Mycosphere 12(1): 670–874 (2021) www.mycosphere.

org ISSN 2077 7019

Article
Doi 10.5943/mycosphere/12/1/8

Diversity in fossil fungal spores

Saxena RK1, Wijayawardene NN2,3, Dai DQ3, Hyde KD4 and Kirk PM5
1
Birbal Sahni Institute of Palaeosciences, 53 University Road, Lucknow–226007, India
2
State Key Laboratory of Functions and Applications of Medicinal Plants, Guizhou Medical University, Guiyang
550014, P.R. China
3
Center for Yunnan Plateau Biological Resources Protection and Utilization, College of Biological Resource and Food
Engineering, Qujing Normal University, Qujing, Yunnan 655011, P.R. China
4
Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand
5
Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS, UK

Saxena RK, Wijayawardene NN, Dai DQ, Hyde KD, Kirk PM 2021 – Diversity in fossil fungal
spores. Mycosphere 12(1), 670–874, Doi 10.5943/mycosphere/12/1/8

Abstract
Diverse types of fungal spores, exhibiting a variety of morphological variations, have been
added to the fossil records from time to time. These may be unicellate (unicellular), dicellate,
tricellate, tetracellate, multicellate, muriform, filiform, spirally coiled and star-like. Similarly, these
may be inaperturate, monoaperturate, diaperturate, triaperturate and multiaperturate. The present
paper documents all published fossil fungal spore genera and species. Assignment of fossil fungal
spores to extant fungal taxa is seldom possible. These are therefore placed into artificial supra-
generic taxa based on morphological characters, e.g. number and nature of cells and characters
associated with size, symmetry, apertures, septa and spore wall. These supra-generic taxa are:
Amerosporae (unicellate spores; 61 genera), Didymosporae (dicellate spores; 21 genera),
Phragmosporae (tri- or pluricellate, transversely septate spores; 41 genera), Dictyosporae
(muriform spores divided by intersecting longitudinal and transverse or diagonal septa; 18 genera),
Helicosporae (one to many celled, spirally coiled spores with curved axis; 9 genera) and
Staurosporae (pluricellate, stellate spores, with more than one axis; 9 genera). In addition, a number
of species of fossil fungal spores, described under 26 modern fungal genera, have also been
included. Genera are arranged in alphabetical order under each of the above groups. Similarly,
species are also arranged under each genus in alphabetical order. Five new species, viz.
Dicellaesporites vermae, Dyadosporites singhii, Fusiformisporites sahii, Diporicellaesporites
tiruchirappalliensis and Multicellites tamilensis, and four new combinations, viz. Hypoxylonites
disciformis (Sheffy & Dilcher), Hypoxylonites lanceolatus (Debi Mukh.), Melanospora primigenia
(Casp.) and Thecaphora mohgaoensis (Chitaley & Yawale) are proposed here. The dominant
genera, both in number and variety, are: Brachysporisporites R.T. Lange & P.H. Sm.,
Dicellaesporites Elsik, Diporicellaesporites Elsik, Diporisporites Hammen, Dyadosporites R.T.
Clarke, Fusiformisporites Rouse, Hypoxylonites Elsik, Inapertisporites Hammen,
Multicellaesporites Elsik, Multicellites Kalgutkar & Janson., Pluricellaesporites Hammen and
Staphlosporonites Sheffy & Dilcher. Fossil fungal spores are known from all parts of the globe and
in the sediments ranging from the Precambrian era to most recent age. Their distribution in time
and space is presented. A comparison of the fossil fungal spores with extant taxa is also made and
affinity of a good number of fossil genera and species could be traced with the extant genera or

Submitted 15 February 2021, Accepted 12 June 2021, Published 2 August 2021

Corresponding Author: Ramesh K. Saxena – e-mail – [email protected],
Nalin N. Wijayawardene – e-mail – [email protected],
Dong-Qin Dai – e-mail – [email protected] 670
higher taxa. Palaeoenvironmental and stratigraphic implications of fungal spores are also discussed.

Keywords – Amerosporae – Classification – Dictyosporae – Didymosporae – Fungi Imperfecti –

Helicosporae – Phragmosporae – Staurosporae

Introduction
The last few decades witnessed a substantial increase in the number of publications dealing
with the study of microfossils, e.g. spores and pollen of vascular plants, dinoflagellate cysts and
fungi, having multiple applications in the field of earth and life sciences, viz. determination of
palaeoenvironment, evolution of morphographic characters and biostratigraphy.
Fossil fungi form the ancestors of extant fungi which evolved relatively early, probably more
than 1500 million years ago. Fossil fungal remnants are found in sedimentary rocks in the form of
spores, mycelia, sporophores and symbiotic associations and are commonly observed in macerated
residues prepared for palynological studies. Fungal remains have been recorded from sediments of
all ages, but are abundant in the Tertiary Period. The fossil records clearly indicate that
Ascomycota, the largest and most diversified group of modern fungi, became established during the
Cretaceous Period and became abundant in the Tertiary Period (Elsik 1968, Jain 1974, Kar &
Saxena 1976, Jansonius 1976, Jain & Kar 1979, Ramanujam 1982, Kalgutkar & Jansonius 2000,
Tripathi 2009, Saxena & Tripathi 2011). The majority of fossil fungal spores, found in
palynological preparations, belong to Ascomycota whereas only a few belong to Basidiomycota. In
members of Ascomycota, ascospores are produced inside the ascus by the sexual morph
(teleomorph) whereas the asexual morph (anamorph) may produce several types of asexual
vegetative spores, named conidiospores (or conidia) which may be unicellate to multicellate and
variously shaped. Saprobic Ascomycota may produce ascomata of macroscopic size and definite
shape.
Fragments of fossil fungi, commonly observed in the palynological preparations, have been
sporadically recorded since Williamson (1878, 1880), Conwentz (1892), Meschinelli (1892, 1902),
Felix (1894), Kidston & Lang (1921), Edwards (1922) and Van der Hammen (1954, 1955, 1956).
Tiffney & Barghoorn (1974) observed that the palaeomycological record is too scanty to allow the
introduction of a firm phylogeny. The paucity of the fungal records before the Cenozoic has often
been cited as a principal reason for lack of interest in their stratigraphic potential. Ramanujam &
Rao (1978) postulated that fungal spores are not suitable for stratigraphical interpretations on a lack
of interest in, and insufficient appreciation of, the spore morphology of modern fungi. Stubblefield
& Taylor (1988) opined that the avoidance of fossil fungi is most likely to be explained by the
difficulties of properly recognizing, manipulating and interpreting them.
However, since the 1950s studies on fossil fungi received more attention with the
amplification of palynological studies (Rouse 1959, 1962, Varma & Rawat 1963, Clarke 1965,
Elsik 1968, 1969, 1990a, Elsik & Jansonius 1974, Kar & Saxena 1976, 1981, Ramanujam & Rao
1978, Ramanujam & Srisailam 1980, Chandra et al. 1984, Glass et al. 1986, Jarzen & Elsik 1986,
Song et al. 1989, Kumar 1990, Saxena 1991, 2006, Saxena & Khare 1992, Kalgutkar 1993, 1997,
Kalgutkar & Sigler 1995 Parsons & Norris 1999, Kagutkar & Jansonius 2000, Gupta 2002, Nandi
et al. 2003, Kar et al. 2010, Saxena & Tripathi 2011, Mukherjee 2012, Guimarães et al. 2013,
Martinez et al. 2016, O’Keefe et al. 2017, Premaor et al. 2018). Although fossil fungi are
encountered in sediments of all ages, their frequency remarkably increases in the Tertiary Period,
which suggests that their proliferation is linked with diversification of angiosperms. Serious efforts
have been made on the study of dispersed fossil fungi from Cretaceous-Tertiary sediments,
including spores, microscopic sporangia, sporophores, hyphae or fragmented mycelia and
mycorrhiza.

Organization of the paper

The main objective of the paper is to document all published genera and species of fossil
fungal spores described to date. For the sake of convenience, the paper is arranged in three parts.

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 The first part contains a general introduction and history of the studies on fossil fungi,
classification of fossil fungal spores, terms related to fossil fungi, geological time and life on earth,
field and laboratory investigation procedures, sources of information and method of data
presentation.
The second part provides an account of the fossil fungal spores. Details of this section are
provided under Materials and Methods. Furthermore, in this part, five new species are proposed.
These forms were earlier described under different genera as informal un-named species.
Specimens of these forms were restudied and careful observations led to the proposal of new
species. The following information is given for each new species: Names of the new species, Index
Fungorum Registration Identifier, Holotype and its Repository details, Diagnosis, Location, Age
and Notes, including comparison and derivation of names. In addition, four new combinations are
proposed. The following information is given for each new combination: Name of the new
combination followed by the author of the basionym, Index Fungorum Registration Identifier, Full
and direct reference to the basionym with full bibliographical details, Holotype and its Repository
details, Diagnosis, Location, Age and Notes, including comparison and derivation of names.
The third part is a discussion on the diagnostic characters of the commonly occurring fossil
fungal spore genera and their relationship with extant fungal taxa, palaeoenvironmental and
biostratigraphic implications of fossil fungi and suggestions for future studies.
Efforts were made to include, in this paper, all genera and species of fossil fungal spores
published worldwide, as far as known to us, including later synonyms and homonyms with their
current names. Even taxa, that were not validly published, have also been included. However, it is
possible that some papers might have escaped our attention. Although, we feel that there are quite a
good number of species that may be later taxonomic synonyms, we did not make any attempt to
point them out because we did not see most of the original material. Similarly, we also refrained
from emending the generic and specific diagnoses for the same reason.

Classification of fossil fungal spores

Since the very beginning of studies on fossil fungi, diverse types of fungal spores have been
added to the fossil records from time to time. The morphological features of these spores are
sometimes distinct to allow identification and comparison with their extant counterparts and help to
assign them to a natural classification system. This, however, is seldom possible. Except for some
distinctive Tertiary forms, fossil fungal spores cannot generally be ascribed to modern taxa and
their classification with extant fungi does not become possible. To overcome this difficulty, these
are described under Artificial System of Classification which is based only on morphological
characters, e.g. characters associated with size, symmetry, apertures, septa and spore wall.
In a classification system proposed by Van der Hammen (1956), fossil fungal spores were
grouped under various morphologic categories having the suffix ‘sporites’. Clarke (1965) proposed
the suffix ‘sporonites’ for naming the fossil fungal spores. Both of these are still being followed.
Elsik (1976a) attempted to prepare a comprehensive applicable taxonomy for the fossil fungal
spores on the basis of morphological features. He considered septal and apertural characters as the
most stable and primary characters, whereas size, shape, symmetry and wall sculpture as the
subsidiary characters. He proposed artificial supra-generic categories of family rank for
classification of fossil fungal spores, viz. Sporae Monocellae, Sporae Dicellae, Sporae Tricellae,
Sporae Tetracellae, Sporae Multicellae and Sporae Cellae Indeterminatae. These categories were
primarily based on the cell number and presence or absence and number of apertures. Under these
categories, artificial genera and species could be conveniently described.
Pirozynski & Weresub (1979) suggested a system named as ‘Saccardoan System’ for
classifying the fungal spore types which are not referable to extant families. Kendrick & Nag Raj
(1979) modified the Saccardoan System to eliminate some of its inconsistencies, and listed
characters of specific importance to separate different groups. This scheme is based on the shape
and number of cells and fungal spores are recognized as Amerospores, Didymospores,

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Phragmospores, Dictyospores, Helicospores, Staurospores (star-like) or Scolecospores.
Characteristic features of each of these groups are as follows:

1. Amerosporae: Spores unicellate, inaperturarate or aperturate; aperturate spores with one pore or
hilum, two or more pores, or variable apertures.
2. Didymosporae: Spores dicellate, inaperturate or aperturate; aperturate spores with one pore or
hilum at the proximal end, or two pores, one each at or near the proximal and distal end.
3. Phragmosporae: Spores tri- or pluricellate, only transversely septate, inaperturate or aperturate;
aperturate spores with a pore or hilum at the proximal end; with or without attenuated distal end, or
with two pores, one each at the proximal and distal end.
4. Dictyosporae: Spores muriform divided by few or many intersecting longitudinal and transverse
or diagonal septa; shapes variable, inaperturate or with a more or less distinct hilum, that may be a
scar or protruding.
5. Helicosporae: Spores uni- or pluricellate; with curved axis (spirally coiled), coiled in one plane
or twisted in three planes.
6. Staurosporae: Spores pluricellate; with more than one axis, or stellate (star-shaped).
7. Scolecosporae: Elongate pluricellate spores with ladder-like septation; length/width ratio of
spore body exceeding 15:1; spores narrow, filamentous, transversely septate, with a pore or hilum
at the proximal end or two pores, one each at the proximal and distal terminations.

This system was included in Wijayawardene et al. (2020a) who provided a current classification of
fungi. Nevertheless, we encourage the readers, palaeomycologists to provide missing information
or new opinions (on classification, generic concepts) according Wijayawardene et al. (2020a) since
the corrections could be done in newly launched website, outlineoffungi.org (Wijayawardene et al.
2020b)

Terms related to fossil fungi

Common terms related to fossil fungi and used in their descriptions are defined below (based
on The A.A.S.P., INC. Workgroup on fossil fungal palynomorphs 1963, Kalgutkar & Jansonius
2000, Saxena & Tripathi 2011, Taylor et al. 2015).
Acrogenous: Conidia growing at the apex of a conidiophore.
Acropetal: Conidia produced in succession forming a chain toward the apex; young conidia
occurring at the tip.
Acervulus (pl. acervuli): A mat of hyphae giving rise to short conidiophores grouped together
forming a specialized mycelial mass. The immersed conidioma consists of a flat layer of
pseudoparenchyma upon which conidia are initiated and produced while still covered by the host
tissues.
Amb (short for Latin ambitus, circuit, orbit): Term used in palynology to refer to the optical section
of a spore or pollen grain, or to the outline of such a grain when seen in polar view.
Amerospore: A one-celled spore.
Amphigenous: A mycelium growing on both sides of the host leaf (as in microthyriaceous fungi).
Apothecium (pl. apothecia): An open ascocarp in which a layer of asci (hyminium) lies exposed on
the surface or hollow part of the disc or variously shaped structure.
Arbuscule: Shrub-like growth; as in a tuft of conidiophores, or the haustoria-like intracellular
development of mycorrhizal fungi.
Ascocarp/Ascophore (= ascoma, pl. ascomata): Any open or closed organ containing asci of
Ascomycota.
Ascospore: A sexual spore produced as a free cell by meiosis and mitotic processes in an ascus.
Ascostroma (pl. ascostromata): Simple type of ascomycetous body consisting of an undifferentiated
mass of tissue forming a stroma, on or in which the asci are developed.
Ascus (pl. asci): An enlarged sac-like cell containing a specific number of ascospores (often four,
typically eight).

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Aseptate (= nonseptate): Lacking cross-walls or septa.
Asexual morph (Anamorph): The asexual vegetative form of an ascomycetous or basidiomycetous
fungus.
Astomate: Without an ostiole.
Basidiophore/ Basidiocarp: A sporophore of Basidiomycota, bearing basidia and basidiospores.
Basidiospore: A sexual spore borne externally on a basidium.
Basidium (pl. basidia): A specialized, club-shaped cell bearing four basidiospores.
Basipetal: Conidia produced in succession, or in a chain, from the base (proximal); older conidia
occurring at the tip.
Blastospore: A spore formed by budding.
Catenate (or catenulate): Produced in chains.
Chlamydospore: 1. A thick-walled, secondary spore developed from hyphae, usually intercalary
that generally functions as a resting spore; 2. An endogenous, multinucleate thick-walled spore,
variable in volume, commonly found in parasitic fungi.
Cleistocarp: Synonymous with cleistothecium.
Cleistothecium (pl. cleistothecia): A closed ascocarp, from which ascospores are liberated by
rupture or decay of the structure.
Coenobium (pl. coenobia): A colony.
Conidiophore: Simple or branched specialized hypha, arising from a vegetative mycelial hypha and
bearing, at its tip or side, one or more conidiogenous cells.
Conidium (pl. conidia): A nonmotile exogenous asexual spore.
Dictyospore: Conidium divided into cells by longitudinal and transverse septa.
Didymospore: Monoseptate (dicellate) conidium.
Dimidiate: Appearing to lack one half of the wall of an ascoma or having one half very much
smaller than the other.
Dolipore: A septum of a dikaryotic basidiomycete hypha which flares out in the middle portion
forming a barrel-shaped structure with open ends.
Ectomycorrhiza: A mycorrhiza in which fungal hyphae grow intercellularly in the host tissue.
Endomycorrhiza: A mycorrhiza in which the fungal hyphae penetrate into the cells of the host; also
called vesicular arbuscular mycorrhiza (VAM).
Foliicolous: Growing on leaves.
Fungi Imperfecti: The Ascomycota are divided into those fungi representing the perfect stage
(teleomorphs) in which sexual spores are produced, or those (anamorphs, representing the
imperfect stage), in which only vegetative spores are produced. Holomorphs are those in which
both sexual and asexual spores have been observed. The group of anamorph fungi are also referred
to as Fungi Imperfecti.
Helicospore: A coiled or helical conidium.
Hilum (pl. hila): A mark or scar on spores appearing like a dot, flat spot or pore, indicating the
point of attachment of the spore to a conidiophore, conidiogenous cell, hypha or sterigma.
Hypha (pl. hyphae): Basic tubular, septate or aseptate, elements of the fungi that may form a
mycelium (thallus).
Hyphomycetous: Relating to the hyphomycetes; moldlike, cobwebby.
Imperfect stage (anamorph): The (conidial) asexual morph of an ascomycetous or basidiomycetous
fungus.
Intramatrical: Hyphae located within the matrix or substratum.
Macronematous: Refers to any conidiophore that is differentiated from the normal hyphal cells.
Miospore: Collective designation for dispersed small or large spores and pollen grains, as found in
palynological preparations, of which it may not be possible to differentiate their biological function
of micro- or macrospore (megaspore).
Muriform (of conidia): Being divided by intersecting septa in more than one plane.
Mycelium (pl. mycelia): Collective term for a mass or group of hyphae or fungal filaments (the
fungal thallus).

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Mycorrhiza (pl. mycorrhizae): The symbiotic association between certain fungi and the roots of
plants.
Ostiole (adj. ostiolate): A pore, often at the end of a neck-like structure, in an ascocarp or a
pycnidium.
Perfect stage (teleomorph): The (gametangial) sexual morph of an ascomycetous or
basidiomycetous fungus.
Perithecium (pl. perithecia): A rounded, oval, pyriform or beaked ascocarp (ascoma), characteristic
of the Pyrenomycetes, with a pore (ostiole) or slit at the top, and within which asci are borne in a
characteristic manner.
Phragmospore: A spore with two or more transverse septa.
Pseudoparenchyma (adj. pseudoparenchymatous): A type of plectenchyma consisting of closely
packed, more or less isodiametric or oval cells resembling the parenchyma of vascular plants.
Pycnidiospore: A conidium borne in a pycnidium.
Pycnidium (pl. pycnidia): An asexual hollow body lined inside with conidiophores bearing conidia.
It may be fully enclosed, or have an opening (ostiole).
Saccardoan system: The grouping of conidia (and dispersed ascospores) proposed by Saccardo,
mostly of the Ascomycota and Fungi imperfecti, based on the number of cells and the organization
of the septa in each spore, as well as the intensity of the pigmentation. The main groups are:
Amerosporae, Didymosporae, Phragmosporae, Dictyosporae, Scolecosporae, Helicosporae and
Staurosporae.
Sclerotium (pl. sclerotia): A resting body composed of a hardened mass of hyphae, from which
stromata or conidiophores may develop.
Scolecospore: An elongated needle- or worm-like spore.
Septate: Provided with (longitudinal or) transverse partitions.
Septum (pl. septa): Internal partition in a hypha or spore.
Sexual morph (Teleomorph): Form of an ascomycetous or basidiomycetous fungus bearing a sexual
organ.
Sporangiospore: A spore produced in a sporangium.
Sporangium (pl. sporangia): A sac-like structure producing spores endogenously.
Sporophore: A spore-bearing structure in fungi; a fungal hypha specialized to bear spores.
Staurospore: stellate conidium.
Stroma (pl. stromata): A compact vegetative tissue of hyphae in which ascomata are formed.
Teleutospore: An old term for ‘teliospore’.
Teliospore: A thick-walled resting spore of the terminal stage of Uredinales and Ustilaginales (rusts
and smuts).
Thallospore: A spore formed on the thallus (mycelium), either singly or in chains within a hypha
and liberated by disintegration of the hyphal wall; or terminal, as the swollen end of a hypha (and
not a distinct structure). It includes blastospores, arthrospores, Chlamydospores and oidia. In part
synonymous with conidia and aleurospores.
Thallus (pl. thalli): General term for the vegetative part of a non-vascular plant, particularly the
Thallophytes; of the fungi, the entire assimilative phase of the individual.
Thyriothecium (pl. thyriothecia): Shield-shaped body (in Hemisphaerales or Microthyriales) that is
oriented not by the mycelium, but by the host, with the generative tissue hanging downward, i.e.
inverted. It may be considered as half a perithecium, with the tip lying beneath.
Vesicular-arbuscular mycorrhiza (VAM): A mycorrhiza in which the fungal hyphae penetrate into
the cells of the host; also called endomycorrhiza.
Uredium (pl. uredia): The sorus of the Uredinales, bearing the spores.
Zoospore: A motile asexual spore.
Zygospore/Zygote: Thick-walled resting spore resulting from the conjugation of isogametes (as in
Zygomycota), or the fusion of similar gametangia.

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Geological time and life on earth
While studying past life, it is essential to know the Geological Time Scale. In geology, time
provides a frame of reference that is essential to the interpretation of extremely diverse types of
study involving all the materials and inhabitants of the earth and all the forces and processes that
have shaped it. Geologic time is often discussed in two forms: 1. Relative time (chronostratic),
referring to the Earth’s geology in a specific order based upon relative age relationships (most
commonly, vertical/ stratigraphic position). These subdivisions are given names, most of which can
be recognized globally, usually on the basis of fossils; and 2. Absolute time (chronometric),
referring to the numerical ages in ‘millions of years’ or some other measurement. These are most
commonly obtained via radiometric dating methods performed on appropriate rock types. In the
present stage of its development, geology is more concerned with relative time and the sequence of
events in the history of the earth than with the actual dating of these events. The development of the
Earth took place during past billions of years. The evolution of life on earth is also a part of the
Earth’s long history. The Geological Time Scale was constructed using the evidences collected
from field observations, fossil records, stratigraphic correlations, radioactive dating,
palaeomagnetic orientation, orbital revolution and rotation of the earth. Several major incidences
happened in the history of the earth, e.g. mass extinctions, appearance of new life forms (genera
and species), mountain-building movements, drifting of continents, spreading of ocean floors,
widespread glaciations, dominance of certain species and massive migration of life between land
and water. These factors are considered while accounting the Geological Time Scale.
The passage of time can be related to a series of events. In order to construct the history of
past events, one must determine how much time elapsed between the events and how long it took
for the events to occur. Time is also marked by certain characteristic or unique set of events. When
we list the events in an order in which they took place, we establish a chronological sequence of all
the events. Time has been flowing since the beginning of the earth. Time will continue to flow long
after the present generation also. To understand the past, we should know the present. To
understand the present, we should know the past. The age of the earth is one important aspect in
Earth Science studies. The relative ages of most of the rocks of the earth’s surface were dated by
following the principles of superposition, fossil correlation and relative dating of atomic clocks in
rocks, the Geologic Time Scale was prepared. Continuous annual deposition of sediments with life
remains in water bodies, can create sequential beds of sediments containing the fossils of animals
and plants. These are called as stratigraphic sequences. These stratigraphic sequences were
correlated to evaluate the evolutionary trend of ancient life and the geological formations.
The geologic time scale is a reference scale for the entire Earth’s history. It helps to
understand the entire history of the earth into workable units. Based on all the available evidences,
the earth is found to be around 4500 million years old since the starting point of the hot universe,
which gave birth to the galaxies. The geologic time of the earth is divided into five major eras, as
follows (oldest to youngest): i. Archean Era, ii. Proterozoic Era (4600–540 Ma.), iii. Palaeozoic Era
(540–245 Ma), iv. Mesozoic Era (245–66 Ma.) and v. Cenozoic Era (66 Ma–Present). The
subdivisions of era are periods and epochs. Various geological time units (eras, periods and epochs)
and significant events and life during the entire history of earth are summarized in Table 1.
Stratigraphic ranges and origins of some major groups of Animals, Plants and Fungi are given in
Fig. 1.

