Cretaceous Research (2001) 22, 565–574doi:10.1006/cres.2001.0277, available online at http://www.idealibrary.com on

Fungal remains from Early CretaceousIntertrappean Beds of Rajmahal Formation inRajmahal Basin, India

Archana Tripathi

Birbal Sahni Institute of Palaeobotany, 53 University Road, Lucknow 226007, India; e-mail: atripathi@bsip.res.in

Revised manuscript accepted 19 June 2001

This paper records fossil fungi from palynologically dated Early Cretaceous Intertrappean Beds of the Rajmahal Basin, Bihar,India. They include microthyriaceous remains, spores and hyphae. One new species, Dendromyceliates rajmahalensis, isreported which bears conidia. The fungal remains suggest that the climate was warm and humid during the deposition of theintertrappean beds. � 2001 Academic Press

K W: fungal remains; Early Cretaceous; Intertrappean Beds; Rajmahal Basin; India.

1. Introduction

The Mesozoic sedimentary strata of the RajmahalBasin are richly fossiliferous. The plant remains re-corded as impressions, compressions and petrifactionsare leaves, fruits, seeds, wood, and spores and pollengrains. Rao (1943) described from the Rajmahal Floraa septate mycelium in the basal part of the coniferseed Nipaniostrobous Sahni. Tiwari & Tripathi (1995)reported the microthyriaceous Phragmothyrites frompalynological preparations of intertrappean sedimentsin the Rajmahal Basin. This latter finding instigated adetailed search for more fossil fungal remains in thebasin. The variety of fossils encountered forms thebasis of the present paper.

2. Material studied

This study is based on subsurface intertrappean bedsin the Rajmahal Basin. The fungal remains wererecovered from the following boreholes (Figure 1):RJNE-32 and RJNE-35 in the northern extension ofHura Coalfield, and RCH-151 in the ChuperbhitaCoalfield.

The intertrappean beds penetrated by these bore-holes have been dated palynologically as Berriasian–Aptian (Tiwari & Tripathi, 1995; Tripathi, 1998,2001). The slides of the specimens illustrated arehoused in the Museum of the Birbal Sahni Institute ofPalaeobotany, Lucknow.

0195–6671/01/050565+10 $35.00/0

3. Systematic palaeontology

The fungal remains recorded and described hereinclude microthyriaceous fruiting bodies, spores andhyphae (Figure 2, Table 1). Some remains are de-scribed under an incertae sedis heading as their affinityis uncertain.

Fruiting bodies

Genus: Parmathyrites Jain & Gupta, 1970

Type species. Parmathyrites indicus Jain & Gupta, 1970

Parmathyrites sp.Figure 3B

Description. Observed specimen incomplete, radius85 �m, nonostiolate; cells aporate; central cells irregu-lar, hexagonal; peripheral cells rectangular, radiating,10–21 �m long, 6–9 �m wide; cell wall >1 �m thick,marginal cells broken, appear to end in robust spines.

Remarks. Only one incomplete specimen has beenobserved. The peripheral spines are broken and theirnature is not clear. Hence, it could not be assigned toany species.

Genus: Paramicrothallites Jain & Gupta, 1970

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566 A. Tripathi

Type species. Paramicrothallites (Microthallites) spinula-tus (Dilcher) Jain & Gupta, 1970

Paramicrothallites sp.Figure 3C

Description. Stromata subcircular, 145.5 �m in diam-eter, ostiolate, margin smooth, undulating; cells apo-rate, rectangular to penta-hexagonal, 2–4�2–4 �m inmaximum diameter; cell wall 1 �m thick. Ostiole 9 �min diameter, not surrounded by specialized cells. Freehyphae not seen.

Remarks. The present specimen differs from Para-microthallites spinulatus (Dilcher) Jain & Gupta (1970)and Paramicrothallites menonii Jain & Gupta (1970)owing to an absence of spines on the margin of thefruiting body and in the nature of the cells. Paramicro-thallites edvensis Rao & Ramanujam (1976) differs inhaving a crenate margin.

Genus: Trichopeltinites Cookson, 1947

Type species. Trichopeltinites pulcher Cookson, 1947

Trichopeltinites fusilis Dilcher, 1965Figure 3G–I

Figure 1. Map showing location of the boreholes studied in the Rajmahal Basin, India.

Figure 2. Lithological logs of sediments of the RajmahalFormation overlying the Dubrajpur Formation(Triassic–Early Cretaceous) in the three boreholes ex-amined and levels of samples yielding fungal remains.The logs show the sequence of trap and intertrappeanbeds (base of Rajmahal Formation not indicated).

