On two enigmatic. infructescences from Permian Gondwana of the Rajmahal Basin*

Usha Bajpai & Hari K. Maheshwari

Bajpai, lIsha &: Maheshwafl. Hari K 1991. On tWu enigmatic infructescences from Permian Gundwana of the Rajmahal Basin l'alaeobolal1ls1 39( I) : 9-19.

Two ne\V dlch()[omou~lybranched infructescences, a gynuclad and an androclad, are reponed from shales associated with the Lalmatla Coalseam in Hlira Coalfield, Rajmahal Basin, Bihar 11le branched infructescences though are found In association with ginkgophyte leaves, yet their affinities seem to be with the pteridospermous groups

Key-words- Frllcl.flcatlon~,G.nkgophytcs, Glossopteris Flora, Gondwana, Pteridosperms (India).

(Isba 13ajpat t- Han K ,l1abesbwan, Birbal Salmi Inslilule oj Palaeobolany, 53 IJniuersiry Road, GPO 130x 106, LucMnOll' JJ6 00 I, India riu

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THE Hura Coalfield is the northernmost outlier of from shales associated with the Lalmatia (II) top coal bearing formations in the Rajmahal Basin. Coal seam In the same collection we have now observed is being exploited in the area since long (Ball, 1877) the presence of Neomariopteris hughesii by the local people who intermittently dig shallow (Feistmantel) Maithy 1974, Glossopteris communis pits to collect coal for their use Raja Rao (1987) Feistmantel 1876, Glossopteris lanceolatus Pant & reports the occurrence of a few coalseams in the Singh 1971, Glossopteris ampla Dana 1849, Barakar Formation. Large scale mining is being done Glossopteris angustijolia Brongnia rt 1831, by the Rajmahal Open Cast Project in the Lalmatia Glossopteris linean's McCoy 1847, Glossopten's retusa bottom (L-II) and Lalmatia top (L-III) seams. Maheshwari 196'5, Glossopteris stricta Bunbury 1861, Plant fossils were reported from the Permian Glossopteris stenoneura Feistmantel 1877, beds near Lohandia by Feistmantel (1880). These Vertebraria indica Royle 1839, Eretmonia-type leaf­ included a few leaves of the genus Glossopteris, scale scales and Arberiella-type sporangia. Recently leaves and seeds. Almost a hundred years later, Maheshwari and Bajpai (in Press) have reported Singh, Srivastava and Maheshwari (1986) recorded a several species of ginkgophyre leaves from shale hete rophyllous spheno phyll- Sphenophyllu m intercalations between Lalmatia bottom and top gondwanensis-and the equisetalean Lelstotheca seams, exposed far away from the main exposures of the II coalseam in the Lalmatia-Simra area. The bed is rich in plant fossils Here we record two 'Contribution to lGCP Project 237: Floras of Gondwanic interesting types of fructifications from the same Continents. beds,

9 10 THE PAlAEOBOTANIST

+ F I v v v N + ~F/V v v v v + v v ~~ '--\V v v v v + \ --.1.....-r...----F~\ ~~\V v + + +\ v + + \ I v v v + + \\ \( /'-. F ( v v o 1 2 3Km I I I ~ \ v 25° "') + + FY,",~~\ v v 65° • "' \ r-, ~~, ~ Traps /' +"::>'" + ++ +"" ..~~,<' ~ ~,,~,~ ~ Precambrian ~~~, /'( + + Haripur + \- ~,\ _ Phulberia CJ Barakar + + + + <' ~,,\ / ~ Burnt out crop ,/ + + ( + " ~b\l Locality + + + + \Yl'l""\ '.V~ + + + .-, \ ./ v + I='"\ \ /(, + + + F "'\ .. .- .J +.J ,PA F F,/ v v + +/ r-\~'-'~-:!-/' + + +( / -=, \ 87° 20 1 I +

Map I-General location of the pit from where the fossils were collected.