Table 1 Geological Time Scale (eras, periods and epochs with their absolute ages) and significant
events and life during the entire history of earth. Ma = mega annum (millions of years).

Eras Periods Epochs Significant events Life on earth

Holocene Palaeolithic and Neolithic cultures Modern plants and animals as
(0.01 Ma to (beginning ca. 10000 BC), Copper, of today, rise of human
Present) Bronze and Iron ages (beginning civilization, major habitat
ca. 3500 BC), man used iron changes and deforestations
implements in 1350 BC, youthful caused by introduction of

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Table 1 Continued.

Eras Periods Epochs Significant events Life on earth

landforms, high relief, humans pests and habitat destruction,
built cities. beginning of agriculture.
Quaternary 1.6 Pleistocene The Era of ice ages, glacial Fluorishing and then
MA to (1.6–0.01 Ma) climate, four great ice advances extinction of many large
Present) (The and retreats, dawn of human stone- mammals, evolution of
Age of man) age cultures, volcanic eruptions modern humans, Cro-Magnon
destroy human populations, man – first appearance of
formation of large scale deserts, present species, Neanderthal
Sahara was formed. man – Paleolithic culture;
Heidelberg man – Paleolithic
culture; Planetary spread of
Homo sapiens over Eurasia;
Extinction of many species
due to ice ages; Extinction of
many large mammals and
birds due to humans.
Cenozoic (66 Pliocene Worldwide elevation continues, Many of the existing
Ma to Present). (05–1.6 Ma) continental uplift and mountain generation of mammals and
Time of building, Ice Age begins, seas recent mollusks appeared,
Recent life. restricted, cool and dry climate. Homo habilis appeared, horses
The Age of and elephants became almost
Mammals modern in appearance, first
known appearance of
hominids (human like
primates), large carnivores
were dominant.
Miocene Moderate Icehouse climate, Modern mammal and bird
(25–05 Ma) extensive glaciation in southern families become recognizable,
hemisphere, orogeny in northern horses and Mastodons diverse,
hemisphere, widespread volcanism first apes appeared, whales,
and basalt flows were seen. apes and grazing mammals
dominated, notable advances
in the horses and elephant
families. Spread of grasslands
as forests contracted.
Tertiary 66– Oligocene Warm but cooling climate. Rapid evolution and
1.6 Ma) Age (38–25 Ma) diversification of fauna
of mammals). especially the mammals, early
ancestral elephants, carnivores
and ungulates become well-
established. Modern flowering
plants diversified.
Eocene Sea Marginal, extensive terrestrial Dawn of mammalian
(55–38 Ma) sedimentation, moderate, cooling dominance, subordinate
climate, reglaciation in South Pole, position for reptiles; archaic
mammals flourished, primitive
whales diversity was seen.
First grasses appeared.
Palaeocene Climate tropical, Alpine orogeny, Mammals diversity into a
(66–55 Ma) Himalayan orogeny. number of primitive lineages
following the extinction of the
dinosaurs. First large
mammals. Modern plants
appear.
Cretaceous – World continent Pangaea begins, This is the last period of the
(144–66 Ma) atmospheric CO2 close to present age of Dinosaurs; First
day levels. Breakup of primates appeared, new types
Gondwanaland and beginning of of insects, ammonites,
Rocky Mountains, widespread belemnites, bivalves,

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Table 1 Continued.
Eras Periods Epochs Significant events Life on earth
epicontinental seas, extensive echinoids and sponges were
chalk deposits. all common. Many new types
of dinosaurs and crocodilians
appeared on land; Modern
sharks appeared in the sea;
Primitive birds appeared
(toothed birds and flying
reptiles), marine pelecypods
became abundant.
Angiosperms appeared,
modern flowering plants
proliferated.
Mesozoic Jurassic – The world’s most famous period is Dominance of dinosaurs, its
(245-66 Ma). (208–144 Ma) the Jurassic period. Atmospheric spread between 208 and 44
Time of CO2 levels 4-5 times more than the Ma, many types of dinosaurs-
Middle life; present day levels (1200-1500 sauropods, carnosaurs and
The Age of ppmv), formation of oilfields of stegosaurs dominated, great
reptiles North Sea happened. Breakup of plant eating dinosaurs were in
Pangaea into Gondwana and existence. Oceans were with
Laurasia happened during this full of fish, squids and coiled
period. ammonites. Appearance of the
first frogs, salamanders,
crocodiles, flying reptiles and
birds were noticed. The first
birds and lizards appeared,
mammals were common,
bivalves, belemnites and
ammonites were abundant.
Sea urchins were common
along with crinoids, starfish,
sponges, terebratulid,
rhynchonellid and
brachiopods. Lush growth of
ferns and palm-like cycads,
gymnosperms also have grown
much.
Triassic – Continent emergent, seas marginal, Dominance of archosaurs on
(245–208 Ma) climate arid, occurrence of land as dinosaurs,
terrestrial deposition, formation of ichthyosaurs, nothosaurs in
salt, gypsum and red beds. oceans and pterosaurs in the
Orogenic movements prevailed in air, appearance of primitive
some parts, Pangaea still in mammals, domination of
existence, altering global climate reptiles and cycads, reduction
and ocean circulation happened. of marine invertebrates, first
mammals and crocodilian
appeared, extreme abundance
of ceratitic ammonoids were
seen, modern corals appeared,
first turtles, lizards, mammals
and dinosaurs. Dicroidium
flora were common on land,
appearance of modern
conifers, cycadeoids.
Permian (286– – World-wide continental uplift and Extinction of Palaeozoic
245 Ma). The orogenic movements, widespread plants and invertebrates (251
Age of aridity on one side and glaciation Ma), reduction in all types of
Amphibians at the other side. Landmasses unite life, almost 95% of life on the
into the super-continent Pangaea, earth became extinct,
creating the Appalachians. This primitive reptiles dominated in

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Table 1 Continued.
Eras Periods Epochs Significant events Life on earth
period records the end of Permo- places, extinction of all
Carboniferous glaciations. trilobites, graptolites and
blastoids, beetles and flies got
evolved, marine life flourished
in warm shallow reefs,
abundance of spiriferid
brachiopods, bivalves,
foraminifers and ammonoids.
Cone-bearing gymnosperms
(the first true seed plants) and
the first true mosses appeared.
Carboniferous – It is known for its coal deposits, Abundant life on land and
(360–286 Ma) limestones and gritstones, water, first land vertebrates,
Hercynian orogenic movements. sea invertebrates, prevalence
Highest ever atmospheric oxygen of foraminifers, bryozoans,
levels seen on earth were in this brachiopods, cephalopods,
period. blastoids, crinoids and corals.
Brachiopods are the zone
fossils of Carboniferous
Period. Lamellibranchs and
winged insects were important
fauna, first Reptiles laid eggs
with shells in this period.
Large primitive trees, swamp
forests with ferns were
existing, Lepidodendron and
Sigillaria were prominent
flora.
Palaeozoic Devonian – Violent volcanic eruptions and Age of fishes, sharks and rays,
(540-245 Ma). (408–360 Ma). crustal movements, folding, fishes move into the open
Time of The Age of mountain-building activities seas, lunged fishes,
Ancient life; fishes prevailed, climate became drier, amphibians appeared in
The Age of sea covered most of the land, earth Devonian, mollusks were
Invertebrates. appeared to look green. abundant. Extinction of
primitive vascular plants
happened, origin of modern
vascular plants with true
leaves, roots and stems, some
plants started to produce seeds
rather than spores.
Silurian (438– – Mild climate, stable and warm Rise of fishes and reef
408 ma) temperature, continents were building corals, abundance of
generally flat and flooded, notable shell-forming sea animals,
mountain building occurred in dominance of sea lilies,
Europe. eurypterids and land
scorpions, invasion of land by
arthropods. Origin of the
earliest vascular plants on
earth, modern group of Algae
and Fungi got evolved, 60% of
marine species were wiped out
at the base of the Silurian
period, first period to see
macrofossils of extensive
terrestrial biota.
Ordovician – Mild climate-Adaptive Radiation All plants and animals still
(505–438 Ma). (Ordovician radiation), shallow restricted to water, first
The Age of seas retreating from land and vertebrates originated as
Graptolites spreading back, first known marine jawless fishes, invertebrates

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Table 1 Continued.

Eras Periods Epochs Significant events Life on earth

transgressions occurred during this dominated, crustaceans,
period, southern continents were trilobites, graptolites,
collected into a single land mass brachiopods, bryozoans,
named as Gondwanaland; Major echinoderms, corals, mollusks
mountain-building activity and cephalopods dominated
happened during this period, sea the water masses. First fungi
levels were high during originated, invasions of land
Ordovician. by plants started.
Cambrian – Prevalence of mild climate, Period of abundant life on
(540–505 Ma). extensive seas, spilling over earth, after the Proterozoic,
The Age of continents. During this period, the shelled marine invertebrates,
trilobites continents had a soil crust and explosive growth of
were resembling like deserts, eukaryotic organisms,
prosperity of life in the oceans than swimming, floating, crawling,
lands. clinging and burrowing sea
animals, appearance of
trilobites, brachiopods,
gastropods, radiolarians,
sponges, echinoderms,
starfish, sea cucumbers, jelly
fish, worms and water
scorpions. Existence of plants
was observed only as algae,
absence of land plants.
Precambrian – – Dry and cold climate to warm and Origin of eukaryotic cells and
(Proterozoic, moist conditions prevailed on the multicellular life, occurrence
2500–540 Ma) earth, atmosphere became enriched of earliest known fossils
in oxygen, tectonic plates were including of soft-bodied
present and began moving. marine invertebrates, origin of
sponges, cnidarians and
annelids. (sea anemones,
segmented flatworms)
Precambrian – – Extensive mountain-building, Origin of life, ancient Life, the
(Archean, formation of banded iron ores and first life forms evolve - one
3960–2500 greenstone belts, existence of celled organisms, especially
Ma) shallow seas, accumulation of free the prokaryotes, bacteria and
oxygen. blue-green algae.
Precambrian – – Rockless Eon, the solidifying of No life
(Hadean, the Earth's continental and oceanic
4600–3960 crusts.
Ma)

Materials & Methods

This section contains:

1. Field investigation procedure;
2. Laboratory investigation procedure;
3. Source of information;
4. Method of data presentation.

1. Field investigation procedure

This includes observation on the exposed rock formations and collection of samples for
microfossil studies. First of all, a reconnaissance survey is carried out in various directions for
understanding the geological setting of the area and lateral persistence of various lithic types and
their order of superposition. In most of the cases, the complete thickness of any rock formation is
not exposed in any one section. A composite section is then compiled with the help of two or more

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sections. While collecting samples from exposures, utmost care is taken to avoid contamination.

Figure 1 – Generalized diagram showing stratigraphic ranges and origins of some major groups of
Animals, Plants and Fungi (Not to scale).

Precise geographic (including GPS readings) and stratigraphic locations along with geological
details are noted in the field diary. Suitable lithologies are identified for collection of samples
because recovery of microfossils, especially with regard to their abundance, depends on the rock
types. Amongst the sedimentary rocks, limestone, carbonaceous shale, lignite, coal, and shaly
siltstone are considered to be rich in microfossil contents whereas coarse textured rocks, weathered
sediments and conglomerates are unlikely to yield microfossils. For collection of samples from
exposed rocks, escarpments, ledges, river or stream cuttings (Fig. 2B), ridges, cliffs (Fig. 3A–C),
are good natural sites. Usually, these sites contain large amounts of weathered rocks and vegetation
or algal scum on its upper surface. Before collection, fresh rocks are exposed by removing the
weathered rocks/ vegetation. Quarries, road cuttings (Fig. 2A, E–H), railway cuttings or open cast
mine faces (Fig. 2C–D) are among the artificial, man-made sites for sample collection. Such sites
are good for sample collection because these are easily accessible and here the rocks are generally

681
not covered with weathered sediments and vegetation. Samples must be collected from measured
stratigraphic sections only.

Figure 2 – Stratigraphic sections. A, E–H Road cutting sections. B Stream cutting section. C–D
Mine face sections. A Stratigraphic section exposing the contact of Matanomadh Formation with
the overlying Naredi Formation, near Matanomadh Village, Kutch District, Gujarat, India (after

682
Saxena 1981). B Intertrappean bed exposed at about 5 km from Naredi on Naliya-Narayan Sarovar
Road, Kutch District, Gujarat, India (after Saxena & Ranhotra 2009). C Mavli Mine section
(NIMCO) at Redi showing Sindhudurg Formation (containing lignite) overlain by red ferruginous
laterite (after Saxena et al. 1992). D Stratigraphic section of Pit no. 1 (Gogte Minerals) at Redi
showing Sindhudurg Formation (containing lignite) overlain by pale red ferruginous laterite (after
Saxena et al. 1992). E–H Road cutting sections showing Kuksho Formation at various locations
along Kargil-Leh Road, Ladakh, India (unpublished photographs by R.K. Saxena).

Spot samples are taken at more or less uniform intervals along the thickness of the bed. The interval
of sampling depends upon the thickness and lithology of the beds. For collection from coal/ lignite
beds or from other beds of homogeneous lithology, channel sampling is done. In this method,
samples are collected through the vertical thickness of any bed or specified thickness after making a
15 cm deep channel at right angle to the bedding plane. Vertical stratigraphic sections are shown in
Fig. 3F–H. The deeply buried (subsurface) sediments are taken with the help of well drilling
equipments by which a core, often hundreds of metres long, can be obtained. The bore cores are
ideal material for microfossil analysis because they provide fresh samples from uninterrupted
stratigraphic sequence without any chance of contamination or mixing (Fig. 3D). Stratigraphic
sections of boreholes are shown in Fig. 3F, H.

2. Laboratory investigation procedure

For recovery of palynofossils and making slides, the following steps are taken: 1. Cleaning
and desegregation, 2. Chemical processing, 3. Density separation, and 4. Mounting of slides and
study of palynofossils (Fig. 4).
Cleaning and desegregation: Rock samples are cleaned thoroughly with water or alcohol to
ensure removal of any kind of extraneous matter. Desegregation of rock samples is done by
breaking them into small pieces with the help of pestle and mortar.
Chemical processing: Chemical processing is carried out in two steps: a. Removal of minerals
(demineralization) and b. Alkali treatment.

a. Removal of minerals (demineralization): For removal of minerals, the rock samples are subjected
to treatment with acids depending upon the kind of rock types. Carbonates, commonly calcites, are
soluble in hydrochloric acid (HCl); sulphates, sulphides and carbon contents are soluble in
concentrated nitric acid (HNO3); and silicates, the most common content of the rocks, are removed
by 40–60% hydrofluoric acid (HF). The steps adopted for processing of different lithologies are as
follows. Shale and siltstone: Crushed samples with few drops of distilled water are kept in 40% HF
for 4–5 days or till the samples become completely pulverized. Containers having pulverized
samples are filled with water; the material is allowed to settle for one hour and the supernatant
water is decanted with the help of siphon tube. After repeating the process twice, samples are
sieved with 400 mesh (37 μm) and the residue left over the mesh is transferred to container. In
carbonaceous shale samples little amount of commercial nitric acid is added and the container is
kept inside the fume hood chamber (Fig. 3E) for 24 hours. After siphoning the supernatant water 4–
5 times, samples are washed again with 400 mesh sieves. Limestone/mudstone: Crushed samples
are first treated with concentrated hydrochloric acid for 24 hours and are then kept in 40%
hydrofluoric acid for 2–3 days. Coal/lignite: Crushed samples are taken in glass containers and after
adding a few drops of water, concentrated nitric acid is added drop by drop. Samples are constantly
stirred and are placed inside the fume-hood chamber for 24–36 hours. The samples, when
pulverized, are washed 2–3 times with water by siphoning the supernatant water. Each sample is
sieved with 400 mesh sieve and the residue left over the mesh is transferred to the container.
b. Alkali treatment: After removal of minerals, samples are treated with solution of sodium
carbonate or potassium hydroxide. The concentration of these solutions (10–20%) and duration of
treatment (2–5 minutes) are decided after checking the demineralized residue under the microscope.
Alkali treatment is essential to remove the humic acids released during the process of

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demineralization. As a result of this treatment, the sample turns dark brown. After alkali treatment,
samples are thoroughly washed with water to remove all traces of alkali.
Density separation: Sequel to the demineralization process, very fine mineral particles mixed
with the macerated residue are generally observed. These are removed by density separation
methods. These methods are also essential to separate and concentrate the microfossil fraction from
the residue. Owing to their specific gravity, minerals and organic particles have different settling
velocities. Spores and pollen, having much less density than the mineral particles, tend to float in
water for a longer time and therefore can easily be separated. The commonly used density
separation methods are: a. Swirling and b. Heavy liquid separation. Steps followed to achieve these
methods are discussed below.

a. Swirling: The macerated residue is mixed with 10–15 ml of water in a medium-sized watch
glass. The watch glass is rotated in clock-wise direction. This action allows the mineral particles
and larger cuticle pieces to settle at the bottom in the centre, whereas the lighter particles, including
spores, pollen and fungi, remain suspended in the water. The upper layer of water is now
transferred to another watch glass with the help of a dropper and is allowed to settle for about half
an hour. The watch glass is again rotated allowing the organic matter to collect in the centre of
watch glass. Water from the watch glass is removed with the help of a dropper. The collected
material, containing spores, pollen and fungal remains, is mounted on slides for observation under
the microscope.
b. Heavy liquid separation: This method is generally used to concentrate and separate the
spore/pollen fraction from the macerated residue. In this method the chemically processed material
is mixed with liquids whose specific gravity is higher than those of the organic particles. Heavy
liquids, when thoroughly mixed with residue, allow the organic fraction to float at the upper part of
it. Minerals being heavier, settle faster than the organic material. Centrifuging of the mixture leads
to better results. Steps followed to separate the organic fraction from the macerated residue are as
below: The residue is transferred to a centrifuge tube and is washed with 10% HCl. Washing with
HCl at this stage prevents precipitation of insoluble impurities when the heavy liquid (ZnCl2 or
ZnBr2) solution is added. The specific gravity of the solution should be about 2. The solution is
thoroughly mixed with residue and is centrifuged at 1500 rpm for 10–20 minutes. This action
permits the settling of inorganic and organic components of the residue at the bottom and top layers
respectively. The topmost layer, rich in spore/pollen contents, is sucked with the help of a dropper
and is washed in sequence with 10% HCl and water.

Mounting of slides and study of palynofossils: Water-free macerated residue is mixed with a
few drops of polyvinyl alcohol solution and is spread uniformly over the cover glass with the help
of a glass rod. The cover glasses are dried in oven for about 30 minutes and are then mounted in
Canada balsam. To make the slides ready for study under microscope, slides are kept in oven at 50–
60°C for 1 to 2 days. Slides prepared out of the productive samples are examined under the
microscope for qualitative and quantitative assessment. Distinguishable morphotypes are identified
and described under the artificial system of classification. Frequency of palynotaxa is determined
by counting 200 palynofossil specimens in each sample. However, in case of poorly productive
samples only 100 specimens may be counted.

3. Source of information
The data, presented here, have been obtained from the various studies on fossil fungi
published during over one hundred years. Kalgutkar & Jansonius (2000) published a synopsis of
fossil fungi and tried to streamline taxonomic status of many fossil fungal genera and species. They
provided descriptions for ca. 950 validly published species, attributed to approximately 300 genera
and introduced twelve new genera and ca. 350 new combinations. Transfers of species to more
appropriate genera resulted in 31 later homonyms, for which they provided new names. They also
validated one genus and several species. In order to include all records of fossil fungal remains

684
from the Indian Tertiary sediments, published till 2005, three catalogues were published
(Lakhanpal et al. 1976, Saxena 1991, 2006). Besides, a monographic study was carried out by
Saxena & Tripathi (2011) with the objective to synthesize the available information on Indian fossil
fungi. This incorporates description of 152 genera and 388 species of fossil fungi, including 15 new
species and 12 new combinations, with comments wherever required. In addition to the above
monographic studies, data have been gathered from scores of publications, containing information

Figure 3 – A–C Cliff sections. D Bore cores. E Fume hood chamber. F, H Borehole sections. G
Mine face section. A Sawai Bay Formation (mudstone) exposed in the type section, Car Nicobar

685
Island, Andaman & Nicobar Islands, India (after Chandra & Saxena 1998). B Hard, arenaceous,
greyish yellow limestone (Kakana Formation) exposed near Mus Jetty, Car Nicobar Island,
Andaman & Nicobar Islands, India (after Ghosh et al. 2004). C Bedded. arenaceous, shelly
limestone (Kakana Formation) exposed near Kakana Village, Car Nicobar Island, Andaman &
Nicobar Islands, India (after Ghosh et al. 2004). D Sequential arrangement of rocks in bore-cores
(after Tripathi 1995). E Fume hood chamber (after Tripathi 1995). F Rıo Foyel Section, showing
stratigraphic position of studied samples, El Foyel Group (Palaeogene) of Nirihuau Basin,
Argentina (after Martínez et al. 2016). G Mavli Mine section at Redi, Sindhudurg District,
Maharashtra, India (after Saxena 2000). H Lithological profile showing stratigraphic position of
selected samples in the wells, Miocene deposits of the Pelotas Basin, Brazil (after Premaor et al.
2018).

