Fungal taxa from the Miocene sediments of Mizoram, northeast India

Abstract

The fungal elements comprising 33 dispersed genera and 35 species were recovered from an exposure near the Tlangsam village on the Indo-Myanmar border from the Dulte Formation (Miocene), near Champhai, eastern Mizoram, India. The fungal spores are most common and represented by 20 genera and 20 species comprising the amerospores, didymospores, phragmospores, dictyospores, scolecospores, helicospores and staurospores. The microthyriaceous ascostromata contribute 4 genera and 4 species and those of the hemisphaerials 2 genera and 2 species to the assemblage; the remaining 7 genera and 9 species belong to the other fungal entities. The assemblage has 14 new genera and 19 new species. Endomycorrhizal, aquatic fungi and other hyphomycetes are frequently found. Most of the fungal elements indicate a warm, humid, tropical subtropical climate during the time of deposition, though a few have temperate affiliation.

Full text

Fungal taxa from the Miocene sediments of Mizoram, northeast India
Ratan Kar
a
, B.D. Mandaokar
a,
⁎, R.K. Kar
b
a
Birbal Sahni Institute of Palaeobotany, 53, University Road, Lucknow-226 007, India
b
B-1/42, Sector J, Aliganj Scheme, Lucknow-226024, India
abstractarticle info
Article history:
Received 23 July 2009
Received in revised form 11 September 2009
Accepted 12 September 2009
Available online 3 October 2009
Keywords:
fungi
Dulte Formation
Miocene
Mizoram
India
The fungal elements comprising 33 dispersed genera and 35 species were recovered from an exposure near
the Tlangsam village on the Indo-Myanmar border from the Dulte Formation (Miocene), near Champhai,
eastern Mizoram, India. The fungal spores are most common and represented by 20 genera and 20 species
comprising the amerospores, didymospores, phragmospores, dictyospores, scolecospores, helicospores and
staurospores. The microthyriaceous ascostromata contribute 4 genera and 4 species and those of the
hemisphaerials 2 genera and 2 species to the assemblage; the remaining 7 genera and 9 species belong to the
other fungal entities. The assemblage has 14 new genera and 19 new species. Endomycorrhizal, aquatic fungi
and other hyphomycetes are frequently found. Most of the fungal elements indicate a warm, humid, tropical
subtropical climate during the time of deposition, though a few have temperate affiliation.
© 2009 Elsevier B.V. All rights reserved.
1. Introduction
The fungi evolved along with the algae in very ancient times.
Taylor and Taylor (1993) recorded fungal entities from the Bitter
Springs Formation (Precambrian) of Australia. Tiffney and Barghoorn
(1974) reported fungal remains from the Ordovician marine bryozo-
ans. However, the first record of terrestrial fungi (probably Ascomy-
cetes) comes from the Late Silurian rocks of Sweden (Sherwood-Pike
and Gray, 1985). Kidston and Lang (1921) described various fungal
remains from the classic Rhynie chert (Devonian) of England. The
Carboniferous Period witnessed a prolific diversity in the fungi along
with the pteridosperms and many remains of Ascomycetes and other
groups were described by Stubblefield et al. (1983, 1985).Taylor and
White (1989) noted the occurrence of fossil fungi (Endogonaceae)
from the Triassic rocks of Antarctica.
In India, the earliest records of fungi are from the Early Cretaceous
when Tripathi (2001) described the fungal remains from the Rajmahal
Intertrappean beds. Kar et al. (2003) dealt with a fungal megafossil
viz. Lithopolyporales zeerabadensis from the Lameta beds (Maastrich-
tian) of Madhya Pradesh. Sharma et al. (2005) noted the fossil remains
of mycorrhizal and epiphyllous fungi from the coprolites of the
dinosaurs (Isisaurus) recovered from the same Formation. Kar et al.
(2004b) observed fossil plant pathogen viz. Protocolletotrichum from
the Deccan Intertrappean beds (Maastrichtian) of India.
The fungal remains obtained from the varied sedimentary deposits
provide valuable information regarding the palaeoenvironmental
reconstruction along with the spores and the pollen grains. The role of
fossil fungi in delimiting the stratigraphy was discussed by Graham
(1962) and Elsik (1974). However, their relevance in palynostratigra-
phy is rather limited dueto their long vertical range (Pirozynski, 1978).
For instance, the fossil remains of microthyriaceous ascostromata have
been recovered from the Maastrichtian to Recent (Elsik, 1968; Sharma
et al., 2005). Despite this drawback, their occurrence in the sedimentary
deposits provide reliable information to unravel the depositional
environment, nature of substrata and the host plants upon which the
fungi flourished (Lange, 1978). In rare instances, they also help to
decipher the food habits of the animals (Sharma et al. 2005).
The fossil fungal studies thus provide reasonable insights to
decipher the palaeoclimatic condition of deposition during the Late
Cretaceous to Tertiary as the fungal flora have hardly changed in this
period. Their usefulness becomes more imperative when the
pteridophytic spores and pollen of angiosperms and gymnosperms
are absent or scanty in the sediments.
The fungal hyphae, spores and microthyriaceous ascostromata are
found in good numbers in the Miocenesediments of Mizoram, whereas,
the spores and the pollen grains of the higher plants are comparatively
rare and monotonous.For this reason, the presentstudy was undertaken
so that by the presence of fungal elements the palaeoclimatic condition
of deposition could be deduced and the antiquity of some of the lesser
known water fungi could be traced (Kar et al., 2006).
2. Stratigraphy
Srivastava et al. (1979) took a traverse on the Seling–Champhai–Tyao
River sections in the eastern part of Mizoram and divided the 7165 m
thick sediments into two informal subdivisions —thelowerDulteandthe
upper Keifang Formations and placed them under the Bhuban Subgroup
Review of Palaeobotany and Palynology 158 (2010) 240–249
⁎Corresponding author. Tel.: +91 522 2740011; fax: +91 522 2740098.
E-mail address: bdmandokar@yahoo.com (B.D. Mandaokar).
0034-6667/$ –see front matter © 2009 Elsevier B.V. All rights reserved.
doi:10.1016/j.revpalbo.2009.09.004
Contents lists available at ScienceDirect
Review of Palaeobotany and Palynology
journal homepage: www.elsevier.com/locate/revpalbo

(Miocene). The Dulte Formation is exposed on Seling–Champhai road
section and comprises predominantly massive–thinly bedded claystone,
subordinate sandstone and sandstone–claystone alternations. The
claystone according to Srivastava et al. (1979) is greenish grey,
moderately hard, micaceous, non-calcareous and occasionally silty. This
formation is more arenaceous at the base and argillaceous at the top. The
approximate thickness of this Formation is 5275 m.
