Chandra and Surange ( 1979) produced a monograph on (b) Margin of the lamina the "Revision of the Indian species of Glossopteris" in which (c) Apex the species of Glossopteris have been classified on the basis (d) Base of external characters alone. These authors considered that (e) Nature of midrib the size, shape, midrib, lateral veins and their behaviour are (f) Venation pattern characters of diagnostic value in differentiating one species from another. (a) Shape of the leaf Maheshwari and Tewari (1992) are of view that the genus Glossopteris should be classified on the basis of both The leaves vary in shape, having a varied length/width morphographical and cuticular characters. Among ratio. The leaves may be : morphographical features. they have taken shape, nature of I. Linear, e.g., G. forlllosa apex, base and margin, nature of midrib, density of lateral 2. Linear-Ianceolate, e.g., G. allgustifolia veins, and shape and size of meshes for speciation. Among 3. Lanceolate. e.g.. G. indica cuticular characters, shape and arrangement of cells, cell walls, 4. Lanceolate-spathulate, e.g., G. indica stomatal type, orientation and distribution of stomata, type of 5. Spathulate, e.g., G. browniallQ guard cells, and stomatal index have been considered useful 6. Oblong, e.g., G. emargillata criteria. 7. Obovate, e.g., G. rerusa In the present study, the speciation of genus Glossopleris 8. Cordate, e.g., G. sahllii is based on morphography and cuticular features. Here. 9. Obcordale, e.g., G. spat/lliiaia morphographical characters have critically analysed in order In modern plants, however. it has been observed that the shape of the leaf shows considerable variation within the same

/' ...... PLATE 3

Glo.lsapterisdanae Maheshwari & Tewari 1992, Specimen no. BSIP­ 3. Glossupteri.l· raniganjensil' Chandra & Surange 1979. Specimen no. 38830 (35/5004), Jharkhand Colliery. West Bokaro Coallielcl (Vena­ BSIP-38849 (13/4998), Sirka Colliery, South Karanpura Coalfield (Ve­ tion x 3). nation x 3). 2. Glossopteris karanpuraensis Kulkarni 1971. Specimen no. BSIP­ 4. Glo,sopteris clarkei Feistmantcl 1878. Specimen no 13SIP-38827 (<)J 38838 (2115004A), Jharkhand Colliery, West Bokaro Coallkld (Ve­ 4752). Gidi·C Colliery. South Karanpura Coalfield (Venation x 3) naLion x 3).

PLATE 4 See Page No. 340

I. A twig of modem plam Morus alba showing variation in shape of the of the same plant. leaves. 3. A twig of modern plant Ixora arlJorea of showing variation in shape 2.4. A modem plant Mille/lia sp. showing variation of leaves in two branch of the lea ves.

PLATES See Page No. 34 I

1·4. A modem Algae Gratelaupia indica (Red Algae) showing variation in shape of the thallus struclUre within the same species. Collected from Okha, Gujarat, January 1999 by DB Sahoo. --i :r: [T1 ~ r }> o[T1 oOl --i }> Z Vl --i ~41