Figure 4 – Showing various steps taken for extraction of fossil fungi from the rock samples.

on fossil fungi from all parts of the globe, published in various journals and conference proceedings
(Van der Hammen 1954, 1955, 1956, Rouse 1959, 1962, Potonié & Sah 1960, Playford 1962,
Varma & Rawat 1963, Ramanujam & Ramachar 1963 1980, Clarke 1965, Dilcher 1965, Mathur
1966, Srivastava 1968, Taugourdeau 1968, Elsik 1968, 1969, 1990a, b, 1992, 1999, Mathur &
Mathur 1969, Dutta & Sah 1970, Jain & Gupta 1970, Trivedi & Verma 1970, Sheffy & Dilcher
1971, Lange & Smith 1971, 1975, Kar et al. 1972, Jain et al. 1973, Venkatachala & Rawat 1973,
Biradar & Mahabale 1974, Elsik & Dilcher 1974, Elsik & Jansonius 1974, Sah & Kar 1974,
Salujha et al. 1974, Jansonius 1976, Kar & Saxena 1976, Paradkar 1976, Chitaley & Yawale 1978,
Ke & Shi 1978, Kemp 1978, Ramanujam & Rao 1978, Smith 1978, Dueñas 1979, Jain & Kar 1979,
Pirozynski & Weresub 1979, Kar 1979, 1990, Ramanujam & Srisailam 1980, Salard-Cheboldaeff
& Locquin 1980, Zhang 1980, Ediger 1981, Huang 1981, Singh & Saxena 1981, Barlinge &
Paradkar 1982, Saxena & Singh 1983, Ambwani 1982, 1983, Chandra et al. 1984, Pathak &
Banerjee 1984, Gupta 1984, 1985, 2002, Song 1985, Stubblefield et al. 1985, Varma & Patil 1985,
Glass et al. 1986, Jarzen & Elsik 1986, Saxena & Sarkar 1986, Singh et al. 1986, Norris 1986,

686
1997, Saxena & Bhattacharyya 1987, 1990, Kalgutkar & Sweet 1988, Saxena et al. 1988, Ediger &
Alisan 1989, Martínez-Hernández & Tomasini-Ortiz 1989, Song et al. 1989, 1999, Elsik et al.

4. Method of data presentation

The taxa are placed into the following spore groups: 1. Amerosporae, 2. Didymosporae, 3.
Phragmosporae, 4. Dictyosporae, 5. Helicosporae, 6. Staurosporae, and 7. Fossil fungal spore
species assigned to modern genera. Under each of these, the genera are arranged in alphabetical
order. The species are also arranged under each genus in alphabetical order, as follows.
The first paragraph includes i. Name of the genus followed by its author(s) and full
bibliographical details, i.e. name of journal, volume number, page number and year of publication,
ii. Index Fungorum Registration Identifier, iii. Type species followed by its author(s) and year of
publication, and iv. Current name (if it is now different from the original name). The name of the
legitimate genus is in bold capital letters whereas the same of the illegitimate one is in non-bold
capital letters. In case of an illegitimate genus name, its current name is provided in bold capital
letters.
The second paragraph presents Original Diagnosis (with reference to its author and year of
publication in parenthesis). This is followed by Emended Diagnosis, if any (with reference to its
author and year of publication in parenthesis). In case of more than one emended diagnosis, all are
given in chronological order with similar details. The subsequent paragraphs pertain to
Classification, Number of species known and Notes, if any.
Similarly, all the species described under each genus are arranged in alphabetical order. Each
species is provided with the following information: i. Name of the species followed by its author(s)
and year of publication, ii. Index Fungorum Registration Identifier, iii. Synonym(s), if any, iv.
Location, v. Age, and vi. Notes, if any. The name of the legitimate species is in bold letters whereas
the same of the illegitimate one is in non-bold letters. In case of illegitimate species name, its
current name is provided in bold letters. Drawings of holotypes are provided for the type species of
each genus, new species and new combinations and some commonly occurring fungal spore
species.

Genera and species of fossil fungal spores

1. Amerosporae

1.1. Genus: AMEPIOSPORA Sal.-Cheb. & Locq., C. r. Congr. natn. Socs. sav. Paris, sect. sci., fasc.
1. (Paleobotanique) 105: 185 (1980); Index Fungorum Registration Identifier: 25592; Type: A.
ellipsoidea Sal.-Cheb. & Locq. 1980; Current name: BASIDIOSPORITES Elsik 1968 fide
Kalgutkar & Jansonius (2000).
Original Diagnosis: Smooth, elliptical amerospores (Salard-Cheboldaeff & Locquin 1980).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Five (all the species, including type species, have been transferred
to other genera).
Notes: The characters mentioned in the generic diagnosis are not sufficient to separate it from
other unicellular inaperturate fungal spore genera (e.g. Basidiosporites Elsik, Lacrimasporonites
R.T. Clarke, Monoporisporites Hammen). Hence, Kalgutkar & Jansonius (2000) transferred
Amepiospora ellipsoidea, the type species of Amepiospora, to Basidiosporites. This makes
Amepiospora a later synonym of Basidiosporites. The other species of Amepiospora have been
transferred to Monoporisporites and Lacrimasporonites, as given ahead.
1.1.1. Species: A. ellipsoidea Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107672; Current name: Basidiosporites ellipsoideus (Sal.-Cheb. & Locq.) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.1.2. Species: A. fusoidis Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107718; Current name: Lacrimasporonites fusoides (Sal.-Cheb. & Locq.) Kalgutkar &

687
 Janson. 2000 fide Kalgutkar & Jansonius (2000); Notes: Orthographic correction was made
by Kalgutkar & Jansonius 2000.
1.1.3. Species: A. globosa Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107719; Current name: Monoporisporites neoglobosus Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.1.4. Species: A. macrospora Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107720; Current name: Monoporisporites macrosporus (Sal.-Cheb. & Locq.) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.1.5. Species: A. triangularis Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107721; Current name: Monoporisporites triangularis (Sal.-Cheb. & Locq.) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).

1.2. Genus: ASYREGRAAMSPORA Sal.-Cheb. & Locq., C. r. Congr. natn. Socs. sav. Paris, sect.
sci., fasc. 1. (Paleobotanique) 105: 191 (1980); Index Fungorum Registration Identifier: 25593;
Type: A. reticulata Sal.-Cheb. & Locq. 1980.
Original Diagnosis: Amerospores, more or less reticulate, slightly asymmetrical,
approximately 20 × 15 µm (Salard-Cheboldaeff & Locquin 1980).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
Notes: Jansonius & Hills (1981) corrected spelling of the type species from “Asyreamspora
reticulata” (as given in the protologue by Salard-Cheboldaeff & Locquin 1980: 185) to
Asyregraamspora reticulata (as per spelling of the generic name used by Salard-Cheboldaeff &
Locquin 1980: 191).
1.2.1. Species: A. reticulata Sal.-Cheb. & Locq. 1980 (Fig. 5A); Index Fungorum Registration
Identifier: 107752; Location: Coast of Equatorial Africa, Gulf of Guinea; Age: Early
Miocene.

1.3. Genus: BASIDIOSPORITES Elsik, Pollen et Spores 10(2): 273 (1968); Index Fungorum
Registration Identifier: 21028; Type: B. fournieri Elsik 1968.
Synonym: Amepiospora Sal.-Cheb. & Locq. 1980 fide Kalgutkar & Jansonius (2000), Index
Fungorum Registration Identifier: 25592.
Original Diagnosis: Monoporate, unicellate, psilate fungal spores with the pore offset from
one apex. No basal attachment area evident. Shape variable, generally elongate in some fashion
(Elsik 1968).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Four.
Notes: The spores of Basidiosporites are characterized by a pore offset from one apex. Since
pore in Amepiospora ellipsoidea Sal.-Cheb. & Locq. 1980 and Monoporisporites ovalis Sheffy &
Dilcher 1971 is offset from one apex, Kalgutkar & Jansonius (2000) transferred them to
Basidiosporites.
1.3.1. Species: B. ellipsoideus (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483241; Basionym: Amepiospora ellipsoidea Sal.-Cheb. & Locq.
1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age: Early
Eocene-Early Miocene.
1.3.2. Species: B. fournieri Elsik 1968 (Fig. 5B); Index Fungorum Registration Identifier: 309461;
Location: Strip mine approximately 11 km south-west of Rockdale, Milam County, Texas,
U.S.A.; Age: Palaeocene (Rockdale lignite); Notes: Elsik (1968) suggested affinity of this
genus with Basidiomycota.
1.3.3. Species: B. ovalis (Sheffy & Dilcher) Kalgutkar & Janson. 2000 (Fig. 5C); Index Fungorum
Registration Identifier: 483267; Basionym: Monoporisporites ovalis Sheffy & Dilcher 1971;
Location: Puryear clay pit, 800 m south of Puryear, Tennessee, Henry County, U.S.A.; Age:

688
 Middle Eocene (Claiborne Formation); Notes: The specimen shows some darkening
(thickening?) in median region of spore.
1.3.4. Species: B. sadasivanii Anil Chandra et al. 1984 (Fig. 5D); Index Fungorum Registration
Identifier: 106626; Location: Sediment core no. 1 (Lat. 17°57.9'N: Long. 70°46.0'E), Arabian
Sea; Age: Late Quaternary; Notes: Chandra et al. (1984) differentiated this species from
Basidiosporites fournieri Elsik 1968 by its larger size, placement of pore in the middle of the
longer axis and thickened pore margin. The species epithet is in honour of T.S. Sadasivan,
Department of Botany, Madras University, Chennai, India.

1.4. Genus: BIPORIPSILONITES Kalgutkar & Janson., AASP Contributions Series (Dallas) 39:
37 (2000); Index Fungorum Registration Identifier: 28612; Type: B. belluloides (Z.C. Song)
Kalgutkar & Janson. 2000.
Original Diagnosis: Generally small to medium-sized unicellate fungal spores of more or less
elongate fusiform to barrel-shaped outline; generally with a plane of symmetry through the equator;
spore wall generally smooth, occasionally with some subdued sculpture, and of medium thickness;
two terminal pores, forming pore chambers subtended by a basal septum, and enclosed by thin wall
material that further thins centrifugally; septa, thin or thick, may show a central perforation and/or
small septal folds; the terminal pore itself may be closed by very thin wall material, or ruptured to
gaping and broad (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Eleven.
Notes: Spores in this genus can be differentiated from Diporisporites Hammen by pore
chambers. The name of the genus is derived from Latin bi-, or two, porate, and smooth (Greek
psilos) character of the wall.
1.4.1. Species: B. anceps (G. Norris) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483269; Basionym: Striadiporites anceps Norris 1997; Location: Mackenzie River
delta, Canada; Age: Palaeocene-Eocene; Notes: Kalgutkar & Jansonius (2000) interpreted the
annular thickenings as septa (or septal bases) and for that reason transferred the species to the
new Biporipsilonites, which is characterized precisely by an oval shape, and one (or two) sets
of septa underlying each of the terminal pores.
1.4.2. Species: B. belluloides (Z.C. Song) Kalgutkar & Janson. 2000 (Fig. 5E); Index Fungorum
Registration Identifier: 483268; Basionym: Diporicellaesporites belluloides Z.C. Song 1985;
Location: Jiandingshan, Huangshi and Huatugou, Qaidam Basin, Qinghai Province, China;
Age: Palaeocene-Late Eocene (Ke & Shi 1978); Early Miocene-Late Miocene (Song 1985).
1.4.3. Species: B. bellulus (P. Ke & Z.Y. Shi) ex Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483270; Basionym: Diporicellaesporites bellulus P. Ke & Z.Y. Shi
1978: 49, pl. 5, fig. 4 (nom. inval.), lectotype was designated by Kalgutkar & Jansonius 2000;
Location: Panshan, Liaoning Province; Laoshanggulin and Beidagang, Tianjin Municipality,
Coastal region of Bohai, China; Age: Eocene-Oligocene.
1.4.4. Species: B. fusiformis (Anil Chandra et al.) Kalgutkar & Janson. 2000 (Fig. 5F); Index
Fungorum Registration Identifier: 483271; Basionym: Diporisporites fusiformis Anil
Chandra et al. 1984; Location: Sediment core no. 1 (Lat. 17°57.9'N: Long. 70°46.0'E),
Arabian Sea; Age: Late Quaternary.
1.4.5. Species: B. karii (Anil Chandra et al.) Kalgutkar & Janson. 2000 (Fig. 5G); Index Fungorum
Registration Identifier: 483272; Basionym: Multicellaesporites karii Anil Chandra et al.
1984; Location: Sediment core no. 2 (Lat. 18°35.2'N: Long. 69°17.2'E), Arabian Sea; Age:
Late Quaternary; Notes: The species epithet is in honour of Dr. R.K. Kar, Birbal Sahni
Institute of Palaeosciences, Lucknow, India.
1.4.6. Species: B. krempii (C.P. Varma & Rawat) Kalgutkar & Janson. 2000 (Fig. 5H); Index
Fungorum Registration Identifier: 483273; Basionym: Psilodiporites krempii C.P. Varma &
Rawat 1963; Location: Western and eastern India, including oil exploration areas in West
Bengal and Assam, India; Age: Early to Late Eocene.

689
1.4.7. Species: B. maximus (Z.C. Song & H.C. Luo in Z.C. Song et al.) Kalgutkar & Janson. 2000;
Index Fungorum Registration Identifier: 483274; Basionym: Diporisporites maximus Z.C.
Song & H.C. Luo in Z.C. Song et al. 1989; Location: Qingfeng County of Henan Province,
China; Age: Late Eocene-Middle Oligocene (Shahejie Formation).
1.4.8. Species: B. mollis (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483275; Basionym: Multicellaesporites mollis P. Ke & Z.Y. Shi 1978;
Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
1.4.9. Species: B. mustus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483276; Basionym: Multicellaespoites mustus P. Ke & Z.Y. Shi 1978;
Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
1.4.10. Species: B. padappakkarensis (P. Kumar) Kalgutkar & Janson. 2000 (Fig. 5I); Index
Fungorum Registration Identifier: 483277; Basionym: Diporicellaesporites
padappakkarensis P. Kumar 1990; Location: Padappakkara, Kollam District, Kerala, India;
Age: Early-Middle Miocene (Quilon Beds).
1.4.11. Species: B. verus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483278; Basionym: Multicellaesporites verus P. Ke & Z.Y. Shi 1978;
Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.

1.5. Genus: BIPORISPORITES P. Ke & Z.Y. Shi, Early Tertiary Spores and Pollen Grains from
the Coastal Region of Bohai (Beijing): 45 (1978); Index Fungorum Registration Identifier: 21029;
Type: B. rotundus P. Ke & Z.Y. Shi 1978.
Original Diagnosis: Spores one-celled, spherical. Diporate, pores situated at same end of
spore. Spore wall of medium thickness, surface psilate or scabrate (Ke & Shi 1978).
Emended Diagnosis: Fungal spore-like bodies, unicellate, equilateral, heteropolar, oblate,
circular to oval in shape, one end (defined as apical) with a central boss-like thickening flanked by
two pores at its proximal end and prominent lumina in the spore wall laterally adjacent to the
constricted median part of the boss. Lateral arm-like thickenings arise from the distal end of the
boss. Spore wall smooth, single layered, unornamented or bearing variable ornament in the form of
concentric or reticulate thickenings that become subdued in the area immediately surrounding the
central boss (Norris 1997).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Two.
Notes: Ke & Shi (1978) distinguished spores in this genus from those in Diporisporites
Hammen 1954 by the two pores situated at the same end of the spore outline.
1.5.1. Species: B. praestigiatus G. Norris 1997; Index Fungorum Registration Identifier: 483787;
Location: Mackenzie River delta, Canada; Age: Palaeocene-Eocene; Notes: The name of the
species epithet is derived from Latin praestigiator = juggler, for the apparently outstretched
arms and circling objects.
1.5.2. Species: B. rotundus P. Ke & Z.Y. Shi 1978 (Fig. 5J); Index Fungorum Registration
Identifier: 115624; Location: Panshan, Liaoning Province, Coastal region of Bohai, China;
Age: Eocene-Oligocene.

1.6. Genus: CADYEXINIS Stach, Palaeontontographica 102(4-6): 86 (1957); Index Fungorum

Registration Identifier: 21040; Type: C. vulgaris Stach 1957.
Original diagnosis: Spores 1-aperturate; elongated; exine psilate, smooth (Stach 1957, vide
Huang 1981).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Three.

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1.6.1. Species: C. taiwanensis T.C. Huang 1981; Index Fungorum Registration Identifier: 115726;
Location: Taiwan; Age: Miocene.
1.6.2. Species: C. tenuis Stach 1957; Index Fungorum Registration Identifier: 114311; Location:
Germany; Age: Carboniferous.
1.6.3. Species: C. vulgaris Stach 1957 (Fig. 5K); Index Fungorum Registration Identifier: 114312;
Location: Germany; Age: Carboniferous.

1.7. Genus: CERVICHLAMYDOSPORA R. Kar et al., Review of Palaeobotany & Palynology

(Amsterdam) 158(3-4): 247 (2010); Index Fungorum Registration Identifier: 541645; Type: C.
nigra R. Kar et al. 2010.
Original Diagnosis: Chlamydospores subcircular, 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 (Kar et al. 2010).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
1.7.1. Species: C. nigra R. Kar et al. 2010 (Fig. 5L); Index Fungorum Registration Identifier:
542235; Location: Tlangsam, Mizoram, India; Age: Miocene (Bhuban Formation).

1.8. Genus: CUPULISPORONITES Z.C. Song & Liu Cao, Monograph, State-Antarctic Committee,
China (Beijing) 3: 38 (1994); Index Fungorum Registration Identifier: 28632; Type: C. megaporus
Z.C. Song & Liu Cao 1994; Current name: ANATOLINITES Elsik et al. 1990 fide Kalgutkar &
Jansonius (2000).
Original Diagnosis: One-celled spore, dolium- or cupula-shaped in outline, with a nearly
equal width in the two terminal ends; one pore, the structure and form of pore variable; exine solid,
laevigate or with weak ornamentation (Song & Cao 1994).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One (the single species has been transferred to Anatolinites Elsik
et al. 1990).
1.8.1. Species: C. megaporus Z.C. Song & Liu Cao 1994; Index Fungorum Registration Identifier:
483763; Current name: Anatolinites megaporus (Z.C. Song & Liu Cao) Janson. et al. 1998
fide Jansonius et al. (1998).

1.9. Genus: DIPORISPORITES Hammen, Bol. Geol. (Bogota) 2(1): 83 (1954); Index Fungorum
Registration Identifier: 21084; Type: D. elongatus Hammen 1954 (holotype was designated by Van
der Hammen 1955).
Synonym: Scabradiporites Y.K. Mathur 1966 fide Kalgutkar & Jansonius (2000), Index
Fungorum Registration Identifier: 21292.
Original Diagnosis: Fungal spores “with two small pores” (Van der Hammen 1954).
Emended Diagnosis: Diporate fungal spore of one cell. Pores on opposite ends of the grains.
Pores may be modified, i.e. with atrium, annulus or septum forming pore chamber. Shape variable.
Ornamentation variable (Elsik 1968).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: 34 (but we accept only 17 species as legitimate because 16 species
have been transferred to other genera and one species, viz. D. minutus Hammen 1954, was not
validly published).
Notes: Potonié (1960: 113) formulated the following generic diagnosis: Shape approximately
fusiform, with two opposite small pores at the tapered ends, smaller than in Diporites, in part with
an annulus; exolamella more or less smooth.
1.9.1. Species: D. anklesvarensis (C.P. Varma & Rawat) Elsik 1968 (comb. inval.); Index
Fungorum Registration Identifier: 313250; Current name: Foveodiporites anklesvarensis
C.P. Varma & Rawat 1963 fide Kalgutkar & Jansonius (2000).

691
1.9.2. Species: D. barrelis A. Gupta 2002 (Fig. 5M); Index Fungorum Registration Identifier:
540470; Location: Dadahu Road Section, Sirmaur District, Himachal Pradesh, India; Age:
Eocene (Subathu Formation).
1.9.3. Species: D. bhavnagarensis R.K. Saxena 2009 (Fig. 5N); Index Fungorum Registration
Identifier: 515013; Basionym: Diporisporites granulatus B. Samant 2000; Location: Near
Bhavnagar, Cambay Basin, Gujarat, India; Age: Early Eocene (Kharsalia Clay Formation).
1.9.4. Species: D. communis P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115643; Location: Panshan, Liaoning Province; Tanggu, Tianjin Municipality; Kenli,
Shandong Province, Coastal region of Bohai, China; Age: Eocene-Oligocene; Notes: Elsik
(1968) distinguished spores in this species from those in Diporisporites hammenii Elsik 1968
by their larger spore size and broadly rounded outline.
1.9.5. Species: D. conspicuus Ramanujam & K.P. Rao 1978; Index Fungorum Registration
Identifier: 115062; Current name: Foveodiporites conspicuus (Ramanujam & K.P. Rao)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.9.6. Species: D. curvatus Ramanujam & K.P. Rao 1978; Index Fungorum Registration Identifier:
115063; Current name: Hypoxylonites curvatus (Ramanujam & K.P. Rao) Elsik 1990a fide
Elsik (1990a).
1.9.7. Species: D. elegans P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier: 115642;
Current name: Foveodiporites elegans (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.9.8. Species: D. ellipsoides (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483322; Basionym: Psiammopomopiospora ellipsoides Sal.-Cheb. &
Locq. 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Late Eocene-Oligocene; Notes: Elsik (1968) suggested affinity of this species with
Basidiomycota.
1.9.9. Species: D. elongatus Hammen 1954 (Fig. 5O); Index Fungorum Registration Identifier:
330110; Location: Eastern Cordilleras, Colombia, South America; Age: Maastrichtian.
1.9.10. Species: D. elsikii R.K. Saxena 2000 (Fig. 5P); Index Fungorum Registration Identifier:
519769; Location: Mavli Mine at Redi, Sindhudurg District, Maharashtra, India; Age:
Miocene (Sindhudurg Formation).
1.9.11. Species: D. fusiformis Anil Chandra et al. 1984; Index Fungorum Registration Identifier:
106781; Current name: Biporipsilonites fusiformis (Anil Chandra et al.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
1.9.12. Species: D. giganticus R.K. Kar in R.K. Saxena 2012 (Fig. 5Q); Index Fungorum
Registration Identifier: 519770; Basionym: Diporisporites giganticus R.K. Kar 1990 (nom.
inval.); Location: Tripura-Assam, North-east India; Age: Miocene-Pliocene (Surma and
Tipam groups).
1.9.13. Species: D. granulatus (Rouse) Elsik 1968 (comb. inval.) fide Kalgutkar & Jansonius
(2000); Index Fungorum Registration Identifier: 631277; Current name: Diporisporites
pergranulatus Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.9.14. Species: D. granulatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115641; Current name: Foveodiporites granulatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
1.9.15. Species: D. granulatus B. Samant 2000; Index Fungorum Registration Identifier: 529671;
Current name: Diporisporites bhavnagarensis R.K. Saxena 2009 fide Saxena (2009).
1.9.16. Species: D. gunniae (C.P. Varma & Rawat) Elsik 1968 (comb. inval.) fide Kalgutkar &
Jansonius (2000); Index Fungorum Registration Identifier: 313253; Current name:
Foveodiporites gunniae (C.P. Varma & Rawat) Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
1.9.17. Species: D. hammenii Elsik 1968 (Fig. 5R); Index Fungorum Registration Identifier:
313254; Location: Strip mine approximately 11 km south-west of Rockdale, Milam County,
Texas, U.S.A.; Age: Palaeocene (Rockdale lignite).