The Keifang Formation, on the other hand, is less thick (1890 m)
and is mostly arenaceous with minor intercalation of claystone and
sandstone–claystone. The sandstone is generally massive, sometimes
bedded and the thin clay beds generally alternate with sandstone.
3. Materials and methods
The fungal elements enumerated here were recovered from a 12 m
thick section exposed near the Tlangsam village (Lat. 23˚28′and Long.
93˚25′). The village is situated on the India–Myanmar border and is
about 50 km east of Champhai (Fig. 1). It belongs to the Dulte
Formation of the Bhuban Subgroup (Srivastava et al.,1979). The
exposure rests on a 0.2 m gravel bed and the remaining part shows an
alternation of claystones and sandstones, sometimes minor shale beds
are also interspersed. The systematic collection of samples was done
from the clay and the shale beds but mostly the shales yielded the
palynofossils. The lithology and the position of the yielding samples
are shown in Fig. 1. The maceration procedure was done by as usual.
The concerned slides are deposited at the repository of the Birbal
Sahni Institute of Palaeobotany, Lucknow.
4. Previous palynological work
Mandaokar (2000) worked out the palynology and palaeoenvir-
onment of deposition of some samples from Ramrikawn, near Aizawl.
Fig. 1. Location map of the study area and litholog showing position of productive samples.
241R. Kar et al. / Review of Palaeobotany and Palynology 158 (2010) 240–249

He inferred a near shore environment of deposition for the presence
of Nypa pollen in the sediments and ascribed a Lower Miocene age for
the assemblage. Mandaokar (2002) suggested a deltaic condition of
deposition for the Dulte Formation on the basis of palynomorphs viz.,
Striatriletes, Polyporina and Spinizonocolpites. Kar et al. (2005)
reported a few endomycorrhizal fungi comprising auxiliary cells,
chlamydospores and sporocarps from the samples studied here.
Besides, there are many fungal elements which they have not
mentioned. These taxa are systematically studied here. The fungal
elements are associated with pteridophytic spores, and pollen of
angiosperms and gymnosperms. The characteristic taxa are Striatri-
letes susannae, Pinuspollenites crestus, Abiespollenites cognatus, Hibis-
ceaepollenites splendus and Palaeomalvaceaepollis mammilatus. These
taxa support a Miocene age for the assemblage.
5. Fungal taxa
The fungal spores, ascostroma and other entities are common in
the samples and they contribute more than 25% to the total
assemblage of the palynomorphs. The genera and species recorded
in the samples are as under:
Archaeoglomus Sharma et al., 2005
Archaeoglomus globatus Sharma et al., 2005
Archaeoglomus minutus sp. nov.
Ceratohirudispora gen. nov.
Ceratohirudispora miocenica sp. nov.
Ceratohirudispora triradiata sp. nov.
Cervichlamydospora gen. nov.
Cervichlamydospora nigra sp. nov.
Chlamydospora gen. nov.
Chlamydospora dichotoma sp. nov.
Circinoconites gen. nov.
Circinoconites arthrus sp. nov.
Cucurbitariaceites Kar et al., 1972
Cucurbitariaceites bellus Kar et al., 1972
Dicellaesporites (Elsik 1968)Sheffy & Dilcher 1971
Dicellaesporites elongatus Kumar 1990
Dictyomykus gen. nov.
Dictyomykus ellipticus sp. nov.
Dictyostromata gen. nov.
Dictyostromata perfecta sp. nov.
Diporicellaesporites Elsik 1968
Diporicellaesporites stacyi Elsik 1968
Diporisporites (van der Hammen 1954)Elsik 1968
Diporisporites psilatus Kumar 1990
Dyadosporites (van der Hammen 1954)Clarke 1965
Dyadosporites ellipsus Clarke 1965
Euthythyrites Cookson, 1947
Euthythyrites bifidus sp. nov.
Fusiformisporites Rouse 1962
Fusiformisporites crabbii Rouse 1962
Inapertisporites (van der Hammen 1954)Sheffy & Dilcher 1971
Inapertisporites vulgaris Sheffy & Dilcher 1971
Kutchiathyrites Kar 1979
Kutchiathyrites eccentricus Kar 1979
Lithomucorites gen. nov.
Lithomucorites miocenicus sp. nov.
Lithosporocarpia gen. nov.
Lithosporocarpia cephala sp. nov.
Meliostroma gen. nov.
Meliostroma tlangsamensis sp. nov.
Monoporisporites (van der Hammen 1954)Sheffy & Dilcher 1971
Monoporisporites ovalis Sheffy & Dilcher 1971
Multicellaesporites Elsik 1968
Multicellaesporites nortonii Elsik 1968
Mycozygosporangia gen. nov.
Mycozygosporangia laevigata sp. nov.
Notothyrites Cookson, 1947
Notothyrites setiferus Cookson=Trichothyrites setiferus (Cook-
son) Saxena & Misra 1990
Palaeogigaspora gen. nov.
Palaeogigaspora excellensa sp. nov.
Parmathyrites Jain & Gupta 1970
Parmathyrites indicus Jain & Gupta 1970
Phragmothyrites (Edwards 1922)Kar & Saxena, 1976
Plate II. (Bar =10 μm). (see on page 244)
1. Teliosporites globatus sp. nov.
2. Dictyomykus ellipticus sp. nov.
3. Teliosporites hirsutus sp. nov.
4. Tetradigita stellata sp. nov.
5. Cervichlamydospora nigra sp. nov.
6. Dictyostroma perfecta sp. nov.
7. Trichopeltinites folius sp. nov.
8. Euthythyrites bifidus sp. nov.
9. Stauromyca radiata sp. nov.
10. Circinoconites arthrus sp. nov.
Plate I. (Bar =10 μm).
1. Lithomucorites miocenicus sp. nov.
2. Archaeoglomus globatus Sharma, et al., 2005
3. Archaeoglomus minutus sp. nov.
4. Palaeogigaspora excellensa sp. nov.
5. Mycozygosporangia laevigata sp. nov.
6. Lithosporocarpia cephala sp. nov.
7. Ceratohirudispora miocenica sp. nov.
8. Ceratohirudispora triradiata sp. nov.
9. Chlamydospora dichotoma sp. nov.
10, 11. Meliostroma tlangsamensis sp. nov.
242 R. Kar et al. / Review of Palaeobotany and Palynology 158 (2010) 240–249

Plate I.
243R. Kar et al. / Review of Palaeobotany and Palynology 158 (2010) 240–249

Plate II (caption on page 242).
