- Email: [email protected]
Fungal remains from the Miocene Quilon Beds of Kerala State, South India
Review of Palaeobotany and Palynology, 62 (1990): 13-28
13
Elsevier Science Publishers B.V., Amsterdam-- Printed in The Netherlands
Fungal Remains from the Miocene Quilon Beds of Kerala State, South India PRAMOD KUMAR Birbal Sahni Institute of Palaeobotany, Lucknow - - 226007 (India)
(Received April 13, 1988; revised and accepted October 2, 1989)
Abstract
Kumar, P., 1990. Fungal remains from the Miocene Quilon Beds of Kerala State, South India. Rev. Palaeobot. Palynol., 62: 13-28. Fossil fungal remains are described from the Quilon Beds (Lower-MiddleMiocene) exposed at Padappakkara, Quilon district in the Kerala State, South India. The fungal spores are represented by 15 genera and 23 species, of which 2 genera viz., Elsikisporonites and Hypoxylonsporites and 18 species are new. A tropical humid climate with heavy precipitation probably prevailed during the time of deposition.
Introduction
During the past three decades much attention has been paid to the study of fossil fungal remains in India. Rao (1959), Jain and Gupta (1970), J a i n (1974), Rao and Ramanujam ( 1 9 7 5 , 1976), Ramanujam and Rao (1978), Jain and Kar (1979), Ramanujam and Srisailam ( 1 9 8 0 ) have made valuable contributions to the study of fungal spores and fruiting bodies from Quilon Beds of Kerala State. An Upper Miocene age (Paulose and Narayanaswami, 1968) pertains to limestones overlying carbonaceous shales of Lower-Middle Miocene age (Rao and Ramanujam, 1982). This dating is also supported by Jacob and Sastri (1952) and Dey (1962). Recently Raha and Rajendran (1983) have suggested a Late Eocene-Oligocene age for the lower part of the sequence (Quilon Beds) on the basis of the foraminiferal assemblage, The present paper deals with the study of fungal spores from the slides previously studied by Jain and Gupta (1970). The geology of 0034-6667/90/$03.50
the area, the material, collected by K.P. J a i n from a clay mine section near Kanjantheria House, Padappakkara (11 km north east of Quilon) and the maceration technique employed for the recovery of fungal entities have already been described by Jain and Gupta (1970), Jain and Kar (1979) and Ramanujam (1982b).The slides are stored in the Museum of the Birbal Sahni Institute of Palaeobotany, Lucknow. The locus typicus for the material is Padappakkara, Quilon District, Kerala State, South India and the stratum typicum is Quilon Beds (Lower-Middle Miocene).
Stratigraphy of the area The marine carbonate and the continental clastic facies of the Tertiary sequence occurring in the coastal region of Kerala have been named as the Quilon and Warkalli Beds (Younger) respectively by King (1882) and Bruce Foote (1883).The entire Tertiary sequence of Kerala rests unconformably upon Archaean Gneisses
© 1990Elsevier Science Publishers B.V.
14
P. KUMAR
76"35"
76 °
40'
,.9
55"
55'
--%+,/ .)
"
'~l~
JJ/
i~
Ba"
SCALE
1:2500000
Map 1. Showing the location of exposure at Padappakkara Quilon, Kerala. and itself is overlain by Recent to sub-Recent marine and estuarine sediments. Description of new taxa Genus
Inapertisporites
van der Hammen,
1954 e m e n d . S a x e n a e t B h a t t a c h a r y y a ,
1987
Type species: Inapertisporites variabilis van der Hammen, 1954
Inapertisporites crenulatus K u m a r sp. nov. (Plate I, 1, 2; Fig.l)
Holotype: P l a t e
I,
2;
S1.
No.
3848;
Size
25 x 28 ~m.
Description: F u n g a l spores unicellular, inaperturate, l e n t i c u l a r in shape. Size 17-25 x 2528 pm in diameter. Spore wall differentially thickened, + 2 ~m thick, e q u a t o r i a l m a r g i n w a v y s i m u l a t i n g crested outline, crests + 2 ~m high and _ 3 ~tm broad in optical section. B o t h the surfaces smooth. A polar t h i n n e r area, 9--10 ~tm in diameter. Specimen studied: 15. Comparison: Inapertisporites crenulatus K u m a r
FUNGAL REMAINS FROM MIOCENE QUILON BEDS
15
does not appear similar to any of the species of this genus due to its crenulated margin, Recently, Jarzen and Elsik (1986) described fungal spores as Geastrum-type (Plate I, 12-14) from the Recent sediments of Luangwa Valley, Zambia, which are comparable with I. crenulatus Kumar. They mentioned, as already informed by Dring (1964), t h a t it grows on soil or vegetable debris in damp places.
differs from all the known species of this genus in having hyaline, hollow, rounded verrucaelike sculptures at the peripheral margin of the spores. Derivation of specific name: Based on peripheral verrucae.
Exesisporites sp. Q (Plate I, 3) Description: Fungal spore unicellate, psilate,
Inapertisporites sp. P. (Plate I, 17) Description: Fungal spore unicellular and sickle-shaped with tapering ends. Size 28-30 x 7-8 ~m. The spore wall _ 1 ~m thick, smooth and hyaline. A thick exinal line seen in the photograph is caused by the oblique preservation only. Specimens studied: 3. Comparison: The studied specimen is different from all the known species in having a sickleshaped body. Affinity: It resembles the conidia of modern taxon Chloridiella (Kendrick and Carmichael in Ainsworth et al., 1973).
broadly oval and monoporate. Size 27 x 21 ~m. Minute pore visible in a darker oval area. Spore wall smooth, __l.6~m thick. Darker subcircular patch in the central region of the spore measures 9-11 ~m in diameter. Comparison: The studied specimen closely compares with Exesisporites neogenicus Elsik (1969, plate 1, figs.3, 5, 6) in possessing a comparative minute pore in the darker region but differs in its broadly oval shape and thicker spore wall. G e n u s Lacrimasporo~ites e m e n d . Elsik, 1968
Clarke,
1965
Type species: Lacrimasporonites levis Clarke, Genus Exesisporites Elsik, 1969
1965
Type species: Exesisporites neogenicus Elsik,
Lacrimasporonites
1969
(Plate I, 6; Fig.3)
Exesisporites
verrucatus
Kumar
sp.
nov.
(Plate I, 4; Fig.2)
niger
Kumar
sp.
nov.
Holotype: Plate I, 6; S1. No. 3841; Size 35 x 14 ~m.
Description: Fungal spore unicellular, monopoHolotype: Plate I, 4; S1. No. 3841; Size 28 x 30 ~m.
Description: Fungal spores lenticular in shape and unicellate. Size 25-27 x 28-30 ~m. Monoporate, pore + 1 pm in diameter surrounded by + 1.6 ~m thick, diffuse margin. Spore wall two layered ___2~m thick, outer layer forming 16-18 hyaline, hollow, rounded verrucae-like sculptures at peripheral margin of spore, measuring 2-2.5 ~m high and 1.6-2.5 ~m broad at base, rather evenly placed, imparting angularity to the ambitus, Specimens studied: 4. Comparison: Exesisporites verrucatus Kumar
rate and elliptical with acutely rounded ends. Size 30-35 x 13-15 ~m. Single pore at one end. Spore wall smooth, _ 0.75 ~m thick, slightly thicker around the pore. Specimens studied: 10.
Comparison: Lacrimasporonites
singularis
Sheffy et Dilcher (1971) differs from L. niger Kumar in being smaller (10.6 x 16.4 ~m) and oval to oblong in shape and having a pore at one side of longitudinal centre. L. basidii Elsik (1968) is smaller (9-14 x 6-8 ~m) and has a spatulate shape with a distinctly two layered spore wall. Lacrimasporonites sp. Elsik et Dilcher (1974) is smaller and has thinner wall
~
r~
C3o
NI
d~
.~l
-
C
q
FUNGAL REMAINS FROM MIOCENE QUILON BEDS
Fig.1. D i a g r a m m a t i c s k e t c h o f Inapertisporites erenulatus K u m a r sp. nov. S h o w i n g e r e n u l a t e m a r g i n a n d a p o l a r t h i n n e r s u b c i r c u l a r a r e a at o n e face in t h e s p o r e wall.
around the pore. L. magnus Saxena et Singh (1982) is much longer (120gm) and has a broader pore (27 ~tm diam.) with thickened pore margin. Affinity: Elsik (1968) suggested that such fungal spores could be related to Coprinus, Cortinarius and Stropharia of Basidiomycetes (Graham, 1962). Derivation of specific name: Diagnostic for its dark wall (Latin niger = black).
17
Fig.2. D i a g r a m m a t i c s k e t c h of Exesisporites verrueatus K u m a r sp. nov. S h o w i n g h y a l i n e v e r r u c a e at p e r i p h e r a l m a r g i n of spore a n d a pore w i t h t h i c k e n e d outline.
G e n u s Diporisporites van der H a m m e n , 1954 e m e n d . Elsik, 1968
Type species: Diporisporites elongatus van der Hammen, 1954
Diporisporitespsilatus Kumar sp. nov. (Plate I, 18; Fig.4)
PLATE I All p h o t o m i c r o g r a p h s a r e f r o m u n r e t o u c h e d n e g a t i v e s a n d m a g n i f i e d ca. 1000 x . C o - o r d i n a t e s of e a c h s p e c i m e n s t u d i e d u n d e r E r g a v a l T r i n o c u l a r M i c r o s c o p e No. 663567 g i v e n in p a r e n t h e s i s . 1, 2. Inapertisporites crenulatus K u m a r sp. nov.: S1. Nos. 3844 (111.9 x 20), 3848 ( h o l o t y p e - l l 2 x 6.7). 3. Exesisporites sp. Q: S1. No. 3841 (101.0 × 14.8). 4. Exesisporites verrucatus K u m a r sp. nov.; S1. No. 3841 (holotype-108.1 x 14.9). 5. Imprimospora ramanujamii K u m a r sp. nov.; S1. No. 3842 (holotype-106.7 × 5). 6. Lacrimasporonites niger K u m a r sp. nov.; S1. No. 3842 (holotype-100.2 x 14.8). 7, 8. Elsikisporonites tubulatus K u m a r , sp. nov.; S1. Nos. 3843 (holotype-96.7 x 12.3), 3844 (90.4 × 7.3). 9. DiceUaesporites elongatus K u m a r sp. nov.; S1. No. 3843 ( h o l o t y p e - 8 7 × 18.1). 10. Psidimobipiospora scabratus K u m a r sp. nov.; S1. No. 3841 (holotype-93.4 × 16.1). 11. Brachysporisporites tenuis K u m a r sp. nov.; S1. No. 3844 (holotype-82.8 × 16.2). 12. Dicellaesporites keralensis K u m a r sp. nov.; S1. No. 3844 (holotype-81.8 × 16.8). 13. Hypoxylonsporites ater K u m a r sp. nov.; S1. No. 3847 ( h o l o t y p e - l l l . 6 x 15.4). 14. Multicellaesporites indicus K u m a r sp. nov.; S1. No. 3843 ( h o l o t y p e 108.7 × 6). 15. Diporicellaesporites concavus K u m a r sp. nov.; Sl. No. 3849 ( h o l o t y p e - 1 1 7 × 15.6). 16. Dyadosporites novus K u m a r sp. nov.; S1. No. 3847 (holotype-101.5 × 10.1). 17. Inapertisporites sp. P; Sl. No. 3845 (101 x 4.7). 18. Diporisporites psilatus K u m a r sp. nov.; S1. No. 3842 (holotype-108.8 x 12). 19. Dyadosporites dubius K u m a r sp. nov. S1. No. 3843 ( h o l o t y p e - 1 0 0 x 7.3). 20. Diporicellaesporites sp. K. S1. No. 3854 (104.1 × 21.2). 21. Diporicellaesporites concavus K u m a r sp. nov.; S1. No. 3848 (114.6 × 22.8). 22. Fusiformisporites acutus K u m a r sp. nov.; S1. No. 3844 ( h o l o t y p e - 1 1 4 x 17.7). 23. Hypoxylonsporites miocenicus K u m a r sp. nov.; S1. No. 3845 (holotype-114.5 x 12.3). 24. Tetraploa sp. A; S1. No. 3842 (110 x 12.8). 25. Diporicellaesporites padappakkarensis K u m a r sp. nov.; S1. No. 3846 (holotype-101.6 × 7.1). 26. Tetraploa sp. B; S1. No. 3842 (85.0 × 13).
18
P. KUMAR
' 10tim ' Fig.3. D i a g r a m m a t i c s k e t c h of Lacrimasporonites K u m a r sp. nov. S h o w i n g slit like pore at one end.
niger
Comparison: Involutisporonites Clark emend. Elsik (1968) differs in being multicellate. Palaeocirrenalia Ramanujam et Srisailam (1980) and Colligerites Jain et Kar (1979) are nonaperturate and multicellate. Affinity: Resembles the conidiogenous cells produced by Helicosporium (Kendrick et Carmichael in Ainsworth et al., 1973). Derivation of Generic name: The generic name recognizes Dr. William C. Elsik of Exxon Company USA, Houston, USA. Elsikisporonites
10 fl.Lm
~
Holotype: Plate I, 18, S1. No. 3842; Size 19 x 10 pm.