692
1.9.18. Species: D. harrisii (C.P. Varma & Rawat) Elsik 1968 (comb. inval.); Index Fungorum
Registration Identifier: 313255; Current name: Foveodiporites harrisii (C.P. Varma &
Rawat) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.9.19. Species: D. incurvus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115640; Location: Cangxian, Hebei Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
1.9.20. Species: D. krempii (C.P. Varma & Rawat) Elsik 1968 (comb. inval.); Index Fungorum
Registration Identifier: 313256; Current name: Biporipsilonites krempii (C.P. Varma &
Rawat) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.9.21. Species: D. major Anil Chandra et al. 1984 (Fig. 5S); Index Fungorum Registration
Identifier: 106782; Location: Sediment core no. 1 (Lat. 17°57.9'N: Long. 70°46.0'E), Arabian
Sea; Age: Late Quaternary.
1.9.22. Species: D. maximus Z.C. Song & H.C. Luo in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 485270; Current name: Biporipsilonites maximus (Z.C. Song & H.C.
Luo in Z.C. Song et al.) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000); Notes:
Kalgutkar & Jansonius (2000) transferred D. maximus to Biporipsilonites because its
holotype has pore chambers at both ends.
1.9.23. Species: D. minutiporatus Hammen 1954; Index Fungorum Registration Identifier: 330111;
Current name: Foveodiporites minutiporatus (Hammen) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000); Notes: Kalgutkar & Jansonius (2000) transferred D. maximus
to Foveodiporites because its holotype has pore chambers at both ends and has foveolate
spore wall.
1.9.24. Species: D. minutus Hammen 1954 (nom. inval.) fide Kalgutkar & Jansonius (2000); Index
Fungorum Registration Identifier: 330112; Location: Eastern Cordilleras, Colombia, South
America; Age: Maastrichtian. Notes: Van der Hammen 1954 did not validly publish the
species because he did not provide any figure of the type, though he published a brief
description.
1.9.25. Species: D. naviculoides (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483323; Basionym: Psiammopomopiospora naviculoides Sal.-Cheb.
& Locq. 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Late Eocene-Oligocene; Notes: Salard-Cheboldaeff & Locquin (1980) suggested affinity of
this genus with Basidiomycota.
1.9.26 Species: D. oblongatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115639; Location: Tanggu, Tianjin Municipality; Kenli and Linyi, Shandong Province,
Coastal Region of Bohai, China; Age: Eocene-Oligocene.
1.9.27. Species: D. pergranulatus Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483324; Basionym: Diporites granulatus Rouse 1962; Location: Terminal Dock,
the city of Vancouver, British Columbia; Age: Late Cretaceous-Middle Eocene (Burrard
Formation).
1.9.28. Species: D. piercei (C.P. Varma & Rawat) Elsik 1968 (comb. inval.); Index Fungorum
Registration Identifier: 313257; Current name: Foveodiporites piercei (C.P. Varma & Rawat)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.9.29. Species: D. pisciculatus G. Norris 1997; Index Fungorum Registration Identifier: 483788;
Current name: Inapertisporites pisciculatus (G. Norris) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.9.30. Species: D. planus Mart.-Hern. & Tom.-Ort. 1989 ex Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483801; Basionym: Diporisporites planus Mart.-Hern. &
Tom.-Ort. 1989 (nom. inval.), lectotype designated by Kalgutkar & Jansonius 2000;
Location: Piedras Negras, Coahuila State, Mexico; Age: Maastrichtian; Notes: Martínez-
Hernández & Tomasini-Ortizin (1989) did not designate a type specimen thus Diporisporites
planus Mart.-Hern. & Tom.-Ort. 1989 was not validly published. Kalgutkar & Jansonius
(2000) designated the lectotype in order to validate the species.

693
1.9.31. Species: D. psilatus P. Kumar 1990 (Fig. 5T); Index Fungorum Registration Identifier:
126555; Location: Padappakkara, Kollam District, Kerala, India; Age: Early-Middle Miocene
(Quilon Beds).
1.9.32. Species: D. reticulatus (C.P. Varma & Rawat) Elsik 1968 (comb. inval.); Index Fungorum
Registration Identifier: 313258; Current name: Striadiporites reticulatus C.P. Varma &
Rawat 1963 fide Kalgutkar & Jansonius (2000).
1.9.33. Species: D. sirmaurensis A. Gupta 2002 (Fig. 5U); Index Fungorum Registration Identifier:
540471; Location: Dadahu Road Section, Sirmaur District, Himachal Pradesh, India; Age:
Eocene (Subathu Formation).
1.9.34. Species: D. varias (Y.K. Mathur) Kalgutkar & Janson. 2000 (Fig. 5V); Index Fungorum
Registration Identifier: 483325; Basionym: Scabradiporites varias Y.K. Mathur 1966;
Location: Matanomadh, western Kutch, Gujarat, India; Age: Palaeocene (Supratrappeans);
Notes: Kalgutkar & Jansonius (2000) considered that Scabradiporites Y.K. Mathur 1966
(Mathur 1966) is superfluous thus the type species, Scabradiporites varias, was transferred to
Diporisporites. This makes Scabradiporites a later synonym of Diporisporites.

1.10. Genus: DREMUSPORA Sal.-Cheb. & Locq., C. r. Congr. natn. Socs. sav. Paris, sect. sci.,
fasc. 1. (Paleobotanique) 105: 191 (1980); Index Fungorum Registration Identifier: 25594; Type:
D. cucurbitaria Sal.-Cheb. & Locq. 1980.
Original Diagnosis: Spores with cells arranged as the parts in a mandarin orange (Salard-
Cheboldaeff & Locquin 1980).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
Notes: Kalgutkar & Jansonius (2000) remarked “The particular structure of the grouped
spores makes the fungal nature of this taxon questionable. The summary diagnosis and description
do not help to clarify its affinity; a reexamination of the holotype is needed to answer these
questions”. From the general organization, the holotype appears to be a fungal spore tetrad.
1.10.1. Species: D. cucurbitaria Sal.-Cheb. & Locq. 1980 (Fig. 5W); Index Fungorum Registration
Identifier: 107941; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa;
Age: Oligocene.

1.11. Genus: EXESISPORITES Elsik, Trans. Gulf Coast Assoc. Geol. Societies 19: 516 (1969);
Index Fungorum Registration Identifier: 21100; Type: E. neogenicus Elsik 1969.
Original Diagnosis: Unicellular, aseptate, psilate, monoporate fungal spores of circular
outline with lenticular to spherical? shape. The centrally located pore in most specimens is
surrounded by a dark circular patch which is interpreted as a thickened wall. This polar area is
occasionally found free of the spore (Elsik 1969).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Four.
Notes: The centrally located pore in Exesisporites is generally surrounded by a dark circular
patch which is interpreted as a thickened wall. Exesisporites is differentiated from
Monoporisporites Hammen 1954, Basidiosporites Elsik 1968 and Lacrimasporonites R.T. Clarke
1965 on the character of the pore or hilum and the orientation of the spores in mounted residues. It
is further differentiated from Reticulatisporonites Elsik 1968 by the lack of reticulate
ornamentation. Glass et al. (1986) cited possible affinity of Exesisporites to the extant fungus
Nigrospora Zimm.
1.11.1. Species: E. annulatus Kalgutkar 1993; Index Fungorum Registration Identifier: 483879;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes:
Kalgutkar (1993) commented that “The spores resemble Exesisporites neogenicus Elsik 1969
in general morphology but show greater variation in size, are somewhat larger, and have a
definite ring around the pore”. The species is named after Latin, annulatus, ring, referring to
the ring of thickening around the pore.

694
1.11.2. Species: E. neogenicus Elsik 1969 (Fig. 5X); Index Fungorum Registration Identifier:
107966; Location: Northern Gulf of Mexico; Age: Miocene-Pleistocene; Notes: This species
is characterized by the presence of somewhat thickened polar area around the minute pore.
1.11.3. Species: E. psilatus R.K. Saxena 2000 (Fig. 5Y); Index Fungorum Registration Identifier:
519806; Location: Mavli Mine at Redi, Sindhudurg District, Maharashtra, India; Age:
Miocene (Sindhudurg Formation).
1.11.4. Species: E. verrucatus P. Kumar 1990 (Fig. 5Z); Index Fungorum Registration Identifier:
126559; Location: Padappakkara, Kollam District, Kerala, India; Age: Early-Middle Miocene
(Quilon Beds); Notes: Exesisporites verrucatus differs from all the known species of this
genus in having hyaline, hollow, rounded verruca-like sculptures at the peripheral margin of
the spore. The species epithet is based on the peripheral verrucae.

1.12. Genus: FOLIOPOLLENITES Sierotin, Sporae dispersae im Rhat und Lias von
Grossbellhofen (Mittelfranken). Inaugural-dissertation (Berlin): 55 (1961); Index Fungorum
Registration Identifier: 25760; Type: F. spinosus Sierotin 1961.
Original Diagnosis: Shaped like a tapered leaf; whole surface is covered with numerous
spines of a little over 1 μm long; exine two layered, a little more than 1 μm thick; surface
chagrinate; size 27 × 21 μm (Sierotin 1961).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Four ((but we accept only three species as legitimate as F.
elongates was not validly published).
Notes: Potonié (1966) quoted a letter from Sierotin that “possibly this is a microfossil that can
be assigned elsewhere” (than in the flowering plants).
1.12.1. Species: F. elongatus Z.C. Song in Z.C. Song et al. 1999 (nom. inval.) fide Kalgutkar &
Jansonius (2000); Index Fungorum Registration Identifier: 483821; Location: China; Age:
Late Cretaceous/Tertiary; Notes: The species name was not validly published because the
author did not specify where the holotype was deposited, and did not provide a Latin
description or English translation. The illustration is that of a smooth, fusiform amerosporae
(28 × 11 µm) with one broadly rounded end, and the other one tapered to a point.
1.12.2. Species: F. qaidamensis Z.C. Song 1985; Index Fungorum Registration Identifier: 637474;
Location: Qigequan, Qaidam Basin, Qinghai Province, China; Age: Late Eocene-Late
Oligocene; Notes: The peculiar echinate ornamentation distinguishes this species from the
other known species of Foliopollenites.
1.12.3. Species: F. spinosus Sierotin 1961 (Fig. 5AA); Index Fungorum Registration Identifier:
114750; Location: Germany; Age: Early Jurassic.
1.12.4. Species: F. taiwanensis T.C. Huang 1981; Index Fungorum Registration Identifier: 115775;
Location: Taiwan; Age: Miocene.

1.13. Genus: FOVEODIPORITES C.P. Varma & Rawat, Grana Palynologica 4(1): 133 (1963);
Index Fungorum Registration Identifier: 21105; Type: F. anklesvarensis C.P. Varma & Rawat
1963.
Synonym: Punctodiporites C.P. Varma & Rawat 1963 fide Kalgutkar & Jansonius (2000),
Index Fungorum Registration Identifier: 28617.
Original Diagnosis: Pollen grains diporate. Exine foveolate (Varma & Rawat 1963).
Emended Diagnosis: Monocellate diporate fungal spores of mostly medium (ca 15–60 μm)
size; overall shape fusiform to elliptic, but characteristically somewhat lob-sided, with one side of
the outline more convex than the other; spore wall relatively thin, externally essentially smooth,
internally smooth, or with punctate, granulate, foveolate or similar sculpture; pores terminal,
complex, consisting of a thin collar and separated from the spore interior by one or two septa (the
latter forming a pore chamber); pore regions often with darker pigmentation (Kalgutkar &
Jansonius 2000).
Classification: Fungi Imperfecti, Amerosporae.

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 Number of species known: Eleven.
Notes: Although originally described as pollen, these forms are now generally recognized as
fungal spores.
1.13.1. Species: F. anklesvarensis C.P. Varma & Rawat 1963 (Fig. 5AB); Index Fungorum
Registration Identifier: 105891; Location: Western and eastern India, including oil
exploration areas in West Bengal and Assam; Age: Early-Middle Eocene, Late Eocene-
Oligocene, Early Miocene; Notes: Kalgutkar & Jansonius (2000) observed that the presence
of punctate or foveolate spore walls in this species is but a minor morphological variation.
Specimens with either punctate or foveolate walls, and with walls showing both types of
ornamentation, were seen (Kalgutkar 1993).
1.13.2. Species: F. bimucronatus (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483363; Basionym: Diporites bimucronatus Sal.-Cheb. &
Locq. 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene.
1.13.3. Species: F. conspicuus (Ramanujam & K.P. Rao) Kalgutkar & Janson. 2000 (Fig. 5AC);
Index Fungorum Registration Identifier: 483364; Basionym: Diporisporites conspicuus
Ramanujam & K.P. Rao 1978; Location: Kannur, Kerala, India; Age: Miocene (Quilon and
Warkalli beds); Notes: The conspicuous pores with atria and the thin, folded spore wall are
the characteristic features of this species.
1.13.4. Species: F. elegans (P. Ke & Z.Y. Shi) Kalgutkar & Janson 2000; Index Fungorum
Registration Identifier: 483365; Basionym: Diporisporites elegans P. Ke & Z.Y. Shi 1978;
Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
1.13.5. Species: F. endogranulosus (Kemp) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483366; Basionym: Diporicellaesporites endogranulosus Kemp 1978;
Location: Deep Sea Drilling Site 254, the Ninetyeast Ridge, Indian Ocean; Age: Late Eocene-
Oligocene; Notes: This species shows close structural similarity to Foveodiporites
anklesvarensis C.P. Varma & Rawat, and for this reason Kalgutkar & Jansonius (2000)
transferred it to Foveodiporites.
1.13.6. Species: F. foedus (G. Norris) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483367; Basionym: Punctodiporites foedus G. Norris 1997; Location: Imperial
ADGO F–28 Well, Mackenzie River delta, Canada; Age: Palaeocene-Eocene; Notes: This
species is named after Latin foedus, foul, horrible.
1.13.7. Species: F. granulatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483368; Basionym: Diporisporites granulatus P. Ke & Z.Y. Shi 1978;
Location: Panshan, Liaoning Province; Kenli, Shandong Province, Coastal region of Bohai,
China; Age: Eocene-Oligocene; Notes: Ke & Shi (1978) stated that further research is needed
to determine whether these fossils are unicellular organisms or fungal spores.
1.13.8. Species: F. gunniae (C.P. Varma & Rawat) Kalgutkar & Janson. 2000 (Fig. 5AD); Index
Fungorum Registration Identifier: 483369; Basionym: Psilodiporites gunniae C.P. Varma &
Rawat 1963; Location: Western and eastern India, including oil exploration areas in West
Bengal and Assam; Age: Middle-Late Eocene; Notes: This species has been named after Mrs.
Gunni Erdtman.
1.13.9. Species: F. harrisii (C.P. Varma & Rawat) Kalgutkar & Janson. 2000 (Fig. 5AE); Index
Fungorum Registration Identifier: 483370; Basionym: Punctodiporites harrisii C.P. Varma &
Rawat 1963; Location: Western and eastern India, including oil exploration areas in West
Bengal and Assam; Age: Early Eocene-Early Oligocene.
1.13.10. Species: F. minutiporatus (Hammen) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483371; Basionym: Diporisporites minutiporatus Hammen 1954;
Location: Magdalena Valley, Eastern Cordellera, Colombia, South America. Age:
Maastrichtian.

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1.13.11. Species: F. piercei (C.P. Varma & Rawat) Kalgutkar & Janson. 2000 (Fig. 5AF); Index
Fungorum Registration Identifier: 483372; Basionym: Granodiporites piercei C.P. Varma &
Rawat 1963; Location: Western and eastern India, including oil exploration areas in West
Bengal and Assam; Age: Middle-Late Eocene.

1.14. Genus: FUSIDIPOROSPORONITES Z.C. Song in Z.C. Song et al., Early Tertiary Sporo-
Pollen Assemblages from the Dongpu Region: 43, pl. 4, fig. 18, (1989); Index Fungorum
Registration Identifier: 637475; Type: F. minutaestriatus Z.C. Song in Z.C. Song et al. 1989.
Original Diagnosis: One-celled spore, narrowly fusiform in outline, tapering to form a short
tube at each terminal end; two pores, situated on the tips of the short tubes; pore simple,
unstructured, exine (i.e. spore wall) moderately thick, ornamentation various, but commonly thin
and weak (Song et al. 1989).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
Notes: According to Song et al. (1989), ornamentation and shape of the type species,
Fusidiporosporonites minutaestriatus, is similar to that in Lacrimasporonites minutaestriatus P. Ke
& Z.Y. Shi 1978 [Current name: Monoporisporites minutaestriatus (P. Ke & Z.Y. Shi) Kalgutkar
& Janson. 2000], but the latter has only one pore and one short terminal tube. Fusidiporosporonites
minutaestriatus is characterized by numerous transverse, fine striae. This feature separates it from
Striadiporites, which is characterized by longitudinal ridges (that may form a reticulum). This
genus can also be distinguished from Fusiformisporites on the basis of the transverse striation, and
the more elongated terminal tubes.
1.14.1. Species: F. minutaestriatus Z.C. Song in Z.C. Song et al. 1989 (Fig. 5AG); Index
Fungorum Registration Identifier: 637476; Location: Shenxian county of Shandong Province,
China; Age: Late Eocene-Middle Oligocene (Shahejie Formation).

1.15. Genus: GEOTRICHITES Stubblef. et al., Mycologia 77(1): 12 (1985); Index Fungorum
Registration Identifier: 25685; Type: G. glaesarius Stubblef. et al. 1985.
Original Diagnosis: Hyphae aerial, erect or decumbent, septate, sometimes branching
dichotomously. Conidiophores micronematous. Conidia holoarthric, aseptate, produced by
schizolytic disarticulation, variable in size, oblong or truncate with obtuse ends (Stubblefield et al.
1985).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
Notes: According to Stubblefield et al. (1985), Geotrichites conforms to the extant genus
Geotrichum Link ex Pers. in most aspects, although it is impossible to determine whether the
mycelium is dry or slimy in the fossilized state.
1.15.1. Species: G. glaesarius Stubblef. et al. 1985 (Fig. 5AH); Index Fungorum Registration
Identifier: 105328; Location: Dominican Republic; Age: Late Oligocene or Early Miocene.

1.16. Genus: GRAAMSPORA Sal.-Cheb. & Locq., C. r. Congr. natn. Socs. sav. Paris, sect. sci.,
fasc. 1. (Paleobotanique) 105: 191 (1980); Index Fungorum Registration Identifier: 25590; Type:
G. pezizaffinis Sal.-Cheb. & Locq. 1980; Current name: UNCINULITES Pampal. 1902 fide
Kalgutkar & Jansonius (2000).
Original Diagnosis: Amerospores, more or less verrucose-spinose; approximately 20 × 15 μm
(Salard-Cheboldaeff & Locquin 1980 fide Jansonius & Hills 1981)
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One (the single species has been transferred to Uncinulites Pampl.
1902).
1.16.1. Species: G. pezizaffinis Sal.-Cheb. & Locq. 1980, p. 184, pl. 1, fig. 3. Current name:
Uncinulites pezizaffinis (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).

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1.17. Genus: GRANODIPORITES C.P. Varma & Rawat, Grana Palynologica 4(1): 135 (1963);
Type: G. erdtmanii C.P. Varma & Rawat 1963, p. 135, fig. 13. Current name:
BANKSIEAEIDITES Cookson 1950 (angiosperm pollen) fide Kalgutkar & Jansonius (2000).
Original Diagnosis: Diporate pollen grains with granular exine (Varma & Rawat 1963).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Three (the type species, G. erdtmanii C.P. Varma & Rawat 1963,
belongs to angiospermous pollen whereas the remaining two species have been transferred to other
fungal spore genera).
Notes: Since the type species (G. erdtmanii) has been transferred to Banksieaeidites Cookson
(an angiospermous pollen) by Kalgutkar & Jansonius (2000), Granodiporites became a later
synonym of Banksieaeidites. The remaining two fungal species have been transferred to other
genera.
1.17.1. Species: G. erdtmanii C.P. Varma & Rawat 1963; Index Fungorum Registration Identifier:
519816; Current name: Banksieaeidites erdtmanii (C.P. Varma & Rawat) Kalgutkar &
Janson. 2000 (angiospermous pollen) fide Kalgutkar & Jansonius (2000).
1.17.2. Species: G. piercei C.P. Varma & Rawat 1963; Index Fungorum Registration Identifier:
483374; Current name: Foveodiporites piercei (C.P. Varma & Rawat) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
1.17.3. Species: G. sahnii C.P. Varma & Rawat 1963; Index Fungorum Registration Identifier:
483375; Current name: Dyadosporites sahnii (C.P. Varma & Rawat) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).

1.18. Genus: GRAPHIOLITES Fritel, Mém. Soc. geol. Fr. Paleont. 40: 12 (1910); Index
Fungorum Registration Identifier: 21122; Type: G. sabaleos Fritel 1910.
Original Diagnosis: In this genus are included fossil forms of which the external features and
habitat agree with those of the extant Graphiola (Exobasidiales,Basidiomycota), i.e. fungi
specialized in parasitizing palm fronds, where they form peridia consisting of deep, round or oval
cupules; these cupules may occur dispersed, or be arranged in longitudinal series parallel to the
nervature of the fronds on which they grow. When the peridia are oval, the longer axis is always
aligned longitudinally (Fritel 1910).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
Notes: Fritel (1910), for the first time, reported fossil spores of Graphiola and placed them in
a new Graphiolites. Kalgutkar & Jansonius (2000) stated that Graphiola was assigned to Fungi
Imperfecti (hyphomycetes, Amerosporae) by Poiteau (1824). Hughes (1953) recognized that this
genus currently is considered to be in the Basidiomycota (Graphiolaceae, Ustilaginales). However,
he also stated that this position generally is regarded as dubious.
1.18.1. Species: G. sabaleos Fritel 1910 (Fig. 5AI); Index Fungorum Registration Identifier:
101378; Location: Cessoy, Seine-et-Marne, France; Age: Sparnacian (Early Eocene).

1.19. Genus: HAPLOGRAPHITES Félix, Zeitschr. Deutsche Geol. Gesell. 46: 275 (1894); Index
Fungorum Registration Identifier: 21125; Type: H. cateniger Felix 1894 (lectotype was designated
by Jansonius & Hills 1976).
Original Diagnosis: Individual conidia average 15–17 μm in length, 9–11 (but up to 14) μm
in width; colour mostly dark amber-brown, occasionally lighter; cell wall very thick (Felix 1894).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Two.
Notes: Felix (1894) proposed Haplographites to accommodate fossil fungal spores showing
similarity to spores of the living representatives of the grouping Haplographieae of Saccardo
(particularly those of Haplographium Berk. & Broome and Dematium Pers.) that their affinity to
this grouping appears probable.

698
1.19.1. Species: H. cateniger Félix 1894 (Fig. 6A); Index Fungorum Registration Identifier:
232186; Location: Perekeschkul, near Baku, Azerbaijan; Age: Eocene.
1.19.2. Species: H. xylophagus Félix 1894; Index Fungorum Registration Identifier: 218003;
Location: Tarnow, Galicia; Age: Tertiary.