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,..., 342 THE PALAEOBOTANIST species (PI. 4'1-4; PI. 5'1-4). Therefore. this character/ feature (ii) Veins arise at acute angles from the midrib. Degree of has only secondary importance. emergence of the veins is species specific. This situation occurs in majority of species. (b) Margin of the lamina: (b) Course of veins: Almost invariably the leaf margin is continuous and smooth. sole exception being G. rei lisa. (i) Veins almost straight, e.g., G. illlerJllillens Feistmantel; G. jonesii Walkom, G. JIlilchelii Walkom. (c) Apex: (ii) Veins arched. e.g., G. indica Schimper, G. COIlllllWlis Feistmantel, G. bmwniana Brongniart. Apex may be acute, acuminate, oblllse. retuse or driptip. (iii) Veins arched and sinuous. e.g., G. verticil/ala This character is important only when wide differences are Thomas. present, otherwise this character is of little importance, e.g.. G. eJllarginala (emarginate apex) and G. relUsa (retuse apex­ (c) Vein density: cum-notched margin) The per centimeter concentration of the veins has also (d) Base: been given importance. For example, in both G. laeniopleroides and G. euryJl.eurn the secondary veins are The leaves of a species may either be sessile or petiolate. almost perpendicular to the midrib but in the latter case the their being no intra-specific variation. As such this character number of veins per centimeter is much less as compared to whenever preserved is usually important for species the former This difference is verified by the flexuous nature delimitation when taken together with other characters. of the secondary veins in the latter. unlike the former where the veins follow an almost straight course. (e) Nature of midrib: (d) Vein anastomoses: The midrib which forms the main vascular supply of the leaf shows considerable variation in various species. It may (i) In some species the veins anastomose infrequently, continue from the base of the leaf up to the apex or may e.g., G. laeniopleroides, G. inlerJllillens. dissolve into secondary veins after some distance from the (ii) In most species veins anastomose frequently, e.g.. base, in which case it is said to be evanescent. The midrib G. brownianC/, G. forlllosa Feistmantel, G. elongala Dana, may look to be solid and stout or flat and insignificant. This etc. character seems to be taxon specific. In case where the midrib is flat, vertically running striations resembling the secondary (e) Frequency of veins dichotomies: veins may be found but these never anastomose and in this sense are different from the median veins of the genus (i) In some species the secondary veins dichotomise just Gangalllopteris McCoy. Sometimes the midrib is pitted once or twice and then follow an almost parallel course, e,g., throughout the length. These pits possibly represent bases of G. parnllela Feistmantel. outgrowths, such as, hairs or spines. Earlier workers mistook (ii) Secondary veins dicholOmise several times and hence this feature as representing the fertile stage. the concentration of veins at the margins is much higher than near the midrib, e.g., G. browniana, G. indica, G. Iinearis (f) Venation pattem: (Pi. 3·]-4) Bunbury, etc.

The most important character in specific delimitation as (f) Shape of meshes: already emphasised by Arber (1905) and Maheshwari (1966) is the nature of the secondary veins and the meshes formed by The shape of the meshes formed by the secondary veins them. depends considerably on the number of dichotomies, concentration of the veins, as well as the number of (a) Angle of origin: anastomoses. Meshes may be: broad and open, e.g., G. conspicua Feistmantel (i) Veins arise almost perpendicular to the midrib, e.g.. narrow and elongate, e.g., G. comJllunis Feistmantel Glossopteris taeniopleroides Feistmantel, G. euryneura elongate-polygonal. e.g., G. browniana Brongniart Maheshwari. polygonal, e.g., G. e!ongara Dana. G. formosa trapezoid, e.g., G. lortuosa Zeiller SINGH- TAXONOMY AND DIVERSITY OF THE GENUS GLOSSOPTERtS 343