Description: Fungal spores unicellate, elongate, fusiform and diporate. Size 19-40x 10 221~m. Pore at each end of the spore, simple, slightly sunken and ___2 pm wide. Spore wall smooth, 1 pm thick. Specimens studied: 7. Comparison: Diporisporites hammenii Elsik (1968) is smaller (11-14 x 6-8 Wn) and has bulging pores. D. conapicua Ramanujam et Rao (1978) differs from D.psilatus Kumar in being bigger size (58 × 37 Wn), thickened pore margins and distinct atria. D. curvatus Ramanujam et Rao (1978) possesses more curved sides and thicker spore wall. Diporisporites sp. Hopkins (1969) is longer (55-60 ~m) than the present species. Diporisporites elongatus van der Hammen (1954) shows curved spore wall near the pores, hence, it is different from the present species.
Elsikisporonites
Kumar sp.
nov.
(Plate I, 7, 8; Fig.5)
Fig.4. D i a g r a m m a t i c sketch of Oiporisporites psilatus K u m a r sp. nov. S h o w i n g s l i g h t l y s u n k e n pore at end.
Genus
tubulatus
Kumar
gen.
Holotype:
Plate I, 7; SI. No. 3843; Size 36.8 x 38.4 pm (ambitus). Description: Fungal spores monoporate, nonseptate, tubular in shape and coiled. Cell broadest in the middle region, 10-12 ~m wide, gradually tapering towards the ends. Pore at free outer end, small, nozzle-like, 1.5 Ilm wide. Spore wall 1 ~m thick, smooth, slightly folded, hyaline.
Specimens studied: 8. G e n u s Hypoxylonsporites K u m a r gen. n o v .
Type species: Hypoxylonsporites miocenicus Kumar gen. et sp. nov.
nov.
Type species: Elsikisporonites tubulatus Kumar sp. nov.
I
I 10,/t..1,rn
Diagnosis: Fungal spores monoporate, nonseptate, tubular and coiled. Pore a t o u t e r end, nozzle-like. Spore wall smooth and hyaline,
Fig.5. D i a g r a m m a t i c s k e t c h of Elsikisporonites tubulatus K u m a r gen. et sp. nov. S h o w i n g t u b u l a r coiled cell w i t h a nozzle-like pore at free uncoiled end.
FUNGAL REMAINS FROM MIOCENE QUILON BEDS
Diagnosis: Fungal spores unicellular, oval to elliptical in shape with acutely rounded ends. A longitudinal slit-like aperture may be running end to end. Spore wall single layered, smooth and may be differentially coloured. Affinity: The form genus Hypoxylonsporites Kumar shows close affinity with extant genus Hypoxylon (See Miller, 1961). Specimens of Hypoxylonsporites also show resemblance with Recent Endocalyx. Remarks: Venkatachala and Rawat (1973) recorded a specimen similar to the present new genus from the subsurface Tertiary sediments of Cauvery Basin, S. India. Sah (1967) reported such fungal spores from the Neogene sediments of Rusizi Valley (Burundi). Elsik and Dilcher (1974) also recorded similar elongate type of fungal spores having a linear fold, slit or scar, from Middle Eocene clay beds in the Lawrence Clay Pit, Henry County, Tennessee. Miller (1961, fide Elsik and Dilcher, 1974) compared such specimens with the cosmopolitan saprophytic fungi Hypoxylon (Bulliard, 1791) Fries (1849). Elsik (1969) figured similar spores from the Late Neogene deposits in northern Gulf of Mexico and referred to them as Hypoxylon spp. (Plate I, 1, 2). Subramanian (1971) stated that conidia of Endocalyx thwaitesii Berkeley et Broome (1850) and E. melanoxanthus (Berkeley et Broome) Petch (See Hughes, 1953) of Hyphomycetes are one-celled, flattened-oval with a longitudinal germ slit. During the course of the present study, few fungal spores similar to Hypoxylon have been recorded from the Miocene Quilon deposits at Padappakkara. It is suggested that such fossil spores recorded by the earlier workers should be referred to Hypoxylonsporites gen. nov.
19
i
lo/a m Fig.6. Diagrammatic sketch of Hypoxylonsporites rniocenicus Kumar sp. nov. Showing a longitudinal slit extending from one end to the other.
tends from one end to the other. Spore wall single-layered, + 1.6 ~m thick and smooth.
Specimens studied: 16. Hypoxylonsporites ater Kumar sp. nov. (Plate I, 13; Fig.7) Holotype: Plate I, 13; S1. No. 3847; Size 29 × 18 ~m. Description: Fungal spores unicellular, broadly oval in shape, ends tapered. Size 25-30x 18-20 ~m. Longitudinal slit runs from end to end. Spore wall single layered, _ 1.6 ~m thick and smooth. Specimens studied: 6.
Comparison: Hypoxylonsporites miocenicus Kumar differs from H. ater Kumar in having elongate shape with narrowly rounded ends.
Hypoxylonsporites rniocenicus Kumar sp. nov. (Plate I, 23; Fig.6) Holotype: Plate I, 23; S1. No. 3845; Size 32 x 13 ~m. Description: Fungal spores unicellular, elliptic t o spindle-shaped with narrowly rounded ends. Size 21-55 x 9-26 ~m. A longitudinal slit ex-
I
20ill m
I
Fig.7. D i a g r a m m a t i c s k e t c h of Hypoxylonsporites ater K u m a r sp. nov. S h o w i n g l o n g i t u d i n a l slit w i t h g r e a t e s t
width of spore in the middle region.
20
P. KUMAR
Genus Dicellaesporites Elsik 1968 e m e n d . S h e l l y et Dilcher, 1971
Type speices: Dicellaesporites popovii Elsik, 1968
Dicellaesporites elongatus Kumar sp. nov. (Plate I, 9; Fig.8)
Holotype: Plate I, 9; S1. No. 3843; Size 43 x 14 ~m.
Diagnosis: Spores dicellate, fusiform and inaperturate. Size 41-50 x 14-17 pm. Cells gradually tapering at either end. Both spore wall and septum __1 ~m thick. Spore wall smooth, light coloured and folded. Septum centrally porate,
Specimens studied: 8. Comparison: The present species differs from D.popovii Elsik (1968) in having narrowly rounded ends. D. ellipticus Jain et Kar (1979) differs in having larger size and ornamented wall. Affinity: The present species shows resemblance with the conidiogenous axis of Dactylaria Saccardo (1880; fide Ainsworth et al., 1973).
Dicellaesporites keralensis Kumar sp. nov.
Holotype: Plate I, 12; S1. No. 3844; Size 21 x 16 pm.
Description: Spores dicellate, inaperturate and broadly oval in shape. Size 18-22 x 13-16 pm. Broadly rounded at free ends but slightly constricted at the septal region, cells unequal in size, 1 0 - 1 3 x l l - 1 4 ~ m . Both septum and spore wall _ 1 ~m thick and smooth. Specimens studied: 11. Comparison: D. popovii Elsik (1968) lacks the constriction at the septum. D. eUipticus Jain et Kar (1979) differs in having microreticulate spore wall and bigger size range. D. disphaericus Shaffy et Dilcher (1971) is distinct in having scabrate spore wall, well pronounced opaque equatorial septum and constriction. G e n u s Imprimospora Norris, 1986
Type species: Imprimospora tankensis Norris, 1986
Imprimospora ramanujamii Kumar sp. nov. (Plate I, 5; Fig.10)
Synonymy 1980 3-celled striate spore type: Ramanujam et Srisailam, pl.2, fig.28, p.131.
Holotype: Plate I, 5; S1. No. 3842; Size
(Plate I, 12; Fig.9)
32 x 18 pm.
Description: Fungal spores tricellate, inapertu-
' Fig.8. Diagrammatic sketch of Dicellaesporites elongatus Kumar sp. nov. Showing dicellate fusiform cell with a centrally slitted septum.
rate and fusiform. Size 32-40 x 18-21 ~m. Septa 1.6-2 ~m thick, with or without a central pore. Central cell large, 11-13 x 16-19 ~m, terminal cells small, 9-10~m long with narrowly
d
Fig.9. Diagrammatic sketch Dicellaesporites keralensis Kumar sp. nov. Showing dicellate spore without any slit.
Fig.10. Diagrammatic sketch of Imprimospora ramanujamii Kumar sp. nov. Showing tricellate spore with central larger cell and longitudinal ribs converging at end.
FUNGAL REMAINS FROM MIOCENE QUILON BEDS
21
rounded ends. Spore wall _ 1.5 ~m thick and ornamented with longitudinal ribs ( ± 2 ~ m wide) that run full length but are more pronounced. Furrows 1-2 ~m wide. Specimens studied: 7. Comparison: Imprimospora ramanujamii Kumar closely resembles 3-celled ribbed spore type described by Ramanujam and Srisailam (1980, pl.2, fig.28). I. tankensis Norris 1986 (Fide Jansonius and Hills, 1987: C.F.4444) differs in having thinner (0.5 ~) spore wall with uniseptate body. Derivation of specific name: The species is named after Dr. C.G.K. Ramanujam, Hyderabad, India.
Comparison: Fusiformisporites acutus Kumar differs from F. elongatus Ramanujam et Rao (1978) in having larger size and acute ends. F. keralensis Ramanujam et Rao (1978) differs from the present species in having rhomboidal shape with cells wider at the septal region. F.pseudocrabbii Elsik (1968) is different in having finer ribs made of rows of punctae parallel to the coarse ribs, F. microstriatus Hopkins (1969) differs in having smaller size (42-49 ~m in length) and finer ribs.
Genus Fusiformisporites Rouse, 1962
1965
Type species: Fusiformisporites crabii Rouse,
Dyadosporites novus Kumar sp. nov. (Plate I,
1962
16; Fig.12)
G e n u s Dyadosporites v a n d e r H a m m e n ex C l a r k e , 1965
Type species: Dyadosporites ellipsus Clarke,
Fusiformisporites
acutus
Kumar
sp.
nov.
(Plate I, 22; Fig.ll)
Holotype: Plate I, 16; S1. No. 3847; Size 32 x 16 ~m.
Description: Fungal spores dicellate and dipoHolotype: Plate I, 22, S1. No. 3844; Size 96x21 ~m.
Description: Fungal spores dicellate, inaperturate and elongated-fusiform. Size range 90-112 x 20-24 ~m generally tapering towards acute ends. Septum _ 1 ~m thick. Spore wall 1-1.6 ~m thick and ribbed. Ribs _ 1 ~m wide, wider 1.6-2.5~m at septal region, closely spaced, alternate or continuously running end to end, sometimes bifurcating and converging at the ends, furrows very narrow. Specimens studied: 5.
I
20/U.m
rate. Size 28-32×13-161Jm. Cells fusiform and gradually taper towards ends. Apical pore 0.5 ~m and simple. Septum 2-3 ~m thick, wedge-shaped or denticulate with a central pore, dark and opaque. Spore wall 1.6 ~tm thick, smooth, slightly constricted at septal region and showing a brief collar at one end (probably basal e n d ) o f the spore.
Specimens studied: 6. Comparison: Dyadosporites ellipsus
Clarke (1965) differs in having longer cells (38-55 ~m) and bigger pores (4-5 ~m diam.).
I
Fig.ll. Diagrammatic sketch of Fusifermisporites acutus Kumar sp. nov. Showing dicellate, acute ends with thick longitudinal ribs continuous or bifurcating and converging at ends.
P KUMAR
22
I
10Nrn
I
Fig.12. Diagrammatic sketch of Dyadosporites novus Kumar sp. nov. Showing dieellate and diporate spore.
t 10/U.m
'
lOpm
Fig.14. Diagrammatic sketch of Psidimobipiospora scabratus Kumar sp. nov. Showing dicellate, diporate spore with scabrate wall.
'
Fig.13. Diagrammatic sketch of Dyadosporites dubius Kumar sp. nov. Showing dicellate, diporate and bracket-
Description: F u n g a l spores cylindrical, dicel-
shaped spore,
late and diporate. Size r a n g e 35-45 x 13-16 pm. Cells r e c t a n g u l a r in shape, each cell 19 x 13-14 pm, 4- equal in size with a pore at free end. P o r e 4-6 I~m wide, simple and slightly sunken. S e p t u m _+3pm thick, opaque and simple. Spore wall ___2pm thick, t h i n n e r a r o u n d pore, surface smooth to finely s c a b r a t e or echinate, slightly c o n s t r i c t e d at septal region. Specimens studied: 10.
Dyadosporites dubius K u m a r sp. nov. (Plate I, 19; Fig.13)
Holotype: P l a t e I, 19; S1. No. 3843; Size 16 x 13 pm.