MATERIAL

Plant fossils were collected by our colleagues ,1.5 m SANDSTONE Shri H. N. Boral and late Dr V. K Singh from an abandoned shallow pit, dug by local villagers for FERRUGINOUS SANDY SHALE coal, near the village Haripur (Map 1), in the 1.2,,, WITH GLOSSOPTERIS LEAVES southern part of the Hura Coalfield. Subsequently we AND TRACE FOSSIL LAYERS + + .25m SHALE WITH GLOSSOPTERIS LEAVES collected additional specimens from the localiry and - - --- _. 3 m UNFOSSILIFEROUS SHALE Ii. <> .. Ii. e <> ". .. observed the following sequence in this particular <> <> ".6 <> Ii. <> 6 " .6 m GRITTY FERRUGINOUS SANDSTONE li 6 Ii. <> ,0 ...... 6. ° pit (Text-fig_ 1) 1 WITH FEW GLOSSOPTERIS LEAVES The rype and figured specimens have been deposited in the Repository of Birbal Sahni Institute ~1_'-'-'-'-'-'-'- SANDSTONE/SHALE ALTERNATIONS of Palaeobotany, Lucknow. 3 m RICH IN PLANT FOSSILS INCLUDING GINKGOPSID LEAVES; OCCASIONALLY DESCRIPTION TRACE FOSSILS SEEN Veekaystngbta gen, nov,

Type species- Veekaysinghia durgavatiae sp. 12 m COAL nov. D{agnosis-Seed-bearing axis, repeatedly branched on a basic dichotomous pattern; ultimate ,1 m SANDSTONE; BASE NOT SEEN branches sometimes apparently pinnate, bear aphyllous, orthotropous seeds, usually compactly Text-figure I-A generalized litholog of the face of the pit from arranged and only on one side of the axis. which the fossils have been collected. 'IPI J];j ;)lj1 UO U;);)S SI J];;)I plsd08)fu,8 ]; JO lJ];d !];Jld]; : I x '! L£9£ ou LI;)Wl:!;)dS 'lJ];dJ;)lUnOJ :;)d,\IOIOH 'z Jlelj IPI J;)/6ol LII U;);)S oSle SI lItril/l(qdO/l;lqdS JO ;)1/61 V J];;)I p1Sd08)fU18 ]; Aq p;)ddep;),\o ...... 'I 'I ," I"til 'lIlJd "lj I 10 1.1 rei I·' \\"I ,>lj II • "_ ~l)~' Oll 1I.'1l11 "'d' _'d.\ \"1\ 'II 1 ,. 'll d' I.) 1I.'" ,1/ '1/" 111:-11111' 1'''<71(111<.1 /I,~.l.l \- .. ' 1 ~;u n8H 12 THE PALAEO BOTANIST

Figure 3- Veekaysmghta durgavaliae gen et sp nov.. a pan of the holorype enlarged to show 'pinnate' ultimate branches bearing closely placed seeds on one side of (he axis (upper left of the figure); specimen no. 36570, • 2.5.

Description-It is rather an unusual dichotomy have been observed (Figures 3, 4). Fine infructescence for the Gondwana. Most of the hand striations run all along the axis (Figure 5). The specimens show only portions of the infructescence. infructescences are monogynons on which Only one specimen (Figure 1), and its counterpart orthotropus seeds are compactly arranged (Figures (Figure 2), seem to represent almost the complete 6·11). The seeds are aphyllous, in not being organ. The specimens are profusely branched on a associated with a sporophyll or scale, sessile and basic dichotomous pattern. The ultimate branches almost invariably si tuated on one side of the axis, seem to be lateral (Figure 3), probably due to usually overlapping one another (Figures 8, 9). suppressed dichotomy, i.e., helicoid branching. Seeds are obovate in shape, 35·4 mm long and The complete infructescence was atleast up to up to 2.5 mm broad at the Widest, neither micropyle 18 cm long with first dichotomy of the axis occurring nor hilum is seen. As the seeds are preserved only as about 6 cm from the base. Up to five levels of impressions, details of wall layers are not clearly BAJPAI & MAHESHWARI-ENIGMATIC INFRlJCTESCENCES FROM PERMIAN GONDWANA 13