Description. Stromata strap- ribbon- or fan-shaped,nonostiolate, simple, unbranched, margin irregularlylobed 126.5–218 �m long; one specimen very broad,about 200 �m; one layered; cells rectangular, 4–15 �mlong, 3–6 �m broad; cell wall 1–1.5 �m thick; radialwall thicker than the horizontal walls, smooth; cellsaporate, appear to be structured or filled with horizon-tal bar-like projections. Individual hyphae radiating,dichotomizing; branching prominent towards margin.

Fungal remains from the Rajmahal Formation, India 567

Genus: Phragmothyrites Edward, 1922

Type species. Phragmothyrites eocaenicus Edward, 1922

Phragmothyrites eocaenicus Edward, 1922Figure 3D

Description. Stromata circular to subcircular in shapewith entire margin, nonostiolate, 73�79–79�93 �min diameter. Hyphae radially arranged, dichotomouslybranched, forming pseudoparenchymatous aporatecells arranged in concentric rings except in centralregion where they are polygonal and irregularly ar-ranged; rectangular cells 6–12 �m long, 3–70 �mwide; polygonal cells 6–12 �m in diameter.

Remarks. Some specimens have been encountered thatare similar in all respects to the material described

above except that the peripheral cells are globularand not rectangular. Specimens with eccentric ringsand asymmetrical growth have also been recorded,and many incomplete specimens have been found inwhich the central portion of the stromata is not clearand, therefore, could only be identified to genericlevel. This was despite the fact that other details areclear: smooth, undulating margin; aporate peripheralcells that are rectangular or globular to hexagonal, andarranged in radiating pattern.

Phragmothyrites sp.Figure 3A

Table 1. Distribution of fungal remains in intertrappean beds of the Rajmahal Basin, Bihar, India.

Borehole RJNE-32

Taxa/Sample depth (m) 259.00 255.00 250.00 249.00 246.00 244.00 223.00 219.40 155.40 95.50

Phragmothyrites eocaenicus + + + + + + +Phragmothyrites sp. + +Trichopeltinites fusilis + + + + +Paramicrothallites sp. +Dendromyceliates rajmahalensis + +Paramicrothallites sp. +Multicellaesporites sp. A +Microthyriaceous germling +Spore Type B +Hyphae Type A +Hyphae Type B +Hyphae Type C +Setae Type A +

Borehole RJNE-35

Taxa/Sample depth (m) 160.00 158.10 110.50 110.00 44.00

Trichopeltinites fusilis +Phragmothyrites eocaenicus +Multicellaesporites sp. A +Multicellaesporites sp. B +Pluricellaesporites sp. A + +Spore Type A +Hyphae Type B +

Borehole RCH-151

Taxa/Sample depth (m) 18.00 16.00

Multicellaesporites sp. A +Phragmothyrites eocaenicus + +

Remarks. The single specimen found is illustratedhere. It has elongate cells. Its fragmentary naturerenders the central region unclear. It resembles the

568 A. Tripathi

Fungal remains from the Rajmahal Formation, India 569

specimen described by Kar et al. (1972, pl. 2, fig. 24)as Phragmothyrites sp.

Microthyriaceous germlingFigure 3E

Description. Germling 62 �m in maximum diameter,irregularly lobed; no cells and no septa seen; wall�1 �m thick.

Fungal spores

Genus: Multicellaesporites Elsik, 1968 emend. Sheffy &Dilcher, 1971

Type species. Multicellaesporites nortonii Elsik, 1968

Multicellaesporites sp. AFigure 3L, M

Description. Fungal spore filamentous in shape, com-prises 7–12 cells; length 30.5–133.5 �m, width4–15 �m; cells broader towards apical end and narrowtowards base; inaperturate; wall laevigate, <1 �mthick; septa prominent, complete, <1 �m thick.

Remarks. The specimens are similar to those describedby Singh et al. (1986) as Multicellaesporites sp. A.However, they differ in having a larger size range, agreater number of cells, and cells that are folded alongthe long axis of the spore resulting from collapse of thecell wall. Some specimens have a long slender stalk atthe narrow end (Figure 3J).

Multicellaesporites sp. BFigure 3N

Description. Spore filamentous in shape, 21–30.5 �mlong, 3–5 celled; cells 6–8 �m wide, thin walled,broader at one end, become narrow towards base;inaperturate; wall laevigate, <1 �m thick; septaprominent, complete, �1 �m thick.

Remarks. The single specimen of this fungal spore ispresent at the ends of aseptate, smooth-walled hyphaethat are 3–5 �m wide.