1.20. Genus: HYPOXYLONITES Elsik, Palaeontographica Abt. B 216(5-6): 143 (1990a), Index
Fungorum Registration Identifier: 25556; Type: H. brazosensis Elsik 1990a.
Synonyms: Hypoxylonsporites P. Kumar 1990 fide Saxena 1992, Index Fungorum
Registration Identifier: 25448; Xylariasporites Debi Mukh. 2012 fide hoc loco, Index Fungorum
Registration Identifier: 588470.

Figure 5 – A–AI Amerosporae. A Asyregraamspora reticulata Sal.-Cheb. & Locq. 1980, Bar = 8
μm. B Basidiosporites fournieri Elsik 1968. 1980, Bar = 8 μm. C Basidiosporites ovalis (Sheffy &

699
Dilcher) Kalgutkar & Janson. 2000. 1980, Bar = 10 μm. D Basidiosporites sadasivanii Anil
Chandra et al. 1984, Bar = 10 μm. E Biporipsilonites belluloides (Z.C. Song) Kalgutkar & Janson.
2000, Bar = 8 μm. F Biporipsilonites fusiformis (Anil Chandra et al.) Kalgutkar & Janson. 2000,
Bar = 20 μm. G Biporipsilonites karii (Anil Chandra et al.) Kalgutkar & Janson. 2000, Bar = 5 μm.
H Biporipsilonites krempii (C.P. Varma & Rawat) Kalgutkar & Janson. 2000, Bar = 20 μm. I
Biporipsilonites padappakkarensis (P. Kumar) Kalgutkar & Janson. 2000, Bar = 20 μm. J
Biporisporites rotundus P. Ke & Z.Y. Shi 1978, Bar = 8 μm. K Cadyexinis vulgaris Stach 1957,
Bar = 40 μm. L Cervichlamydospora nigra R. Kar et al. 2010, Bar = 10 μm. M Diporisporites
barrelis A. Gupta 2002, Bar = 10 μm. N Diporisporites bhavnagarensis R.K. Saxena 2009, Bar =
20 μm. O Diporisporites elongatus Hammen 1954, Bar = 10 μm. P Diporisporites elsikii R.K.
Saxena 2000, Bar = 30 μm. Q Diporisporites giganticus R.K. Kar 2012, Bar = 35 μm. R
Diporisporites hammenii Elsik 1968, Bar = 5 μm. S Diporisporites major Anil Chandra et al. 1984,
Bar = 10 μm. T Diporisporites psilatus P. Kumar 1990, Bar = 20 μm. U Diporisporites
sirmaurensis A. Gupta 2002, Bar = 10 μm. V Diporisporites varias (Y.K. Mathur) Kalgutkar &
Janson. 2000, Bar = 7 μm. W Dremuspora cucurbitaria Sal.-Cheb. & Locq. 1980, Bar = 8 μm. X
Exesisporites neogenicus Elsik 1969, Bar = 12 μm. Y Exesisporites psilatus R.K. Saxena 2000, Bar
= 20 μm. Z Exesisporites verrucatus P. Kumar 1990, Bar = 10 μm. AA Foliopollenites spinosus
Sierotin 1961, Bar = 12 μm. AB Foveodiporites anklesvarensis C.P. Varma & Rawat 1963, Bar =
10 μm. AC Foveodiporites conspicuus (Ramanujam & K.P. Rao) Kalgutkar & Janson. 2000, Bar =
10 μm. AD Foveodiporites gunniae (C.P. Varma & Rawat) Kalgutkar & Janson. 2000, Bar = 10
μm. AE Foveodiporites harrisii (C.P. Varma & Rawat) Kalgutkar & Janson. 2000, Bar = 10 μm.
AF Foveodiporites piercei (C.P. Varma & Rawat) Kalgutkar & Janson. 2000, Bar = 10 μm. AG
Fusidiporosporonites minutaestriatus Z.C. Song 1989, Bar = 12 μm. AH Geotrichites glaesarius
Stubblef. et al. 1985, Bar = 5 μm. AI Graphiolites sabaleos Fritel 1910, Bar = 5 μm.

Original Diagnosis: Oval to elongate, aseptate, bilateral, psilate fungal spores bearing an
elongate scar, slit or furrow. At the level of transmitted light microscopy, at least one species is
faintly sculptured. The elongate furrow is parallel to the axis and can be of various lengths. Apices
rounded to pointed; usually of similar shape but some species have an attachment scar at one end;
apices can also be thickened or otherwise modified. The spore wall in most specimens is generally
rigid (Elsik 1990a).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: 55 [but we accept only 53 species as legitimate because H.
elongatus (Rouse) Elsik 1990a has been transferred to Inapertisporites Hammen 1954, and H.
elongatus Sal.-Cheb. & Locq. 1980 (nom. inval.) is considered to be a later taxonomic synonym of
Hypoxylonites africanus Sal.-Cheb. & Locq. ex Kalgutkar & Janson. 2000]).
Notes: Hypoxylonites Elsik 1990a is characterized by one elongate scar or furrow which is
straight and parallel to the long axis of the spore. The Hypoxylonites morphotype is possibly
produced by many living fungi; most are in Xylariaceae (Ascomycota). The spore type is also
produced by a few hyphomycetes. Elsik (1990a) cited a number of papers giving further details on
possible affinities, and comparisons with similar spores produced by extant fungal species
(Kalgutkar & Jansonius 2000). Salard-Cheboldaeff & Locquin (1980) assigned three species (viz.
H. ellipsoideus, H. elongatus and H. xylarioides) to the generic name “Hypoxylonites”, but did not
provide a diagnosis for this genus. Thus, the generic name was not validly published in Salard-
Cheboldaeff & Locquin 1980, and neither were the species then assigned to it. Kumar (1990)
proposed Hypoxylonsporites (Type: H. miocenicus) with the following diagnosis “Fungal spores
unicellular, oval to ellipsoidal 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.”
Hypoxylonites Elsik (March 1990) and Hypoxylonsporites Kumar (May 1990) are identical in all
essential characters and therefore the latter is a later taxonomic synonym of Hypoxylonites. The
above observation was made by Saxena (1992) who transferred the species described under
Hypoxylonsporites Kumar to Hypoxylonites Elsik. Both of these genera have affinity with the

700
extant Hypoxylon of the family Xylariaceae. The genus is named after its resemblance to the
ascospores of extant Hypoxylon (Bull.) Fries.
1.20.1. Species: H. africanus Sal.-Cheb. ex Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483399; Basionym: Hypoxylonites elongatus Sal.-Cheb. & Locq.
1980 (nom. inval.); Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa;
Age: Late Eocene-Oligocene-Early Miocene.
1.20.2. Species: H. armentroutii Elsik 1990a; Index Fungorum Registration Identifier: 130348;
Location: The Green River Section, Washington, U.S.A.; Age: Late Eocene; Notes: The
species epithet is in honour of John M. Armentrout, Mobil Technology Company, Dallas,
Texas, U.S.A.
1.20.3. Species: H. asymetricus (Sal.-Cheb. & Locq.) Elsik 1990a; Index Fungorum Registration
Identifier: 130342; Basionym: Inapertisporites asymetricus Sal.-Cheb. & Locq. 1980;
Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age: Early Eocene-
Early Miocene (Salard-Cheboldaeff & Locquin 1980); Early Eocene (Elsik 1990a); Notes:
Salard-Cheboldaeff & Locquin (1980) assigned its affinity with Ascomycota. Elsik (1990a)
published an emended description, as follows: “Aseptate, psilate fungal spores ca. 12 × 24
μm overall. Outline reniform in side view, one side less convex and almost straight. Outline
elliptical in top view, ends narrowly rounded. Spore wall ca. 0.5 µm at the ends of the spore,
ca. 0.5–1.0 µm elsewhere except perhaps along the straighter side. The originally described
spore wall of two layers is an expression of the overlapping edges of the furrow, which
traverses most of the length of the more convex side of the spore”.
1.20.4. Species: H. ater (P. Kumar) R.K. Saxena 1992 (Fig. 6B); Index Fungorum Registration
Identifier: 483400; Basionym: Hypoxylonsporites ater P. Kumar 1990; Synonym:
Hypoxylonites ater (P. Kumar) Kalgutkar & Janson. 2000 fide Saxena & Tripathi 2011;
Location: Padappakkara, Kollam District, Kerala, India; Age: Early-Middle (Quilon Beds).
1.20.5. Species: H. bhubanensis Nandi & Subhra Banerjee in R.K. Saxena 2012 (Fig. 6C); Index
Fungorum Registration Identifier: 519748; Basionym: Hypoxylonites bhubanensis Nandi &
Subhra Banerjee in Nandi et al. 2003; Location: Renkte Kawn-Sherlui Road, Mizoram, India;
Age: Neogene (Bhuban, Bokabil Tipam and Dupitila formations); Notes: Nandi & Banerjee
in Nandi et al. (2003) did not validly publish Hypoxylonites bhubanensis because they did not
mention where the holotype was deposited. Saxena (2012) validated it by providing the
missing validating information.
1.20.6. Species: H. brazosensis Elsik 1990a (Fig. 6D); Index Fungorum Registration Identifier:
130349; Location: Brazos County, Texas, U.S.A.; Age: Late Middle Eocene (Yegua
Formation); Notes: Elsik (1990a) differentiated this species on the basis of slightly
asymmetrical shape and the longitudinal furrow reaching the apices along the more convex
side of the spore. The species epithet is derived from Brazos River, Brazos County, Texas.
1.20.7. Species: H. chaiffetzii Elsik 1990a (Fig. 6E); Index Fungorum Registration Identifier:
130350; Location: The Gulf Coast, U.S.A.; Age: Neogene; Notes: The species epithet is in
honour of Michael Chaiffetz.
1.20.8. Species: H. chuittensis Elsik 1990a; Index Fungorum Registration Identifier: 130351;
Location: Southern Alaska; Age: Neogene; Notes: The species epithet is derived from its
occurrence in strata penetrated by the Superior No. 1, Chuitt River, Alaska.
1.20.9. Species: H. claibornensis Elsik 1990a; Index Fungorum Registration Identifier: 130352;
Location: Claiborne Group, Henry County, Tennessee, U.S.A.; Age: Middle Eocene; Notes:
The species epithet is derived from its occurrence in Claiborne Group.
1.20.10. Species: H. curvatus (Ramanujam & K.P. Rao) Elsik 1990a (Fig. 6F); Index Fungorum
Registration Identifier: 130343; Basionym: Diporisporites curvatus Ramanujam & K.P. Rao
1978; Location: Alleppey, Alappuzha District, Kerala, India; Age: Miocene (Quilon and
Warkalli beds).

701
1.20.11. Species: H. disciformis (Sheffy & Dilcher) R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde
& P.M. Kirk comb. nov. (Fig. 6G). This new combination is described under the section
“New species and new combinations”.
1.20.12. Species: H. edigeri Elsik 1990a; Index Fungorum Registration Identifier: 130353;
Location: Turkey; Age: Late Eocene or Oligocene; Notes: The species epithet is in honour of
V.S. Ediger. Turkish Petroleum Corporation, Research Center, Ankara, Turkey.
1.20.13. Species: H. ellipsoideus Sal.-Cheb. & Locq. ex Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 108077; Location: Coast of Equatorial Africa, Gulf of
Guinea, Cameroon, Africa; Age: Late Eocene-Oligocene-Early Miocene; Notes: Salard-
Cheboldaeff & Locquin (1980) assigned affinity of this species to Ascomycota.
1.20.14. Species: H. elongatioides Elsik 1990a; Index Fungorum Registration Identifier: 130354;
Location: Tennessee, U.S.A.; Age: Middle Eocene; Notes: The species epithet is derived
from its similarity to Hypoxylonites elongatus (Rouse) Kalgutkar & Janson. 2000.
1.20.15. Species: H. elongatus (Rouse) Elsik 1990a; Index Fungorum Registration Identifier:
130344; Current name: Inapertisporites elongatus Rouse 1962 fide Kalgutkar & Jansonius
(2000).
1.20.16. Species: H. elongatus Sal.-Cheb. & Locq. 1980 (nom. inval.) fide Kalgutkar & Jansonius
(2000); Index Fungorum Registration Identifier: 107682; Current name: Hypoxylonites
africanus Sal.-Cheb. & Locq. ex Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius
(2000).
1.20.17. Species: H. elsikii Nandi & Shubhra Banerjee in R.K. Saxena 2012 (Fig. 6H); Index
Fungorum Registration Identifier: 519749; Synonym: Hypoxylonites elsikii Nandi & Shubhra
Banerjee in Nandi et al. 2003; Location: Renkte Kawn-Sherlui Road, Mizoram, India; Age:
Tertiary (Siju, Kherapara, Bhuban, Bokabil and Dihing formations).
1.20.18. Species: H. eocenicus Elsik 1990a (Fig. 6I); Index Fungorum Registration Identifier:
130355; Location: The Green River Section, Washington, U.S.A.; Age: Late Eocene; Notes:
The species epithet is after its occurrence in Eocene strata.
1.20.19. Species: H. eopleistocenicus Elsik 1990a; Index Fungorum Registration Identifier:
130356; Location: The Gulf Coast, U.S.A.; Age: Late Miocene; Notes: The species epithet is
derived from its similarity to Hypoxylonites pleistocenicus.
1.20.20. Species: H. felixii Elsik 1990a (Fig. 6J); Index Fungorum Registration Identifier: 130357;
Location: Mid Creek Section, Bristol Bay, Alaska; Age: Miocene; Notes: The species epithet
is in memory of Johannes Felix.
1.20.21. Species: H. foldexinus Elsik 1990a; Index Fungorum Registration Identifier: 130358;
Location: The Gulf Coast, U.S.A.; Age: Neogene; Notes: The species epithet is derived from
the folds of the spore wall.
1.20.22. Species: H. foyelensis Bianchin., Alej. Martínez & R.K. Saxena in Alej. Martínez et al.
2016; Index Fungorum Registration Identifier: 812333; Location: Rio Foyel section, Nirihuau
Basin, Argentina; Age: Palaeogene (El Foyel Group); Notes: The specific epithet is after the
El Foyel Group, from which samples were collected.
1.20.23. Species: H. fusiformis Elsik 1990a (Fig. 6K); Index Fungorum Registration Identifier:
130359; Location: The Gulf Coast, U.S.A.; Age: Neogene; Notes: The specific epithet is after
its fusiform outline.
1.20.24. Species: H. gulfensis Elsik 1990a (Fig. 6L); Index Fungorum Registration Identifier:
130360; Location: The Gulf Coast, U.S.A.; Age: Neogene; Notes: The specific epithet is after
the Gulf, i.e. Gulf of Mexico.
1.20.25. Species: H. horowitzii Elsik 1990a; Index Fungorum Registration Identifier: 130361;
Location: Borehole Ashquelon 2, at the depth of 1685 m; Israel; Age: Neogene; Notes: The
specific epithet is in honour of Aharon Horowitz.
1.20.26. Species: H. kumarii Kalgutkar & Janson. 2000 (Fig. 6M); Index Fungorum Registration
Identifier: 483402; Basionym: Hypoxylonsporites miocenicus Kumar 1990; Location:
Padappakkara, Kollam District, Kerala, India; Age: Lower-Middle Miocene (Quilon Beds).

702
1.20.27. Species: H. lammonsii Elsik 1990a; Index Fungorum Registration Identifier: 130362;
Location: The Gulf Coast, U.S.A.; Age: Neogene; Notes: The specific epithet is in honour of
J.M. Lammons.
1.20.28. Species: H. lanceolatus (Debi Mukh.) R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde &
P.M. Kirk comb. nov. (Fig. 6N). This new combination is described under the section "New
species and new combinations".
1.20.29. Species: H. lineatus Elsik 1990a; Index Fungorum Registration Identifier: 130363;
Location: The Gulf Coast, U.S.A.; Age: Neogene; Notes: The specific epithet is after the long
straight furrow, i.e. like a line.
1.20.30. Species: H. magnus Elsik 1990a (Fig. 6O); Index Fungorum Registration Identifier:
130364; Location: Socony-Vacuum Borehole no. 2, Isla de Cubagua, Venezuela; Age:
Middle Miocene; Notes: The specific epithet is for its large size.
1.20.31. Species: H. megaexinus Elsik 1990a; Index Fungorum Registration Identifier: 130365;
Location: The Gulf Coast, U.S.A.; Age: Late Miocene; Notes: The specific epithet is after the
relative thickness of the spore wall.
1.20.32. Species: H. minimus Nandi & Subhra Banerjee in R.K. Saxena 2012 (Fig. 6P); Index
Fungorum Registration Identifier: 519750; Basionym: Hypoxylonites minimus Nandi &
Subhra Banerjee in Nandi et al. 2003 (nom. inval.); Location: Renkte Kawn-Sherlui Road,
Mizoram, India; Age: Tertiary (Siju, Bhuban, Bokabil, Tipam and Dupitila formations);
Notes: The specific epithet indicates small size of spores.
1.20.33. Species: H. minutus Elsik 1990a; Index Fungorum Registration Identifier: 130366;
Location: The Gulf Coast, U.S.A.; Age: Late Miocene.
1.20.34. Species: H. miocenicus Elsik 1990a (Fig. 6Q); Index Fungorum Registration Identifier:
130367; Location: The Gulf Coast, U.S.A.; Age: Late Miocene; Notes: The specific epithet
indicates its occurrence in the Miocene.
1.20.35. Species: H. neogenicus Nandi & Subhra Banerjee in R.K. Saxena 2012 (Fig. 6R); Index
Fungorum Registration Identifier: 519753; Basionym: Hypoxylonites neogenicus Nandi &
Subhra Banerjee in Nandi et al. 2003; Location: Renkte Kawn-Sherlui Road, Mizoram, India;
Age: Neogene (Bhuban, Bokabil, Tipam, Dupitila and Dihing formations).
1.20.36. Species: H. oblongus Elsik 1990a; Index Fungorum Registration Identifier: 130368;
Location: The Sespe sand facies offshore California, U.S.A.; Age: Late Eocene; Notes: The
specific epithet indicates its oblong shape.
1.20.37. Species: H. ovalis Elsik 1990a (Fig. 6S); Index Fungorum Registration Identifier: 130369;
Location: The Green River Section, Washington, U.S.A.; Age: Eocene; Notes: The specific
epithet indicates its oval shape.
1.20.38. Species: H. ovaloides Elsik 1990a; Index Fungorum Registration Identifier: 130370;
Location: Strata in the Socony-Vacuum Borehole # 2, Isla de Cubagua, Venezuela Age:
Middle Eocene; Notes: The specific epithet indicates its similarity to Hypoxylonites ovalis.
1.20.39. Species: H. pirozynskii Elsik 1990a; Index Fungorum Registration Identifier: 130371;
Location: The Gulf Coast, U.S.A.; Age: Neogene Notes: The specific epithet is in honour of
Kris A. Pirozynski.
1.20.40. Species: H. pirozynskioides Elsik 1990a; Index Fungorum Registration Identifier: 130372;
Location: Claiborne Group of Henry County, Tennessee, U.S.A., Also reworked into
Neogene sediments offshore Louisiana, U.S.A.; Age: Middle Eocene; Notes: The specific
epithet indicates its similarity to Hypoxylonites pirozynskii.
1.20.41. Species: H. pleistocenicus Elsik 1990a; Index Fungorum Registration Identifier: 130373;
Location: Beaumont Clay, Harris County, Texas, U.S.A.; Age: Late Pleistocene; Notes: The
specific epithet indicates its occurrence in Pleistocene sediments.
1.20.42. Species: H. pulvinatus (Sheffy & Dilcher) Elsik 1990a; Index Fungorum Registration
Identifier: 130345; Basionym: Inapertisporites pulvinatus Sheffy & Dilcher 1971; Location:
Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.(Sheffy & Dilcher
1971); Northern Thrace Basin, Turkey (Ediger & Alisan 1989); Age: Middle Eocene

703
 (Claiborne Formation) (Sheffy & Dilcher 1971); Middle?-Late Eocene to Late Oligocene,
Miocene-Pliocene (Ediger & Alisan 1989); Notes: The diagnosis of this species was emended
by Ediger & Alisan (1989) and Elsik (1990a).
1.20.43. Species: H. ramanujamii Elsik 1990a; Index Fungorum Registration Identifier: 130374;
Location: Possibly reworked in the Gulf Coast, U.S.A.; Green River section, Washington,
U.S.A.; Age: Neogene of the Gulf Coast, U.S.A.; Late Eocene of the Green River section,
U.S.A.; Notes: The specific epithet is in honour of C.G.K. Ramanujam, Department of
Botany, University College of Science, Saifabad, Osmania University, Hyderabad, India
1.20.44. Species: H. subrotundus Nandi & Subhra Banerjee in R.K. Saxena 2012 (Fig. 6T); Index
Fungorum Registration Identifier: 519752; Basionym: Hypoxylonites subrotundus Nandi &
Subhra Banerjee in Nandi et al. 2003; Location: Renkte Kawn-Sherlui Road, Mizoram, India;
Age: Tertiary (Siju, Bhuban, Bokabil, Tipam and Dupitila formations).
1.20.45. Species: H. subuliformis Elsik 1990a (Fig. 6U); Index Fungorum Registration Identifier:
130375; Location: Strata in the Socony-Vacuum Borehole No. 2, Isla de Cubagua,
Venezuela; Age: Miocene; Notes: The specific epithet is after Latin subula = awl; and forma
= shape.
1.20.46. Species: H. sulekii Elsik 1990a (Fig. 6V); Index Fungorum Registration Identifier:
130376; Location: Socony-Vacuum Borehole no. 2, Isla de Cubagua, Venezuela; the Gulf
Coast, U.S.A., The Pacific Coast, U.S.A.; Age: Middle Miocene strata in Venezuela;
Neogene of the Gulf Coast, U.S.A.; Neogene of the Pacific Coast, from northern California to
offshore Baja California, U.S.A.; Notes: The specific epithet is in honour of John A. Sulek.
1.20.47. Species: H. tennesseensis Elsik 1990a; Index Fungorum Registration Identifier: 130377;
Location: The Gulf Coast and the Mississipi Embayment, U.S.A.; Age: Middle Eocene;
Notes: The specific epithet indicates its occurrence in Tennessee.
1.20.48. Species: H. thindii Nandi & A. Sinha in R.K. Saxena 2012 (Fig. 6W); Index Fungorum
Registration Identifier: 519751; Basionym: Hypoxylonites thindii Nandi & A. Sinha in Nandi
et al. 2003; Location: Renkte Kawn-Sherlui Road, Mizoram, India; Age: Tertiary (Siju,
Kherapara, Bhuban, Bokabil, Tipam, Dupitila and Dihing formations).
1.20.49. Species: H. truncatus Elsik 1990a; Index Fungorum Registration Identifier: 130378;
Location: The Gulf Coast, U.S.A.; Age: Neogene; Notes: The specific epithet indicates its
truncated outline.
1.20.50. Species: H. vicksburgensis Elsik 1990a; Index Fungorum Registration Identifier: 130379;
Location: South Texas, U.S.A.; Age: Oligocene (Vicksburg Group); Notes: The specific
epithet indicates its occurrence in the Vicksburg Group.
1.20.51. Species: H. vittatoides Elsik 1990a (Fig. 6X); Index Fungorum Registration Identifier:
130380; Location: Imperial Nuktak C–22 well, Mackenzie River delta, Mackenzie District,
Northwest Territories, Canada; Age: Eocene; Notes: The holotype is Inapertisporites sp. cf. I.
vittatus Sheffy & Dilcher 1971 in Norris 1986: 18, pl. 1, fig. 1. The specific epithet indicates
its resemblance to Hypoxylonites vittatus Sheffy & Dilcher 1971.
1.20.52. Species: H. vittatus (Sheffy & Dilcher) Elsik 1990a; Index Fungorum Registration
Identifier: 130346; Basionym: Inapertisporites vittatus Sheffy & Dilcher 1971; Location:
Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.; Age: Middle
Eocene (Claiborne Formation); Notes: Elsik (1990a) published an emended description of
this species. The specific name refers to the opaque band formed by one layer of the cell wall.
1.20.53. Species: H. washingtonensis Elsik 1990a; Index Fungorum Registration Identifier:
130381; Location: The Green River section, Washington, U.S.A.; Age: Late Eocene; Notes:
The specific epithet indicates its occurrence in Washington.
1.20.54. Species: H. wolfei Elsik 1990a; Index Fungorum Registration Identifier: 130382;
Location: The Green River section, Washington, U.S.A.; Brazos County, Texas, U.S.A.; Age:
Late Eocene of the Green River section; earliest Late Eocene of Caddell Formation of Texas,
U.S.A.; Notes: The specific epithet is in honour of Jack A. Wolfe.