Figure 4- Veekaysmghia durgUl·alw<: gen el sp nov a gynoclad assoCiated with ,everal glllkgorsid leaves A well preserved leaf of Saporlaea is seen on (he left side, specimen no 36574, x L decipherable At some places one can see Comparison-The striking features of the indications of two integuments and an almost described gynoclad are a dichotomous branching rounded nucellus (Figures 12, 13 l, which is lip to 2 pattern, and orthotropus seeds compactly arranged mm in diametre Under incident light cellular on ultimate branches. The infructescence compares outlines are faintly discernible The cells are with the gynoclad Utkalia dichotoma described from recranguloid and arranged end·to·end. the Hinjir (= Kamthi 1 Formation exposed in the Diagnosis oj species-As for the genus. Hinjirida Ghati near Handapa, Mahanadi Basin Derivation oj generic name-After late Vi nay (Chandra, 1984 l. [n both the infructescences, the Kumar Singh, our associate on the Rajmahal Basin primary axis is dichotomously branched and the studies. ultimate axes bear aphyltous seeds. However, in Derivation oj species name-After Durgavati, Utkalia dichotoma further branching is on an the valiant Queen of the principality of Gondwana, alternate pattern, and the ultimate branches are central India, circa 16th Century A.D. (Chopra, reported to bear a Single terminal seed. An 1973 ). androclad organizationally similar to Utkalia has 14 THE PAIAEOBOTANIST

Stapbidiopbora secunda Harris 1935 consists of an axis that bears a few sessile seeds in its upper part and only on one side as in Veekaysinghia durgavatifle. According to Harris (1935), the seeds were probably borne alternately but as a result of twisting of axis came to be located on one side. On the basis of similarity in cuticular features, Stapbidiophora is attributed to the leaves of Hartzia tenuis. Caytonia is a monogynon that bore 'fruits' opPositely or sub-oppositely on a dorsiventral rachis. It is attributed to the leaf Sagenopteris ;Ilongwith the androclad Caytonanthus on the basis of regular association (Sporne, 1965). The ginkgoalean female 'cone' reported from Molteno Formation exposed at Matatiele in the Karoo Basin (Anderson & Anderson, 1983, pI. 24, fig. I) superficially resembles our specimens in being a gynoclad. Details of the African fossil are, however, not yet known. Holotype-Specimen no. 36570, Birbal Sahni Institute of Palaeobotany, Lucknow; Early Permian, Barakar Formation, shales associated with Lalmatia Coalseam, near village Haripur, Hura Coalfield, Rajmahal Basin, India.

Btrbalsabnta gen. nov. Type species-Birbalsahnia divyadarshanii sp. figure 5- \ ('('k'UI'\IIIJ..:h/tI dl/l).~(4U'/I(H'gt'n vi ...p Ill1\ :1 p.lrl (l! nov. the gynodad enlarged l() ~h()wa dichotomy of [he axi~and Diagnosis-And roclad, main axis dichotomously panly exposed seeds, apparenlly attached on one side of the axis; specimen no. 36S71, x 4. branched, subsequent branches lateral, alternate to one another, ultimate branches thin, terminally been reported from the Upper Triassic Molteno bearing pollen-sacs, attachment of pollen-sacs on Formation (Anderson & Anderson, 1983, pI. 26, fig. ultimate branches not discernible; pollen-sacs 1) apparen tIl' tetra-loc ulaf. The corystospermaceous gynoclads referred to Description- The incomplete pollen ·sac· the genus Umkomasia Thomas 1933 are also bearing organ (Figure 14) occurs in the same profusely branched. However, the branches arise sediment layers that have yielded ginkgopsid leaves laterally in the axils of bracts and bear cupules in and the branched seed-bearing organ Veekaysinghia. imparipinnate manner (Holmes, 1987). This The preserved length of the main axis is about 4 em; gynoclad has been linked with DicrOidium leaves on width is S-6 mm. However, it is not possible to the basis of constant association in the sediments. determine whether the main axis is the principal Wankiea bondii, a gynoclad from the Early axis of the androclad vI' it is a branch of the principal Permian of Zambia (Lacey & Huard-Moine, 1966), axis. The two-level dichotomy (If the main branch differs in having opposite/subopposite lateral gives an apparent Jecussate pattern (Figure 17). branches and a terminal cluster of 4-6 'seeds'. It has Subsequent and ultimate branches are alternate been suggested that Wankiea may be a (Figure 1S). Quite often ultimate branches are seen microsporangiate fructification of Pecopteris arcuata only on one side and that too at irregular intervals (Huard-Moine, 1964). (Figure 17). Pollen-sacs could have been terminal to Seeds of Ampborispermum ellipticum Harris the ultimate branches; however, actual attachment is 1932 aI'e arranged in a manner that simulates the not seen. They are found either Singly or in groups arrangement in our specimens. However, in the of 3-7 (Figures 15, 16). Pollen-sacs are oblong, former, the main axis and branches are unknown probably tetra-locular. No pollen grain was and the terminal branches have only been inferred recovered from the pollen·sacs. Probably all the (Harris, 1981). pollen was shed before preservation. Scanning BAJPAl & MAHESHWARI-ENIGMATIC INFRUCTESCENCES FROM PERMIAN GONDWANA 15