Multicellaesporites sp. CFigure 3O

Description. Fungal spore elongate with one end widerthan the other; size 36�12 �m; 5 cells, widest in themiddle; basal cell small, maximum diameter 6 �m;wall <1 �m thick, laevigate; pore not seen; septadistinct, complete; cell wall outline notched nearsepta.

Remarks. The shape of the spore is different fromknown species; hence, it is described under opennomenclature.

Genus: Pluricellaesporites van der Hammen, 1954emend. Sheffy & Dilcher, 1971

Type species. Pluricellaesporites typicus van derHammen, 1954.

Pluricellaesporites sp.Figure 3P

Description. Fungal spore elongate, 4–15 globularcells; dimensions 6–21.5 �m long, 9–24.5 �m wide;wall <1 �m thick; septa thin and porate; few have athin area which may represent a pore; cell-wall outlinenotched adjacent to the septa.

Spore Type AFigure 3J, K

Description. Spores long, tapering at ends, consist ofthree cells of different sizes; apical and basal cellssmaller and conical; sometimes apical cell detachedfrom the middle cell; apical cell 6 �m long; basal cell17–24 �m long, pointed at base; middle cell rectangu-lar with flat ends, 18–36 �m in length. Wall smooth,laevigate, <1 �m thick.

Spore Type BFigure 3F

Figure 3. A, Phragmothyrites sp., BSIP Slide 12544. B, Parmathyrites sp., BSIP Slide 12540. C, Paramicrothallites sp., BSIPSlide 12540. D, Phragmothyrites eoacenicus, BSIP Slide 12546. E, microthyriaceous germling, BSIP Slide 12552. F, sporeType B, BSIP Slide 12552a. G–I, Trichopeltinites fusilis. G, BSIP Slide 12547; H, BSIP Slide 12542; I, BSIP Slide 12545.J, K, spore Type A, BSIP Slide 12550. L, M, Multicellaesporites sp. A, BSIP Slide 12548. N, Multicellaesporites sp. B, BSIPSlide 12549. O, Multicellaesporites sp. C, BSIP Slide 12540. P, Pluricellaesporites sp., BSIP Slide 125543. Scale barrepresents 10 �m.

Description. Specimens filamentous, 20–30 celled;cells 6–12.5 �m wide, 4–10 �m long; apical cell coni-cal, 5–7.5 �m wide, 5 �m long, tip rounded; wall

570 A. Tripathi

smooth �1 �m thick; septa 1–2 �m thick, incompletein places.

Remarks. Ramanujam & Rao (1978) described formsunder Multicellaesporites? which are similar to thespecimens described here; however, no details ofapical portion is available in their specimens.

Fungal hyphae

Genus: Dendromyceliates Jain & Kar, 1979

Type species. Dendromyceliates splendus Jain & Kar,1979

Dendromyceliates rajmahalensis sp. nov.Figure 4F–K

Holotype. Figure 4G, BSIP Slide No. 12540.Isotype. Figure 4F, BSIP Slide No. 12540.Derivation of name. From Rajmahal Formation.Type locality. Borehole RJNE-32, depth 95.50 m,Rajmahal Basin, Bihar, India; Rajmahal Formation,Early Cretaceous.

Description. Fungal hyphae dark brown, 3–3.5 �mwide, <1 �m thick, smooth, dichotomously branched,rounded tips, septate; septa thin; cells aporate. Freeends of hyphae bear globular, round-headed, smoothstructures: conidia separated by thin septa. Inplaces hyphae bear small (<0.5 �m long) baculateprojections.

Comparison. This species differs from the type speciesof the genus in the nature of branching of the hyphae(many times instead of 3–4 times), and in havinghyphae with rounded tips and showing globularstructures (conidia) at some of the free ends.

Remarks. In incomplete specimens it is sometimesdifficult to locate the principal hyphae because thehyphae are profusely branched.

Incertae sedis

Hyphae Type AFigure 4A, E

Description. Hyphae stout, septate, linear, 5–6 �mwide, branching at ends; wall bears fine baculateprojections <0.5 �m in height. The terminal or freeends bear two globular, balloon-shaped smooth struc-tures (conidia) 24–5�9 �m in size.

Remarks. Small (diameter 3–4.5 �m) alete, ovatespores are associated with the hyphae. These arelinearly arranged in groups of four. They are laevigatewith a central fold, and have thin (<0.5 �m) walls.

Hyphae Type BFigure 4D

Description. Hyphae 4–6 �m wide, thin walled, septate;wall constricted at the terminal septum; terminal isinflated and 24–26 �m long, 4–10 �m wide.