244 R. Kar et al. / Review of Palaeobotany and Palynology 158 (2010) 240–249

Phragmothyrites eocaenicus (Edwards 1922)Kar & Saxena 1976
Pluricellaesporites (van der Hammen 1954)Elsik 1968
Pluricellaesporites longicollus Sheffy & Dilcher 1971
Protocolletotrichum Kar et al., 2004b
Protocolletotrichum deccanensis Kar et al., 2004b
Stauromyca gen. nov.
Stauromyca radiata sp. nov.
Teliosporites gen. nov.
Teliosporites globatus sp. nov.
Teliosporites hirsutus sp. nov.
Tetradigita gen. nov.
Tetradigita stellata sp. nov.
Tetraploa Berkeley & Broome, 1877
Tetraploa sp.
Trichopeltinites Cookson, 1974
Trichopeltinites folius sp. nov.
However, only the new genera and species are described here.
5.1. Description of taxa
Genus:Lithomucorites gen. nov.
Type species:Lithomucorites miocenicus sp. nov.
Derivation of name:Litho (L.)=rock, mucorites from the order
Mucorales.
Generic Diagnosis: Fungal sporangia, apophysate, flask shaped,
subcircular–circular in shape, size range 25–52×22–49 μm, some-
times with sporangiophore. Wall about 1 μm thick, closely ornamen-
ted with bacula–pila and verrucae, translucent–light brown, some
could be made up of calcium oxalate, 2–5μm in length, closely placed
on both sides to provide negative reticulum on surface view; no slit on
sporangia observed.
Lithomucorites miocenicus sp. nov.
Pl. 1, Fig. 1
Holotype:Plate I,Fig. 1. Size 32 ×28 μm (without sporangiophore).
BSIP no. 13342 (P28).
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Diagnosis: Sporangia subcircular with serrated margin due to heavy
ornamentation, mostly occur in dispersed condition, 32–28 ×26–
40 μm, flask shaped, sporangia wall about 1 μm thick, ornamented
with bacula–pila, and verrucae, pila–bacula 3–5μm height, closely
placed to form pseudoreticulate structure.
Remarks: The species described here seems to belong to the class
Zygomycetes for the presence of coenocytic hyphae, asexual repro-
duction by means of sporangiophores and absence of flagellate cells
(Alexopoulos et al., 1996).
Genus:Archaeoglomus Sharma et al., 2005
Type species:Archaeoglomus globatus Sharma et al., 2005
Archaeoglomus globatus Sharma, et al., 2005
Pl. 1, Fig. 2. BSIP no.13349 (Q42).
Description: Auxiliary cells generally oval, sometimes subcircular,
28–46×26–44 μm, occur at terminal end of mycelium, wall about
1μm thick, laevigate.
Archaeoglomus minutus sp. nov.
Pl. 1. Fig. 3
Holotype:Plate I, Fig. 3. Size 20 ×18 μm. BSIP no.13340 (H/40).
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Diagnosis: Auxiliary cells develop at terminal ends of hyphae, hyphae
laterally branched bear one cell at each end, 10–12×8–20 μm, wall
about 1 μm thick, psilate.
Remarks: The species referrable to the order Glomales are also called
as vesicular–arbuscular mycorrhizae (VAM) and are found in 70% of
all the plant families (Sieverding, 1991).
Genus:Palaeogigaspora gen. nov.
Type species:Palaeogigaspora excellensa sp. nov.
Derivation of name:Palaeo (L.)=old, Gigaspora —a genus of the
family Gigasporaceae.
Generic Diagnosis: Chlamydospores solitary, grow at tip of hyphae,
oval, 30–51×28–47 μm, dark brown, wall about 2 μm thick, laevigate.
Hyphae tubular, nonseptate, hyaline.
Remarks: The specimens assigned to Palaeogigaspora could either
be vesicles or azygospores. Both these are formed at the terminal
end of the hyphae and are difficult to distinguish in the dispersed
state.The vesicle illustrated by Alexopoulos et al. (1996, Fig. 5-22 B)
is oval in shape and closely resembles some specimens (Pl.1,
Fig. 4 ).
Palaeogigaspora excellensa sp. nov.
Pl. 1, Fig. 4
Holotype:Plate I, Fig. 4. Size 45 ×36 μm. BSIP no.13349 (N19/2)
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Diagnosis: Chlamydospores generally oval with equally broad lateral
ends, sometimes subcircular, 31–45×28–42 μm; spore wall up to
2μm thick, laevigate; mycelia nonseptate, hyaline, laterally branched.
Genus:Chlamydospora gen. nov.
Type species:Chlamydospora dichotoma sp. nov.
Derivation of name: After the chlamydospores.
Generic Diagnosis: Chlamydospores oval, 12–25×10–22 μm, dark
brown, hyphae dichotomously branched to produce two spores, wall
up to 2 μm thick, laevigate, hyphae septate.
Chlamydospora dichotoma sp. nov.
Pl. 1, Fig. 9
Holotype:Plate I, Fig. 9. Size 24 ×16 μm. BSIP no.13338 (O27)
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Diagnosis: Chlamydospores oval, 16–28×12–24 μm, found in pairs
due to dichotomous division of hyphae, spores in terminal position,
equal in size and shape, spore wall robustly built, about 2 μm thick,
thicker at basal end, psilate, sometimes intrastructured.
Genus:Lithosporocarpia gen. nov.
Type species:Lithosporocarpia cephala sp.nov.
Derivation of name:Litho (L.) =rock, sporocarp = fruiting body,
cephalus=head.
Generic Diagnosis: Sporocarps subcircular–circular, 22–48× 20–
45 μm, often with chlamydospores, chlamydospores stalked, subcir-
cular; sporocarp wall up to 2 μm thick, hyphae forming reticulation on
both sides.
Remarks:Alexopoulos et al. (1996) observed that the sporocarp is
generally a hypogeous fruiting structure which contains either the
zygospores, sporangia or chlamydospores. It is mostly covered with
hyphae which enmeshed to form a reticulate structure.
Lithosporocarpia cephala sp. nov.
Pl. 1, Fig .6
Holotype:Plate I, Fig. 6. Size 32 ×30 μm. BSIP no. 13350 (R15/1)
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Diagnosis: Sporocarps generally subcircular with uneven margin due
to projection of hyphae, 28–30 ×26–32 μm, hyphae forms regular
reticulation on both surfaces, meshes mostly square in shape,
sometimes rectangular. Chlamydospore present, one chlamydospore
found on each sporocarp, chlamydospore subcircular, 10–22×8–
20 μm, dark brown, laevigate, with a small stalk and globular head.
Genus:Mycozygosporangia gen. nov.
Type species:Mycozygosporangia laevigata sp. nov.
Derivation of name:Mykes (Gr.)=fungus, zygos (Gr.) =yoke.