704
1.20.55. Species: H. xylarioides Sal.-Cheb. & Locq. ex Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483403; Basionym: Hypoxylonites xylarioides Sal.-Cheb.
& Locq. 1980 (nom. inval.); Location: Coast of Equatorial Africa, Gulf of Guinea,
Cameroon, Africa; Age: Late Eocene-Oligocene-Early Miocene; Notes: Salard-Cheboldaeff
(1980) assigned its affinity with Ascomycota (Hypoxylon).

1.21. Genus: HYPOXYLONSPORITES P. Kumar, Review of Palaeobotany & Palynology

(Amsterdam) 63(1–2): 18 (1990); Index Fungorum Registration Identifier: 25448, Type: H.
miocenicus Kumar 1990; Current name: HYPOXYLONITES Elsik 1990a fide Saxena (1992).
Original 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 (Kumar 1990).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Two (since Hypoxylonsporites P. Kumar 1990 is a later synonym
of Hypoxylonites Elsik 1990a, both the species have been transferred to Hypoxylonites Elsik
1990a).
Notes: Hypoxylonsporites shows close affinity with the extant Hypoxylon and Endocalyx.
Similar specimens have been reported earlier from subsurface Tertiary sediments of Cauvery Basin,
India (Venkatachala & Rawat 1973), Neogene sediments of Rusizi Valley, Burundi (Sah 1967),
Middle Eocene clay beds in the Lawrence Clay Pit, Henry County, Tennessee, U.S.A. (Elsik &
Dilcher 1974). Elsik (1969) figured similar spores from the Late Neogene deposits in the northern
Gulf of Mexico and referred to them as Hypoxylon spp. Elsik’s (March 1990) paper has priority
over that of Kumar (May 1990), and hence Hypoxylonsporites is a later taxonomic synonym of
Hypoxylonites fide Saxena 1992.
1.21.1. Species: H. ater P. Kumar 1990; Index Fungorum Registration Identifier: 126561; Current
name: Hypoxylonites ater (P. Kumar) R.K. Saxena 1992 fide Saxena (1992).
1.21.2. Species: H. miocenicus P. Kumar 1990; Index Fungorum Registration Identifier: 126562;
Current name: Hypoxylonites kumarii Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius
(2000).

1.22. Genus: IMPRIMOSPORA G. Norris, Bull. geol. Suev. Can: 22 (1986); Index Fungorum
Registration Identifier: 25449; Type: I. tankensis G. Norris 1986; Current name:
PALAEOAMPHISPHAERELLA Ramanujam & Srisailam 1980 fide Kalgutkar & Jansonius
(2000).
Original Diagnosis: Unicellate, ovoidal, equilateral, isopolar fungal spores with a central
region of parallel striae or fissures parallel to the apical line. Pore or furrow present near one end of
spore (Norris 1986).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Two (both the species have been transferred to other genera).
Notes: Imprimospora is identical to Palaeoamphisphaerella Ramanujam & Srisailam 1980 in
diagnostic and general morphology and therefore is considered to be a later synonym of
Palaeoamphisphaerella.
1.22.1. Species: I. ramanujamii P. Kumar 1990; Index Fungorum Registration Identifier: 126563;
Current name: Kumarisporites ramanujamii (P. Kumar) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.22.2. Species: I. tankensis G. Norris 1986; Index Fungorum Registration Identifier: 126574;
Current name: Palaeoamphisphaerella tankensis (G. Norris) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).

1.23. Genus: INAPERTISPORITES Hammen, Bol. Geol. (Bogota) 2(1): 83 (1954); Index
Fungorum Registration Identifier: 560987; Type: I. variabilis Hammen 1954 (lectotype was
designated by Van der Hammen 1955).

705
 Synonyms: Inapertisporites Rouse 1959 (also a later homonym), Index Fungorum
Registration Identifier: 21139; Triporisporonites Sheffy & Dilcher 1971, Index Fungorum
Registration Identifier: 21346.
Original Diagnosis: Fungal spores without performed aperture (Van der Hammen 1954).
Emended Diagnoses: Diagnosis of Inapertisporites was emended by Rouse (1959), Elsik
(1968), Sheffy & Dilcher (1971), Ediger (1981) and Saxena & Bhattacharyya (1987), as follows:
Spores free or grouped, anisopolar and inaperturate. Shape circular, oval or elliptical; outline often
uneven because of wrinkles or folds. Ornamentation variable, ranging from laevigate to
pseudoreticulate. Size-range 5–100 µm (Rouse 1959); Inaperturate, psilate, fungal spores. One cell,
no septa. Shape variable (Elsik 1968); Fungal or algal spores unicellate, nonseptate, and
inaperturate. Shape globular or non-globular; outline smooth or often uneven because of wrinkles
or folds. Ornamentation variable. Size range 5–11 µm (Sheffy & Dilcher 1971); Unicellular fungal
spores without preformed aperture; shape variable, mostly irregularly rounded; one or more cells
usually randomly clustered; exine not too thick, folded or cracked, scabrate or usually pitted
(Ediger 1981); Inaperturate algal or fungal spores, unicellate, nonseptate. Shape and size variable,
outline smooth or often uneven because of wrinkles or folds. Spore wall ornamentation variable
(Saxena & Bhattacharyya 1987).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: 71 (but we accept only 60 species as legitimate because eleven
species have been transferred to other genera by various authors).
Notes: Rouse (1959) incorrectly considered the generic name Inapertisporites Hammen 1954
as not validly published. He stated that “Although a generic diagnosis and type species were not
presented (i.e. by Van der Hammen 1954), the name is considered appropriate for reference to
fossil spores that appear to be fungal, algal or possibly bryophytic in affiliation, and is therefore
conserved.” Actually, Van der Hammen (1954) had validly published the name of this genus with a
diagnosis. A type species was not required for valid publication until 1958; Van der Hammen
(1955) designated a type species, Inapertisporites variabilis. This is one of three species that Van
der Hammen (1954) validly published. Consequently, Rouse (1959) created a later homonym and a
taxonomic synonym of Inapertisporites Hammen 1954. Triporisporonites Sheffy & Dilcher 1971 is
a later taxonomic synonym of Inapertisporites. Ediger (1981) considered Microsporonites to be a
later synonym and proposed to transfer its type species as “I. cacheutensis” (R.K. Jain) V.S. Ediger
1981. However, he did not validly publish this new combination because he did not provide full
and direct reference of the basionym.
1.23.1. Species: I. argentinus (R.K. Jain) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483404; Basionym: Reticulatasporites argentinus R.K. Jain 1968; Location: Minas
de Petroleo, about 32 km south-west of Mendoza, western Argentina; Age: Middle Triassic;
Notes: Jain (1968) stated that this species is quite distinct from the others in this genus in
having the fewest and largest brochi.
1.23.2. Species: I. asymetricus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108084; Current name: Hypoxylonites asymetricus (Sal.-Cheb. & Locq.) Elsik 1990a fide
Elsik (1990a).
1.23.3. Species: I. cacheutensis (R.K. Jain) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483405; Basionym: Reticulatasporites cacheutensis R.K. Jain 1968;
Location: Minas de Petroleo, about 32 km south-west of Mendoza, western Argentina; Age:
Middle Triassic; Notes: Reticulatasporites cacheutensis broadly resembles other species of
Reticulatasporites described earlier from various Permo-Carboniferous strata. However, it
differs from them in having larger brochi with broader muri.
1.23.4. Species: I. cephalus Anil Chandra et al. 1984 (Fig. 6Y); Index Fungorum Registration
Identifier: 106915; Location: Sediment core no. 1 (Lat. 17°57.9'N: Long. 70°46.0'E), Arabian
Sea; Age: Late Quaternary; Notes: The species epithet in the original publication is spelled as
“cephalu” which has been corrected here.

706
1.23.5. Species: I. chandrae R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 6Z); Index Fungorum
Registration Identifier: 519898; Location: Sediment core no. 4 (Lat. 21°10.0'N: Long.
70°26.9'E), Arabian Sea; Age: Late Quaternary; Notes: The species epithet is in honour of Dr.
Anil Chandra, Birbal Sahni Institute of Palaeosciences, Lucknow, India.
1.23.6. Species: I. circularis Sheffy & Dilcher 1971 (Fig. 6AA); Index Fungorum Registration
Identifier: 111543; Location: Puryear clay pit, Tennessee, Henry County, Texas, U.S.A.; Age:
Middle Eocene (Claiborne Formation); Notes: The species epithet indicates its circular shape.
1.23.7. Species: I. clarkei Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483406; Basionym: Monoporisporites globosus R.T. Clarke 1965; Location: Canon City coal
field, Fremont County, Colorado, U.S.A.; Age: Late Cretaceous; Notes: The species epithet is
in honour of Professor Robert T. Clarke.

Figure 6 – A–AI Amerosporae. A Haplographites cateniger Félix 1894, Bar = 8 μm. B

Hypoxylonites ater (P. Kumar) R.K. Saxena 1992, Bar = 10 μm. C Hypoxylonites bhubanensis

707
Nandi & Subhra Banerjee 2012, Bar = 5 μm. D Hypoxylonites brazosensis Elsik 1990a, Bar = 10
μm. E Hypoxylonites chaiffetzii Elsik 1990a, Bar = 10 μm. F Hypoxylonites curvatus (Ramanujam
& K.P. Rao) Elsik 1990a, Bar = 5 μm. G Hypoxylonites disciformis (Sheffy & Dilcher) R.K.
Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk comb. nov., Bar = 5 μm. H Hypoxylonites
elsikii Nandi & Shubhra Banerjee 2012, Bar = 5 μm. I Hypoxylonites eocenicus Elsik 1990a, Bar =
10 μm. J Hypoxylonites felixii Elsik 1990a, Bar = 5 μm. K Hypoxylonites fusiformis Elsik 1990a,
Bar = 5 μm. L Hypoxylonites gulfensis Elsik 1990a, Bar = 10 μm. M Hypoxylonites kumarii
Kalgutkar & Janson. 2000, Bar = 10 μm. N Hypoxylonites lanceolatus (Debi Mukh.) R.K. Saxena,
Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk comb. nov., Bar = 20 μm. O Hypoxylonites magnus
Elsik 1990a, Bar = 10 μm. P Hypoxylonites minimus Nandi & Subhra Banerjee 2012, Bar = 3 μm.
Q Hypoxylonites miocenicus Elsik 1990a, Bar = 10 μm. R Hypoxylonites neogenicus Nandi &
Subhra Banerjee 2012, Bar = 5 μm. S Hypoxylonites ovalis Elsik 1990a, Bar = 5 μm. T
Hypoxylonites subrotundus Nandi & Subhra Banerjee 2012, Bar = 5 μm. U Hypoxylonites
subuliformis Elsik 1990a, Bar = 10 μm. V Hypoxylonites sulekii Elsik 1990a, Bar = 10 μm. W
Hypoxylonites thindii Nandi & A. Sinha 2012, Bar = 5 μm. X Hypoxylonites vittatoides Elsik
1990a, Bar = 10 μm. Y Inapertisporites cephalus Anil Chandra et al. 1984, Bar = 10 μm. Z
Inapertisporites chandrae R.K. Saxena & S.K.M. Tripathi 2011, Bar = 5 μm. AA Inapertisporites
circularis Sheffy & Dilcher 1971, Bar = 7 μm. AB Inapertisporites crenulatus P. Kumar 1990, Bar
= 10 μm. AC Inapertisporites cystoides Ambwani 1982, Bar = 10 μm. AD Inapertisporites
deccanii (Chitaley & Yawale) Kalgutkar & Janson. 2000, Bar = 10 μm. AE Inapertisporites edigeri
Kalgutkar & Janson. 2000, Bar = 10 μm. AF Inapertisporites ellipticus Anil Chandra et al. 1984,
Bar = 20 μm. AG Inapertisporites giganteus Z.C. Song 1985, Bar = 10 μm. AH Inapertisporites
globatus S.C.D. Sah & R.K. Kar 1974, Bar = 10 μm. AI Inapertisporites granulosus Anil Chandra
et al. 1984, Bar = 10 μm.

1.23.8. Species: I. communis Z.C. Song & G.X. Li in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 637477; Location: Qingfeng county of Henan Province, China; Age:
Late Eocene-Middle Oligocene (Shahejie Formation).
1.23.9. Species: I. conicus Z.C. Song & G.X. Li in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 637478; Location: Qingfeng county of Henan Province, China; Age:
Late Oligocene (Dongying Formation); Notes: This species can be distinguished from other
species of Inapertisporites by its pointed protuberant end. The species epithet is after conical
protuberance at one end of the spore.
1.23.10. Species: I. crenulatus P. Kumar 1990 (Fig. 6AB); Index Fungorum Registration Identifier:
126564; Location: Padappakkara, Kollam District, Kerala, India; Age: Early-Middle Miocene
(Quilon Beds); Notes: Kumar (1990) stated that this species is comparable with fungal spores
described as Geastrum-type (Jarzen & Elsik 1986) from Recent sediments of Luangwa
Valley, Zambia. The species epithet is after crenulated margin of the spore.
1.23.11. Species: I. cystoides Ambwani 1982 (Fig. 6AC); Index Fungorum Registration Identifier:
483900; Location: Kotta-Bommuru, near Rajamahendravaram, East Godavari District,
Andhra Pradesh, India; Age: Early Eocene (Deccan Intertrappean Series).
1.23.12. Species: I. deccanii (Chitaley & Yawale) Kalgutkar & Janson. 2000 (Fig. 6AD); Index
Fungorum Registration Identifier: 483407; Basionym: Ustilago deccanii Chitaley & Yawale
1978; Location: Mohgaon Kalan, Chhindwara District, Madhya Pradesh, India; Age: Late
Cretaceous, Maastrichtian (Deccan Intertrappean Series); Notes: Chitaley & Yawale (1978)
opined that the characters present in these spores lead to their placement under the smut
family Ustilaginaceae. Comparison was made with Ustilago (Pers.) Roussel and
Sphacelotheca de Bary because of some resemblances with them (Chitaley & Yawale 1978).
Kalgutkar & Jansonius (2000) transferred this species to Inapertisporites, because they
thought that, in a dispersed condition, individual spores could not possibly be associated with
the smut fungi.

708
1.23.13. Species: I. dilcheri Anil Chandra et al. 1984; Index Fungorum Registration Identifier:
106916; Current name: Diporicellaesporites dilcheri (Anil Chandra et al.) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000); Notes: The species epithet is in honour of
Professor David L. Dilcher, Department of Botany, Indiana University, Bloomington,
Indiana, U.S.A.
1.23.14. Species: I. disciformis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111544; Current name: Hypoxylonites disciformis (Sheffy & Dilcher) R.K. Saxena,
Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk comb. nov. This new combination is described
under the section "New species and new combinations"
1.23.15. Species: I. edigeri Kalgutkar & Janson. 2000 (Fig. 6AE); Index Fungorum Registration
Identifier: 483408; Basionym: Inapertisporites rotundus V.S. Ediger 1981; Location: Thrace
Basin, Turkey (Ediger 1981); Northern Thrace Basin, Turkey (Ediger & Alisan 1989); Age:
Late Eocene-Oligocene, Miocene-Pliocene (Ediger 1981); Middle?-Late Eocene to Late
Oligocene, Miocene-Pliocene (Ediger & Alisan 1989); Notes: Inapertisporites edigeri
Kalgutkar & Janson. 2000 is a replacement name of Inapertisporites rotundus V.S. Ediger
1981.
1.23.16. Species: I. elencantense Sepúlveda 1980; Index Fungorum Registration Identifier: 483916;
Current name: Monoporisporites elencantensis (Sepúlveda) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.23.17. Species: I. ellipticus Anil Chandra et al. 1984 (Fig. 6AF); Index Fungorum Registration
Identifier: 106917; Location: Sediment core no. 1 (Lat. 17°57.9'N: Long. 70°46.0'E), Arabian
Sea; Age: Late Quaternary.
1.23.18. Species: I. elongatus Rouse 1959; Index Fungorum Registration Identifier: 109864;
Synonym: Hypoxylonites elongatus (Rouse) Elsik 1990a fide Kalgutkar & Jansonius (2000);
Location: Terminal Dock, British Columbia, Canada; Age: Late Cretaceous & Middle Eocene
(Burrard Formation).
1.23.19. Species: I. giganteus Z.C. Song 1985 (Fig. 6AG); Index Fungorum Registration Identifier:
519814; Location: Dafengshan, Qaidam Basin, Qinghai Province, China; Age: Middle-Late
Miocene.
1.23.20. Species: I. globatus S.C.D. Sah & R.K. Kar 1974 (Fig. 6AH); Index Fungorum
Registration Identifier: 519813; Location: Palana, Bikaner District, Rajasthan, India; Age:
Early Eocene (Palana lignite).
1.23.21. Species: I. globosus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108086; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age: Early
Eocene-Early Miocene.
1.23.22. Species: I. globulosus Rouse 1962; Index Fungorum Registration Identifier: 110211;
Current name: Monoporisporites globulosus (Rouse) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.23.23. Species: I. granulatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115663; Location: Panshan, Liaoning Province; Kenli, Shandong Province, Coastal region of
Bohai, China; Age: Eocene-Oligocene.
1.23.24. Species: I. granulosus Anil Chandra et al. 1984 (Fig. 6AI); Index Fungorum Registration
Identifier: 106918; Location: Sediment core no. 5 (Lat. 24°04.5'N: Long. 69°26.0'E), Arabian
Sea; Age: Late Quaternary.
1.23.25. Species: I. hammenii Anil Chandra et al. 1984 (Fig. 7A); Index Fungorum Registration
Identifier: 106919; Location: Sediment core no. 4 (Lat. 21°10.0'N: Long. 70°26.9'E), Arabian
Sea; Age: Late Quaternary; Notes: The species epithet is in honour of Dr. T. Van der
Hammen, who made commendable contributions to Tertiary palynology.
1.23.26. Species: I. ibrahimii V.S. Ediger & Alisan 1989; Index Fungorum Registration Identifier:
125506; Location: Northern Thrace Basin, Turkey; Age: Middle?-Late Eocene to Late
Oligocene, Miocene-Pliocene; Notes: The species epithet is in honour of Professor Ahmet
Can Ibrahim.

709
1.23.27. Species: I. indicus A. Gupta 2002 (Fig. 7B); Index Fungorum Registration Identifier:
540593; Location: Dadahu Road Section (left bank of Giri River), Sirmaur District, Himachal
Pradesh, India; Age: Early Eocene (Subathu Formation).
1.23.28. Species: I. irregularis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111545; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation).
1.23.29. Species: I. karii R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 7C); Index Fungorum
Registration Identifier: 519943; Location: Sediment core no. 1 (Lat. 17°57.9'N: Long.
70°46.0'E), Arabian Sea; Age: Late Quaternary; Notes: The species epithet honours Dr. R.K.
Kar, Birbal Sahni Institute of Palaeosciences, Lucknow, India.
1.23.30. Species: I. kedvesii Elsik 1968 (Fig. 7D); Index Fungorum Registration Identifier: 315797;
Location: Strip mine 11 km south-west of Rockdale, Milam County, Texas, U.S.A.; Age:
Palaeocene (Rockdale lignite).
1.23.31. Species: I. laevigatus Rouse 1959; Index Fungorum Registration Identifier: 106195;
Location: South-eastern British Columbia, Canada; Age: Early Cretaceous (Kootenay
Formation).
1.23.32. Species: I. longissimus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111546; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation).
1.23.33. Species: I. major P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier: 115676;
Location: Kenli, Shandong Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
1.23.34. Species: I. maximus H.P. Singh & R.K. Saxena 1981 (Fig. 7E); Index Fungorum
Registration Identifier: 483889; Location: Gagret-Bharwain Road Section, Una District,
Himachal Pradesh, India; Age: Pliocene-Pleistocene (Upper Siwalik).
1.23.35. Species: I. minutus Hammen 1954 (Fig. 7F); Index Fungorum Registration Identifier:
332523; Location: Magdalena Valley, Eastern Cordellera, Colombia, South America; Age:
Maastrichtian.
1.23.36. Species: I. miocenicus H.P. Singh et al. 1986; Index Fungorum Registration Identifier:
131932; Current name: Quilonia miocenica (H.P. Singh et al.) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.23.37. Species: I. multiporus J.T.F. Guim. et al. 2013; Index Fungorum Registration Identifier:
637479; Location: BOP2 outcrop, central and coastal Amazon Region, North Brazil; Age:
Miocene (Barreiras Formation); Notes: Guimarães et al. (2013) stated taxonomic affinity of
this species with Ustilago (Ustilaginaceae). The so called pores on the spore wall are likely to
be fungal scars, comparable to Fungal Scar Type 1 of Elsik (1968: 266, pl. 1, figs. 1–4). The
species epithet indicates numbers of pores along the spore body.
1.23.38. Species: I. nodulus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111547; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation).
1.23.39. Species: I. novus A. Gupta 2002 (Fig. 7G); Index Fungorum Registration Identifier:
540594; Location: Jamtah Road Section, Sirmaur District, Himachal Pradesh, India; Age:
Early Eocene (Subathu Formation).
1.23.40. Species: I. obpyriformis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111549; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation).
1.23.41. Species: I. obscurus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111550; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation).
1.23.42. Species: I. ovalis Sheffy & Dilcher 1971 (Fig. 7H); Index Fungorum Registration
Identifier: 111548; Location: Puryear clay pit, Tennessee, Henry County, U.S.A.; Age:
Middle Eocene (Claiborne Formation).