Figures 6-9- Veekaysinghia durgavatiae gen. et sp. nov. 6,7 Part and counterpart of anorher gynoclad shoWIng closely placed seeds, specimen nos. 36576, 36577, x 1. 8. Disjoinred axes showing seeds in lateral view; specimen no. 36575, x 1 9 Disjointed axes, apparently bearing seeds only on one side, specimen no. 36573, x 1. electron micrograph of broken surface of a pollen­ Den'vation of species name-After Professor sac shows that probably hyaline silica (chalcedony) Divya Darshan Pant. has infiltrated the pollen-sac. At places, degradation Comparison-The infructescence is comparable of pollen-sac surface could be seen clearly A few to the peltaspermaceous androclad Anteusia Harris minute rounded bodies and filaments (fungal) have 1937 from the Triassic. The latter. however, is also been observed. Another fertile axis is present bipinnate, with alternate primary branching, and less which shows opposite to sub-oPPosite branches on regularly arranged secondary branches. Pollen-sacs both sides of the axis are with longitudinal line of dehiscence. Diagnosis of species-As for the species. Pteruchus Thomas 1933, the androclad of the Den'vation of generic name-After Professor Corystospermaceae, known from the Triassic of the Birbal Sahni (1891-1949)_ Gondwana Supercontinent differs in having a central 16 THE PALAEOBOTANIST

Figures 10-13- Veekaystnghia durgavatiae gen. et sp. nov. Portions of gynoclads enlarged to show attachment and nature of seeds. A nucellus is seen in one seed each in figures 12 and 13. 10.11. Specimen no. 36574, x 3; 12,13. specimen no. 36572, x 4. axis which bears short lateral branches, generally in Holotype-Specimen no. 36'578, Birbal Sahni one plane The pollen-sacs are borne on the Institute of Palaeobotany, Lucknow; Early Permian, underside of peltate head_ The dehiscence again is Barakar Formation, shales associated with Lalmatia longitudinal Coalseam, near village Haripur, Hura Coalfield, Caytonanthus Harris 1937 (CaytOniaceae), from Rajmahal Basin, India. the Jurassic, is a dorsiventral axis bearing opposite/sub-opposite pinnae_ The pinnae are AFFINITIES branched irregularly and each terminal branchlet Veekaysinghia is a very unusual type of bears a single 'anther' comprising four chambers, fructification for the Gondwana. It is no doubt a thus comparing closely with the pollen-sac of the present infructescence. ..-+ Antholithus Brongnian 1822 comprises a Figures 14-17- Birbalsahnia divyadarshanil gen. et sp. nov. 14. HoJotype: specimen no. 36578, x 1 15. The lower right dichotomously branched axis, bearing terminal portion of the holotype enlarged to show the 'foliose' nature clusters of pollen-sacs According to Nathorst (1908) of the branches, x 4. 16,17 Upper and lower portions, the genus Antholithus is affiliated to a ginkgoalean respectively of the holotype enlarged to show the branching or cycadalean group. pattern and clusters of pollen-sacs, x 7.5 BAJPAl & MAHESHWARI-ENIGMATIC INFRUCTESCENCES FROM PERMIAN GONDWANA 17