Generic Diagnosis: Zygosporangia light–dark brown, 22–48×20–
46 μm, subcircular–circular in shape, often attached with two
opposite hyphae, sporangia wall 2–4μm thick, laevigate.
Remarks: It is very difficult to distinguish a zygosporangium resulting
from the sexual conjugation and an azygospore formed by asexual
means. However, the attachment of two hyphae opposite to each
other with the sporangium leads to the speculation that the sexual
245R. Kar et al. / Review of Palaeobotany and Palynology 158 (2010) 240–249

conjugation took place resulting in the formation of the sporangium.
In the asexual method only one hyphal remnant is observed.
Mycozygosporangia laevigata sp. nov.
Pl. 1, Fig. 5
Holotype:Plate I, Fig, 5. Size 24 ×48 μm. BSIP no. 13351 (U23/1).
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Diagnosis: Zygosporangium subcircular in shape, 23–48 ×21–46 μm,
two hyphae often attached opposite to each other, look like ordinary
mycelia, tubular. Spore wall 2–4μm thick, psilate, often weakly
intrastructured.
Genus:Ceratohirudispora gen. nov.
Type species:Ceratohirudispora miocenica sp. nov.
Derivation of name: After the genera Ceratosporium and Hirudinaria
of the hyphomycetes.
Generic Diagnosis: Hyphomycetaceous fungi, conidiophore small,
growth terminated by production of apical conidium; conidium
enlarges laterally in opposite direction to produce two–three arms,
conidia 5–10 celled, septa up to 2 μm thick, with broad base and
narrow tip.
Remarks:Ceratosporium Schweinitz (1832) and Hirudinaria Cesati
(1856) produce same type of conidia. The two conidia that are
generally developed by the conidiophore are placed opposite to each
other laterally. In the dispersed state it is not possible to distinguish
one from the other (Subramanian, 1971).
Ceratohirudispora miocenica sp. nov.
Pl. 1, Fig. 7
Holotype:Plate I, Fig. 7. Size 35 ×8 μm (one arm). BSIP no. 13348 (S37/
2).
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Diagnosis: Conidia of imperfect fungi, two armed, V-shaped, laterally,
opposite to each other forming an acute angle, arms more or less equal
in size, 24–38×5–9μm; conidia septate, septa 2 μm thick, septa 6–8;
conidia broader in middle, narrow at base and apex, wall about 1 μm
thick, laevigate.
Ceratohirudispora triradiata sp. nov.
Pl. 1, Fig. 8
Holotype:Plate I, Fig. 8. Size 38 ×8 μm. BSIP no. 13338 (L37).
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Diagnosis: Hyphomycetaceous conidia, three armed providing a
triradiate appearance, arms unequal in length and size, septate,
septa more or less 1 μm thick, 4–8 celled, conidia wall 1 μm thick,
laevigate.
Genus:Meliostroma gen. nov.
Type species:Meliostroma tlangsamensis sp. nov.
Derivation of name: After the order Meliolales, stroma (Gr.) =
mattress.
Generic Diagnosis: Fungal stromata with mycelia, 25–38×22–36 μm,
subcircular–circular in shape, dark brown, hyphae forming reticula-
tion on both sides, stromata bear capitate and mucronate hyphopodia.
Remarks: The various members of this order grow on the leaves and
stems often producing a thick, dark, velvety cover causing black
mildew disease.
Meliostroma tlangsamensis sp. nov.
Pl.1 , Figs. 10,11
Holotype:Plate I, Fig.11. Size 24 ×36 μm. BSIP no.13340 (L27).
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram ,
India.
Derivation of name: After the village Tlangsam, Mizoram.
Diagnosis: Fungal stromata subcircular–circular, 26–36×24–32 μm,
generally with hyphae of two types–two celled, globular, capitate
hyphopodia and one celled, bottle shaped mucronate hyphopodia.
Stromata reticulate on both sides forming square–rectangular
meshes.
Remarks:Luttrell (1989, p.194) illustrated a stroma with the capitate
and mucronate hyphopodia which resembles the present specimens.
The specimens described by Dilcher (1965) from U.S.A. and
Ramanujam and Rao (1978) from Kerala have both longer capitate
hyphopodia and thus are easily separated from the present
specimens.
Genus:Teliosporites gen. nov.
Type species:Teliosporites globatus sp. nov.
Derivation of name:Telos (Gr.)=end, spora (Gr.) =seed.
Generic Diagnosis: Fungal teliospores, always in mass, 21–32 ×19–
28 μm , generally surrounded by gelatinous translucent sheath, up to
2μm thick, spores laevigate, sterile cells may be associated with fertile
ones, pseudobaculate appearance on surface view.
Teliosporites globatus sp. nov.
Pl. 2, Fig. 1
Holotype:Plate II,Fig. 1. Size 28×19 μm. BSIP no. 13352 (× 32/4).
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Diagnosis: Spore mass subcircular–oval, 21–28×19–27 μm, margin
undulated due to the presence of gelatinous covering; 12–18 spores
joined together, individual spore subcircular, 8–11×6–9μm, spore
wall up to 2 μm thick, laevigate, light brown; gelatinous sheath often
disappears during maceration, about 2 μm thick, translucent, weakly
granulose, grana less than 1 μm high, sparsely placed.
Remarks: The spore balls of smut fungi illustrated by Alexopoulos
et al. (1996, figure 21-7 C, E) resemble the present species in shape
and absence of sterile cells. The SEM photo of the spore ball of The-
caphora mexicana by Duràn (1987) and reproduced by Alexopoulos
et al. (1996) also resembles in subcircular shape but is reticulate.
Teliosporites hirsutus sp. nov.
Pl. 2, Fig. 3
Holotype:Plate II, Fig. 3. Size 27 ×25 μm. BSIP no. 13352 (× 32/2).
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Diagnosis: Teliospores always occur in mass, 12–24 spores joined
together to form subcircular shape, 18–31×16–29 μm. Cells of two
kinds —fertile and sterile, fertile cells bigger 12–18×10–16 μm,
sterile cells 4–8×3–7μm; spore wall about 1 μm thick, laevigate;
fertile cells 12–28 in number, sterile cells numerous, covering fertile
cells, closely placed to form retibaculate pattern.
Genus:Circinoconites gen. nov.
Type species:Circinoconites arthrus sp. nov.
Derivation of name: After the genus Ciracinoconis, arthron (Gr.)=
joint.
Generic Diagnosis: Fungal conidia, conidia acrogenous, strongly
spiralled, spirals 30–39×25–31 μm; solitary, coiled, not in chains or
slime, 8–14 septate, fist-shaped, dark brown, constricted at septa, cells
increasing in diameter from base to apex, dissimilar, spirally arranged.