Description: F u n g a l spores diporate, dicellate and b r o a d l y oval or bracket-shaped (outline). Size 15-18 x 12-13 i~n. P o r e s 4-6 pm wide, simple, one at e i t h e r end. Spore wall ___1.6pm thick, smooth, g r a d u a l l y t h i n n i n g out t o w a r d s the pore. S e p t u m 4-2 pm thick, opaque, Specimens studied: 20. Comparison: Dyadosporites ellipsus Clarke (1965) differs in having elliptical shape and larger size (23-27 x 38-55 ~lm). D. novus K u m a r differs in being narrow apical pores and larger cells,
Genus Psidimobipiospora Locqvin e t Salard-Cheboldaeff in Salard-Cheboldaeff et Locquin, 1980
Comparison:
Psidimobipiospora
dyadospora
Sal-Cheb. et L o c q u i n 1980 (Fide J a n s o n i u s and Hills, 1981, G. F i l e c a r d 3900) differs in h a v i n g t r u n c a t e d extremities and g r a n u l o s e spore wall. Genus Multicellaesporites emend. Kumar
Elsik,
1968,
Type species: Multicellaesporites nortonii Elsik, 1968
Emended Description: F u n g a l spores multicelPsidimobipiospora
dyadospora
Salard-Cheb.
et Locquin, 1980
Psidimobipiospora scabratus K u m a r sp. nov. (Plate I, 10; Fig.14)
Holotype: P l a t e I, 10; S1. No. 3841; Size 40 x 14 tim.
late, elongate. A l o n g i t u d i n a l slit or f u r r o w present. Spore wall smooth or o r n a m e n t e d or differentially c o l o u r e d or t h i c k e n e d . Remarks: Elsik (1968) diagnosed the genus " i n a p e r t u r a t e " , w h e r e a s the type species (pl.2, fig.15) was diagnosed as " t h e grain split l o n g i t u d i n a l l y " . The p r e s e n t s t u d y reveals t h a t the genus has a t e n d e n c y to b e a r a l o n g i t u d i n a l
23
FUNGAL REMAINS FROM MIOCENE QUILON BEDS
slit or furrow. Hence, it has been accordingly emended,
Multicellaesporites elsikii ( R a m a n u j a m et Srisailam, 1980) comb. nov.
Basionym: Staphlosporonites elsikii Ramanujam et Srisailam 1980; Botanique, 9(1-4): p.122, pl.1, figs.6, 7.
ally-coloured or thickened. The middle cell hyaline, 14-19 ~m long, whereas the terminal cells c o m p a r a t i v e l y d a r k e r and smaller in size (8-9 ~m long). Specimens studied: 20. Comparison: It differs from the o t h e r species in h a v i n g differentially t h i c k e n e d wall and shorter aperture.
Genus Brachysporisporites S m i t h , 1971
Synonymy 1959 Multicellular phragmospora Rao, pl.1, fig.14 1968 Hyphae Type-1 Elsik, pl.3, figs.13, 15 1968 Hyphae (?) Elsik, pl.3, Fig.18 1978 MulticeUaesporites ? Ramanujam et Rao, pl.1, Fig.20 1980 Staphlosporonites elsikii Ramanujam et Srisailam, pl.1, figs.6, 7 1981 Inapertisporites globulosus Rouse, 1962 in Ediger, 1981; pl.1, fig.6
Lange
et
Type species: Brachysporisporites pyriformis Lange et Smith, 1971
Brachysporisporites
tenuis K u m a r
sp.
nov.
(Plate I, 11; Fig.16)
Affinity:
Synonym:
dematiaceous h y p h o m y c e t e s Annellophorella (Ellis, 1971) and to some e x t e n t with the
43x30~m.
conidia of Stigmina compacta.
Description: F u n g a l spores di- or multicellate
R a m a n u j a m and Srisailam (1980) showed its strong similarity with the conidia of
Multicellaesporites
indicus K u m a r
sp.
nov.
3843;
Size
(Plate I, 14; Fig.15)
Holotype: Plate
I,
14; S1. No.
32 x 16 ~m.
Description:
F u n g a l spores tricellate, elongated, elliptical or spindle-shaped with narrowly r o u n d e d to a n g u l a r ends. Size r a n g e 31-38 x 11-18 pm. A longitudinal slit present in hyaline middle region. Spore wall smooth, less t h a n 1 pm thick in central region and ± 2 ~m at the terminal and septal regions and differenti-
Fig.15. Diagrammatic sketch of Multicellaesporites indicus Kumar sp. nov. Showing longitudinal slit restricted to hyaline middle cell with comparatively darker terminal ends.
1978 Spore Type B Ramanujam et Rao; pl.1, figs.18, 19
Holotype: P l a t e
I,
11; S1. No.
3844;
Size
and balloon-shaped. Size 38-43 x 22-30 pm. Cells unequal, small cell hyaline and thin, m e a s u r i n g 4-10 x 3-6 ~m broad. L a r g e r apical cell or cells forming a dome-shaped, thick and dark coloured to opaque structure 23-40 x 14-32 ~m. Septa 2-3 ~m thick, smooth to wedge-shaped. Specimens studied: 10.
Fig.16. Diagrammatic sketch of Brachysporisporites tenuis Kumar sp. nov. Showing balloon-shaped multicellate spore with hyaline smaller cell.
24
Comparison:
P. KUMAR
Brachysporisporites
pyriformis
Lange et Smith (1971) is smaller (36--41 x 15-18 pro) and pyriform.
in
size
G e n u s Diporicellaesporites Elsik, 1968
Type species: Diporicellaesporites stacyi Elsik, 1968
Diporicellaesporites concavus Z u m a r sp. nov. (Plate I, 15, 21; Fig.17)
Holotype:
Plate I, 15; S1. No. 3849; Size 38.4 x 22.4 pm. Description: Fungal spores multicellate, diporate and fusiform in shape. Size range 38 42 x 22-24 pm. Spore gradually tapering at both ends. Central cells dark in colour and larger in size (8--13 x 13-21 ~m), apical cells smaller, hyaline, porate and delimited by a concave d a r k e r outline (septum). Spore wall thickest at the central septal region and smooth. Septum straight or disc-shaped, septurn of central cell _+3 pm thick with a central pore. Terminal pores _+2 ~m wide, pore margin thin and hyaline. Specimens studied: 15. Comparison: Diporicellaesporites concavus Kumar differs from all o t h e r k n o w n species having differentially coloured t h i c k e n e d spore wall and concave septa,
Diporicellaesporites padappakkarensis K u m a r sp. nov. (Plate I, 25; Fig.18)
I
I 10/t£m Fig.18. Diagrammatic sketch of DiporicellaesporitespadappakkarensisKumar sp. nov. Showing multicellate unequal cells, pore offset, slit-like and partially divided into two pore cells at the terminal hyaline ends.
Holotype:
Plate 30.4 x 19.2 pm.
I,
25; S1. No.
3846;
Size
Description: Fungal spores multicellate and elongated-elliptical in size. Size 30-35 x 19-22 ~m. Cells unequal in size, the central cell large, 19 x 17 ~m and terminal cells splitting, resulting in a beak-shaped structure, 4-6 pm long. Both ends with single pore, offset, slitlike and measuring 1 2 × 1 - 5 ~ m , partially divided into two pore cells. Septa three, comparatively thicker t h a n the spore wall, disc-shaped with central pore. Spore wall +1.6 ~m thick, smooth. Wall of central cell dark coloured and terminal cells hyaline. Specimens studied: 5. Comparison: Diporicellaesporites concavus Kumar differs in having undivided apical cells at poral region. Remarks: Beak-shaped s t r u c t u r e of the hyaline cells might have developed during the compression for accommodating the thicker wall of central cell.
Diporicellaesporites sp. K (Plate I, 20) Description: This fungal spore is tetracellate,
' 10/trm I Fig.17. Diagrammatic sketch of Diporicellaesporites concavus Kumar sp. nov. Showing tetracellate, diporate spore apical cells hyaline and porate.
diporate and fusiform in shape, its size is 3 0 . 5 x 1 3 ~ m . The central cells are slightly constricted at middle septal region and slightly curved at one side. Terminal cells are smaller in size, tapering with a slightly pouting pore. The septa are nearly as thick as spore wall. The spore wall is 1.0 ~m thick and smooth.
Specimens studied: 1. Comparison: Diporicellaesporites sp. Ramanu-
FUNGAL REMAINS FROM MIOCENE QUILON BEDS
jam et Rao (1978) is different from Diporicellaesporites sp. K in having larger size (81 ~n) and stalk-like basal cells. D. stacyii Elsik (1968) differs in having thick septa, granulate to punctate inner surface of cells and pores with slitted margins. Genus
25
broad, its terminal cell gradually tapering. The septa 1-3 ~m thick, simple to wedge-shaped. The spore wall _ 1 ~m thick, smooth. Specimens studied: 1. Comparison: Tetraploa sp. B differs from T. sp. A in size (36-48 x 6-11 ~m), having a septate main body and narrower appendages.
Tetraploa B e r k e l e y et B r o o m e , 1850 Palaeoclimate
Type species: Tetraploa aristata Berkeley et Broome, 1850 Tetraploa sp. A (Plate I, 24) Description: Fungal conidia slightly triangular in shape with nonseptate body. Size 40-46 × 8-10 pm. Conidium 14 x 10 ~tm. The body terminating into four long processes, Processes 3-4 septate, 32-35 × 3-5 ~m, hyaline, + 1.0 ~m thick and smooth, Specimens studied: 1. Comparison: The present species differs from Type 89: Tetraploa aristata Berkeley et Broome (1850) by its smooth processes and triangular body which in the later is verrucate. Tetraploa (Frasnacritetrus) sp. Singh et Saxena (1981) described from the Upper Siwalik of Himachal Pradesh, closely compares with the present species in having psilate appendages, but differs in exhibiting nonseptate appendages and quadrangular body. Frasnacritetrus josettae Taugourdeau (1968) is different from Tetraploa sp. A in having a quadrangular body and bigger size. Remarks: The author considers that Taugourdeau described as Frasnacritetrus a fungal conidium comparable to Tetraploa Berkeley et Broome and not an acritarch type sporomorph. Tetraploa sp. B (Plate I, 26) Description: Fungal spore branched, multicellate; main body four to five celled and fusiform in shape, i.e. size 60 × 20 ~tm. Apical cell termihating into four appendages, ca. 150 pm long. Cells of main body rectangular, broader than long, 19-21 × 9-11 ~tm. Cells of appendages are longer than broad, 17-27 pm long and 6-8 ~tm
Certain sporomorphs in the Padappakkara sediments are useful in determining the palaeoclimate. The form taxa which are closely related to modern ones (Table I) are most helpful because their modern environmental conditions are known. Elsik (1969) pointed out that the abundant Exesisporites in the northern Gulf of Mexico during the Pliocene might be indicative of a warm climate. The record of Grallomyces, Spegazzinia from Quilon Beds of Kerala, clearly indicates that a tropical, humid climate prevailed during the Neogene time in this region (Ramanujam and Rao, 1978; Ramanujam and Srisailam, 1980). Helicosporous hyphomycetes are usually found on moist wood (Subramanian, 1971). The presence of helicoidal fungal forms (e.g. Elsikisporonites) indicates the same climatic conditions during the sedimentation of Quilon Beds. Lange (1976) discussed the value of the fossil TABLEI Affinities of fossil fungal remains with modern taxa Fossil taxa
Modern taxa
Inapertisporessp. P
I. crenulatus
Chloridiella Geastrum saccatum G. lageniforma
Lacrimasporonites
Basidiomycetes (Coprinus,
Elsikisporonites
Cortinarius, Stropharia) Helicosporium
Hypoxylonsporites Dicellaesporiteselongatus
Hypoxylon Dactylaria
Spegazzinites Multicellaesporiteselsikii
Spegazzinia AnnellophoreUa,Stigmina
Tetraploa
compacta Tetraploa
Fusiformisporites Brachysporisporites
Cookeinia and Porodiella Brachysporium
26
epiphyllous microthyriaceous fungi and their germlings as an indicator of palaeohabitats. He pointed out that abundance of such germlings in the deposits could be a good indicator of moist tropical type of vegetation. This is apparent in the Quilon Beds, Kerala, as was also recorded by Jain and Gupta (1970), Ramanujam and Rao (1972), Rao and Ramanujam (1976) and Jain and Kar (1979). Rao and Ramanujam (1975) studied the palynoflora from these Quilon beds and recorded spores and pollen grains which belonged to pteridophytes and angiosperms. They mentioned that the occurrence of such families points towards tropical humid conditions, viz., Gleicheniaceae, Schizaeaceae, Polypodiaceae, Aroideae, Palmae, Lecythidaceae, Araliaceae, Symplocaceae, Rhizophoraceae, Sonneratiaceae, Bombacaceae, Caesalpiniaceae, Combretaceae, Sapotaceae, *Dipterocarpaceae, *Ctenolophonaceae, Meliaceae, *Olacaceae, Anacardiaceae and Droseraceae. The abundance of ferns (families marked with an asterisk) and Duabanga of Sonneratiaceae indicates heavy precipitation also. The climate during the Neogene of Kerala was inferred as tropical humid. There is not much of a change from the past to the present climatic conditions, Raha and Rajendran (1983) suggested that the deposition of continental facies both below and above the Quilon Formation is indicative of major transgressive and regressive phases, The fungal remains and the miospores (Rao and Ramanujam, 1978, 1982) that occur in these sediments further suggest that the deposits are from fresh water lake or ponds where lowland vegetation thrived.