Figures 14-17 18 THE PALAEOBOTANIST gynoclad as each ultimate branch bears a number of shoot' habit was also a later feature (Archangelsky, compactly arranged obovate bodies-the seeds. The 196'5, p. 136). According to Meyen (1987, p. 1'5'5) seeds have an sclerotesta surrounded by a flat independent status of the Family Karkeniaceae is border, the sarcotesta. Some of the specimens even still inopportune. Thus, it seems so far no fossil show a third region, the nucellus (Figures 12, 13). infructescence can definitely be aSSigned to the No subtending bract or sporophyll has been Ginkgoales. observed so far. The arrangement of the seeds is also The basic pinnate branching pattern and the very peculiar as all the seeds are apparently on one flattened axis of Birhalsahnia suggest modified side. There are two explanations. One, the seeds pinna in which the lamina got completely reduced. were spirally disposed but due to twisting of the axis The faint impressions of cells seen over the flattened now occupy same plane; second, the ultimate axis also indicate a modified pinna. The Early branches are fertile pinnules with reduced lamina, Permian androclad Callipterianthus also has each bearing seeds in a linear row. modified pinnae with a completely reduced lamina In the latter case, each gynoclad may be (Meyen, 1987, p. 149) compared to a modified frond,' thus indicating Thus the data at hand seems to indicate the affinities with Pteridosperms. However, so far there presence of a Pteridosperm·related group in the is no supporting evidence, like the presence of a assemblage. This group does not have affinities with cupule, or a manoxylic wood, at this level except for the true glossopterids, dominant elements of the occurrence of Sphenopteris· and Callipteris-type Permian vege~ationof the Gondwana of foliage. The Permian Gondwana Sphenopteris Supercontinent. fronds are now known to be definite ferns (Neomariopten·s/Damudopteris). Callipteris-type of REFERENCES frond is a new find and though its taxonomic status is yet to be ascertained, it does occur in the same Archangelsky, S. 196'). Fossil Ginkgoales from the Tic6 Flora, horizon as Veekaysinghia. Santa Cruz PrOVince, Argentina. Bull. Br. Mus. nat. His!. The flora at this level is dominated by (Ceol) 10 : 121·137. ginkgophyte leaves, such as Rhipidopsis, Ginkgoites, Anderson, J. M. & Anderson, H. M. 198'). Palaeoflora of southern Africa. Prodromus of South Afric.:an megafloras, Devonian 10 and Saportaea. Though the fructification has not Lower Cretaceous. A. A. Balkema, ROlwrdam. been found in organic connection, yet a ginkgophyte Ball, v. 1877 Geology of the Rajmahal Hills. Mem geol. SlIrv. affinity can also not be negated. The female India 13. ginkgoalean 'cone' recorded from Late Triassic Brongnia r :, Adolphe 1822 Sur la classification et la distri· bution des vegetaux fossiles en general et sur ceux des Molteno Formation (Anderson & Anderson, 1983, pI. terrains de sediment superieur en paniculier. Mem. Mus. 24, fig. 1) also has a superficial resemblance. Hist. nat Paris 8 : 203- 348. Staphidiophora secunda Harris 193'5 from the Chandra, Shaila 1984. Utkalia dicholOma gen. et sp. nov.-a fossil Rhaetic of Greenland, which on the basis of fructification from the Kamthi Formation of Orissa, India. association is believed to belong to ginkgoalean leaf Palaeobotanist 31 : 208·212. Chopra, P. N. (EdilOr) 1973. The Cazeteer of India. Indian Union Hartzia Harris, shows a comparable arrangement 2, History and Culture. Publications Division, Government of and organization of seeds. India, New Delhi. So far only two fossil seed-bearing organs were Feistmantel, Ollokar 1880. The fossil flora of the Lower Gondwana definitely assigned to the ginkgophytes. Both are as 2. The flora of the Damuda and Panchet divisions. Mem. geol Surv. India Palaeont. indica, ·ser. 12, 3: 1·77. removed from each other as are from Veekaysinghia Florin, RUdolph 1949. The morphology of Trichopilys hetero­ or from extant Ginkgo. Florin (1949) reasoned in morpha Sapona, a seed plant of Palaeozoic age and the evo· detail to regard the Palaeozoic taxon Trichopitys lution of female flowers in the Cinkgoinae. Acta hort. Ber· heteromorpha de Saporta 187'5 as a ginkgophyte, giani 15 : 79·109. albeit a primitive one. Meyen (1987, p. 148) does Harris, T. M. 1932. The fossil flora of Scoresby Sound, East Green· not agree with Florin's interpretation. According to land·} Medd. Cnt>nland 85(')): 1·133. Harris, T M. 193'). The fossil flora of Scoresby Sound, East Green him the Family Trichopityaceae should be placed land·4. Ginkgoales, Coniferales, Lycopodiales and isolated under the Order Peltaspermales. Archangelsky fructifications. Medd. Cnt>nland 112( I) : 1·176. (196'5) suggests that the Early Cretaceous taxon Harris, T M. 1937 The fossil flora of Scoresby Sound, East Green· Karkenia incurua "may well be an intermediate type land·,). Medd. Cnt>nland 112(2): 1·114. of female structure ("flower") between Trichopitys Harris, T. M. 1981. Amphorispermum, an enigmatic assemblage. Palaeobotanist 28-29: 210·217. Ginkgo". and In Karkenia the' inverted, collar-less Holmes, B. K. 1987. New COlyslOsperm ovulate fructifications ovules/seeds are arranged compactly on the axis. from the Middle Triassic of eastern Australia. Alcheringa 11 . Collar has been assumed to be a secondary feature 16')·173 of the Ginkgo ovule (Florin, 1949). Probably 'short Huard·Moine, D. 1964. Presence de Pecopteris arcuata Halle dans BAJPAI & MAHESHWARI-ENIGMATIC INFRUCfESCENCES FROM PERMIAN GONDWANA 19