Remarks: The coiled conidia generally occur amongst the aquatic fungi
and Ingold (1942, 1975) and Shearer (1992, 1993) worked intensively
on them. Besides, there are also aeroaquatic or helicosporous fungi
which also produce helicoid conidia (van Beverwijk, 1953).
Circinoconites arthrus sp. nov.
Pl. 2, Fig. 10
Holotype:Plate II, Fig. 10. Size 44 ×27 μm. BSIP no. 13164 (U21/4).
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Diagnosis: Conidia arise from tip of conidiophores, helicoid, looking
like fist, 42–30×23–27 μm made up of 10–16 cells, cells increase in
size from base to top, basal cell, rectangular, 6–8×4–6μm3–4, lower
cells straight, rest cells coiled, septate, septa up to 2 μm thick,
constriction more marked in middle region, individual cells rectan-
gular–wedge shaped, terminal cell oval–subcircular, 11–18× 12–
16 μm; spore wall about 1 μm thick, mostly laevigate, sometimes
weakly intrastructured.
Genus:Tetradigita gen. nov.
Type species:Tetradigita stellata sp. nov.
Derivation of name:Tetra (L.) four, digita (L.) =finger, stellata (L.) =
star.
Generic diagnosis: Fungal conidia, blastic in development, conidium
long, tubular, bends in middle to produce generally two lateral arms,
simultaneously giving a stellate appearance, arms equal–unequal in
246 R. Kar et al. / Review of Palaeobotany and Palynology 158 (2010) 240–249

length (5–30×2–4μm), sometimes with further branching, spore
wall 1 μm thick, laevigate, translucent–light brown, mostly not
septate.
Remarks: Ever since Ingold (1942) described aquatic hyphomycetes
on decaying alder leaves from England, many workers throughout the
world started the investigation in earnest on the aquatic fungi and
accumulated a lot of information (Subramanian, 1971).
Tetradigita stellata sp. nov.
Pl. 2 , Fig. 4
Holotype:Plate II, Fig. 4. Size 62×52 μm. BSIP no. 13164 (F33).
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Diagnosis: Conidiogenous cells like ordinary hyphae, conidia develop
before septation of conidiogenous cells, filamentous, straight in young
stage, slightly curved at maturity to bear two arms laterally at the
same time; arms 14–28 ×2–4μm, tubular, generally without septa, 8–
10 times longer than broad, arms gradually tapering at terminal end,
generally 4 armed, sometimes 5, arm may be branched laterally or
dichotomously at base; wall roughly 1 μm thick, laevigate.
Genus:Dictyomykus gen. nov.
Type species:Dictyomykus ellipticus sp.nov.
Derivation of name:Dictyon (Gr.)=net, mykes (Gr.) =fungus.
Generic Diagnosis: Conidiophores small, closely placed to form a
sporodochium like cluster; conidia elliptical, basal part narrowed than
terminal, 30–48×7–12 μm, branches 4–8, parallel, develop from a
basal cell, laterally fused, multiseptate, septa margin not constricted,
incurved at tip, spore wall laevigate.
Dictyomykus ellipticus sp.nov.
Pl. 2, Fig. 2
Holotype:Plate II, Fig. 2. Size 40×13 μm. BSIP no. 13343 (O24/2).
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Diagnosis: Conidiophores inconspicuous, 4–6 celled, closely adhered
to each other; conidia borne singly, elliptical in shape, basal part
generally narrower than terminal end, may also be equally broad at
two ends, 30–48×9–14 μm, branches 5–8, equal in length, fused
laterally except at tip, transversely septate, septa 10–18, distinct,
parallel to each other, no appreciable constriction at margin, terminal
cell markedly curved, spore wall about 1 μm thick, laevigate.
Genus:Trichopeltinites Cookson, 1947
Type species:Trichopeltinites pulcher Cookson, 1947
Trichopeltinites folius sp. nov.
Pl. 2, Fig. 7
Holotype:Plate II, Fig. 7. Size 110 ×25 μm BSIP no.13165 (L20/1).
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Derivation of name:Folium (L.)=leaf
Diagnosis: Ascostromata leaf-like with lobes, 90–110×20–25 μm,
margin undulated, hyphae radially arranged one layered thick, more
or less parallel to each other, raised, often branched, transverse septa
few, anastomose to form pseudoparenchymatous cells particularly on
fruiting bodies, fruiting bodies subcircular, 8–10 μm, alternately
placed, internal structure not visible due to hyphae.
Genus:Euthythyrites Cookson, 1947
Type species:Euthythyrites oleinitis Cookson, 1947
Euthythyrites bifidus sp. nov.
Pl. 2, Fig. 8
Holotype:Plate II, Fig. 8. Size 60–32 μm. BSIP no. 13338 (P47).
Type Locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Derivation of name:Bifidus (L.)=split into two parts.
Diagnosis: Ascostromate, 51–60×30–45 μm, dimidiate, nonostiolate,
dark brown, generally one celled thick, darker in central region,
hyphae radially arranged, anastomose to form square–rectangular
pseudoparenchymatous cells, branches semicircular at ends, gener-
ally divided into two, sometimes an incipient third also visible,
mycelia absent.
Remarks: The specimens are rare. Cookson (1947) commented that
the extant species of this genus grows on the oleaceous leaves.
Genus:Dictyostromata gen. nov.
Type species:Dictyostromata perfecta sp. nov.
Derivation of name:Dictyon (L.)=net, stroma (Gr.) =mattress, per-
fectus (L.)=perfect.
Generic Diagnosis: Stromata fan-shaped except attachment zone;
attachment slightly pointed, haustorium hyaline, nonseptate, tubular,
32–49×22–38 μm, lateral arms more or less equal, hyphae radially
and transversely anastomose to form pseudoreticulation on both
sides.
Dictyostromata perfecta sp. nov.
Pl. 2, Fig. 6
Holotype:Plate II, Fig. 6. Size 37 ×23 μm. BSIP no. 13341 (L52/3).
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Diagnosis: Stromata with two lateral sides divergent from each other,
outer margin convex, slightly undulated due to pseudoreticulation,
35–42 × 23–37 μm; haustorium present or absent, 4–7×2–3μm,
hyaline, no septa observed, stromata generally conical at attachment
zone; radial hyphae stronger than transverse hyphae, anastomose to
develop pseudoreticulation, meshes square–rectangular; faint at basal
region.
Remarks: Some of the extant hyphomycetes viz., Mycoenterolobium
Goos (1970),Tretopileus Dodge (1942) and Trichodocheum von
Sydow (1927) approximate the species described here by their free
stromata. Of them, Mycoenterolobium platysporum Goos (1970) as
illustrated by Kendrick and Carmichael (1973; Fig. 67A) closely
resembles in semicircular shape and presence of a solitary haustori-
um. However, the present species is distinguished by its divergent
lateral sides and fan-like shape.