P. KUMAR
fungal spores from carbonaceous clays of Padappakkara. Fungal remains from the Palaeogene and Neogene beds of India have been studied by Rao (1959), Kar et al. (1972), Biradar and Mahabale (1974), Kar and Saxena (1976), Varma and Rawat (1963), Jain and Gupta (1970), Ramanujam and Rao (1978), Ramanujam and Srisailam (1980), Jain and Kar (1979), Ramanujam (1982a, b). On comparison of the present study with the previous ones, it can be seen that genera like
DiceUaesporites,Diporicellaesporites, Brachysporisporites, Involutisporonites, Psidimobipiospora, Imprimospora, MulticeUaesporites, Meliola, Pluricellaesporites, Basidiosporites, Staphlosporonites and Inapertisporites have been recorded from Palaeogene as well as Neogene beds. Therefore such long ranging taxa at the generic level are not useful for stratigraphic consideration. Characteristic forms recorded from Palaeogene strata (Kar et al., 1972; Mahabale, 1969; Biradar and Mahabale, 1974) but so far not reported from the Neogene are: Cucurbitariaceites, Diplodia and Tetracoccosporium eocenum (the last two genera being extant taxa). These forms were also absent in the assemblage studied here. Some forms that have been recorded from Neogene sediments but are absent in Palaeogene sediments are: Quilonia, Diploneurospora,
Comparison with other assemblages
Spegazzinites, Foveo letisporonites, Warkallisporonites, Exesisporites, Hypoxylonsporites (= Hypoxylon), Brachysporisporites tenuis, Retihelicosporonites. Ramanujam (1982a) stated that Fusiformisporites, Palaeocirrenalia-like coiled spores and Colligerites-like helical spores are also present
Jain and Gupta (1970) described some fruiting bodies and fungal spores from Quilon carbonaceous clay beds exposed at Padappakkara. Later, Rao and Ramanujam (1976), Ramanujam and Rao (1978) and Jain and Kar (1979) studied fruiting bodies and fungal spores from the Neogene sediments of the Kerala Coast. The present study is based on
in Miocene beds of Neyveli Lignite, South Arcot, Madras. Some palynologists seem to hold the opinion that fungal spores may not be helpful in resolving stratigraphic problems, but their importance for stratigraphy has been highlighted by Elsik (1968), Graham (1962) and Jansonius (1976). This has encouraged the author to undertake this study. Elsik (1977),
FUNGALREMAINSFROMMIOCENEQUILONBEDS R a m a n u j a m a n d R a o (1978), R a m a n u j a m a n d S r i s a i l a m (1980) a n d R a m a n u j a m (1982a, b)
suggested that ornamented diporate and foveolate fungal spores are not known earlier than Eocene and Miocene respectively; reticulate helicoidal s p o r e s a r e known from Miocene onwards. Ramanujam further stated that spore types with highly sculptured walls are encount e r e d c o n s i s t e n t l y in N e o g e n e s e d i m e n t s . F o r m s like Foveoletisporonites miocenicus (fov e o l a t e d i p o r a t e ) , Retihelicosporonites elsikii
(reticulate helicoidal), Exesisporites, *CoUigererites ( h e l i c o i d a l spores), *Spegazzinites (sculpt u r e d forms), Hypoxylonsporites l o n g i t u d i n a l l y slitted forms) along with
some ornamented
forms, viz., Exesisporites verrucatus sp. nov., Psidimobipiospora scabratus sp. n o v . m a y be s t r a t i g r a p h i c a l l y s i g n i f i c a n t e l e m e n t s . T h e fungal s p o r e s m a r k e d b y a s t e r i s k s a n d o r n a m ented forms are therefore further indications t h a t t h i s a s s e m b l a g e m i g h t h a v e b e e n deposited d u r i n g t h e N e o g e n e P e r i o d ( M i o c e n e ) . Conclusion
(1) On the basis of palaeontological and palynological
evidence,
a Lower to Middle
M i o c e n e a g e is a s s i g n e d t o t h e Q u i l o n c a r b o n a c e o u s c l a y beds. (2) T h e p a r t i c u l a r a s s e m b l a g e o f f u n g a l remains and miospores indicates that Quilon c a r b o n a c e o u s c l a y beds h a v e b e e n d e p o s i t e d in an area having ponding environments. (3) T h e m o s t a b u n d a n t f u n g a l e n t i t i e s a r e suggestive of humid climate. Acknowledgements T h e author is grateful to Dr. B.S. Venkatachala, Director, B.S.I.P., L u c k n o w for valuable suggestions a n d for critically reading the manuscript. I a m thankful to Dr. R.K. S a x e n a for his kind help. References Ainsworth, G.C., Sparrow, F.K. and Sussman, A.S. (Editorn),1973.The Fungi, An Advanced Treatise.Vol IV, A, Academic Press, N e w York, N.Y., pp.1-509.
27 Berkely, M.J. and Broome, C.E., 1850. Notices of British fungi. Ann. Mag. Nat. Hist., 2(5): 459. Biradar, N.V. and Mahabale, T.S., 1974. On the occurrence of unperfect fungus, Tetracocosporium obtained from a fossil wood belonging to the Deccan Intertrappean series. Palaeobotanist, 21: 223-226. Bruce Foote, R., 1883, On the geology of South Travancore. Rec. Geol. Surv. India, 16(1): 20-35. Clarke, A.T., 1965. Fungal spores from Varmeju Formation coal beds (Upper Cretaceous) of Central Colorado, Mt. Geo1.,2:85-93. Dey, A.K., 1962. The Miocene mollusca from Quilon, Kerala (India). Mere. Geol. Surv. India Palaeontol. Indica, N.S., 36: 1-129. Dring, D.M., 1964. Gasteromycetes of the West Tropical Africa. Mycol. Pap., 98: 1-60. Ellis, M.B., 1971. Dematiaceous Hyphomycetes. Commonw. Mycol. Inst. Kew, Surv.: 508 pp. Elsik, W.C., 1968. Palynology of a Palaeocene Rockdale lignite Milam Country, Texas - - 1. Morphology and Taxonomy. Pollen Spores, 10: 263-314. Elsik, W.C., 1969. Late Neogene palynomorph diagram, northern Gulf of Mexico. Gulf Coast Assoc. Geol. Soc. Trans., 19: 509-528. Elsik, W.C., 1977. Morphology phylogeny of dispersed fossil fungal spores - - Intimations. Second Int. Mycol. Congr., Univ. South Florida, Tampa, Abs.: 169. Elsik, W.C. and Jansonius, J., 1974. New genera of Palaeogene fungal spores. Can. J. Bot., 52: 953-958. Graham, A., 1962. The role of fungal spores in palynology. J. Palaeontol., 36: 60-68. Jacob, J. and Sastri, V.V., 1952. Miocene foraminifera from Chavara near Quilon, Travancore. Rec. Geol. Surv. India, 82: 342-353. Jain, K.P., 1974. Fossil Fungi. An Aspects and Appraisal of Indian Palaeobotany, B.S.I,P., Lucknow, pp.38-46. Jain, K.P. and Gupta, R.C., 1970. Some fungal remains from the Tertiaries of Kerala Coast. Palaeobotanist, 18: 177-182. Jain, K.P. and Kar, R.K., 1979. Palynology of Neogene sediments around Quilon and Varkala, Kerala Coast, South India - - 1. Fungal remains. Palaeobotanist, 26: 105-118. Jansonius, J., 1976. Palaeogene fungal spores and fruiting bodies of the Canadian Arctic. Geosci. Man, 15: 129-132. Jansonius, J. and Hills, L.V,, 1976-1987. Genera File of Fossil spores, Spec. Publ. Dep., Geol. Univ. Calgary, Canada and Supplements. Jarzen,D.M. and Elsik,W.C., 1986. Fungal palynomorphs recovered from recent deposits, Luangwa Valley, zambia. Palynology, 10: 35-60. Kar, R.K. and Saxena, R.K., 1976. Algal and fungal microfossils from Matanomadh Formation (Palaeocene), Kutch, India. Palaeobotanist,23: 1-15. Kar, R.K., Singh, R.Y. and Sah, S.C.D. 1972. Some algal and fungal remains from Tura Formation of Garo Hills, Assam. Palaeobotanist, 19: 146-154. Kendrick, W.B. and Carmichael, J.W., 1973. Hyphomycetes. In: G.C. Ainsworth, F.K. Sparrow and A.S. Sussman (Editors),The Fungi, An Advanced Treatise. Vol. IV, A, Academic Press,N e w York, N.Y., pp.323-501.
28 King, W., 1882. General sketch of the geology of Travancore State. The Warkali beds and associated deposits at Quilon in Travancore. Rec. Geol. Surv. India, 15(2): 93-102. Lange, R.T., 1976. Fossil epiphyllous "germlings" their living equivalents and their palaeohabitat indicator value. N. Jahrb. Geol. Palaeontol. Abh., 151: 142 165. Mahabale, T.S., 1969. On a fossil species of Diplodia from the Deccan Intertrappean Series, M.P., India. Palaeobotanist, 17(3): 295 297. Miller, J.H., 1961. A monograph of the world species of Hypoxylon. Univ. Georgia Press, Athens, 158 pp. Paulose, K.V. and Narayanaswami, S., 1968. The Tertiaries of Kerala Coast. Mem. Geol. Soc. India, 2: 300--308. Raha, P.K. and Rajendran, C.P., 1983. Basal Tertiary ferruginous Sandstones in the west coast of southern India and their geological significance. Q.J. Geol. Min. Metall. Soc. India, 56(4): 208-216. Raju, D.S.N., 1978. Contribution to the Neogene stratigraphy of two areas of Kerala basin with special reference to Myogypsinidae (Abstract). VII Indian Colloq. Micropaleontol. Stratigr. Madras. Ramanujam, C.G.K., 1982a. Recent advances in the study of fossil fungi. Recent advances in Cryptogamic Botany Pt. II: Fossil Cryptogams. Palaeobot. Soc. Lucknow.: 287 301. Ramanujam, C.G.K., 1982b. Tertiary palynology and Palynostratigraphy of southern India. Palaeontol. Soc. India, Spec. Publ., 1:57 64. Ramanujam, C.G.K. and Rao, K.P., 1978. Fungal spores from the Neogene strata of Kerala in South India. IVth Int. Palynol. Conf., Lucknow (1976 77), 1: 291 304. Ramanujam, C.G.K. and Srisailam, K., 1980. Fossil fungal
P. KUMAR spores from the Neogene beds around Cannanore in Kerala State. Botanique, 9(1 2): 119-138. Rao, A.R., 1959. Fungal remains from some Tertiary deposits of India. Palaeobotanist, 7:43 46. Rao, K.P. and Ramanujam, C.G.K., 1975. A palynological approach to the study of Quilon beds of Kerala State in South India. Curr. Sci., 44(20): 730-732. Rao, K.P. and Ramanujam, C.G.K., 1976. A further record of microthyriaceous fungi from the Neogene deposits of Kerala in South India. Geophytology, 6: 98-104. Rao, K.P. and Ramanujam, C.G.K., 1982. Palynology of the Quilon beds of Kerala State in South India - - II Pollen of Dicotyledons and Discussions. Palaeobotanist, 30(1): 68-100. Saccardo, P.A., 1880. Conspectus generum fungorum italiac inferiorum nempe and Sphaeropsideas. Melonconicas et Hyphomyceteas peritinentium, Systematic sporologico dispositorum. Michelia, 2:1 39. Sah, S.C.D., 1968. Palynology of an Upper Neogene profile from Rusizi Valley (Burundi). Ann. Mus. R. Afr. Cent. Ser. Sci. Geol., 88(57): 1 173. Saxena, R.K. and Sarkar, S., 1986. Morphological study of Frasnacriterus Taugourdeau emend, from the Tertiary sediments of Himachal Pradesh, India. Rev. Palaeobot. Palynol., 46: 209-225. Subramanian, C.V., 1971. "Hyphomycetes". Indian Counc. Agr. Res. New Delhi. 1 899. Varma, C.P. and Rawat, M.S., 1963. A note on some diporate grains recovered from Tertiary horizons of India and their potential marker value. Grana Palynol. 4(1): 130 139. Venkatachala, B.S. and Rawat, M.S., 1973. Palynology of the Tertiary sediments in the Cauvery Basin 2. Oligocene Miocene palynoflora from the sub-surface. Palaeobotanist, 20(2): 238 263.
