GEOLOGICA CARPATHICA, APRIL 2005, 56, 2, 149154
Fossil flora (Costatheca and Spermatites) from the Upper
Maastrichtian Deccan Intertrappean Beds of India
RANAJIT KUMAR KAR
, ASHOK SAHNI
, KRISHNA AMBWANI
and DEBI DUTTA
Birbal Sahni Institute of Palaeobotany 53, University Road, 226 007 Lucknow, India
Centre of Advanced Study in Geology, Panjab University, 160 014 Chandigarh, India
(Manuscript received June 17, 2003; accepted in revised form June 16, 2004)
Abstract: Costatheca and Spermatites are recorded from the Maastrichtian Deccan Intertrappean Beds of Mohgaon-
Kalan and Padwar, Madhya Pradesh, India. These forms are found in association with typical Late Maastrichtian palyno-
logical species of aquatic pteridophytic sporomorphs namely, Azolla cretacea, Ariadnaesporites sp., Gabonisporites
vigourouxii and angiospermic pollen Aquilapollenites bengalensis. Specimens of Spermatites are compared with the
megasporangium of Azolla pinnata. It is observed that the indusium of A. pinnata resembles Spermatites in shape, size
and ornamentation. The views of other scientists regarding the affinity of Costatheca and Spermatites have also been
Key words: Deccan Intertrappean, Late Cretaceous, Costatheca, Spermatites.
Miner (1932, 1935) investigated the Upper Cretaceous coals of
Western Greenland and amongst other forms reported
Chrysotheca and Spermatites presumably seed cuticles of
angiosperms. Later, Binda & Nambudiri (1983) and Friis
(1985) changed the name of Chrysotheca to Costatheca as the
previous name was already occupied for Chrysophycean algae.
These two genera were later found to be of stratigraphic impor-
tance as they are confined to Cretaceous strata. Schemel (1950),
Binda (1968), Gunther & Hills (1972), Collin (1973), Binda &
Nambudiri (1983), Knobloch (1986) and Kovach & Dilcher
(1988) studied these two genera and commented on their mor-
phology and affinity. Batten & Zavattieri (1996) extensively
studied Miners material of these two genera, critically assessed
the various species and provided detailed descriptions.
The two genera were described and recorded only from the
Northern Hemisphere until Sahni et al. (1999), reported them
as casts from the Maastrichtian Lameta Formation near Ja-
balpur, India. Kar & Singh (1986: pl. 12, Fig. 5) illustrated a
Spermatites-like specimen from the Cretaceous Mahadek For-
mation of Meghalaya. However, it was designated as ?anther
type-1. The present paper includes descriptions of Costatheca
and Spermatites represented in the Maastrichtian Deccan In-
tertrappean Beds of Madhya Pradesh (India) and discusses
about their probable affinities.
Material and methods
Costatheca and Spermatites were collected from the Deccan
Intertrappean Beds of Mohgaon-Kalan and Padwar (Chhind-
wara District and Jabalpur District respectively, Madhya
Pradesh, India) (Fig. 1). At Mohgaon-Kalan, the intertrappean
bed was observed by Kar & Srinivasan (1998) in an unlined
water well situated 0.5 km west of the village (79°11\E,
22°1\N). The intertrappean bed is about 1 m thick and is sand-
wiched between two basaltic flows (Fig. 2). The lower sedi-
ment is made up of buff-coloured, hard chert of 0.25 m thick
with occasional wood fragments; it is overlain by green shale
of 0.25 m thick containing fresh water pelecypod. The upper
bed consists of carbonaceous shale of more or less the same
thickness. Kar & Srinivasan (1998) described typical Late
Cretaceous palynomorphs including aquatic pteridophytes
like Azolla cretacea, Ariadnaesporites sp., Gabonisporites vi-
gourouxii and angiosperm namely, Aquilapollenites bengalen-
sis from this shale. Costatheca and Spermatites were also ex-
amined from this unit.
The palynological investigation of Padwar Intertrappean
Bed was carried out by Prakash et al. (1990) and Sahni et al.
(1996). In a dug out abandoned well at Padwar, Kar et al.