Cuticular characteristics: (PI. 1-2) or less straight, dichotomising and anastomosing variously to form meshes of various shapes and sizes, density of veins It is not often that leaves with a carbonified crust (leaf usually lesser near midrib than near the margin. compressions) are present. Very seldom the carbonified crust, Leaf cuticle usually hypostomatic, rarely amphistomatic, on maceration, yields well-preserved cuticles. But whenever either undifferentiated or differentiated into vein and mesh satisfactorily preserved pieces of cuticle are recovered these areas on both the surfaces, cells over veins narrow, elongated, provide valuable secondary data for species delimitation. Some rectangular or squarish, arranged end-to-end in linear rows; of the characters of the cuticle that have been taken into cells in mesh areas varying in shape and size and arranged consideration for speciation are: irregularly, lateral walls of cells usually straight, sometimes (i) The presence of stomata on one (hypostomatic) or undulate or sinuous, surface walls mostly unspecialised. both (amphistomatic) the surfaces of the leaf. In the species sometimes granulate or papillate, trichomes generally absent; of the genus Glossopteris amphistomatic cuticle is unusual. stomata haplocheilic, present only in mesh areas, distributed (ii) Shape and arrangement of cells-The vein and mesh and oriented irregularly, stomatal index variable, stomatal areas may be decipherable on the cuticle through arrangement apparatus mostly monocyclic, rarely dicyclic or amphicyclic, of the cells. The cells over the veins are usually narrow, guard cells sunken or normal, subsidiary cells unspecialised elongate, rectangular or squarish and arranged end-to-end in or with papillae overhanging guard cells. Cells over midrib longitudinal rows. The cells in the mesh areas are polygonal, squarish or rectangular, arranged in rows, stomata usually rectanguloid or very rarely trianguloid and squarish and do absent over midrib. not show a regular arrangement. (iii) Cell walls-The anticlinal and periclinal walls of Habit of Glossopteris the cells may be straight, slightly undulate or sinuous. The surface walls may be smooth or papillate; the number of The habit of the plant which bore Glossopteris leaves is papillae may be numerous or only one. When the papillae are Iiule known. Seward (1931, p. 247) proposed its habit as many in number they are usually small and rounded. "shrubs". Plumstead (1958, p. 92) opined that "it is probable (iv) Distribution and orientation of the stomata-The that they were deciduous, woody plants with an arborescent Stomata occur only in mesh areas and are usually haplocheilic habit, and that the leaves, flowers and fruits were borne on and anamocytic (irregular number of subsidiary cells). The short shoots, a few of which developed into long shoots to stoma may have only one ring of encircling cells (monocyclic), form branches". Earlier it was suggested that the genus rarely two rings (dicyclic); sometimes the encircling cells may Vertebraria is one of the stems which bore such leaves partly cover the stoma. Usually the stomata do not exhibit (Dolianiti, 1954; Surange & Maheshwari, 1962; Pant. 1962). any regularity in distribution and orientation. These leaves were borne on stems either in tight spirals (v) Types of guard cells-The guard cells may be normal (Surange & Maheshwari, 1962, text-figure 8) or in pairs or sunken. The nature of the stomatal pore usually can not be (Dolianiti, 1954, figure I). However, it has now been very well deciphered in fossil cuticles. conclusively shown that Vel1ebraria is a root axis (Schopf, (vi) Stomatal index-It refers to the number of stomata 1965). It is possible that all Glossopteris-bearing plants were in a unit area. The formula that is used to work out the stomatal not arborescent. In the Talchir and Karharbari Formations no index is: petrified wood has yet been recovered. The climate was Stomatal Number of stomata relatively cool and not much coal was formed. During this = x 100 index Number Number of period it is probable that Glossopteris-bearing plants were of + epidermal shrubby in nature. In the late Early and Late Permian petrified stomata cells wood is assigned to the Glossopteris-bearing plants on the It is said to be highly variable within a species thus its basis of association. In one case this association seems to be diagnostic value, remains doubtful. indoubitable. In the shales associated with the topmost seam of the Raniganj Formation in the Raniganj and Jharia Generic Diagnosis (from Maheshwari & Coalfields, and below the Kumarpur and Mahuda Sandstone Tewari, 1992): Members, respectively, one finds an almost pure association of the leaf Glossopteris shailae Bajpai and the wood Leaves simple, sessile or petiolate; entire or slightly Araucarioxylon kU/1/Qlpurensis Singh & Bajpai (Bajpai, 1987; notched in the upper half; shape, size, apex and base variable; Bajpai & Singh, 1986; Bajpai & Tewari, 1990). Banerjee el midrib prominent, flat or elevated, persistent or evanescent, al. (1991) have reported a Glossopteris plant in-silu from the when flat, with longitudinally running parallel striations or Barakar (=Karharbari) Formation of Saharjuri Coal field. The pits or sometimes both, anastomoses of striations absent; lateral specimen is said to show branched stems bearing Glossopteris veins arching from the midrib at acute angles, arched or more leaves and vertical Vel1ebraria roots with spreading branches. .." w ~. l:> l:> PERMIAN TRIASSIC I INDIA AUSTRALIA S. AFRICA I S.AMERICA ASIA 9 N.s. Wales 5. I I 0-s A M T K B B R M N V Q T C N Orang Z B A T 0 N E a a a a a u e u a a a River n 0 ~'" T DI r n w c e p t colony m a g d n ~ A A c h a r e e m e a & b z e q "~ R G ha k e g e C 0 n a a n a, u ~ ;;; Glossopteris species C A a n a a r s n C T b -, ~. T S b n I ) 0 w P n "0'" I C a M J, S a a a I a e n a () "~. C A e e n 0 n a n A R a K e d n c '"~ s a y v h 3 '" u m e S a e "' I e a Q.'" C/l e h I "0 "'() (1) -l e :c ro ~ »r Glossopteris acuminara + ro aCt/fa 0 G. + + o:J G. ampla + ? + + + + + + + + 0 »-l G. angus/ifolia + + + + + + + + + + + + z G. angusta + + cr. -l G. anthrophylloides + G. arberi + + G. asansolensis + G. barakarensis + + G. bengalensis + G. browniana + + + + + + + + + + + + + ? + + G. brongniartii + G. bunburyana + G churiensis + G. clarkei + + G. COIIIIIIUllis + + + + + + + + + + + G. contracta + G. cordifonllis + G cordata + + G dallludica + + + + + + G danae (G. danae) + G daenaeoides + G dil'ergens + G decipiens + + + + + + ? G. dhenkanalensis + G. deogarhensis + G. ednoe + G. efegons + + G. emorginara + + G. eUl)lneura + + G. formosa + + + + G. feis/l/lGlI/efii + G. fibrosa cf. G. frondosa + G. ganga/llopleroides + G. gigas + + G. giridihensis + C/) G. ghusikensis + z G. gondwanensis + Cl :r: G. gopadensis + I G. harrisii + --l :> G. hingridaensis + x G. ingens + + 0z G. inlermedia + + cf. 0:s: G. indica + + + + + + + ? + + + + + + + + -< :> G. inequalis + z G. ill/er/llillens + + c G. isolaleralis + c < G. jayall/iensis + en ;;0 G. karanpuraensis + C/) ~ G. karharbariensis + + -< G. kusllllliae + 0 -n G. kanllhiensis + --l :r: G. Lanceolallls + en G. leplOneura ? + + Cl en G. longijolia + + z c G. longicaulis + + + + + C/) G. linearis + + £? () G. /IIaCUlala + v, v, G. manjuiae + a ~ G. major + + -; !"'1 ;" G. maheshwari + ;:; G. lIlohudaensis + G. lI1usaefolia + G. nauliyalii + G. nephroedicus cr G. obovala sp. nov. + G.ovara + G.obscura + + G.oldha/lli + + G. orbicularis + +