13
Elsevier Science Publishers B.V., Amsterdam-- Printed in The Netherlands
Fungal Remains from the Miocene Quilon Beds of Kerala State, South India PRAMOD KUMAR Birbal Sahni Institute of Palaeobotany, Lucknow - - 226007 (India)
(Received April 13, 1988; revised and accepted October 2, 1989)
Abstract
Kumar, P., 1990. Fungal remains from the Miocene Quilon Beds of Kerala State, South India. Rev. Palaeobot. Palynol., 62: 13-28. Fossil fungal remains are described from the Quilon Beds (Lower-MiddleMiocene) exposed at Padappakkara, Quilon district in the Kerala State, South India. The fungal spores are represented by 15 genera and 23 species, of which 2 genera viz., Elsikisporonites and Hypoxylonsporites and 18 species are new. A tropical humid climate with heavy precipitation probably prevailed during the time of deposition.
Introduction
During the past three decades much attention has been paid to the study of fossil fungal remains in India. Rao (1959), Jain and Gupta (1970), J a i n (1974), Rao and Ramanujam ( 1 9 7 5 , 1976), Ramanujam and Rao (1978), Jain and Kar (1979), Ramanujam and Srisailam ( 1 9 8 0 ) have made valuable contributions to the study of fungal spores and fruiting bodies from Quilon Beds of Kerala State. An Upper Miocene age (Paulose and Narayanaswami, 1968) pertains to limestones overlying carbonaceous shales of Lower-Middle Miocene age (Rao and Ramanujam, 1982). This dating is also supported by Jacob and Sastri (1952) and Dey (1962). Recently Raha and Rajendran (1983) have suggested a Late Eocene-Oligocene age for the lower part of the sequence (Quilon Beds) on the basis of the foraminiferal assemblage, The present paper deals with the study of fungal spores from the slides previously studied by Jain and Gupta (1970). The geology of 0034-6667/90/$03.50
the area, the material, collected by K.P. J a i n from a clay mine section near Kanjantheria House, Padappakkara (11 km north east of Quilon) and the maceration technique employed for the recovery of fungal entities have already been described by Jain and Gupta (1970), Jain and Kar (1979) and Ramanujam (1982b).The slides are stored in the Museum of the Birbal Sahni Institute of Palaeobotany, Lucknow. The locus typicus for the material is Padappakkara, Quilon District, Kerala State, South India and the stratum typicum is Quilon Beds (Lower-Middle Miocene).
Stratigraphy of the area The marine carbonate and the continental clastic facies of the Tertiary sequence occurring in the coastal region of Kerala have been named as the Quilon and Warkalli Beds (Younger) respectively by King (1882) and Bruce Foote (1883).The entire Tertiary sequence of Kerala rests unconformably upon Archaean Gneisses
© 1990Elsevier Science Publishers B.V.
14
P. KUMAR
76"35"
76 °
40'
,.9
55"
55'
--%+,/ .)
"
'~l~
JJ/
i~
Ba"
SCALE
1:2500000
Map 1. Showing the location of exposure at Padappakkara Quilon, Kerala. and itself is overlain by Recent to sub-Recent marine and estuarine sediments. Description of new taxa Genus
Inapertisporites
van der Hammen,
1954 e m e n d . S a x e n a e t B h a t t a c h a r y y a ,
1987
Type species: Inapertisporites variabilis van der Hammen, 1954
Inapertisporites crenulatus K u m a r sp. nov. (Plate I, 1, 2; Fig.l)
Holotype: P l a t e
I,
2;
S1.
No.
3848;
Size
25 x 28 ~m.
Description: F u n g a l spores unicellular, inaperturate, l e n t i c u l a r in shape. Size 17-25 x 2528 pm in diameter. Spore wall differentially thickened, + 2 ~m thick, e q u a t o r i a l m a r g i n w a v y s i m u l a t i n g crested outline, crests + 2 ~m high and _ 3 ~tm broad in optical section. B o t h the surfaces smooth. A polar t h i n n e r area, 9--10 ~tm in diameter. Specimen studied: 15. Comparison: Inapertisporites crenulatus K u m a r
FUNGAL REMAINS FROM MIOCENE QUILON BEDS
15
does not appear similar to any of the species of this genus due to its crenulated margin, Recently, Jarzen and Elsik (1986) described fungal spores as Geastrum-type (Plate I, 12-14) from the Recent sediments of Luangwa Valley, Zambia, which are comparable with I. crenulatus Kumar. They mentioned, as already informed by Dring (1964), t h a t it grows on soil or vegetable debris in damp places.
differs from all the known species of this genus in having hyaline, hollow, rounded verrucaelike sculptures at the peripheral margin of the spores. Derivation of specific name: Based on peripheral verrucae.
Exesisporites sp. Q (Plate I, 3) Description: Fungal spore unicellate, psilate,
Inapertisporites sp. P. (Plate I, 17) Description: Fungal spore unicellular and sickle-shaped with tapering ends. Size 28-30 x 7-8 ~m. The spore wall _ 1 ~m thick, smooth and hyaline. A thick exinal line seen in the photograph is caused by the oblique preservation only. Specimens studied: 3. Comparison: The studied specimen is different from all the known species in having a sickleshaped body. Affinity: It resembles the conidia of modern taxon Chloridiella (Kendrick and Carmichael in Ainsworth et al., 1973).
broadly oval and monoporate. Size 27 x 21 ~m. Minute pore visible in a darker oval area. Spore wall smooth, __l.6~m thick. Darker subcircular patch in the central region of the spore measures 9-11 ~m in diameter. Comparison: The studied specimen closely compares with Exesisporites neogenicus Elsik (1969, plate 1, figs.3, 5, 6) in possessing a comparative minute pore in the darker region but differs in its broadly oval shape and thicker spore wall. G e n u s Lacrimasporo~ites e m e n d . Elsik, 1968
Clarke,
1965
Type species: Lacrimasporonites levis Clarke, Genus Exesisporites Elsik, 1969
1965
Type species: Exesisporites neogenicus Elsik,
Lacrimasporonites
1969
(Plate I, 6; Fig.3)
Exesisporites
verrucatus
Kumar
sp.
nov.
(Plate I, 4; Fig.2)
niger
Kumar
sp.
nov.
Holotype: Plate I, 6; S1. No. 3841; Size 35 x 14 ~m.
Description: Fungal spore unicellular, monopoHolotype: Plate I, 4; S1. No. 3841; Size 28 x 30 ~m.
Description: Fungal spores lenticular in shape and unicellate. Size 25-27 x 28-30 ~m. Monoporate, pore + 1 pm in diameter surrounded by + 1.6 ~m thick, diffuse margin. Spore wall two layered ___2~m thick, outer layer forming 16-18 hyaline, hollow, rounded verrucae-like sculptures at peripheral margin of spore, measuring 2-2.5 ~m high and 1.6-2.5 ~m broad at base, rather evenly placed, imparting angularity to the ambitus, Specimens studied: 4. Comparison: Exesisporites verrucatus Kumar
rate and elliptical with acutely rounded ends. Size 30-35 x 13-15 ~m. Single pore at one end. Spore wall smooth, _ 0.75 ~m thick, slightly thicker around the pore. Specimens studied: 10.
Comparison: Lacrimasporonites
singularis
Sheffy et Dilcher (1971) differs from L. niger Kumar in being smaller (10.6 x 16.4 ~m) and oval to oblong in shape and having a pore at one side of longitudinal centre. L. basidii Elsik (1968) is smaller (9-14 x 6-8 ~m) and has a spatulate shape with a distinctly two layered spore wall. Lacrimasporonites sp. Elsik et Dilcher (1974) is smaller and has thinner wall
~
r~
C3o
NI
d~
.~l
-
C
q
FUNGAL REMAINS FROM MIOCENE QUILON BEDS
Fig.1. D i a g r a m m a t i c s k e t c h o f Inapertisporites erenulatus K u m a r sp. nov. S h o w i n g e r e n u l a t e m a r g i n a n d a p o l a r t h i n n e r s u b c i r c u l a r a r e a at o n e face in t h e s p o r e wall.
around the pore. L. magnus Saxena et Singh (1982) is much longer (120gm) and has a broader pore (27 ~tm diam.) with thickened pore margin. Affinity: Elsik (1968) suggested that such fungal spores could be related to Coprinus, Cortinarius and Stropharia of Basidiomycetes (Graham, 1962). Derivation of specific name: Diagnostic for its dark wall (Latin niger = black).
17
Fig.2. D i a g r a m m a t i c s k e t c h of Exesisporites verrueatus K u m a r sp. nov. S h o w i n g h y a l i n e v e r r u c a e at p e r i p h e r a l m a r g i n of spore a n d a pore w i t h t h i c k e n e d outline.
G e n u s Diporisporites van der H a m m e n , 1954 e m e n d . Elsik, 1968
Type species: Diporisporites elongatus van der Hammen, 1954
Diporisporitespsilatus Kumar sp. nov. (Plate I, 18; Fig.4)
PLATE I All p h o t o m i c r o g r a p h s a r e f r o m u n r e t o u c h e d n e g a t i v e s a n d m a g n i f i e d ca. 1000 x . C o - o r d i n a t e s of e a c h s p e c i m e n s t u d i e d u n d e r E r g a v a l T r i n o c u l a r M i c r o s c o p e No. 663567 g i v e n in p a r e n t h e s i s . 1, 2. Inapertisporites crenulatus K u m a r sp. nov.: S1. Nos. 3844 (111.9 x 20), 3848 ( h o l o t y p e - l l 2 x 6.7). 3. Exesisporites sp. Q: S1. No. 3841 (101.0 × 14.8). 4. Exesisporites verrucatus K u m a r sp. nov.; S1. No. 3841 (holotype-108.1 x 14.9). 5. Imprimospora ramanujamii K u m a r sp. nov.; S1. No. 3842 (holotype-106.7 × 5). 6. Lacrimasporonites niger K u m a r sp. nov.; S1. No. 3842 (holotype-100.2 x 14.8). 7, 8. Elsikisporonites tubulatus K u m a r , sp. nov.; S1. Nos. 3843 (holotype-96.7 x 12.3), 3844 (90.4 × 7.3). 9. DiceUaesporites elongatus K u m a r sp. nov.; S1. No. 3843 ( h o l o t y p e - 8 7 × 18.1). 10. Psidimobipiospora scabratus K u m a r sp. nov.; S1. No. 3841 (holotype-93.4 × 16.1). 11. Brachysporisporites tenuis K u m a r sp. nov.; S1. No. 3844 (holotype-82.8 × 16.2). 12. Dicellaesporites keralensis K u m a r sp. nov.; S1. No. 3844 (holotype-81.8 × 16.8). 13. Hypoxylonsporites ater K u m a r sp. nov.; S1. No. 3847 ( h o l o t y p e - l l l . 6 x 15.4). 14. Multicellaesporites indicus K u m a r sp. nov.; S1. No. 3843 ( h o l o t y p e 108.7 × 6). 15. Diporicellaesporites concavus K u m a r sp. nov.; Sl. No. 3849 ( h o l o t y p e - 1 1 7 × 15.6). 16. Dyadosporites novus K u m a r sp. nov.; S1. No. 3847 (holotype-101.5 × 10.1). 17. Inapertisporites sp. P; Sl. No. 3845 (101 x 4.7). 18. Diporisporites psilatus K u m a r sp. nov.; S1. No. 3842 (holotype-108.8 x 12). 19. Dyadosporites dubius K u m a r sp. nov. S1. No. 3843 ( h o l o t y p e - 1 0 0 x 7.3). 20. Diporicellaesporites sp. K. S1. No. 3854 (104.1 × 21.2). 21. Diporicellaesporites concavus K u m a r sp. nov.; S1. No. 3848 (114.6 × 22.8). 22. Fusiformisporites acutus K u m a r sp. nov.; S1. No. 3844 ( h o l o t y p e - 1 1 4 x 17.7). 23. Hypoxylonsporites miocenicus K u m a r sp. nov.; S1. No. 3845 (holotype-114.5 x 12.3). 24. Tetraploa sp. A; S1. No. 3842 (110 x 12.8). 25. Diporicellaesporites padappakkarensis K u m a r sp. nov.; S1. No. 3846 (holotype-101.6 × 7.1). 26. Tetraploa sp. B; S1. No. 3842 (85.0 × 13).
18
P. KUMAR
' 10tim ' Fig.3. D i a g r a m m a t i c s k e t c h of Lacrimasporonites K u m a r sp. nov. S h o w i n g slit like pore at one end.
niger
Comparison: Involutisporonites Clark emend. Elsik (1968) differs in being multicellate. Palaeocirrenalia Ramanujam et Srisailam (1980) and Colligerites Jain et Kar (1979) are nonaperturate and multicellate. Affinity: Resembles the conidiogenous cells produced by Helicosporium (Kendrick et Carmichael in Ainsworth et al., 1973). Derivation of Generic name: The generic name recognizes Dr. William C. Elsik of Exxon Company USA, Houston, USA. Elsikisporonites
10 fl.Lm
~
Holotype: Plate I, 18, S1. No. 3842; Size 19 x 10 pm.