Hyphae Type CFigure 4C

Description. Hyphae 6–9 �m wide, septate; septa�1 �m thick. The ends of hyphae are trifurcated,resulting in three balloon-shaped elongate structures18–33.5 �m long, 6–9 �m wide.

Setae Type AFigure 4B

Description. Setae dark brown, ribbon like, 170–188 �m long, 3–5 �m wide at base, with pointed tips,unbranched, occur in a bunch, base not distinct.

Figure 4. A, E, hyphae Type A, BSIP Slide 12540. B, setae Type A, BSIP Slide 12541. C, hyphae Type C, BSIP Slide12542. D, hyphae Type B, BSIP Slide 12542. F–K, Dendromyceliates rajmahalensis sp. nov., BSIP Slide 12540. F,isotype; G, holotype showing conidia (arrow) at free ends of hyphae; H, nature of dichotomous branching; I, showingseptate hyphae (arrow). J, K, same specimen in different focus showing complete hyphae from base to ultimate branches.Scale bar represents 10 �m for A–G, I; 50 �m for H, J. K.

Discussion

The presence of fungal spores and microthyriaceousremains are well documented from Tertiary depositsin India (Ramanujam & Rao, 1973, 1978; Jain, 1974;Kar & Saxena, 1976; Ramanujam, 1982; Reddy et al.,1982; Varma & Patil, 1985; Patil & Ramanujam,1988; Tripathi, 1988; Saxena & Misra, 1990; Rao,1995, 2000; Kumar, 1996; Chandra & Kumar, 1997;Rao & Nair, 1998; Mandoakar, 2000; Kumar et al.,2000; Saxena, 2000). Banerjee & Misra (1968) re-ported fruiting bodies of fossil fungi from UpperCretaceous deposits in southern India. The EarlyCretaceous record of microthyriales is from theRajmahal Basin (Tiwari & Tripathi, 1995), and ofspores from the western Bengal Basin (Bera &

Fungal remains from the Rajmahal Formation, India 571

572 A. Tripathi

Figure 5. Stratigraphic occurrences of fossil microthyriaceous and other fungal taxa (recorded during present study) in India,and northern and southern continents.

Banerjee, 1997). As described above, the presentassemblage of fungal remains recovered from thesubsurface sediments of the Rajmahal Basin is repre-sented by seven genera, four of which belong tomicrothyriaceous fungi.

From the available published data it is noted thatthe occurrence of fruiting bodies of fossil fungi ismeagre in Cretaceous deposits (Elsik, 1978). Thepre-Tertiary records are from Antarctica (Cranwell,1968; Dettmann & Thomson, 1987), northwesternAlberta (Singh, 1971), South America (Hammen,1954), West Africa off the Senegal Coast(Venkatachala et al., 1998). Most of these weredescribed as fungal spores and fruiting bodies butnot assigned to genera. The genera to which thespecimens described here are attributed are well docu-mented and illustrated from the Tertiary deposits theworld over (Edwards, 1922; Johnson, 1949; Hunger,1952 in Elsik, 1978; Chinguryayeva, 1953, 1956, inElsik, 1978; Martin & Rouse, 1966; Cranwell, 1968;Ebtehadj, 1969 in Elsik, 1978; Hopkins, 1969; Sheffy& Dilcher, 1971; Elsik & Dilcher, 1974; Song &Zhichen, 1998; Dutra & Batten, 2000; Palamarczuk& Barreda, 2000). The reporting of fossil fungalremains from the intertrappean beds in the basin is

significant because it extends the range of severalmicrothyriaceous and spore taxa from the Tertiaryback to the Early Cretaceous (Figure 5).

The occurrence of epiphyllous microthyriaceousfruiting bodies suggests that the climate was warm andhumid at the time of deposition of the sediments.This ties in with a similar palaeoclimatic conclusionthat was drawn by Tripathi (1995) on the basis ofpteridophytic spores recorded from the intertrappeansediments of the Rajmahal Basin.

Acknowledgements

Sincere thanks are extended to the Director, BirbalShani Institute of Palaeobotany, Lucknow for permis-sion to publish the results. I acknowledge with grati-tude the Geological Survey of India for permission tocollect the borehole core material and for the helpextended during the collection. Sri Chandrpal isthanked for processing and preparing the palynologi-cal slides and Sri Pradeep Mohan for preparing thephotographs. I am grateful for the useful commentsmade by Prof. C. G. K. Ramanujam on an earlierdraft of the paper, and by the anonymous referees andProf. D. J. Batten.

Fungal remains from the Rajmahal Formation, India 573

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