(1998) detected two layers of trap, one layer of volcanic ash
and two intertrappean beds in the well. They divided the pa-
lynological assemblage recovered from this well into a lower
Aquilapollenites bengalensis Zone and an upper Algae domi-
nant Zone. The former zone is also very rich in Costatheca
and Spermatites and most of the specimens illustrated here
come from this horizon (Fig. 3). The samples were macerated
in commercial HNO
followed by a wash in 5% KOH solution
and passed through a 400 meshed sieve. The residues were
collected and the slides were prepared with polyvinyl alcohol
solution and when dried mounted in Canada balsam. The
slides and stub studied are deposited at the repository of the
Birbal Sahni Institute of Paleobotany, Lucknow.
150 KAR et al.
Genus: Costatheca (Miner) Hall, 1967.
Type Species: Costatheca diskoensis (Miner) Hall, 1967,
p. 1298, Figs. 13
Locality: Mohgaon-Kalan, Chhindwara District, Madhya
Horizon: Deccan Intertrappean, Maastrichtian, Late Cre-
Description: The specimens are oval with more or less
equally broad lateral ends. The size range is 594704 × 321
378 µm. The specimens are adorned with two distinct ribs,
they are 23 cells thick, parallel to longer axis. The cells on re-
mainder of the surface are transversely oriented, irregular in
size and shape, 610 µm thick. The margin of the cells is thin
and muri are up to 3 µm thick.
Comparison: The species described here resembles Cos-
tatheca diskoensis (Miner) Hall (1967) in size range and gen-
eral organization; however, C. diskoensis has 6 vertical ribs
whereas the present specimens have only two. C. dakotaensis
(Miner) Hall (1967) has pronounced midribs and transverse
cellular outlines while C. levis Vangerow (1954) has more or
less psilate cell wall.
Number of specimens studied: Four. Sample No.
H2, 5a; Sample No. Pad. 1.1, Sample No. Pad. 2.2.
Fig. 1. Geological map showing the localities of Mohgaon-Kalan and Padwar in Madhya Pradesh.
Fig. 2. Lithological profile of unlined water well at Mohgaon-
Kalan, Chhindwara District, Madhya Pradesh. Lower 0.25 m shale
bears Costatheca and Spermatites.
FOSSIL FLORA FROM THE UPPER MAASTRICHTIAN DECCAN INTERTRAPPEAN BEDS 151
Fig. 3. Lithological profile in an abandoned dug out well at Pad-
war, Jabalpur District,0 Madhya Pradesh. Samples Nos. P2P8
contain Costatheca and Spermatites.
Genus: Spermatites Miner, 1935.
Type Species: Spermatites ellipticus Miner, 1935.
Spermatites ellipticus Miner, 1935, p. 609, Pl. 19, Fig. 46
Locality: Padwar, Jabalpur District, Madhya Pradesh.
Horizon: Deccan Intertrappean, Maastrichtian, Upper
Description: The specimens are oval-elliptical in shape;
the size is very variable, ranging from 432825×250270 µm.
The apical part is mostly cutinized, generally forms a cap with
protruding end (Fig. 5.1,5), cutinized layer may be dissolved
due to maceration. The lower part of the specimens when de-
tached forms a circular opening, the marginal cells of this area
may also be cutinized. The cell walls have two layers the
outer wall is thin and mostly laevigate while the inner layer is
thick and reticulate in structure; cell walls are 112 µm thick.
The meshes are arranged parallel to longer axis, squarish-rect-
angular in outline and up to 2 µm thick.
Comparison: The specimens described here as Sperma-
tites ellipticus Miner (1935) closely resemble those figured by
Miner (1935, pl. 19, Fig. 46) in oblong-elliptical shape,
square-rectangular cells and presence of apical cap. S. arcua-
tus Miner (1935) is easily distinguished from S. ellipticus
Miner (1935) by its bean shape. S. elongatus Miner (1935) is
oblong-elliptical in shape and S. orbicularis Miner (1935) is
oval-subcircular in disposition. The characteristic feature of S.
polyporous Miner (1935) is its lateral position of the chalazal
Number of specimens studied: Twenty seven. Sample
Nos. Pad. 2.2, Pad. 1.3, Pad. 4.5, Pad. 6.2, Pad. 7.2, Pad. 8.6.