Description: Fungal spores unicellate, elongate, fusiform and diporate. Size 19-40x 10 221~m. Pore at each end of the spore, simple, slightly sunken and ___2 pm wide. Spore wall smooth, 1 pm thick. Specimens studied: 7. Comparison: Diporisporites hammenii Elsik (1968) is smaller (11-14 x 6-8 Wn) and has bulging pores. D. conapicua Ramanujam et Rao (1978) differs from D.psilatus Kumar in being bigger size (58 × 37 Wn), thickened pore margins and distinct atria. D. curvatus Ramanujam et Rao (1978) possesses more curved sides and thicker spore wall. Diporisporites sp. Hopkins (1969) is longer (55-60 ~m) than the present species. Diporisporites elongatus van der Hammen (1954) shows curved spore wall near the pores, hence, it is different from the present species.
Elsikisporonites
Kumar sp.
nov.
(Plate I, 7, 8; Fig.5)
Fig.4. D i a g r a m m a t i c sketch of Oiporisporites psilatus K u m a r sp. nov. S h o w i n g s l i g h t l y s u n k e n pore at end.
Genus
tubulatus
Kumar
gen.
Holotype:
Plate I, 7; SI. No. 3843; Size 36.8 x 38.4 pm (ambitus). Description: Fungal spores monoporate, nonseptate, tubular in shape and coiled. Cell broadest in the middle region, 10-12 ~m wide, gradually tapering towards the ends. Pore at free outer end, small, nozzle-like, 1.5 Ilm wide. Spore wall 1 ~m thick, smooth, slightly folded, hyaline.
Specimens studied: 8. G e n u s Hypoxylonsporites K u m a r gen. n o v .
Type species: Hypoxylonsporites miocenicus Kumar gen. et sp. nov.
nov.
Type species: Elsikisporonites tubulatus Kumar sp. nov.
I
I 10,/t..1,rn
Diagnosis: Fungal spores monoporate, nonseptate, tubular and coiled. Pore a t o u t e r end, nozzle-like. Spore wall smooth and hyaline,
Fig.5. D i a g r a m m a t i c s k e t c h of Elsikisporonites tubulatus K u m a r gen. et sp. nov. S h o w i n g t u b u l a r coiled cell w i t h a nozzle-like pore at free uncoiled end.
FUNGAL REMAINS FROM MIOCENE QUILON BEDS
Diagnosis: Fungal spores unicellular, oval to elliptical in shape with acutely rounded ends. A longitudinal slit-like aperture may be running end to end. Spore wall single layered, smooth and may be differentially coloured. Affinity: The form genus Hypoxylonsporites Kumar shows close affinity with extant genus Hypoxylon (See Miller, 1961). Specimens of Hypoxylonsporites also show resemblance with Recent Endocalyx. Remarks: Venkatachala and Rawat (1973) recorded a specimen similar to the present new genus from the subsurface Tertiary sediments of Cauvery Basin, S. India. Sah (1967) reported such fungal spores from the Neogene sediments of Rusizi Valley (Burundi). Elsik and Dilcher (1974) also recorded similar elongate type of fungal spores having a linear fold, slit or scar, from Middle Eocene clay beds in the Lawrence Clay Pit, Henry County, Tennessee. Miller (1961, fide Elsik and Dilcher, 1974) compared such specimens with the cosmopolitan saprophytic fungi Hypoxylon (Bulliard, 1791) Fries (1849). Elsik (1969) figured similar spores from the Late Neogene deposits in northern Gulf of Mexico and referred to them as Hypoxylon spp. (Plate I, 1, 2). Subramanian (1971) stated that conidia of Endocalyx thwaitesii Berkeley et Broome (1850) and E. melanoxanthus (Berkeley et Broome) Petch (See Hughes, 1953) of Hyphomycetes are one-celled, flattened-oval with a longitudinal germ slit. During the course of the present study, few fungal spores similar to Hypoxylon have been recorded from the Miocene Quilon deposits at Padappakkara. It is suggested that such fossil spores recorded by the earlier workers should be referred to Hypoxylonsporites gen. nov.
19
i
lo/a m Fig.6. Diagrammatic sketch of Hypoxylonsporites rniocenicus Kumar sp. nov. Showing a longitudinal slit extending from one end to the other.
tends from one end to the other. Spore wall single-layered, + 1.6 ~m thick and smooth.
Specimens studied: 16. Hypoxylonsporites ater Kumar sp. nov. (Plate I, 13; Fig.7) Holotype: Plate I, 13; S1. No. 3847; Size 29 × 18 ~m. Description: Fungal spores unicellular, broadly oval in shape, ends tapered. Size 25-30x 18-20 ~m. Longitudinal slit runs from end to end. Spore wall single layered, _ 1.6 ~m thick and smooth. Specimens studied: 6.
Comparison: Hypoxylonsporites miocenicus Kumar differs from H. ater Kumar in having elongate shape with narrowly rounded ends.
Hypoxylonsporites rniocenicus Kumar sp. nov. (Plate I, 23; Fig.6) Holotype: Plate I, 23; S1. No. 3845; Size 32 x 13 ~m. Description: Fungal spores unicellular, elliptic t o spindle-shaped with narrowly rounded ends. Size 21-55 x 9-26 ~m. A longitudinal slit ex-
I
20ill m
I
Fig.7. D i a g r a m m a t i c s k e t c h of Hypoxylonsporites ater K u m a r sp. nov. S h o w i n g l o n g i t u d i n a l slit w i t h g r e a t e s t
width of spore in the middle region.
20
P. KUMAR
Genus Dicellaesporites Elsik 1968 e m e n d . S h e l l y et Dilcher, 1971
Type speices: Dicellaesporites popovii Elsik, 1968
Dicellaesporites elongatus Kumar sp. nov. (Plate I, 9; Fig.8)
Holotype: Plate I, 9; S1. No. 3843; Size 43 x 14 ~m.
Diagnosis: Spores dicellate, fusiform and inaperturate. Size 41-50 x 14-17 pm. Cells gradually tapering at either end. Both spore wall and septum __1 ~m thick. Spore wall smooth, light coloured and folded. Septum centrally porate,
Specimens studied: 8. Comparison: The present species differs from D.popovii Elsik (1968) in having narrowly rounded ends. D. ellipticus Jain et Kar (1979) differs in having larger size and ornamented wall. Affinity: The present species shows resemblance with the conidiogenous axis of Dactylaria Saccardo (1880; fide Ainsworth et al., 1973).
Dicellaesporites keralensis Kumar sp. nov.
Holotype: Plate I, 12; S1. No. 3844; Size 21 x 16 pm.
Description: Spores dicellate, inaperturate and broadly oval in shape. Size 18-22 x 13-16 pm. Broadly rounded at free ends but slightly constricted at the septal region, cells unequal in size, 1 0 - 1 3 x l l - 1 4 ~ m . Both septum and spore wall _ 1 ~m thick and smooth. Specimens studied: 11. Comparison: D. popovii Elsik (1968) lacks the constriction at the septum. D. eUipticus Jain et Kar (1979) differs in having microreticulate spore wall and bigger size range. D. disphaericus Shaffy et Dilcher (1971) is distinct in having scabrate spore wall, well pronounced opaque equatorial septum and constriction. G e n u s Imprimospora Norris, 1986
Type species: Imprimospora tankensis Norris, 1986
Imprimospora ramanujamii Kumar sp. nov. (Plate I, 5; Fig.10)
Synonymy 1980 3-celled striate spore type: Ramanujam et Srisailam, pl.2, fig.28, p.131.
Holotype: Plate I, 5; S1. No. 3842; Size
(Plate I, 12; Fig.9)
32 x 18 pm.
Description: Fungal spores tricellate, inapertu-
' Fig.8. Diagrammatic sketch of Dicellaesporites elongatus Kumar sp. nov. Showing dicellate fusiform cell with a centrally slitted septum.
rate and fusiform. Size 32-40 x 18-21 ~m. Septa 1.6-2 ~m thick, with or without a central pore. Central cell large, 11-13 x 16-19 ~m, terminal cells small, 9-10~m long with narrowly
d
Fig.9. Diagrammatic sketch Dicellaesporites keralensis Kumar sp. nov. Showing dicellate spore without any slit.
Fig.10. Diagrammatic sketch of Imprimospora ramanujamii Kumar sp. nov. Showing tricellate spore with central larger cell and longitudinal ribs converging at end.
FUNGAL REMAINS FROM MIOCENE QUILON BEDS
21
rounded ends. Spore wall _ 1.5 ~m thick and ornamented with longitudinal ribs ( ± 2 ~ m wide) that run full length but are more pronounced. Furrows 1-2 ~m wide. Specimens studied: 7. Comparison: Imprimospora ramanujamii Kumar closely resembles 3-celled ribbed spore type described by Ramanujam and Srisailam (1980, pl.2, fig.28). I. tankensis Norris 1986 (Fide Jansonius and Hills, 1987: C.F.4444) differs in having thinner (0.5 ~) spore wall with uniseptate body. Derivation of specific name: The species is named after Dr. C.G.K. Ramanujam, Hyderabad, India.
Comparison: Fusiformisporites acutus Kumar differs from F. elongatus Ramanujam et Rao (1978) in having larger size and acute ends. F. keralensis Ramanujam et Rao (1978) differs from the present species in having rhomboidal shape with cells wider at the septal region. F.pseudocrabbii Elsik (1968) is different in having finer ribs made of rows of punctae parallel to the coarse ribs, F. microstriatus Hopkins (1969) differs in having smaller size (42-49 ~m in length) and finer ribs.
Genus Fusiformisporites Rouse, 1962
1965
Type species: Fusiformisporites crabii Rouse,
Dyadosporites novus Kumar sp. nov. (Plate I,
1962
16; Fig.12)
G e n u s Dyadosporites v a n d e r H a m m e n ex C l a r k e , 1965
Type species: Dyadosporites ellipsus Clarke,
Fusiformisporites
acutus
Kumar
sp.
nov.
(Plate I, 22; Fig.ll)
Holotype: Plate I, 16; S1. No. 3847; Size 32 x 16 ~m.
Description: Fungal spores dicellate and dipoHolotype: Plate I, 22, S1. No. 3844; Size 96x21 ~m.
Description: Fungal spores dicellate, inaperturate and elongated-fusiform. Size range 90-112 x 20-24 ~m generally tapering towards acute ends. Septum _ 1 ~m thick. Spore wall 1-1.6 ~m thick and ribbed. Ribs _ 1 ~m wide, wider 1.6-2.5~m at septal region, closely spaced, alternate or continuously running end to end, sometimes bifurcating and converging at the ends, furrows very narrow. Specimens studied: 5.
I
20/U.m
rate. Size 28-32×13-161Jm. Cells fusiform and gradually taper towards ends. Apical pore 0.5 ~m and simple. Septum 2-3 ~m thick, wedge-shaped or denticulate with a central pore, dark and opaque. Spore wall 1.6 ~tm thick, smooth, slightly constricted at septal region and showing a brief collar at one end (probably basal e n d ) o f the spore.
Specimens studied: 6. Comparison: Dyadosporites ellipsus
Clarke (1965) differs in having longer cells (38-55 ~m) and bigger pores (4-5 ~m diam.).
I
Fig.ll. Diagrammatic sketch of Fusifermisporites acutus Kumar sp. nov. Showing dicellate, acute ends with thick longitudinal ribs continuous or bifurcating and converging at ends.
P KUMAR
22
I
10Nrn
I
Fig.12. Diagrammatic sketch of Dyadosporites novus Kumar sp. nov. Showing dieellate and diporate spore.
t 10/U.m
'
lOpm
Fig.14. Diagrammatic sketch of Psidimobipiospora scabratus Kumar sp. nov. Showing dicellate, diporate spore with scabrate wall.
'
Fig.13. Diagrammatic sketch of Dyadosporites dubius Kumar sp. nov. Showing dicellate, diporate and bracket-
Description: F u n g a l spores cylindrical, dicel-
shaped spore,
late and diporate. Size r a n g e 35-45 x 13-16 pm. Cells r e c t a n g u l a r in shape, each cell 19 x 13-14 pm, 4- equal in size with a pore at free end. P o r e 4-6 I~m wide, simple and slightly sunken. S e p t u m _+3pm thick, opaque and simple. Spore wall ___2pm thick, t h i n n e r a r o u n d pore, surface smooth to finely s c a b r a t e or echinate, slightly c o n s t r i c t e d at septal region. Specimens studied: 10.
Dyadosporites dubius K u m a r sp. nov. (Plate I, 19; Fig.13)
Holotype: P l a t e I, 19; S1. No. 3843; Size 16 x 13 pm.