Genus:Cervichlamydospora gen. nov.
Type species:Cervichlamydospora nigra sp. nov.
Derivation of name:Cervix (L.)=neck, nigrum (L.) =blacken
Generic Diagnosis: Chlamydospores sub-circular, dark brown-black,
originate from neck of hyphae; solitary, 14–24 ×12–22 μm, many
hyphae adhere together at base, branch out laterally at tip; hyphae
wall laevigate–granulose, grana up to 1 μm thick, sparsely placed.
Cervichlamydospora nigra sp nov.
Pl. 2, Fig. 5
Holotype:Plate II, Fig. 5. Size 17 ×30 μm. BSIP no. 13351 (D43/1).
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Diagnosis: Chlamydospore sub-circular, solitary, dark brown–black
with a constriction at margin, margin entire like a shield, smooth, 13–
23×12–21 μm; arise from neck of hyphal strand, strand composed of
8–13 hyphae, closely adhered at base and region of chlamydospore
attachment, separate and loose in rest part, central hypha strongly
built, nonseptate or rarely septate; hypha wall about 1 μm thick,
generally laevigate, sometimes weakly granulose.
Genus:Stauromyca gen. nov.
Type species:Stauromyca radiata sp. nov.
Derivation of name:Staure (L.)=star, mykes (L.) =fungus.
Generic Diagnosis: Hyphomycetaceous fungi, conidiophores un-
branched, tubular, septate; conidia solitary, globular, generally four
armed, basal cell septate, swollen at base, arms of variable length,
mostly nonseptate, tubular, light brown.
Remarks: The swollen conidia and the four arms of the present genus
seems to be an adaptation to the aquatic environment like Jaculispora
submersa Hudson & Ingold (1960).
Stauromyca radiata sp. nov.
Pl. 2 , Fig. 9
Holotype:Plate II, Fig. 9. Size 15 ×13 μm, without the conidiophore
and the arms. BSIP no. 13163 (R19/1).
Type locality: Tlangsam, Bhuban Formation, Miocene, Mizoram, India.
Diagnosis: Conidiophores long, tubular, straight to bent, 12–15× 2–
3μm, wall less than 1 μm thick, laevigate, 3 septa distinct, straight,
conidia triangular, folded, attachment cell of conidiophore slightly
swollen, 12–18×10–16 μm, four arms divergent, straight or bent,
unequal in length, 8–16×3–4μm, bulged at base, nonseptate except
one at base, wall about 1 μm thick, laevigate.
247R. Kar et al. / Review of Palaeobotany and Palynology 158 (2010) 240–249

6. Discussion
The fungal assemblage comprises 33 genera and 35 species. Of
these, 20 genera and 20 species belong to spores, 4 genera and 4
species to the microthryriaceous ascostromata, 2 genera and 2 species
to ascostromata of the hemisphaerials and the rest 7 genera and 9
species to other fungal entities. Amongst the spores, all types viz.,
amerospores, didymospores, phragmospores, dictyospores, scolecos-
pores, helicospores and staurospores are observed (Saccardo, 1886).
The helicospores represented by the genus Circinoconites and the
staurospores designated by the genera Tetradigita and Stauromyca are
of special interest as they are probably aquatic fungi. The aquatic fungi
mostly belong to the Hyphomycetes (Deuteromycetes) and family
Pleosporaceae (Ascomycetes).
The aquatic fungi belong to many groups of Hyphomycetes e.g.
Nectria of Pyrenomycetes, Hymenoscyphus of Discomycetes and Mas-
sarina of Loculoascomycetes (Shearer, 1992, 1993). Bandoni (1981)
recorded some species with tetraradiate conidia which are equally
adept to terrestrial habitat. Webster (1987) reported some of them as
phylloplane dwellers and can very well exist in sea water. Ingold
(1942) observed that most of the water fungi live luxuriantly in the
running water, whereas the aeroaquatic or the helicosporous aquatic
Hyphomycetes occur in stagnant fresh water or sometimes also in
marine condition. Kar et al. (2006) reported the occurrence of fossil
aquatic fungi from the Miocene sediments of Mizoram.
The endomycorrhizal fungi are quite common in the samples and
are represented by Archaeoglomus, Palaeogigaspora, Chlamydospora
and probably also Lithosporocarpia. These vesicular–arbuscular my-
corrhizae (VAM) not only supply extra amount of water and minerals
to the host plants from the soil but also protect them from the other
fungi, bacteria, cyanobacteria, algae and Actinomycetes (Linderman,
1992). Powell and Bagyaraj (1984), Sieverding (1991) and Bethlen-
falvay and Linderman (1992) observed that VAM also offer resistance
to some root pathogens and nematodes by building antibiotic
substances.
Of all the VAM, Archaeoglomus is most common and occurs in
between the two cells and in the cells without causing any change in
the external morphology of the roots. Taylor and White (1989) and
Simon et al. (1993) estimated an age of 353–462 millionyears for the
VAM, in general, by means of DNA sequence information and
415millionyears for Glomus in particular. It seems that since its
inception, Glomus was associated with the roots of plants as evident
from the Rhynie chert of Devonian age (Pirozynski and Malloch,
1975). Sieverding (1991) calculated that the VAM are present in the
roots of about 70% of the extant terrestrial plants and they even do not
spare the algae.
The epiphyllous fungi are also frequently found, being represented
by the genera of microthyriaceous ascostromata viz., Phragmothyrites,
Trichothyrites and Kutchiathyrites. Eriksson and Hawksworth (1993)
observed that the shield like ascostromata occur in a number of
unrelated families of Loculoascomycetes. In Trichothyrium, Actinopeltis
and Microthyrium, the hyphae are arranged in radiating rows. In Tri-
chopeltinites Cookson (1947) and Euthythyrites Cookson (1947), the
epiphyllous fungi form leaf like ascostromata by means of longitudinal
extension of the radiating hyphae. The various species are mostly
found in the tropical–subtropical climate, though a few can even grow
in the temperate region. Due to their hard structure, they are often
found as fossils and even can withstand the digestive mechanism of
the saurischian dinosaurs as they were recovered from their
coprolites (Kar et al., 2004a). They are also good indicator of
palaeoclimate and their occurrence indicates the presence of broad
leaved trees and tropical–subtropical climate.
Most of the endomycorrhizal fungi, the Meliolales and the
epiphyllous fungi live in the tropical–subtropical climate in the
present day. As the assemblage is dominated by these fungi, it is very
likely that the same type of climate also prevailed during the time of
deposition. However, some of the aquatic fungi viz. Tetradigita and the
conidiophorus genus Dictyomykus indicate towards a cooler climate.