Since their introduction Costatheca and Spermatites attract-
ed the attention of a number of paleobotanists. They were
thought to be algal (Deák & Combaz 1967), bryophytic (Ar-
nold 1932), pteridophytic (Vangerow 1954), pteridospermic
(Harris 1954), angiospermic (Miner 1935; Binda 1968 etc.)
and even animal in origin (Tiffney 1983). It is obviously very
debatable and we would take Spermatites first because it is
Deák & Combaz (1967) thought that Spermatites could be
an alga resembling Polysiphonia because their morphological
characters broadly resemble each other. Arnold (1932) postu-
lated that Spermatites might be an antheridium of bryophyte.
Miner (1935) refuted this assumption stating that amongst the
present day bryophyte no comparable form is known.
Vangerow (1954) thought that Spermatites might be related
to some aquatic pteridophytes. Batten & Zavattieri (1996) re-
covered many heterosporous pteridophytes namely, Arcellites
disciformis, Ariadnaesporites ariadnae, Minerisporites bore-
alis and others, along with Costatheca and Spermatites and
other palynomorphs. Rao (1935) observed that the cells of the
upper part of the indusium in megasporangium of A. pinnata
become hardened and lignified so that when the lower part has
decayed these dark coloured cells remain as a conical cap at
152 KAR et al.
Fig. 5. 15 Spermatites ellipticus Miner, 1935. Sample No. Pad. 2,
Slide No. 3, R49; Sample No. Pad. 2, Slide No. 3, G23/4; Sample No.
Pad. 2, Slide No. 3, R49; Sample No. Pad. 2, Slide No. 3, Q31; Sam-
ple No. Pad. 2, Slide No. 2, Q53. Scale bar = 0.1 mm.
the top of the spore. Microscopic study of the plants reveals
that the indusium of A. pinnata which is the outer covering of
the megasporangium, very much resembles the genus Sperma-
tites in shape, size and ornamentation. The cutinized cap is
very similar to the micropylar end of seeds and consists of
closely spaced convoluted cell walls forming a cap, which
may be extended to a stalk of varying length (see also Tryon &
Tryon 1982; Tryon & Lugardon 1991). Hall & Peake (1968)
also mentioned that the cap of Spermatites somewhat resem-
bles the cap of present day Azolla. The indusium generally
dissolves when the megasporocarp is mature except for the
apical part. In the abortive megasporocarp the indusium does
not dissolve and when fossilized may form Spermatites-like
bodies (Fig. 6.1,2).
The sporophylls in Azolla filiculoides are always the lowest
leaves on the branch (McLean & Ivimey-Cook 1951). The
sporophyll has two lobes, the lower lobe divides early into two
at the apical part where the sori develop. The indusium grows
out as a ring-wall below the sorus and covers it completely ex-
cept for a micropyle-like opening at the top. Von Goebel (see
McLean & Ivimey-Cook 1951, p. 550) remarked that this type
of indusium resembles very much an integument surrounding
the ovule of a seed plant. No wonder that the aborted indusium
of Azolla in the fossil condition could apparently be regarded as
an angiospermic seed. The SEM photograph of the megaspo-
rangium of A. pinnata (Fig. 6.3) is broadly comparable to Sper-
matites sp. illustrated by Batten & Zavattieri (1995: Fig. 2C) re-
covered from the Campanian sediments of Alberta, Canada.
Harris (1954) observed that the isolated nucellar cuticles of
the pteridosperm Caytonia approximates Spermatites in mor-
phology. The nucellus has a thick and strong cuticle and in
both micropylar and chalazal regions the cells become small
Fig. 4. 1, 2 Costatheca sp. Sample No. H2, Slide No. 1, U61;
Sample No. Pad. 2, Slide No. 3. 3 Costatheca diskoensis (Mi-
ner) Hall, 1967. Sample No. Pad. 2, Slide No. 3, V45/1. All scale
bars = 0.1 mm.
FOSSIL FLORA FROM THE UPPER MAASTRICHTIAN DECCAN INTERTRAPPEAN BEDS 153
The reproductive structures of Archaefructus described by
Sun et al. (1998, 2002) from the Jurassic sediments of China
apparently resemble Caytonia in gross morphological charac-
ters. But is easily distinguished by its closed carpels along an
adaxial stigmatic crest; in Caytonia it is not conduplicate. Be-
sides it has bisaccate pollen whereas in Archaefructus the pol-
len are non saccate and monosulcate.