G. pandurara + + .J) A G. panIii + + V> G. papillosa + G. parallela + + + + VJ.,.. G. peliolala + 0- G. primaeva + G. pseudocommunis + + G. pseudOlorluosa sp. noy. + G. pseudoSIricia sp. noy. + G. radiata + G. recurva + G. reliculala + G. roylei + G. reclinervis + G. rewanensis + G relifera (=G. elongala) + + + + + + + + G. rhabdolaenoides + + G. reliculum + + G. relusa + + G. sahnii + G. saksenae + G. saslrii +

G. schopfli + --l J: G. searsolensis + rn G. senii cf ~ G. schimperii + r» G. singularis + rn 0 G. spalhiliaia + + OJ G. spalhulo-cordala + + 0 + »--l G. srivaslavae + + + z CIi G. slenoneura + + + --l G. surangei + G. shailae + + G. sublilis + G. syaldiensis + G. slricla + + + ? + + G. laelliopleroides + + G.lalei + G. lenuifolia + G. laenioides + + G. lransversalis + G.lorluosa + + + cf. G. ulkalensis + G. varia + G. venuSIus + G. verticil/ala + G vulgaris + G wallonii + + G. wilkinsoni + + G. zeilleri + + + GLOSSOPTERIS FLORA Talchir Karharbari Barakar Kulli Raniganj Kamlhi Bijori Pali Pachware Hinjir Maiture Hirapur Parsorsa ::TI 00 Neomariopleris lobi/olia +* + + ? IV I Glossopleris angusli/olia + + + + + + + + + ? + ;i- G. arberi +* + + ;:l n' G. barakarensis + + "' -c- G. browniana + + + + + + + + + ? ? §: G. clarkei +* + v c G. comlllUllis + + + + + + + + + ? o' :::l G. dal/ae (=G. damudica) + + + + + + + cf ? 2., G. decipiens + + 'J + ::; 0 G. elol/gala + + + + + + + ? 0; v: G. erJIarginala + + 2., + z 0 ~ G. l'Ulynelira +* + :r: V>'" G. indica + + + + + + + + + + + + I :::l '" G. inlennedia + + --l -V> :t- C c- G. inlermillens + + + X '< O G. karanpuraensis + z :::l 0 G. linl'aris + + <5. :s: ;3" G longi/olia +* + -< :::l :t- O G. major +* + Z 0 0 <5. G.oldhami +* + :;: 0 :: G.obscura +* + <: ;:; +;!: rn G. obovara sp. nov. ;;0 ~ G. pselldoslricra sp. nov. + Vl + =i Glossopleris pseudolortuosa sp. nov +* -< <5. 0 r. G. raniganjensis + + + + + "71 "' G. shailal' +* + --l ~ :r: :::l G. Sll'l/onellra + + + rn :;: '" G.lortuosa +~ + + CJ () r:l 0 Z :::l G. laeniopleroides +* + C/O V G. lenuinervis + + c C! ~. G. vlIlgaris +* + r- () +* v, G. wallonii + ~ () G. zeitleri + +* + "1:; ? -; Gangalllopleris sp. cf. G. buriadica + + +* r" "= G.obovara + + 'r. Palaeoviilaria sp. + + Gonophyl/oidl's (=G. cislel/a) + SCIIIUI/I + + Cordaicarpus + + +* Palllophyl/um gidiensl' sll. nov. +* Euryphyl/lIrJ1 whillianul1l + + Kawi~ophyl/Uln dunpalhril'l/sis + K. barakarl'l/sis sp. nov. +* Ginkgopsid + Vl'rlebraria indica + + + + + + + + + + + .... --J Equiselalean axes + + + + + + + + + + + Incertal' sedis +* 348 THE PALAE0130TANIST