A fungus comparable to Tetradigita was recorded by Ingold (1942)
from alder leaves during the autumn in U.K. Subramanian (1953)
noted the conidiophore resembling the genus Dictyomykus from the
height of 1200 m in the Nilgiri Hills, Tamil Nadu, India. Hait and
Banerjee (1994) also noted some temperate elements in the
Champhai assemblage.
Besides the fungal elements, bisaccate pollen assignable to Pinus
and Abies are often found in the assemblage along with the typical
tropical elements e.g. Striatriletes, Polyporina and Hibisceaepollenites.
It should be stated here that the depositional site was not far from the
Shillong Plateau which was at lofty heights (>2000 m) during the
Miocene time. It is quite possible that the cooler elements in the
assemblage are derived from this plateau.
Acknowledgements
The authors are grateful to Dr. N.C. Mehrotra, Director, Birbal Sahni
Institute of Palaeobotany, Lucknow for encouragement and for
providing the necessary research facilities. We are greatly indebted
to Prof. A.F. Lotter and two anonymous reviewers for their invaluable
comments. We also thank Dr. R.K. Saxena for helpful suggestions
which significantly improved this paper.
References
Alexopoulos, C.J., Mims, C.W., Blackwell, M., 1996. Introductory mycology, Fourth
edition. John Wiley & Sons, Inc., New York.
Bandoni, R.J., 1981. Aquatic hypomycetes in terrestrial litter. In: Wicklow, D.T., Caroll,
G.C. (Eds.), The Fungal Communuty: Its Role in the Ecosystem. Dekker, New York,
pp. 693–703.
Berkeley, M.J., Broome, C.F., 1877. The fungi of Ceylon-1. J. Linn. Soc. London Bot. 11,
494–572.
Bethlenfalvay, G.J., Linderman, R.G., 1992. Mycorrhizae in sustainable agriculture. Am.
Soc. Agrn. Crop. Sci. Soc. Am. Soil Sci. Soc. Am, Madison, WI.
Cesati, C.A., 1856. Klotzsch Herb.Myc. Edit. Nova. 269 pp.
Clarke, R.T., 1965. Fungal spores from Vermejo Formation coal beds (Upper Cretaceous )
of Central Colorado. Mt. Geol. 2, 85–93.
Cookson, I.C., 1947. Plant microfossils from the lignites of Kerguelen archipelago. Rep.
B.A.N.Z. Antarct. Exped. Ser. A 129–142.
Dilcher, D.L., 1965. Epiphyllous fungi from Eocene deposit in western Tennessee, U.S.A.
Palaeontographica 116B, 1–54.
Dodge, B.O., 1942. Heterocaryotic vigor in Neurospora. Bull. Tor. Bot. Club 69, 75–91.
Duràn, R., 1987. Ustilaginales of Mexico. Washington State University Press, Pullman.
Edwards, W.N., 1922. An Eocene microthyriaceous fungus from Mull. Scotland. Trans.
Brit. Mycol. Soc. 8, 66.
Elsik, W.C., 1968. Palynology of Palaeocene Rockdale lignite, Milan County, Texas —1:
morphology and taxonomy. Pollen Spore 10, 263–314.
Elsik, W.C., 1974. Fungal spores and Caenozoic palynostratigraphy. Xerox Handout at
7th Ann. Mtg. A.A.S.P., pp. 1–19.
Eriksson, O.E., Hawksworth, D.L., 1993. Index to notes 969–1529. Sys. Ascomycet. 11,
1–201.
Goos, R.D., 1970. Mycoenterolobium platisporum. Mycologia 62, 171–175.
Graham, A., 1962. The role of fungal spores in palynology. J. Palaeontol. 36, 60–68.
Hait, A.K., Banerjee, M., 1994. Palynology of lignite sediments from Mizoram, eastern
India with remarks on age and environment of deposition. J. Palynol. 30, 113–135.
Hudson, H.J., Ingold, C.T., 1960. Aquatic hyphomycetes from Jamaica. Trans. Brit. Mycol.
Soc. 43, 469–478.
Ingold, C.T., 1942. Aquatic hyphomycetes of decaying older leaves. Trans. Brit. Mycol.
Soc. 25, 339–417.
Ingold, C.T., 1975. Convergent evolution in aquatic fungi: the tetraradiate spore. Biol. J.
Linn. Soc. 7, 1–25.
Jain, K.P., Gupta, R.C., 1970. Some fungal remains from Tertiary of Kerala coast.
Palaeobotanist 18, 177–182.
Kar, R.K., 1979. Palynological fossils from the Oligocene and their biostratigraphy in the
district of Kutch, western India. Palaeobotanist 26, 16–49.
Kar, R.K., Saxena, R.K., 1976. Algal and Fungal microfossils from Matanomadh
Formation (Palaeocene), Kutch, India. Palaeobotanist 26, 1–15.
Kar, R.K., Singh, R.Y., Sah, S.C.D., 1972. On some algal and fungal remains from Tura
Formation of Garo Hills, Assam. Palaeobotanist 19, 146–154.
Kar, R.K., Sharma, N., Agarwal, A., Kar, Ratan, 2003. Occurrence of fossil wood rotters
(polyporales) from the Lameta Formation (Maastrichtian), India. Curr. Sci. 85,
37–40.
Kar, R.K., Sharma, N., Kar, Ratan, 2004a. Occurrence of fossil fungi in dinosaur dung and
its implication on food habit. Curr. Sci. 87, 1053–1056.
Kar, R.K., Sharma, N., Verma, U.K., 2004b. Plant pathogen Protocolletotrichum from a
Deccan intertrappean bed (Maastrichtian), India. Cret. Res. 25, 945–950.
248 R. Kar et al. / Review of Palaeobotany and Palynology 158 (2010) 240–249

Kar, R.K., Mandaokar, B.D., Kar, Ratan, 2005. Mycorrhizal fossil fungi from the Miocene
sediments of Mizoram, northeast India. Curr. Sci. 89, 257–259.
Kar, R.K., Mandaokar, B.D., Kar, Ratan, 2006. Fossil aquatic fungi from the Miocene
sediments of Mizoram, northeast India. Curr. Sci. 90, 291–292.
Kendrick,W.B.,Carmichael,J.W.,1973.Hyphomycetes.In:Ainsworth,G.C.,Sparrow,
F.K., Sussman, A.S. (Eds.), The fungi an advanced treatise IV A —Ataxonomic
review with keys: ascomycetes and fungi imperfecti. Academic Press, New York,
pp. 323–509.