Binda (1968), Binda & Nambudiri (1983) and Nambudiri &
Binda (1991) contemplated that Spermatites resembles the
seed of Juncus. Kovach & Dilcher (1988) also endorsed this
view and commented that the thin outer cuticular layer of seed
of Juncus is comparable to some specimens of Spermatites.
They, however, pointed out the stratigraphic disparity between
the two as no form of Juncus is known from the Upper Creta-
ceous sediments. Huang & Dilcher (1988) remarked that the
two layers that are seen in most specimens of Spermatites may
be interpreted as representing the two integumentary layers
typical of angiosperms ovules in seeds. In Archaeanthus lin-
nenbergeri Dilcher & Crane (1984) Spermatites-like seeds or
aborted ovules isolated by maceration of a follicle fragment
have strongly cutinized pointed tips and rounded bases with a
circular perforation. These were interpreted by them as micro-
pylar and chalazal ends respectively.
Batten & Zavattieri (1995) however, observed that Sperma-
tites must be regarded as a heterogenous taxon because it com-
prises a variety of plant groups. Most of the Spermatites are
devoid of morphological characters that could be compared to
the extant species of angiosperms. Before them, Gunther &
Hills (1972) and Spleeman & Hills (1980) regarded Sperma-
tites as well as Costatheca as merely plant remains of uncer-
tain affinity. Batten & Zavattieri (1996) commented that the
disappearance of Spermatites at the end of the Cretaceous
seems to be more apparent than real as the associated me-
gaspores and angiosperms do not disappear. Considering all
these facts they concluded that at present the evidence is
equivocal to assert that Spermatites were derived from herba-
ceous angiosperms inhabiting hydrophyllous environments.
Costatheca sp. described here is, however, different from
the indusium of Azolla pinnata as cells other than the longitu-
dinal ribs are transversely oriented. Knobloch (1986) observed
that shape, number of ribs and external character of Costathe-
ca is very variable. Miner (1935) postulated that Costatheca
resembles Jungermanniales of bryophyta. Vangerow (1954)
thought that it could be a spore sac. Tiffney (1983) suggested
that Costatheca could be an insect egg. Batten & Zavattieri
(1995) opined that some Costatheca could be insect eggs be-
cause the cuticles of some insect eggs are very similar to Cos-
tatheca. Knobloch (1981) thought that this genus is not an-
giospermous in origin. Binda & Nambudiri (1983), however,
assumed that Costatheca could be related to the seeds of Bu-
tomaceae of angiosperms.
Costatheca and Spermatites are known from wide geo-
graphical regions with a limited stratigraphic range. Most of
the workers think Spermatites is an angiospermic seed while a
few assume it of gymnospermic or pteridophytic in origin.
and isodiametric. The chalaza has a small circular hole and the
micropylar end is occasionally raised. However, the me-
gaspore membrane in most of the modern gymnosperms is
generally heavily cutinized and according to Batten & Zavat-
tieri (1995) Spermatites has no such thickening.
Fig. 6. 1, 2 Indusium of Azolla pinnata R. Brown with megaspo-
rangium. Scale bar = 0.1 mm. 3 SEM of indusium of Azolla pin-
nata R. Brown showing cap and reticulation. Scale bar = 0.5 mm.
4 Cap of the same in SEM. Scale bar = 0.5 mm.
154 KAR et al.
Since it grew in aquatic environments, the external morpholo-
gy of the diverse organisms could have developed more or less
similar forms in response to the same kind of adaptation.
Hence Spermatites could very well be a heterogenous com-
plex comprising different entities.
Acknowledgments: Sincere appreciation is expressed to the
Department of Science & Technology, New Delhi for granting
a project entitled Deccan Intertrappean palynoflora and its
implication for the demarcation of K/T boundary (ESS/CA/
A4-16/96). The authors appreciate Dr. A.K. Sinha, Director,
Birbal Sahni Institute of Paleobotany, Lucknow for providing
infrastructural facilities. Thanks are due to Prof. C.L. Verma,
Botany Department, Lucknow University, Lucknow for sup-
plying Azolla pinnata. We also thank Dr. Ratan Kar for help
in the processing of plates and figures.
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