710
1.23.43. Species: I. pisciculatus (G. Norris) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483409; Basionym: Diporisporites pisciculatus G. Norris 1997;
Location: Mackenzie River delta, Canada; Age: Palaeocene-Eocene.
1.23.44. Species: I. plicatus Z.C. Song 1985; Index Fungorum Registration Identifier: 637480;
Location: Dafengshan, Qaidam Basin, Qinghai Province, China; Age: Middle-Late Miocene.
1.23.45. Species: I. pseudoreticulatus Rouse 1959; Index Fungorum Registration Identifier:
106196; Location: South-eastern British Columbia, Canada; Age: Early Cretaceous
(Kootenay Formation).
1.23.46. Species: I. pulvinatus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111551; Current name: Hypoxylonites pulvinatus (Sheffy & Dilcher) Elsik 1990a fide Elsik
(1990a).
1.23.47. Species: I. punctatus Rouse 1959; Index Fungorum Registration Identifier: 106197;
Location: South-eastern British Columbia, Canada; Age: Early Cretaceous (Kootenay
Formation).
1.23.48. Species: I. punctatus Anil Chandra et al. 1984; Index Fungorum Registration Identifier:
106569; Current name: Inapertisporites udarii A. Gupta 1985 fide Gupta (1985).
1.23.49. Species: I. quadrangularis Anil Chandra et al. 1984 (Fig. 7I); Index Fungorum
Registration Identifier: 106920; Location: Sediment core no. 4 (Lat. 21°10.0'N: Long.
70°26.9'E), Arabian Sea; Age: Late Quaternary.
1.23.50. Species: I. reniformis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111552; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation).
1.23.51. Species: I. reticulatus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111553; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation).
1.23.52. Species: I. rotundus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115675; Location: Lijin and Kenli, Shandong Province, Coastal region of Bohai, China; Age:
Eocene-Oligocene.
1.23.53. Species: I. rotundus V.S. Ediger 1981; Index Fungorum Registration Identifier: 108087;
Current name: Inapertisporites edigeri Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius
(2000).
1.23.54. Species: I. sahii R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 7J); Index Fungorum
Registration Identifier: 519944; Location: Palana, Bikaner District, Rajasthan, India; Age:
Early Eocene (Palana lignite); Notes: The species epithet is in honour of Dr. S.C.D. Sah,
Wadia Institute of Himalayan Geology, Dehradun, India.
1.23.55. Species: I. sahnii Kalgutkar 1997; Index Fungorum Registration Identifier: 437912;
Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories, Canada; Age: Late
Palaeocene-Early Eocene; Notes: The species epithet is in honour of Professor Birbal Sahni.
1.23.56. Species: I. scabridus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111554; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation).
1.23.57. Species: I. sinhae A. Gupta 2002 (Fig. 7K); Index Fungorum Registration Identifier:
540595; Location: Dadahu Road Section, Sirmaur District, Himachal Pradesh, India; Age:
Early Eocene (Subathu Formation); Notes: The species epithet is in honour of Dr. R.K. Kar,
Birbal Sahni Institute of Palaeosciences, Lucknow, India.
1.23.58. Species: I. solidus Z.C. Song & Liu Cao 1994; Index Fungorum Registration Identifier:
483760; Location: King George Island, Antarctica; Age: Late Cretaceous.
1.23.59. Species: I. subcapsularis Sheffy & Dilcher 1971 (Fig. 7L); Index Fungorum Registration
Identifier: 111555; Location: Puryear clay pit, Tennessee, Henry County, U.S.A.; Age:
Middle Eocene (Claiborne Formation).

711
1.23.60. Species: I. subcurvatus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111556; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation).
1.23.61. Species: I. subovoides Sheffy & Dilcher 1971 (Fig. 7M); Index Fungorum Registration
Identifier: 111557; Location: Puryear clay pit, 800 m south of Puryear, Henry County,
Tennessee, U.S.A.; Age: Middle Eocene (Claiborne Formation).
1.23.62. Species: I. subverrucatus A. Gupta 2002 (Fig. 7N); Index Fungorum Registration
Identifier: 540596; Location: Jamtah Road Section, Sirmaur District, Himachal Pradesh,
India; Age: Early Eocene (Subathu Formation).
1.23.63. Species: I. tetradus Rouse 1962; Index Fungorum Registration Identifier: 109865; Current
name: Spegazzinites tetradus (Rouse) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius
(2000).
1.23.64. Species: I. tiwarii A. Gupta 2002 (Fig. 7O); Index Fungorum Registration Identifier:
540597; Location: Dadahu Road Section, Sirmaur District, Himachal Pradesh, India; Age:
Early Eocene (Subathu Formation); Notes: The species epithet is in honour of Dr. R.K. Kar,
Birbal Sahni Institute of Palaeosciences, Lucknow, India.
1.23.65. Species: I. triporatus Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483413; Basionym: Triporisporites minutus Hammen 1954; Location: Magdalena Valley,
Eastern Cordillera, Colombia, South America; Age: Maastrichtian.
1.23.66. Species: I. trivedii Ambwani 1982 (Fig. 7P); Index Fungorum Registration Identifier:
483899; Location: Kotta-Bommuru, near Rajamahendravaram, East Godavari District,
Andhra Pradesh, India; Age: Early Eocene (Deccan Intertrappean Series); Notes: The species
epithet is in honour of Professor B.S. Trivedi, Department of Botany, Lucknow University,
Lucknow, India.
1.23.67. Species: I. typicus Hammen 1954; Index Fungorum Registration Identifier: 332524;
Location: Magdalena Valley, Eastern Cordillera, Colombia, South America; Age:
Maastrichtian.
1.23.68. Species: I. udarii A. Gupta 1985 (Fig. 7Q); Index Fungorum Registration Identifier:
133492; Basionym: Inapertisporites punctatus Anil Chandra et al. 1984; Location: Sediment
core no. 3 (Lat. 19°32.8'N: Long. 71°21.5'E), Arabian Sea; Age: Late Quaternary; Notes: The
species epithet is in honour of Professor Ram Udar, Department of Botany, Lucknow
University, Lucknow, India.
1.23.69. Species: I. variabilis Hammen 1954 (Fig. 7R); Index Fungorum Registration Identifier:
332525; Location: Magdalena Valley, Eastern Cordillera, Colombia, South America; Age:
Maastrichtian.
1.23.70. Species: I. vittatus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111558; Current name: Hypoxylonites vittatus (Sheffy & Dilcher) Elsik 1990 fide Kalgutkar
& Jansonius (2000).
1.23.71. Species: I. vulgaris Sheffy & Dilcher 1971 (Fig. 7S); Index Fungorum Registration
Identifier: 111559; Location: Puryear clay pit, Tennessee, Henry County, U.S.A.; Age:
Middle Eocene (Claiborne Formation).

1.24. Genus: INCERTISPORITES Hammen, Bol. Geol. (Bogota) 2(1): 83 (1954); Index
Fungorum Registration Identifier: 21140; Type: I. polygranulatus Hammen 1954.
Original Diagnosis: Fungal spores of indefinite type. (Van der Hammen 1954).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
1.24.1. Species: I. polygranulatus Hammen 1954 (Fig. 7T); Index Fungorum Registration
Identifier: 332526; Location: Magdalena Valley, Eastern Cordillera, Colombia, South
America; Age: Maastrichtian.

712
1.25. Genus: LACRIMASPORONITES R.T. Clarke, Mountain Geologist 2(2): 87 (1965); Index
Fungorum Registration Identifier: 21146; Type: L. levis R.T. Clarke 1965.
Original Diagnosis: Fungal spores unicellular (amerospores), elliptical (tear-shaped), hilate or
monoporate, cell wall psilate (Clarke 1965).
Emended Diagnoses: Diagnosis of Lacrimasporonites was emended by Elsik (1968) and
Kalgutkar & Jansonius (2000), as follows: Monoporate, nonseptate, psilate fungal spores. Spatulate
to elliptical. Pore apical (Elsik 1968); Unicellate, mostly medium-sized, spatulate to lacrimate,
rarely approaching elliptical, smooth-walled fungal spores; with a flat hilar scar at one end, and a
round pore at the opposite end of the spore (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: 31 (but we accept only six species as legitimate because 22
species have been transferred to other genera by Kalgutkar & Jansonius 2000 and three species, viz.
L. dolium Z.C. Song, Qian & Y.H. Zheng in Z.C. Song et al. 1999, L. ovatus Z.C. Song, Qian &
Y.H. Zheng in Z.C. Song et al. 1999 and L. sondensis S. Soomro et al. 2010, have not been validly
published).
Notes: Clarke (1965) opined that the shape is an important criterion in taxonomic
identification of fungal spores. He restricted Monoporisporites for spherical and subspherical hilate
or monoporate fungal spores and Lacrimasporonites for elliptical hilate or monoporate fungal
spores. Kalgutkar & Jansonius (2000) emended the diagnosis of Lacrimasporonites and proposed
that unicellular spores with a real pore (or pore-shaped hilum), irrespective of their overall shape,
be included in Monoporisporites. Consequently, a large number of species of Lacrimasporonites
were transferred to Monoporisporites and to some other genera by Kalgutkar & Jansonius (2000).
The generic prefix is Latin for a tear, given in reference to the tear-drop shape of the spore.
1.25.1. Species: L. arcuatus Doub. & D. Pons 1973; Index Fungorum Registration Identifier:
485259; Current name: Monoporisporites arcuatus (Doub. & D. Pons) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
1.25.2. Species: L. basidii Elsik 1968; Index Fungorum Registration Identifier: 316200; Current
name: Monoporisporites basidii (Elsik) Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
1.25.3. Species: L. bellus Anil Chandra et al. 1984; Index Fungorum Registration Identifier:
106961; Current name: Monoporisporites bellus (Anil Chandra et al.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
1.25.4. Species: L. buerglii (Hammen) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483415; Basionym: Monoporisporites buerglii Hammen 1954; Location:
Magdalena Valley, Eastern Cordellera, Colombia, South America; Age: Maastrichtian.
1.25.5. Species: L. cupuliformis (Sheffy & Dilcher) D.L.E. Glass et al. 1986; Index Fungorum
Registration Identifier: 360322; Current name: Anatolinites cupuliformis (Sheffy & Dilcher)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.25.6. Species: L. dolium Z.C. Song, Qian & Y.H. Zheng in Z.C. Song et al. 1999 (nom. inval.)
fide Kalgutkar & Jansonius (2000); Index Fungorum Registration Identifier: 483842;
Location: China; Age: Late Cretaceous/Tertiary; Notes: The species name was not validly
published because the author did not specify where the holotype was deposited.
1.25.7. Species: L. fusoides (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483242; Basionym: Amepiospora fusoides Sal.-Cheb. & Locq. 1980;
Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age: Oligocene.
1.25.8. Species: L. globulosus (Rouse) G. Norris 1997; Index Fungorum Registration Identifier:
483785; Current name: Monoporisporites globulosus (Rouse) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.25.9. Species: L. hammenii (Mart.-Hern. & Tom.-Ort.) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483416; Basionym: Monoporisporites hammenii Mart.-
Hern. & Tom.-Ort. 1989; Location: Piedras Negras, Coahuila State, Mexico; Age:
Maastrichtian.

713
1.25.10. Species: L. levis R.T. Clarke 1965 (Fig. 7U); Index Fungorum Registration Identifier:
332866; Location: Canon City Coalfield, Fremont County, Colorado, U.S.A.; Age: Late
Cretaceous.
1.25.11. Species: L. longus R.K. Kar 1979; Index Fungorum Registration Identifier: 112387;
Current name: Didymoporisporonites longus (R.K. Kar) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.25.12. Species: L. magnus Haseld. 1973; Index Fungorum Registration Identifier: 483816;
Current name: Monoporisporites nemagnus Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
1.25.13. Species: L. magnus R.K. Saxena & H.P. Singh 1983; Index Fungorum Registration
Identifier: 485266; Current name: Didymoporisporonites gigas Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
1.25.14. Species: L. major P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115683; Current name: Hilidicellites major (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
1.25.15. Species: L. minutaestriatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration
Identifier: 115682; Current name: Monoporisporites minutaestriatus (P. Ke & Z.Y. Shi)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.25.16. Species: L. niger P. Kumar 1990; Index Fungorum Registration Identifier: 126565;
Current name: Monoporisporites niger (P. Kumar) Kalgutkar & Janson. 2000 fide Kalgutkar
& Jansonius (2000).
1.25.17. Species: L. ovaliformis Anil Chandra et al. 1984; Index Fungorum Registration Identifier:
106962; Current name: Monoporisporites ovaliformis (Anil Chandra et al.) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.25.18. Species: L. ovalis Doub. & D. Pons 1973; Index Fungorum Registration Identifier:
485260; Current name: Monoporisporites doubingerae Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.25.19. Species: L. ovatus Z.C. Song, Qian & Y.H. Zheng in Z.C. Song et al. 1999 (nom. inval.)
fide Kalgutkar & Jansonius (2000); Index Fungorum Registration Identifier: 483843; Notes:
This species name was not validly published because the author did not specify where the
holotype was deposited, and did not provide a Latin description or its English translation.
1.25.20. Species: L. oviformis V.S. Ediger 1981; Index Fungorum Registration Identifier: 108119;
Current name: Monoporisporites oviformis (V.S. Ediger) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.25.21. Species: L. permagnus M.G. Parsons & G. Norris 1999; Index Fungorum Registration
Identifier: 483917; Location: Caribou Hills, Mackenzie River delta, Canada; Age: Eocene;
Notes: The species epithet is derived from Latin permagnus = very large.
1.25.22. Species: L. pseudoabruptus M.G. Parsons & G. Norris 1999; Index Fungorum
Registration Identifier: 483918; Location: Caribou Hills, Mackenzie River delta, Canada;
Age: Eocene.
1.25.23. Species: L. psilatus Z.C. Song & G.X. Li in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 485261; Current name: Monoporisporites perpsilatus Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.25.24. Species: L. reniformis Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108120; Current name: Monoporisporites reniformis (Sal.-Cheb. & Locq.) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.25.25. Species: L. scabratus Z.C. Song & Liu Cao 1994; Index Fungorum Registration Identifier:
483761; Current name: Monoporisporites scabratus (Z.C. Song & Liu Cao) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.25.26. Species: L. singularis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111596; Current name: Monoporisporites singularovalis Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).

714
1.25.27. Species: L. sondensis S. Soomro et al. 2010 (nom. inval.) fide hoc loco; Index Fungorum
Registration Identifier: 637481; Location: Borehole Core DH No. 18 in the Sonda Coal field,
Thatta District, Sindh, Pakistan; Age: Palaeocene; Notes: The species name was not validly
published because the author did not specify where the holotype was deposited.
1.25.28. Species: L. stoughiae Elsik 1968; Index Fungorum Registration Identifier: 509432;
Current name: Saccisporonites stoughiae (Elsik) Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
1.25.29. Species: L. tenuis Z.C. Song & Liu Cao 1994; Index Fungorum Registration Identifier:
483762; Current name: Anatolinites tenuis (Z.C. Song & Liu Cao) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
1.25.30. Species: L. traversii V.S. Ediger & Alisan 1989; Index Fungorum Registration Identifier:
125507; Current name: Monoporisporites traversii (V.S. Ediger & Alisan) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.25.31. Species: L. westii Elsik 1968; Index Fungorum Registration Identifier: 316203; Current
name: Monoporisporites westii (Elsik) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius
(2000).

1.26. Genus: LEPIOTASPORITES T.C. Huang, Taiwania 26: 47 (1981); Index Fungorum
Registration Identifier: 21149; Type: L. taiwanensis T.C. Huang 1981
Original Diagnosis: Fungal spore with terminal hilum.
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One
1.26.1. Species: L. taiwanensis T.C. Huang 1981 (Fig. 7V); Index Fungorum Registration
Identifier: 115785; Location: Taiwan; Age: Miocene (Talu Shale).

1.27. Genus: MAGNOSPORITES Rouse, Micropaleontology 8(2): 210 (1962); Index Fungorum
Registration Identifier: 18487; Type: M. staplinii Rouse 1962.
Original Diagnosis: Spore-like bodies, circular to elliptical in outline and with a fairly thin
wall (ca. 1.5 μm). The wall is a bright yellow and is always folded and contorted. There is no
aperture apparent and no ornamentation on the smooth surface. Size-range 100–170 μm (Rouse
1962).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
Notes: The fungal affinity of these forms is questionable. These structures are apparently
associated with freshwater plants, as they are found only in freshwater deposits. Whether they are
spores, sporangia, or remains of one-celled organisms is entirely open to question.
1.27.1. Species: M. staplinii Rouse 1962 (Fig. 7W); Index Fungorum Registration Identifier:
560989; Location: Terminal Dock, the city of Vancouver, British Columbia, Canada; Age:
Late Cretaceous-Middle Eocene (Burrard Formation); Notes: The species is named in honour
of Professor Frank L. Staplin.

1.28. Genus: MEDIAVERRUNITES Nandi & A. Sinha, Palynology 31(1): 98 (2007); Index
Fungorum Registration Identifier: 548478; Type: M. mulleri Nandi & A. Sinha 2007; Current
name: POTAMOMYCES K. D. Hyde 1995 fide Nuñez Otaño et al. (2017).
Original Diagnosis: Spores aseptate, oval to elliptical, monoaperturate (or sometimes
inaperturate), pore situated at the basal end of the axis, equatorial region ornamented with flat or
slightly elevated verrucae that remain arranged either freely around the equator or merge to form a
shallow, thin to wide, dark to light, shadow-like rim or band, verrucae large or small, apex of
verrucae rounded or slightly connate, spore wall psilate to sculptured (Nandi & Sinha 2007).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Seven (all the species, including type species, have been
transferred to Potamomyces K. D. Hyde 1995).

715
 Notes: Muller (1959) recorded a characteristic unicellular fungal spore, having several
verrucae at the equatorial region, from the Recent shelf sediments of the Orinoco Delta, Venezuela.
He, however, did not assign it to any named taxon. Elsik (1976b) illustrated identical spores and
informally named it Mediaverrusporonites. Jarzen & Elsik (1986) informally used the name
Mediaverrunites to accommodate these spores but they neither proposed it as a genus nor assigned
any species to it. They described Mediaverrunites as follows: “Fungal spores one-celled, with a
single aperture (some specimens appear inaperturate), outline oval to elliptical, pore situated at one
end of axis, 7 μm in diameter; spore colour medium brown; septum lacking, but with a shadow
band, 10–12 μm wide; surface psilate except for shadow band which is ornamented with 8 (9?)
large verrucae, 10–12 μm in diameter; verrucae dark brown to black; spore wall thin <1μm thick;
spore length 62–65 μm, spore width 33–35 μm. ” Nandi & Sinha (2007) were the first to formally
describe Mediaverrunites as a genus and designated Mediaverrunites mulleri Nandi & A. Sinha
2007 as its type. Nandi & Sinha (2007), however, considered Jarzen & Elsik (1986) as the author of
the genus and proposed an emended diagnosis. We interpret that Nandi & Sinha (2007) were the
original author (not the validating author) of Mediaverrunites because earlier authors had no
intention of proposing it as a genus nor they assigned it any species. The genus is therefore solely
ascribed to Nandi & Sinha 2007 and cited as Mediaverrunites Nandi & A. Sinha 2007 (not Jarzen
& Elsik ex Nandi & A. Sinha 2007). The genus name is derived from the Latin media referring to
the central position of the verrucae. Nuñez Otaño et al. 2017 considered Mediaverrunites Nandi &
A. Sinha 2007 to be a later synonym of Potamomyces K.D. Hyde 1995. They transferred all species
Mediaverrunites, viz. M. batii Sancay 2014, M. elsikii Nandi & A. Sinha 2007, M. fournieri Elsik &
Jarzen 2009, M. invaginatus Elsik & Jarzen 2009, M. magnus Elsik & Jarzen 2009, M. mulleri
Nandi & A. Sinha 2007 and M. pontidiensis Sancay 2014 to Potamomyces K.D. Hyde 1995.
1.28.1. Species: M. batii Sancay 2014; Index Fungorum Registration Identifier: 821044; Current
name: Potamomyces batii (Sancay) ex Nuñez Otaño et al. 2017 fide Nuñez Otaño et al.
(2017).
1.28.2. Species: M. elsikii Nandi & A. Sinha 2007; Index Fungorum Registration Identifier:
568744; Current name: Potamomyces elsikii (Nandi & A. Sinha) Nuñez Otaño et al. 2017
fide Nuñez Otaño et al. (2017).
1.28.3. Species: M. fournieri Elsik & Jarzen 2009; Index Fungorum Registration Identifier: 568010;
Current name: Potamomyces fournieri (Elsik & Jarzen) Nuñez Otaño et al. 2017 fide Nuñez
Otaño et al. (2017).
1.28.4. Species: M. invaginatus Elsik & Jarzen 2009; Index Fungorum Registration Identifier:
568011; Current name: Potamomyces invaginatus (Elsik & Jarzen) Nuñez Otaño et al. 2017
fide Nuñez Otaño et al. (2017).
1.28.5. Species: M. magnus Elsik & Jarzen 2009; Index Fungorum Registration Identifier: 568012;
Current name: Potamomyces magnus (Elsik & Jarzen) Nuñez Otaño et al. 2017 fide Nuñez
Otaño et al. (2017).
1.28.6. Species: M. mulleri Nandi & A. Sinha 2007; Index Fungorum Registration Identifier:
568745; Current name: Potamomyces mulleri (Nandi & A. Sinha) Nuñez Otaño et al. 2017
fide Nuñez Otaño et al. (2017).
1.28.7. Species: M. pontidiensis Sancay 2014; Index Fungorum Registration Identifier: 821045;
Current name: Potamomyces pontidiensis (Sancay) ex Nuñez Otaño et al. 2017 fide Nuñez
Otaño et al. (2017).

1.29. Genus: MICROSPORONITES R.K. Jain, Palaeontographica Abt. B 122(1-3): 8 (1968);

Index Fungorum Registration Identifier: 21164; Type: M. cacheutensis R.K. Jain 1968.
Original Diagnosis: Fungal spores very small (10–20 μm in diameter), spherical, inaperturate,
exine thin, psilate, smooth (Jain 1968).
Emended Diagnosis: Thin-walled single celled fungal spores of small diameter, that tend to
occur in irregular clusters, but retain their individual spherical shape (not appressed into semi-
polyhedral shapes) (Kalgutkar & Jansonius 2000).

716
 Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Two.
Notes: Jain (1968) proposed this genus to accommodate very small, spherical, inaperturate,
smooth walled fungal spores which were earlier referred to Sporonites.
1.29.1. Species: M. cacheutensis R.K. Jain 1968 (Fig. 7X); Index Fungorum Registration
Identifier: 114325; Location: Minas de Petroleo, about 32 km south-west of Mendoza,
western Argentina; Age: Middle Triassic.
1.29.2. Species: M. taiwanensis T.C. Huang 1981; Index Fungorum Registration Identifier:
115789; Location: Taiwan; Age: Miocene.