North Karanpura South Karanpura West Bokaro Coaltields D K. R K S R G B S U J P K K T S A T G P a D. a a i c i h a r h u u e a a r 0 h i k Y r r I d u u i a II j d P r a P a II a H k k i i r n III r d \I I i u a t d r e a a g k d a k i a n h 0 r Glossopteris Species a s t a -A u a r h e a I a r \I i a r 0 a & d II a II a d g C

Glossopleris arberi +1 G. angllslijolia -h'• + G. browniana + + G. barakarensis + G clarkei + + G. cOl1lnl/lnis + + + + + + + + + + + + G. danae + + G. decipiens + + + + G. e[ongGla + G. I'l1Iorginara + G. eurynellra + G. indica + + + + + + + + G. inlertnedia + G illlerl1lillens + + G. karanpl/raensis + + + + + G. Iinl'aris + + + G. longijolia + G.l1Iajor + + + + G. oldhal1li + G.obscilra + + + G. obovara sp. nov. + + + + G. pselldolOrlllosa sp. nov. + + + + + G. psel/doslriCla sr. nov. + + + + G. slmilae + + G. Slenonellra + + G. rClniganjensis + G lenllinervis + + G.lo!1l/osa + G. laeniopleroides + G. vulgoris + + + G. walronii + + + + + + + G. z.eilll'ri +

Fig. 3-Colliery-wise distribulion of Glulloplai.\ species SINGH- TAXONOMY AND DIVERSITY OF THE GENUS GUJSSOP7LRIS 14')