une flora fossil dite "a Glossopteris" d'Afrique du Sud. Ann. fossiter Pflanzenreile. Palaeobot. Mitt. 4·6. K. svenska Univ. AR.E.R.S Reims 2 : 123·129. Vetensk· Akad. Handl. 42(6) : 3. Ulcey, W. S. & Huard·Moine, D. 1966. Karroo floras of Rhodesia Raja Rao, C. S. (Editor) 1987. Coalfields of India. 4 (1) Coal and Malawi·Pall 2. The Glossopteris Flora in the Wankie resources of Bihar (excluding Dhanbad District). Bull. geol. Disrrict of Southern Rhodesia In: Anonymous (Ed.)-Sym· Surv. India ser. A( 45) : 1·336. • posium on jloristics and stratigraphy oj Gondwanaland. Singh, V. K., Srivastava, A. K. & Maheshwari, H. K. 1986 Sphenop· Birbal Sahni Instirure of Palaeobotany, Lucknow, pp. 13·25. sids from the Barakar Formation of Hura Tract, Rajmahal Meyen, S. V. 1987 Fundamentals oj palaeobotany. Chapman & Hills, Bihar. Palaeobotanist 35 : 236·241. Hall, London, New York Sporne, K. R. 1965. The morphology oj gymnosperms. Hutchinson Maheshwari, H. K. & Bajpai, Usha (in Press). Ginkgophyte leaves University Library, London from the Permian Gondwana of Rajmahal Basin, India. Thomas, H. H. 1933. On some pteridospermous plants from the Palaeontographica. Mesozoic rocks of South Africa. Phil. Trans. R. Soc. Lond. Narhorst, A. G. 1908. Ober die unrersuchungen kutinisierter 8222 : 193·265.