Kidston, R., Lang, W.H., 1921. On the old redstone plants showing structure, from the
Rhynie Chert Bed, Aberdeenshire. Trans. R. Soc. 52, 855–902.
Kumar, P., 1990. Fungal remains from Miocene Quilon beds of Kerala Sta te, South India.
Rev. Palaeobot. Palynol. 62, 13–28.
Lange, R.T., 1978. Southern Australian Tertiary epiphyllous fungi, modern equivalents
in the Australasian region and their palaeohabitat indicator value. Can. J. Bot. 46,
532–541.
Linderman, R.G., 1992. Vesicular–arbuscular mycorrhizae and soil microbial interac-
tions. In: Bethelenfalvay, G.L., Linderman, R.G. (Eds.), Mycorrhizae in substainable
agriculture American Society of Agrinomy. Madison, WI.
Luttrell, E.S., 1989. Morphology of Meliola floridensis. Mycologia 81, 192–204.
Mandaokar, B.D., 2000. Palynology and palaeoenvironment of the Bhuban Formation
(Early Miocene) of Ramrikawn, near Aizawl, Mizoram. Palaeobotanist 49, 317–324.
Mandaokar, B.D., 2002. An interpretation of the palynology and palaeoecology of the
Early Miocene Dulte Formation, Mizoram, India. Palaeobotanist 51, 113–121.
Pirozynski, K.A., 1978. Fungal spores through the ages —a mycologist's view. Proc. IV.
Int. Palynol. Conf. Lucknow (1976–77), vol. 1, pp. 327–330.
Pirozynski, K.A., Malloch, D., 1975. The origin of land plants: a matter of mycotrophism.
Bio systems 6, 153–164.
Powell, C.L., Bagyaraj, D.J., 1984. VA Mycorrhiza. CRC Press, Boca Raton, Florida.
Ramanujam, C.G.K., Rao, K.P., 1978. Fungal spores from the Neogene strata of Kerala in
South India. Fourth Intn. Palynol. Conf. Lucknow, vol. 1, pp. 291–304.
Rouse, G.E., 1962. Plant microfossils from the Burrad Formation of western British
Columbia. Micropalaeontology 8, 187–218.
Saccardo, P.A., 1886. Sylloge fungorum omnium hucusque cognitorum; Hyphomycetes,
Pavia. Italy 4, 1–807.
Saxena, R.K., Misra, N.K., 1990. Palynological investigation of the Ratnagiri Beds of
Sindhu Durg District, Maharashtra. Palaeobotanist 38, 263–276.
Schweinitz, G.S., 1832. Creatosporium fuscescens Trans. Am. Phil. Soc. 11, 300.
Sharma, N., Kar, R.K., Agarwal, A., Kar, Ratan, 2005. Fungi in dinosaurian (Isisaurus)
coprolites from the Lameta Formation (Maastrichtian) and its reflection on food
habit and environment. Micropaleontology 51, 73–82.
Shearer, C.A., 1992. The role of woody debris. In: Baerlocher, F. (Ed.), Ecological studies
94. The ecology of aquatic hyphomycetes. Springer-Verlag, New York, pp. 77–98.
Shearer, C.A., 1993. A new species of Kirschsteiniothelia (Pleosporales) with an unusual
fissitunicate ascus. Mycologia 85, 963–969.
Sheffy, M.V., Dilcher, D.L., 1971. Mor phology and taxonomy of fungal spores.
Palaeontographica 133B, 34–51.
Sherwood-Pike, M.A., Gray, J., 1985. Silurian fungal remains: probable records of the
class Ascomycetes. Lethaia 18, 1–20.
Sieverding, E., 1991. Vesicular–arbuscular mycorrhiza management in tropical
agrosystems. Technical Corp. Germany, Eschborn, Germany.
Simon, L., Bousquet, J., Lévesque, R.C., Lalonde, M., 1993. Origin and diversification of
endomycorrhizal fungi and coincidence with vascular plants. Science 363, 67–69.
Srivastava, B.P., Ramachandran, K.K., Chaturvedi, J.G., 1979. Stratigraphy of eastern
Mizo Hills. Bull. Oil Nat. Gas Comm. 16, 87–94.
Stubblefield, S.P., Taylor, T.N., Miller, C.E., Cole, G.T., 1983. Studies in Carboniferous fungi
II. The structure and organization of Mycocarpon, Sporocarpon, Dubiocarpon and
Coleocarpon (Ascomycotina). Am. J. Bot. 70, 1482–1498.
Stubblefield, S.P., Taylor, T.N., Miller, C.E., 1985. Studies in Palaeozoic fungi, IV: wall
structure of fossil endogonaceous chlamydospores. Mycologia 77, 88–96.
Subramanian, C.V., 1953. Fungi Imperfecti from Madras-V. Proc. Ind. Acad. Sci. B 38,
36–39.
Subramanian, C.V., 1971. Hyphomycetes an account of Indian species, except
Cercosporae. Indian Council Agricultural Research, New Delhi. 930 pp.
Taylor, T.N., Taylor, E.L., 1993. The biology and evolution of fossil plants. Englewood
Cliffs, New Jersey, pp. 1–982.
Taylor, T.N., White, J.F., 1989. Triassic fungi with suggested affinities to endogonales
(Zygomycotina). Rev. Palaeobot. Palynol. 61, 53–61.
Tiffney, B.H., Barghoorn, E.H., 1974. The fossil record of the fungi. In: Rollins, R.C., Roby,
K., (Eds.) Occasional papers of the Farlow Herbarium of. Cryptogam. Bot. 7, 1–42.
Tripathi, A., 2001. Fungal remains from Early Cretaceous Intertrappean beds of
Rajmahal Formation in Rajmahal Basin, India. Cret. Res. 22, 565–574.
van Beverwijk, A.L., 1953. Helicosporous hyphomycetes. 1. Trans. Br. Mycol. Soc. 36,
111–124.
van der Hammen, T., 1954. El desarrollo de la flora Columbiana en los periodos
geologicos. 1. Maestrichtiano Hasta Terciario mas Inferior. Boln. Geol. Bogota II (1),
49–106.
von Sydow, H., 1927. Die Verwertung der Verwantschaftsverh ältnisse und des
gegenwartigen Entwicklungsganges zur Umgrenzung der Gattungen bei den
Uredineen. Ann. Mycol. Berlin 19, 161–175.
Webster, J., 1987. Convergent evolution and the functional significanceofsporeshapein
aquatic and semi-aquatic fungi. In: Rayner, A.D.M., Brasier, C.M., Moore, D. (Eds.),
Evolutionary biology of the fungi. Cambridge University Press, Cambridge, pp. 191–201.
249R. Kar et al. / Review of Palaeobotany and Palynology 158 (2010) 240–249