Description: F u n g a l spores diporate, dicellate and b r o a d l y oval or bracket-shaped (outline). Size 15-18 x 12-13 i~n. P o r e s 4-6 pm wide, simple, one at e i t h e r end. Spore wall ___1.6pm thick, smooth, g r a d u a l l y t h i n n i n g out t o w a r d s the pore. S e p t u m 4-2 pm thick, opaque, Specimens studied: 20. Comparison: Dyadosporites ellipsus Clarke (1965) differs in having elliptical shape and larger size (23-27 x 38-55 ~lm). D. novus K u m a r differs in being narrow apical pores and larger cells,
Genus Psidimobipiospora Locqvin e t Salard-Cheboldaeff in Salard-Cheboldaeff et Locquin, 1980
Comparison:
Psidimobipiospora
dyadospora
Sal-Cheb. et L o c q u i n 1980 (Fide J a n s o n i u s and Hills, 1981, G. F i l e c a r d 3900) differs in h a v i n g t r u n c a t e d extremities and g r a n u l o s e spore wall. Genus Multicellaesporites emend. Kumar
Elsik,
1968,
Type species: Multicellaesporites nortonii Elsik, 1968
Emended Description: F u n g a l spores multicelPsidimobipiospora
dyadospora
Salard-Cheb.
et Locquin, 1980
Psidimobipiospora scabratus K u m a r sp. nov. (Plate I, 10; Fig.14)
Holotype: P l a t e I, 10; S1. No. 3841; Size 40 x 14 tim.
late, elongate. A l o n g i t u d i n a l slit or f u r r o w present. Spore wall smooth or o r n a m e n t e d or differentially c o l o u r e d or t h i c k e n e d . Remarks: Elsik (1968) diagnosed the genus " i n a p e r t u r a t e " , w h e r e a s the type species (pl.2, fig.15) was diagnosed as " t h e grain split l o n g i t u d i n a l l y " . The p r e s e n t s t u d y reveals t h a t the genus has a t e n d e n c y to b e a r a l o n g i t u d i n a l
23
FUNGAL REMAINS FROM MIOCENE QUILON BEDS
slit or furrow. Hence, it has been accordingly emended,
Multicellaesporites elsikii ( R a m a n u j a m et Srisailam, 1980) comb. nov.
Basionym: Staphlosporonites elsikii Ramanujam et Srisailam 1980; Botanique, 9(1-4): p.122, pl.1, figs.6, 7.
ally-coloured or thickened. The middle cell hyaline, 14-19 ~m long, whereas the terminal cells c o m p a r a t i v e l y d a r k e r and smaller in size (8-9 ~m long). Specimens studied: 20. Comparison: It differs from the o t h e r species in h a v i n g differentially t h i c k e n e d wall and shorter aperture.
Genus Brachysporisporites S m i t h , 1971
Synonymy 1959 Multicellular phragmospora Rao, pl.1, fig.14 1968 Hyphae Type-1 Elsik, pl.3, figs.13, 15 1968 Hyphae (?) Elsik, pl.3, Fig.18 1978 MulticeUaesporites ? Ramanujam et Rao, pl.1, Fig.20 1980 Staphlosporonites elsikii Ramanujam et Srisailam, pl.1, figs.6, 7 1981 Inapertisporites globulosus Rouse, 1962 in Ediger, 1981; pl.1, fig.6
Lange
et
Type species: Brachysporisporites pyriformis Lange et Smith, 1971
Brachysporisporites
tenuis K u m a r
sp.
nov.
(Plate I, 11; Fig.16)
Affinity:
Synonym:
dematiaceous h y p h o m y c e t e s Annellophorella (Ellis, 1971) and to some e x t e n t with the
43x30~m.
conidia of Stigmina compacta.
Description: F u n g a l spores di- or multicellate
R a m a n u j a m and Srisailam (1980) showed its strong similarity with the conidia of
Multicellaesporites
indicus K u m a r
sp.
nov.
3843;
Size
(Plate I, 14; Fig.15)
Holotype: Plate
I,
14; S1. No.
32 x 16 ~m.
Description:
F u n g a l spores tricellate, elongated, elliptical or spindle-shaped with narrowly r o u n d e d to a n g u l a r ends. Size r a n g e 31-38 x 11-18 pm. A longitudinal slit present in hyaline middle region. Spore wall smooth, less t h a n 1 pm thick in central region and ± 2 ~m at the terminal and septal regions and differenti-
Fig.15. Diagrammatic sketch of Multicellaesporites indicus Kumar sp. nov. Showing longitudinal slit restricted to hyaline middle cell with comparatively darker terminal ends.
1978 Spore Type B Ramanujam et Rao; pl.1, figs.18, 19
Holotype: P l a t e
I,
11; S1. No.
3844;
Size
and balloon-shaped. Size 38-43 x 22-30 pm. Cells unequal, small cell hyaline and thin, m e a s u r i n g 4-10 x 3-6 ~m broad. L a r g e r apical cell or cells forming a dome-shaped, thick and dark coloured to opaque structure 23-40 x 14-32 ~m. Septa 2-3 ~m thick, smooth to wedge-shaped. Specimens studied: 10.
Fig.16. Diagrammatic sketch of Brachysporisporites tenuis Kumar sp. nov. Showing balloon-shaped multicellate spore with hyaline smaller cell.
24
Comparison:
P. KUMAR
Brachysporisporites
pyriformis
Lange et Smith (1971) is smaller (36--41 x 15-18 pro) and pyriform.
in
size
G e n u s Diporicellaesporites Elsik, 1968
Type species: Diporicellaesporites stacyi Elsik, 1968
Diporicellaesporites concavus Z u m a r sp. nov. (Plate I, 15, 21; Fig.17)
Holotype:
Plate I, 15; S1. No. 3849; Size 38.4 x 22.4 pm. Description: Fungal spores multicellate, diporate and fusiform in shape. Size range 38 42 x 22-24 pm. Spore gradually tapering at both ends. Central cells dark in colour and larger in size (8--13 x 13-21 ~m), apical cells smaller, hyaline, porate and delimited by a concave d a r k e r outline (septum). Spore wall thickest at the central septal region and smooth. Septum straight or disc-shaped, septurn of central cell _+3 pm thick with a central pore. Terminal pores _+2 ~m wide, pore margin thin and hyaline. Specimens studied: 15. Comparison: Diporicellaesporites concavus Kumar differs from all o t h e r k n o w n species having differentially coloured t h i c k e n e d spore wall and concave septa,
Diporicellaesporites padappakkarensis K u m a r sp. nov. (Plate I, 25; Fig.18)
I
I 10/t£m Fig.18. Diagrammatic sketch of DiporicellaesporitespadappakkarensisKumar sp. nov. Showing multicellate unequal cells, pore offset, slit-like and partially divided into two pore cells at the terminal hyaline ends.
Holotype:
Plate 30.4 x 19.2 pm.
I,
25; S1. No.
3846;
Size
Description: Fungal spores multicellate and elongated-elliptical in size. Size 30-35 x 19-22 ~m. Cells unequal in size, the central cell large, 19 x 17 ~m and terminal cells splitting, resulting in a beak-shaped structure, 4-6 pm long. Both ends with single pore, offset, slitlike and measuring 1 2 × 1 - 5 ~ m , partially divided into two pore cells. Septa three, comparatively thicker t h a n the spore wall, disc-shaped with central pore. Spore wall +1.6 ~m thick, smooth. Wall of central cell dark coloured and terminal cells hyaline. Specimens studied: 5. Comparison: Diporicellaesporites concavus Kumar differs in having undivided apical cells at poral region. Remarks: Beak-shaped s t r u c t u r e of the hyaline cells might have developed during the compression for accommodating the thicker wall of central cell.
Diporicellaesporites sp. K (Plate I, 20) Description: This fungal spore is tetracellate,
' 10/trm I Fig.17. Diagrammatic sketch of Diporicellaesporites concavus Kumar sp. nov. Showing tetracellate, diporate spore apical cells hyaline and porate.
diporate and fusiform in shape, its size is 3 0 . 5 x 1 3 ~ m . The central cells are slightly constricted at middle septal region and slightly curved at one side. Terminal cells are smaller in size, tapering with a slightly pouting pore. The septa are nearly as thick as spore wall. The spore wall is 1.0 ~m thick and smooth.
Specimens studied: 1. Comparison: Diporicellaesporites sp. Ramanu-
FUNGAL REMAINS FROM MIOCENE QUILON BEDS
jam et Rao (1978) is different from Diporicellaesporites sp. K in having larger size (81 ~n) and stalk-like basal cells. D. stacyii Elsik (1968) differs in having thick septa, granulate to punctate inner surface of cells and pores with slitted margins. Genus
25
broad, its terminal cell gradually tapering. The septa 1-3 ~m thick, simple to wedge-shaped. The spore wall _ 1 ~m thick, smooth. Specimens studied: 1. Comparison: Tetraploa sp. B differs from T. sp. A in size (36-48 x 6-11 ~m), having a septate main body and narrower appendages.
Tetraploa B e r k e l e y et B r o o m e , 1850 Palaeoclimate
Type species: Tetraploa aristata Berkeley et Broome, 1850 Tetraploa sp. A (Plate I, 24) Description: Fungal conidia slightly triangular in shape with nonseptate body. Size 40-46 × 8-10 pm. Conidium 14 x 10 ~tm. The body terminating into four long processes, Processes 3-4 septate, 32-35 × 3-5 ~m, hyaline, + 1.0 ~m thick and smooth, Specimens studied: 1. Comparison: The present species differs from Type 89: Tetraploa aristata Berkeley et Broome (1850) by its smooth processes and triangular body which in the later is verrucate. Tetraploa (Frasnacritetrus) sp. Singh et Saxena (1981) described from the Upper Siwalik of Himachal Pradesh, closely compares with the present species in having psilate appendages, but differs in exhibiting nonseptate appendages and quadrangular body. Frasnacritetrus josettae Taugourdeau (1968) is different from Tetraploa sp. A in having a quadrangular body and bigger size. Remarks: The author considers that Taugourdeau described as Frasnacritetrus a fungal conidium comparable to Tetraploa Berkeley et Broome and not an acritarch type sporomorph. Tetraploa sp. B (Plate I, 26) Description: Fungal spore branched, multicellate; main body four to five celled and fusiform in shape, i.e. size 60 × 20 ~tm. Apical cell termihating into four appendages, ca. 150 pm long. Cells of main body rectangular, broader than long, 19-21 × 9-11 ~tm. Cells of appendages are longer than broad, 17-27 pm long and 6-8 ~tm
Certain sporomorphs in the Padappakkara sediments are useful in determining the palaeoclimate. The form taxa which are closely related to modern ones (Table I) are most helpful because their modern environmental conditions are known. Elsik (1969) pointed out that the abundant Exesisporites in the northern Gulf of Mexico during the Pliocene might be indicative of a warm climate. The record of Grallomyces, Spegazzinia from Quilon Beds of Kerala, clearly indicates that a tropical, humid climate prevailed during the Neogene time in this region (Ramanujam and Rao, 1978; Ramanujam and Srisailam, 1980). Helicosporous hyphomycetes are usually found on moist wood (Subramanian, 1971). The presence of helicoidal fungal forms (e.g. Elsikisporonites) indicates the same climatic conditions during the sedimentation of Quilon Beds. Lange (1976) discussed the value of the fossil TABLEI Affinities of fossil fungal remains with modern taxa Fossil taxa
Modern taxa
Inapertisporessp. P
I. crenulatus
Chloridiella Geastrum saccatum G. lageniforma
Lacrimasporonites
Basidiomycetes (Coprinus,
Elsikisporonites
Cortinarius, Stropharia) Helicosporium
Hypoxylonsporites Dicellaesporiteselongatus
Hypoxylon Dactylaria
Spegazzinites Multicellaesporiteselsikii
Spegazzinia AnnellophoreUa,Stigmina
Tetraploa
compacta Tetraploa
Fusiformisporites Brachysporisporites
Cookeinia and Porodiella Brachysporium
26
epiphyllous microthyriaceous fungi and their germlings as an indicator of palaeohabitats. He pointed out that abundance of such germlings in the deposits could be a good indicator of moist tropical type of vegetation. This is apparent in the Quilon Beds, Kerala, as was also recorded by Jain and Gupta (1970), Ramanujam and Rao (1972), Rao and Ramanujam (1976) and Jain and Kar (1979). Rao and Ramanujam (1975) studied the palynoflora from these Quilon beds and recorded spores and pollen grains which belonged to pteridophytes and angiosperms. They mentioned that the occurrence of such families points towards tropical humid conditions, viz., Gleicheniaceae, Schizaeaceae, Polypodiaceae, Aroideae, Palmae, Lecythidaceae, Araliaceae, Symplocaceae, Rhizophoraceae, Sonneratiaceae, Bombacaceae, Caesalpiniaceae, Combretaceae, Sapotaceae, *Dipterocarpaceae, *Ctenolophonaceae, Meliaceae, *Olacaceae, Anacardiaceae and Droseraceae. The abundance of ferns (families marked with an asterisk) and Duabanga of Sonneratiaceae indicates heavy precipitation also. The climate during the Neogene of Kerala was inferred as tropical humid. There is not much of a change from the past to the present climatic conditions, Raha and Rajendran (1983) suggested that the deposition of continental facies both below and above the Quilon Formation is indicative of major transgressive and regressive phases, The fungal remains and the miospores (Rao and Ramanujam, 1978, 1982) that occur in these sediments further suggest that the deposits are from fresh water lake or ponds where lowland vegetation thrived.