Diversity in Glossopteris hydrological cycle. Di versity provides a base for ecologically sound system. Glossopleris {{/Ilpla, G. allgllsli{olia. G. Glossopteris diversified rapidly once established after brOlvllialla. G. COllllllllllis, G. illdica and G. slricra have world­ glaciation. There were two major phases of diversification. wide distribution (Fig. 1). viz: in Barakar and Raniganj Formations. In these periods maximum number of species diversified to occupy almost DISCUSSION & CONCLUSION every kind of terrestrial habitat. Over the whole long period. plant communities have been evolved. Diversification has The species of this geological past genus is often occurred as a result of competition between plants. There are confused with each other due to close similarities in their fragmentary plant fossils of Glossopteris from Talchir appearances. Most of the work published in the past was Formation and comparatively some better were recovered based on either morphological features (Size. Shape. Base. from Karharbari Formation. These plants were successful in Apex) or cuticular features and in this way a large number of unsupporting environment. They probably required several Glossopteris species have been created. In present study both adaptations: mechanical strength to support them in the air to the features together have been taken as rei iable parameters expose light catching surfaces. an anchoring system to prevent during identi fication and speciation. Large number of closely them being blown over, a conducting system to supply water related species have been minimised by making the species to all parts of plants, a system for obtaining mineral nutrients, complexes (e.g., G. arberi G. zeilleri (Thesis Shiv a means of restricting water loss in the desiccating environment Mohan Singh, 1998). Species complexes can be compared to of the air, a means of reducing and dispersing on land. the mountainous terrain where the major hills are named. In The earliest picture of the plant community comes from fact the method of naming the peaks is akin to the way we present study of Barakar Formation of Karanpura and Bokaro name the species. The names of the hills for instance are not Coalfields. These includes Glossopteris, Gallg(//llopteris, referred to the peaks alone but to the whole domain of the hill. Palaeovittaria, Palltophyllul/I, Ellrypllylllllli. Kawiz.ophyllllll1 A small shoulder or a sub peak on the major peak belong to etc. There were many short herbaceous species viz .. the major hill and any two hills can be clearly separated only if Neomariopteris lobi/olia and a new fossil (lilcetae sedis). there is distinct valley between them. Glossopteris, Gangamopteris and Palaeoviffaria were small Species complexes can be similarly considered as shrubs or small trees. In these genera leaves are in spiral and morphometric terrain's with species as the hills. varieties as whorls. Euryphyllul/l, PalltophyllwlI, Kawizophyllil//l were the shoulders Within the species domains. In other words in probably tree habit. The great adaptive radiation in genus this view species are not merely peaks (means) but are Glossopteris reaches its fullest expression in the Raniganj population domains isolated by reproductive valleys. However Formation. Glossopteris illdica, G. allguSIl!o!ia, Vertebraria as not all hills are distinctly isolated, not all morphometric illdica and Equisetalean axes have wide longitudinal peaks would be completely separated offering challenges to distribution from Talchir to Parsora. G. brolVllialla, G. the taxonomists. communis, G. elongata and G. stellollellm have comparatively Arber suggested that as the classification of Glossopteris narrow distribution from KarharbariIBarakar to Hinzir/Maitur is an artificial one it would be beller to maintain comparatively (Figs 1, 2, 3). G. barakarensis, G. decipiells, G. el/largillafa, few species by grouping together those species which differ G. euryneura. G. illtel'lnedia, G. k;:!rallpuraellsis, G. lillearis, in one or two characters but are not sufficiently dissimilar in G. longifolia, G. //Ia)ol; G. oldhami, G. obscura, G. obovara, the aggregaie of their characters. He also doubted the G. ranigan)ensis, G. tortuosa etc. show maximum diversity usefulness of creating varieties or sub-species and in this in Barakar and Raniganj Formations. Glossopteris flora owing connection he is amply supported by Edwards ( 1928, p. 325) to high degree of inaccessibility, had escaped from complete who says .'..... I think that the custom of applying varietal names transformation and hence exhibits high magnitude of bio­ to isolated fossil leaf impressions is to be deprecated.... The diversity at species level. Since the data is lacking at the Ie,,; use of trinomial nomenclature does not appear to add to the of gene and ecosystem, so bio-diversity at this level is avoided. convenience of this artificial classification." While the Probably genome of Glossopteris modified according to time tendency to create varieties or sub-species is to be deprecated and space and different species of Glossopteris ranges from it is equally true that a genus, howsoever artificial has to be Triassic to Parsora. The variation in shape and size withi n critically resolved into various specific components, whatever species of genus Glossopteris was due to mutation. Up to the their numbe~.Seward (1897) has very rightly observed that Parsora "Nature" conserved their germplasm which was the "while endeavouring to avoid dangerous and unscientific basis of diversity. The climax community of this genus practice of needlessly multiplying specific names. we must be naturally remains stable until another environmental change careful to bear in mind the possibility of carrying too far the Occurs. Bio-diversity-rich support area probably supply system of linking together distinct types by a long series of massive amounts of nutrients to livestock to maintain local intermediate forms." Arber was infact more interested in 350 THE PALAEOBOTANIST reducing the number of species by linking together, which Ackflowledgements-I {/III greu/ly illdeiJ/C'u /0 /-IK Mohe.l'h\Vuri resulted in aconglomeration of many disti ncttypes under fewer for C'lIcoliragell/ell! ulldfor cri/ica/ly goillg throllgh thl' 1IlOllliScript. names which sometimes became so unwieldly as to be of no I (1111also thanljllito the ProfAllsillt K Sillhu, Director. IJSII~LIlCkllolV stratigraphical value. His Glossopteris browi/iana may Jor providillg I/('cessw)' researchJacilities. particularly be cited as an example where different and distinct forms were huddled together, e.g., G. pawl/efa, G. lil/earis. REFERENCES G. taeniopferoides. etc. Further researches have. however, shown the Anderson JM & Anderson HM J985. Palaeoflora of southern Africa unsatisfactory nature of Arber's broad based specific : Prodromus ofSouth Afriean megafloras: Devonian to Cretaceous circumscriptions. 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