P. KUMAR
fungal spores from carbonaceous clays of Padappakkara. Fungal remains from the Palaeogene and Neogene beds of India have been studied by Rao (1959), Kar et al. (1972), Biradar and Mahabale (1974), Kar and Saxena (1976), Varma and Rawat (1963), Jain and Gupta (1970), Ramanujam and Rao (1978), Ramanujam and Srisailam (1980), Jain and Kar (1979), Ramanujam (1982a, b). On comparison of the present study with the previous ones, it can be seen that genera like
DiceUaesporites,Diporicellaesporites, Brachysporisporites, Involutisporonites, Psidimobipiospora, Imprimospora, MulticeUaesporites, Meliola, Pluricellaesporites, Basidiosporites, Staphlosporonites and Inapertisporites have been recorded from Palaeogene as well as Neogene beds. Therefore such long ranging taxa at the generic level are not useful for stratigraphic consideration. Characteristic forms recorded from Palaeogene strata (Kar et al., 1972; Mahabale, 1969; Biradar and Mahabale, 1974) but so far not reported from the Neogene are: Cucurbitariaceites, Diplodia and Tetracoccosporium eocenum (the last two genera being extant taxa). These forms were also absent in the assemblage studied here. Some forms that have been recorded from Neogene sediments but are absent in Palaeogene sediments are: Quilonia, Diploneurospora,
Comparison with other assemblages
Spegazzinites, Foveo letisporonites, Warkallisporonites, Exesisporites, Hypoxylonsporites (= Hypoxylon), Brachysporisporites tenuis, Retihelicosporonites. Ramanujam (1982a) stated that Fusiformisporites, Palaeocirrenalia-like coiled spores and Colligerites-like helical spores are also present
Jain and Gupta (1970) described some fruiting bodies and fungal spores from Quilon carbonaceous clay beds exposed at Padappakkara. Later, Rao and Ramanujam (1976), Ramanujam and Rao (1978) and Jain and Kar (1979) studied fruiting bodies and fungal spores from the Neogene sediments of the Kerala Coast. The present study is based on
in Miocene beds of Neyveli Lignite, South Arcot, Madras. Some palynologists seem to hold the opinion that fungal spores may not be helpful in resolving stratigraphic problems, but their importance for stratigraphy has been highlighted by Elsik (1968), Graham (1962) and Jansonius (1976). This has encouraged the author to undertake this study. Elsik (1977),
FUNGALREMAINSFROMMIOCENEQUILONBEDS R a m a n u j a m a n d R a o (1978), R a m a n u j a m a n d S r i s a i l a m (1980) a n d R a m a n u j a m (1982a, b)
suggested that ornamented diporate and foveolate fungal spores are not known earlier than Eocene and Miocene respectively; reticulate helicoidal s p o r e s a r e known from Miocene onwards. Ramanujam further stated that spore types with highly sculptured walls are encount e r e d c o n s i s t e n t l y in N e o g e n e s e d i m e n t s . F o r m s like Foveoletisporonites miocenicus (fov e o l a t e d i p o r a t e ) , Retihelicosporonites elsikii
(reticulate helicoidal), Exesisporites, *CoUigererites ( h e l i c o i d a l spores), *Spegazzinites (sculpt u r e d forms), Hypoxylonsporites l o n g i t u d i n a l l y slitted forms) along with
some ornamented
forms, viz., Exesisporites verrucatus sp. nov., Psidimobipiospora scabratus sp. n o v . m a y be s t r a t i g r a p h i c a l l y s i g n i f i c a n t e l e m e n t s . T h e fungal s p o r e s m a r k e d b y a s t e r i s k s a n d o r n a m ented forms are therefore further indications t h a t t h i s a s s e m b l a g e m i g h t h a v e b e e n deposited d u r i n g t h e N e o g e n e P e r i o d ( M i o c e n e ) . Conclusion
(1) On the basis of palaeontological and palynological
evidence,
a Lower to Middle
M i o c e n e a g e is a s s i g n e d t o t h e Q u i l o n c a r b o n a c e o u s c l a y beds. (2) T h e p a r t i c u l a r a s s e m b l a g e o f f u n g a l remains and miospores indicates that Quilon c a r b o n a c e o u s c l a y beds h a v e b e e n d e p o s i t e d in an area having ponding environments. (3) T h e m o s t a b u n d a n t f u n g a l e n t i t i e s a r e suggestive of humid climate. Acknowledgements T h e author is grateful to Dr. B.S. Venkatachala, Director, B.S.I.P., L u c k n o w for valuable suggestions a n d for critically reading the manuscript. I a m thankful to Dr. R.K. S a x e n a for his kind help. References Ainsworth, G.C., Sparrow, F.K. and Sussman, A.S. (Editorn),1973.The Fungi, An Advanced Treatise.Vol IV, A, Academic Press, N e w York, N.Y., pp.1-509.
27 Berkely, M.J. and Broome, C.E., 1850. Notices of British fungi. Ann. Mag. Nat. Hist., 2(5): 459. Biradar, N.V. and Mahabale, T.S., 1974. On the occurrence of unperfect fungus, Tetracocosporium obtained from a fossil wood belonging to the Deccan Intertrappean series. Palaeobotanist, 21: 223-226. Bruce Foote, R., 1883, On the geology of South Travancore. Rec. Geol. Surv. India, 16(1): 20-35. Clarke, A.T., 1965. Fungal spores from Varmeju Formation coal beds (Upper Cretaceous) of Central Colorado, Mt. Geo1.,2:85-93. Dey, A.K., 1962. The Miocene mollusca from Quilon, Kerala (India). Mere. Geol. Surv. India Palaeontol. Indica, N.S., 36: 1-129. Dring, D.M., 1964. Gasteromycetes of the West Tropical Africa. Mycol. Pap., 98: 1-60. Ellis, M.B., 1971. Dematiaceous Hyphomycetes. Commonw. Mycol. Inst. Kew, Surv.: 508 pp. Elsik, W.C., 1968. Palynology of a Palaeocene Rockdale lignite Milam Country, Texas - - 1. Morphology and Taxonomy. Pollen Spores, 10: 263-314. Elsik, W.C., 1969. Late Neogene palynomorph diagram, northern Gulf of Mexico. Gulf Coast Assoc. Geol. Soc. Trans., 19: 509-528. Elsik, W.C., 1977. Morphology phylogeny of dispersed fossil fungal spores - - Intimations. Second Int. Mycol. Congr., Univ. South Florida, Tampa, Abs.: 169. Elsik, W.C. and Jansonius, J., 1974. New genera of Palaeogene fungal spores. Can. J. Bot., 52: 953-958. Graham, A., 1962. The role of fungal spores in palynology. J. Palaeontol., 36: 60-68. Jacob, J. and Sastri, V.V., 1952. Miocene foraminifera from Chavara near Quilon, Travancore. Rec. Geol. Surv. India, 82: 342-353. Jain, K.P., 1974. Fossil Fungi. An Aspects and Appraisal of Indian Palaeobotany, B.S.I,P., Lucknow, pp.38-46. Jain, K.P. and Gupta, R.C., 1970. Some fungal remains from the Tertiaries of Kerala Coast. Palaeobotanist, 18: 177-182. Jain, K.P. and Kar, R.K., 1979. Palynology of Neogene sediments around Quilon and Varkala, Kerala Coast, South India - - 1. Fungal remains. Palaeobotanist, 26: 105-118. Jansonius, J., 1976. Palaeogene fungal spores and fruiting bodies of the Canadian Arctic. Geosci. Man, 15: 129-132. Jansonius, J. and Hills, L.V,, 1976-1987. Genera File of Fossil spores, Spec. Publ. Dep., Geol. Univ. Calgary, Canada and Supplements. Jarzen,D.M. and Elsik,W.C., 1986. Fungal palynomorphs recovered from recent deposits, Luangwa Valley, zambia. Palynology, 10: 35-60. Kar, R.K. and Saxena, R.K., 1976. Algal and fungal microfossils from Matanomadh Formation (Palaeocene), Kutch, India. Palaeobotanist,23: 1-15. Kar, R.K., Singh, R.Y. and Sah, S.C.D. 1972. Some algal and fungal remains from Tura Formation of Garo Hills, Assam. Palaeobotanist, 19: 146-154. Kendrick, W.B. and Carmichael, J.W., 1973. Hyphomycetes. In: G.C. Ainsworth, F.K. Sparrow and A.S. Sussman (Editors),The Fungi, An Advanced Treatise. Vol. IV, A, Academic Press,N e w York, N.Y., pp.323-501.
28 King, W., 1882. General sketch of the geology of Travancore State. The Warkali beds and associated deposits at Quilon in Travancore. Rec. Geol. Surv. India, 15(2): 93-102. Lange, R.T., 1976. Fossil epiphyllous "germlings" their living equivalents and their palaeohabitat indicator value. N. Jahrb. Geol. Palaeontol. Abh., 151: 142 165. Mahabale, T.S., 1969. On a fossil species of Diplodia from the Deccan Intertrappean Series, M.P., India. Palaeobotanist, 17(3): 295 297. Miller, J.H., 1961. A monograph of the world species of Hypoxylon. Univ. Georgia Press, Athens, 158 pp. Paulose, K.V. and Narayanaswami, S., 1968. The Tertiaries of Kerala Coast. Mem. Geol. Soc. India, 2: 300--308. Raha, P.K. and Rajendran, C.P., 1983. Basal Tertiary ferruginous Sandstones in the west coast of southern India and their geological significance. Q.J. Geol. Min. Metall. Soc. India, 56(4): 208-216. Raju, D.S.N., 1978. Contribution to the Neogene stratigraphy of two areas of Kerala basin with special reference to Myogypsinidae (Abstract). VII Indian Colloq. Micropaleontol. Stratigr. Madras. Ramanujam, C.G.K., 1982a. Recent advances in the study of fossil fungi. Recent advances in Cryptogamic Botany Pt. II: Fossil Cryptogams. Palaeobot. Soc. Lucknow.: 287 301. Ramanujam, C.G.K., 1982b. Tertiary palynology and Palynostratigraphy of southern India. Palaeontol. Soc. India, Spec. Publ., 1:57 64. Ramanujam, C.G.K. and Rao, K.P., 1978. Fungal spores from the Neogene strata of Kerala in South India. IVth Int. Palynol. Conf., Lucknow (1976 77), 1: 291 304. Ramanujam, C.G.K. and Srisailam, K., 1980. Fossil fungal
P. KUMAR spores from the Neogene beds around Cannanore in Kerala State. Botanique, 9(1 2): 119-138. Rao, A.R., 1959. Fungal remains from some Tertiary deposits of India. Palaeobotanist, 7:43 46. Rao, K.P. and Ramanujam, C.G.K., 1975. A palynological approach to the study of Quilon beds of Kerala State in South India. Curr. Sci., 44(20): 730-732. Rao, K.P. and Ramanujam, C.G.K., 1976. A further record of microthyriaceous fungi from the Neogene deposits of Kerala in South India. Geophytology, 6: 98-104. Rao, K.P. and Ramanujam, C.G.K., 1982. Palynology of the Quilon beds of Kerala State in South India - - II Pollen of Dicotyledons and Discussions. Palaeobotanist, 30(1): 68-100. Saccardo, P.A., 1880. Conspectus generum fungorum italiac inferiorum nempe and Sphaeropsideas. Melonconicas et Hyphomyceteas peritinentium, Systematic sporologico dispositorum. Michelia, 2:1 39. Sah, S.C.D., 1968. Palynology of an Upper Neogene profile from Rusizi Valley (Burundi). Ann. Mus. R. Afr. Cent. Ser. Sci. Geol., 88(57): 1 173. Saxena, R.K. and Sarkar, S., 1986. Morphological study of Frasnacriterus Taugourdeau emend, from the Tertiary sediments of Himachal Pradesh, India. Rev. Palaeobot. Palynol., 46: 209-225. Subramanian, C.V., 1971. "Hyphomycetes". Indian Counc. Agr. Res. New Delhi. 1 899. Varma, C.P. and Rawat, M.S., 1963. A note on some diporate grains recovered from Tertiary horizons of India and their potential marker value. Grana Palynol. 4(1): 130 139. Venkatachala, B.S. and Rawat, M.S., 1973. Palynology of the Tertiary sediments in the Cauvery Basin 2. Oligocene Miocene palynoflora from the sub-surface. Palaeobotanist, 20(2): 238 263.