1.30. Genus: MONOPORISPORITES Hammen 1954, Bol. Geol. (Bogota) 2(2): 14 (1954); Index
Fungorum Registration Identifier: 21174; Type: M. minutus Hammen 1954 (designated by Van der
Hammen 1955).
Synonyms: Ornatisporites M.G. Parsons & G. Norris 1999, Index Fungorum Registration
Identifier: 28642; Polyporisporites Hammen 1954, Index Fungorum Registration Identifier: 21259;
Psiammopomopiospora Sal.-Cheb. & Locq. 1980, Index Fungorum Registration Identifier: 25597;
Psiamspora Sal.-Cheb. & Locq. 1980, Index Fungorum Registration Identifier: 25589;
Reticulatisporonites Elsik 1968, Index Fungorum Registration Identifier: 21279.
Original Diagnosis: (Fungal) spore with one small, [(round) Van der Hammen 1955] pore.
(Van der Hammen 1954).
Emended Diagnoses: Diagnosis of Monoporisporites was emended by Elsik (1968), Sheffy &
Dilcher (1971) and Kalgutkar & Jansonius (2000), as follows: Monoporate, nonseptate, psilate
fungal or algal spores. Shape spherical to subspherical (Elsik 1968); Monoporate, aseptate, psilate
to finely punctate fungal or algal spores. Shapes spherical to subspherical, hilate or monoporate
(Sheffy & Dilcher 1971); Monohilate (or monoporate), unicellate, generally small to medium-sized,
round, oval or elongate elliptical fungal spores. Wall generally smooth, but ornamented forms are
included (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: 58 (but we accept only 47 species as legitimate because six
species have been transferred to other genera by various authors and five species, viz. M. biformis
Z.C. Song, Qian & Y.H. Zheng in Z.C. Song et al. 1999, M. gigasus Z.C. Song in Z.C. Song et al.
1999, M. grymaeformis Z.C. Song in Z.C. Song et al. 1999, M. megaporus Z.C. Song in Z.C. Song
et al. 1999 and M. operculatus Z.C. Song in Z.C. Song et al. 1999, have not been validly
published).
Notes: Clarke (1965) provided the following restated (not emended) diagnosis: “Fungal
spores unicellular (amerosporous), spherical to subspherical, hilate or monoporate, cell wall psilate
to finely punctate.” The emended diagnosis by Sheffy & Dilcher (1971) provided combination of
two previously published descriptions, i.e. restated description by Clarke (1965) and emended
description by Elsik (1968). Moreover, the genus was expanded to include the punctate species as
well, viz. Monoporisporites globosus R.T. Clarke 1965, M. buerglii Hammen 1954 and M. minutus
Hammen 1954. Kalgutkar & Jansonius (2000) included spherical and elongate spores in the genus,
because both shapes may be produced by the same genus or species of modern fungi.
Polyporisporites Hammen, Psiammopomopiospora Sal.-Cheb. & Locq. and Reticulatisporonites
Elsik are later taxonomic synonyms of Monoporisporites Hammen (Kalgutkar & Jansonius 2000).
1.30.1. Species: M. abruptus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111658; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation).
1.30.2. Species: M. annulatus Hammen 1954 (Fig. 7Y); Index Fungorum Registration Identifier:
334446; Location: Magdalena Valley, Eastern Cordellera, Colombia, South America; Age:
Maastrichtian.

717
1.30.3. Species: M. aquilus Kalgutkar 1997; Index Fungorum Registration Identifier: 437927;
Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories, Canada; Age: Late
Palaeocene-Early Eocene.
1.30.4. Species: M. arcuatus (Doub. & D. Pons) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483423; Basionym: Lacrimasporonites arcuatus Doub. & D. Pons
1973; Location: Cerrejon basin, Colombia, South America; Age: Palaeocene-Eocene.
1.30.5. Species: M. basidii (Elsik) Kalgutkar & Janson. 2000 (Fig. 7Z); Index Fungorum
Registration Identifier: 483422; Basionym: Lacrimasporonites basidii Elsik 1968; Location:
Strip mine approximately 11 km south-west of Rockdale, Milam County, Texas, U.S.A.;
Age: Palaeocene.
1.30.6. Species: M. bellus (Anil Chandra et al.) Kalgutkar & Janson. 2000 (Fig. 7AA); Index
Fungorum Registration Identifier: 483424; Basionym: Lacrimasporonites bellus Anil
Chandra et al. 1984; Location: Sediment core no. 1 (Lat. 17°57.9'N: Long. 70°46.0'E),
Arabian Sea; Age: Late Quaternary.
1.30.7. Species: M. biformis Z.C. Song, Qian & Y.H. Zheng in Z.C. Song et al. 1999 (nom. inval.)
fide Kalgutkar & Jansonius (2000); Index Fungorum Registration Identifier: 483915; This
species was not validly published because the author did not specify where the holotype is
stored, and did not provide a Latin description or its English translation.
1.30.8. Species: M. buerglii Hammen 1954; Index Fungorum Registration Identifier: 334447;
Current name: Lacrimasporonites buerglii (Hammen) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.30.9. Species: M. circularis R.K. Saxena 2009 (Fig. 7AB); Index Fungorum Registration
Identifier: 515007; Basionym: Monoporisporites hammenii B. Samant & Tapaswi 2000;
Location: Cambay Basin, Surat District, Gujarat, India; Age: Early Eocene (Cambay Shale);
Notes: Monoporisporites circularis R.K. Saxena 2009 is a replacement name of
Monoporisporites hammenii B. Samant & Tapaswi 2000.
1.30.10. Species: M. cupuliformis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111659; Current name: Anatolinites cupuliformis (Sheffy & Dilcher) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
1.30.11. Species: M. dilcheri Mart.-Hern. & Tom.-Ort. 1989; Index Fungorum Registration
Identifier: 483806; Location: Piedras Negras, Coahuila State, Mexico; Age: Maastrichtian.
1.30.12. Species: M. doubingerae Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483425; Basionym: Lacrimasporonites ovalis Doub. & D. Pons 1973; Location:
Cerrejon basin, Colombia, South America; Age: Palaeocene-Eocene.
1.30.13. Species: M. elencantensis (Sepúlveda) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483426; Basionym: Inapertisporites elencantensis Sepúlveda 1980;
Location: Northern Patagonian Cordillera, Chubut Province, Argentina; Age: Middle Eocene
(Andean Andesitic Series).
1.30.14. Species: M. elongatus (Hammen) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483427; Basionym: Polyporisporites elongatus Hammen 1954;
Location: Magdalena Valley, Eastern Cordillera, Colombia, South America; Age:
Maastrichtian.
1.30.15. Species: M. gigasus Z.C. Song in Z.C. Song et al. 1999 (nom. inval.) fide Kalgutkar &
Jansonius (2000); Index Fungorum Registration Identifier: 483838; Notes: The species name
was not validly published because the author did not specify where the holotype was
deposited.
1.30.16. Species: M. globoides (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483428; Basionym: Psiammopomopiospora globoides Sal.-Cheb. &
Locq. 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene.

718
1.30.17. Species: M. globosus R.T. Clarke 1965; Index Fungorum Registration Identifier: 334448;
Current name: Inapertisporites clarkei Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius
(2000).
1.30.18. Species: M. globulosus (Rouse) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483429; Basionym: Inapertisporites globulosus Rouse 1962; Location: Terminal
Dock, the city of Vancouver, British Columbia, Canada; Age: Late Cretaceous-Middle
Eocene (Burrard Formation).
1.30.19. Species: M. grandis Hammen 1954; Index Fungorum Registration Identifier: 334449;
Location: Magdalena Valley, Eastern Cordellera, Colombia, South America; Age:
Maastrichtian.

Figure 7 – A–AI Amerosporae. A Inapertisporites hammenii Anil Chandra et al. 1984, Bar = 20
μm. B Inapertisporites indicus A. Gupta 2002, Bar = 5 μm. C Inapertisporites karii R.K. Saxena &
S.K.M. Tripathi 2011, Bar = 20 μm. D Inapertisporites kedvesii Elsik 1968, Bar = 10 μm.
E Inapertisporites maximus H.P. Singh & R.K. Saxena 1981, Bar = 20 μm. F Inapertisporites

719
minutus Hammen 1954, Bar = 5 μm. G Inapertisporites novus A. Gupta 2002, Bar = 25 μm. H
Inapertisporites ovalis Sheffy & Dilcher 1971, Bar = 7 μm. I Inapertisporites quadrangularis Anil
Chandra et al. 1984, Bar = 5 μm. J Inapertisporites sahii R.K. Saxena & S.K.M. Tripathi 2011, Bar
= 30 μm. K Inapertisporites sinhae A. Gupta 2002, Bar = 5 μm. L Inapertisporites subcapsularis
Sheffy & Dilcher 1971, Bar = 5 μm. M Inapertisporites subovoides Sheffy & Dilcher 1971, Bar = 5
μm. N Inapertisporites subverrucatus A. Gupta 2002, Bar = 10 μm. O Inapertisporites tiwarii A.
Gupta 2002, Bar = 5 μm. P Inapertisporites trivedii Ambwani 1982, Bar = 20 μm. Q
Inapertisporites udarii A. Gupta 1985, Bar = 20 μm. R Inapertisporites variabilis Hammen 1954,
Bar = 8 μm. S Inapertisporites vulgaris Sheffy & Dilcher 1971, Bar = 7 μm. T Incertisporites
polygranulatus Hammen 1954, Bar = 20 μm. U Lacrimasporonites levis R.T. Clarke 1965, Bar = 8
μm. V Lepiotasporites taiwanensis T.C. Huang 1981, Bar = 10 μm. W Magnosporites staplinii
Rouse 1962, Bar = 30 μm. X Microsporonites cacheutensis R.K. Jain 1968, Bar = 3 μm. Y
Monoporisporites annulatus Hammen 1954, Bar = 5 μm. Z Monoporisporites basidii (Elsik)
Kalgutkar & Janson. 2000, Bar = 5 μm. AA Monoporisporites bellus (Anil Chandra et al.)
Kalgutkar & Janson. 2000, Bar = 20 μm. AB Monoporisporites circularis R.K. Saxena 2009, Bar =
10 μm. AC Monoporisporites keralensis Ramanujam & K.P. Rao 1978, Bar = 10 μm. AD
Monoporisporites koenigii Elsik 1968, Bar = 10 μm. AE Monoporisporites meghalayaensis R.K.
Saxena & S.K.M. Tripathi 2011, Bar = 20 μm. AF Monoporisporites minutus Hammen 1954, Bar =
8 μm. AG Monoporisporites neyveliensis Ramanujam & Ramachar 1980, Bar = 5 μm. AH
Monoporisporites niger (P. Kumar) Kalgutkar & Janson. 2000, Bar = 10 μm. AI Monoporisporites
novus Anil Chandra et al. 1984, Bar = 10 μm.

1.30.20. Species: M. grymaeformis Z.C. Song in Z.C. Song et al. 1999 (nom. inval.) fide Kalgutkar
& Jansonius (2000); Index Fungorum Registration Identifier: 483839; Notes: The species
name was not validly published because the author did not specify where the holotype was
deposited, and did not provide a Latin description or its English translation
1.30.21. Species: M. hammenii B. Samant & Tapaswi 2000; Index Fungorum Registration
Identifier: 515006; Current name: Monoporisporites circularis R.K. Saxena 2009 fide
Saxena (2009).
1.30.22. Species: M. hammenii Mart.-Hern. & Tom.-Ort. 1989; Index Fungorum Registration
Identifier: 483805; Current name: Lacrimasporonites hammenii (Mart.-Hern. & Tom.-Ort.)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.30.23. Species: M. keralensis Ramanujam & K.P. Rao 1978 (Fig. 7AC); Index Fungorum
Registration Identifier: 115076; Location: Padappakkara, Kollam District, Kerala, India; Age:
Miocene (Quilon and Warkalli beds).
1.30.24. Species: M. koenigii Elsik 1968 (Fig. 7AD); Index Fungorum Registration Identifier:
317864; Location: Strip mine approximately 11 km south-west of Rockdale, Milam County,
Texas, U.S.A.; Age: Palaeocene.
1.30.25. Species: M. macrosporus (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483244; Basionym: Amepiospora macrospora Sal.-Cheb.
& Locq. 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene.
1.30.26. Species: M. magnus Kalgutkar 1993; Index Fungorum Registration Identifier: 483882;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene.
1.30.27. Species: M. mathurii Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483430; Basionym: Psilainaperturites ovalis Y.K. Mathur 1966; Location: Matanomadh,
western Kutch, India; Age: Palaeocene.
1.30.28. Species: M. megaporus Z.C. Song in Z.C. Song et al. 1999 (nom. inval.) fide Kalgutkar &
Jansonius (2000); Index Fungorum Registration Identifier: 483840; Notes: The species name
was not validly published because the author did not specify where the holotype is stored, and
did not provide a Latin description or its English translation.

720
1.30.29. Species: M. meghalayaensis R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 7AE); Index
Fungorum Registration Identifier: 519946; Location: Sonapur-Badarpur Road Section, Jaintia
Hills, Meghalaya and Cachar District, Assam, India; Age: Miocene (Bokabil Formation).
1.30.30. Species: M. minionoides O’Keefe 2017; Index Fungorum Registration Identifier: 821910;
Location: Tumbes Province, Peru; Age: Miocene (Heath Formation); Notes: The species
epithet refers to the ovoid form and single eye of the fictional cartoon characters known as
‘minions’.
1.30.31. Species: M. minutaestriatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483431; Basionym: Lacrimasporonites minutaestriatus P.
Ke & Z.Y. Shi 1978; Location: Panshan, Liaoning Province, Coastal region of Bohai, China;
Age: Eocene-Oligocene.
1.30.32. Species: M. minutus Hammen 1954 (Fig. 7AF); Index Fungorum Registration Identifier:
334450; Basionym: Monoporisporites minutus A Hammen 1954; left, top row of figures
(lectotype selected by Jansonius & Hills 1976); Location: Magdalena Valley, Eastern
Cordillera, Colombia, South America; Age: Maastrichtian.
1.30.33. Species: M. nemagnus Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483432; Basionym: Lacrimasporonites magnus Haseld. 1973; Location: San
Martin (Graus), Esera Valley, Pyrenees, Spain; Age: Palaeocene-Late Eocene; Notes:
Monoporisporites nemagnus Kalgutkar & Janson. 2000 is a replacement name of
Monoporisporites minutus A Hammen 1954.
1.30.34. Species: M. neoglobosus Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483243; Basionym: Amepiospora globosa Sal.-Cheb. & Locq. 1980; Location:
Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age: Oligocene.
1.30.35. Species: M. neyveliensis Ramanujam & Ramachar 1980 (Fig. 7AG); Index Fungorum
Registration Identifier: 483756; Location: Neyveli, Cuddalore District, Tamil Nadu, India;
Age: Miocene (Neyveli lignite).
1.30.36. Species: M. niger (P. Kumar) Kalgutkar & Janson. 2000 (Fig. 7AH).; Index Fungorum
Registration Identifier: 483433; Basionym: Lacrimasporonites niger Kumar 1990; Location:
Padappakkara, Kollam District, Kerala, India; Age: Early-Middle Miocene; Notes: The
species epithet is derived from its diagnostic dark wall (Latin niger = black).
1.30.37. Species: M. novus Anil Chandra et al. 1984 (Fig. 7AI); Index Fungorum Registration
Identifier: 107032; Location: Sediment core no. 4 (Lat. 21°10.02'N: Long. 70°26.9'E),
Arabian Sea; Age: Late Quaternary.
1.30.38. Species: M. operculatus Z.C. Song in Z.C. Song et al. 1999 (nom. inval.) fide Kalgutkar &
Jansonius (2000); Index Fungorum Registration Identifier: 483841 Notes: The species name
was not validly published because the author did not specify where the holotype was
deposited, and did not provide a Latin description or its English translation
1.30.39. Species: M. ovaliformis (Anil Chandra et al.) Kalgutkar & Janson. 2000 (Fig. 8A); Index
Fungorum Registration Identifier: 483434; Basionym: Lacrimasporonites ovaliformis Anil
Chandra et al. 1984; Location: Sediment core no. 1 (Lat. 17°57.9'N: Long. 70°46.0'E),
Arabian Sea; Age: Late Quaternary.
1.30.40. Species: M. ovalis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111660; Current name: Basidiosporites ovalis (Sheffy & Dilcher) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
1.30.41. Species: M. oviformis (V.S. Ediger) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483435; Basionym: Lacrimasporonites oviformis V.S. Ediger 1981;
Location: Thrace Basin, Turkey; Age: Upper Eocene-Oligocene, Miocene-Pliocene.
1.30.42. Species: M. pannosus (M.G. Parsons & G. Norris) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483436; Basionym: Ornatisporites pannosus M.G. Parsons
& G. Norris 1999; Location: Caribou Hills, Mackenzie River delta, Canada; Age: Late
Palaeocene-Early Eocene.

721
1.30.43. Species: M. perminutus Doub. & D. Pons 1973; Index Fungorum Registration Identifier:
637482; Location: Cerrejon basin, Colombia, South America; Age: Palaeocene-Eocene.
1.30.44. Species: M. perpsilatus Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 83437; Basionym: Lacrimasporonites psilatus Z.C. Song & G.X. Li in Z.C. Song
et al. 1989; Qingteng county and Fanxian county of Henan Province, China; Age: Late
Eocene-Middle Oligocene (Shahejie Formation).
1.30.45. Species: M. psilatus Anil Chandra et al. 1984 (Fig. 8B); Index Fungorum Registration
Identifier: 107033; Location: Sediment core no. 1 (Lat. 17°57.9'N: Long. 70°46.0'E), Arabian
Sea; Age: Late Quaternary.
1.30.46. Species: M. reniformis (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483438; Basionym: Lacrimasporonites reniformis Sal.-Cheb. & Locq.
1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age: Early
Eocene-Early Miocene.
1.30.47. Species: M. rigens Kalgutkar 1997; Index Fungorum Registration Identifier: 437928;
Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories, Canada; Age: Late
Palaeocene-Early Eocene.
1.30.48. Species: M. scabratus (Z.C. Song & Liu Cao) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483439; Basionym: Lacrimasporonites scabratus Z.C. Song & Liu
Cao 1994; Location: King George Island, Antarctica; Age: Late Cretaceous.
1.30.49. Species: M. sheffyi Anil Chandra et al. 1984 (Fig. 8C); Index Fungorum Registration
Identifier: 107034; Location: Sediment core no. 5 (Lat. 24°04.5'N: Long. 69°26.0'E), Arabian
Sea; Age: Late Quaternary; Notes: The species epithet is in honour of Dr. M.V. Sheffy.
1.30.50. Species: M. singhii A. Gupta 2002 (Fig. 8D); Index Fungorum Registration Identifier:
540671; Location: Jamtah Road Section, Sirmaur District, Himachal Pradesh, India; Age:
Eocene (Subathu Formation); Notes: The species epithet honours Dr. H.P. Singh, Birbal
Sahni Institute of Palaeosciences, Lucknow, India.
1.30.51. Species: M. singularis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111661; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation).
1.30.52. Species: M. singularovalis Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483440; Basionym: Lacrimasporonites singularis Sheffy & Dilcher 1971;
Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.; Age:
Middle Eocene.
1.30.53. Species: M. smithii Elsik 1968; Index Fungorum Registration Identifier: 317865;
Location: Strip mine approximately 11 km south-west of Rockdale, Milam County, Texas,
U.S.A.; Age: Palaeocene.
1.30.54. Species: M. sperryi (Elsik) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483441; Basionym: Reticulatisporonites sperryi Elsik 1968; Location: Strip mine
approximately 11 km south-west of Rockdale, Milam County, Texas, U,S,A.; Age:
Palaeocene.
1.30.55. Species: M. stoveri Elsik 1968 (Fig. 8E); Index Fungorum Registration Identifier: 317866;
Location: Strip mine approximately 11 km south-west of Rockdale, Milam County, Texas,
U.S.A.; Age: Palaeocene.
1.30.56. Species: M. traversei (V.S. Ediger & Alisan) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483442; Basionym: Lacrimasporonites traversii V.S. Ediger & Alisan
1989; Location: Northern Thrace Basin, Turkey; Age: Middle?-Late Eocene to Late
Oligocene, Miocene-Pliocene.
1.30.57. Species: M. triangularis (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483245; Basionym: Amepiospora triangularis Sal.-Cheb.
& Locq. 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene.

722
1.30.58. Species: M. westii (Elsik) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483443; Basionym: Lacrimasporonites westii Elsik 1968; Location: Strip mine
approximately 11 km south-west of Rockdale, Milam County, Texas, U.S.A.; Age:
Palaeocene.

1.31. Genus: NAILISPORITES T.C. Huang, Taiwania 26: 47 (1981); Index Fungorum
Registration Identifier: 21190; Type: N. taiwanensis T.C. Huang 1981.
Original Diagnosis: Fungal spore with nail shape at one end.
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
1.31.1. Species: N. taiwanensis T.C. Huang 1981 (Fig. 8F); Index Fungorum Registration
Identifier: 115792; Location: Taiwan; Age: Miocene (Talu Shale).

1.32. Genus: NIGROSPORITES Debi Mukh., International Journal of Geology, Earth and
Environmental Sciences 2(2): 9 (2012); Index Fungorum Registration Identifier: 588464; Type: N.
neyveliensis Debi Mukh. 2012.
Original Diagnosis: Dispersed conidia oval/ lanceolate, somewhat flattened, large, absolutely
opaque, colour black, spore wall hyaline; size range 35–45 × 25–30 μm (Mukherjee 2012).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
Notes: The genus name is derived from Nigrospora, a hyphomycete (Subramanian 1971).
1.32.1. Species: N. neyveliensis Debi Mukh. 2012 (Fig. 8G); Index Fungorum Registration
Identifier: 588477; Location: Neyveli Lignite Mine-I, Cuddalore District, Tamil Nadu, India;
Age: Miocene (Neyveli Lignite); Notes: Mukherjee (2012) opined that this species resembles
Nigrospora (Hyphomycetes) for being black, opaque and lanceolate in shape. The specific
epithet refers to its type locality Neyveli.

1.33. Genus: ORNATISPORITES M.G. Parsons & G. Norris, Palaeontographica Abt. B 250(4-6):
117 (1999); Type: O. pannosus M.G. Parsons & G. Norris 1999; Current name:
MONOPORISPORITES Hammen 1954 fide Kalgutkar & Jansonius (2000).
Original Diagnosis: Monoporate fungal (amero)spores subspherical in shape or with a
somewhat elongate spore body – commonly elliptical, obovate or spatulate in outline. Spore wall
ornamented with scabs (scabrate), verrucae or granula, or a combination of these types of ornament
(Parsons & Norris 1999).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One (the single species has been transferred to Monoporisporites
Hammen 1954).
Notes: The generic name is illegitimate, being a later homonym of Ornatisporites Nagy 1963
and Ornatisporites Górecka 1969.
1.33.1. Species: O. pannosus M.G. Parsons & G. Norris 1999; Index Fungorum Registration
Identifier: 483919; Current name: Monoporisporites pannosus (M.G. Parsons & G. Norris)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).

1.34. Genus: PALAEOAMPHISPHAERELLA Ramanujam & Srisailam, Botanique, Nagpur 9(1-

4): 128 (1980); Index Fungorum Registration Identifier: 21204; Type: P. pirozynskii Ramanujam &
Srisailam 1980.
Synonym: Imprimospora G. Norris 1986 fide Kalgutkar & Jansonius (2000), Index Fungorum
Registration Identifier: 25449.
Original Diagnosis: Spores brownish to dark brown, aseptate, elliptical, oblong or somewhat
rhomboidal, with more or less rounded ends; with equatorial pores, placed equidistantly; surface
psilate to scabrate (Ramanujam & Srisailam 1980).
Classification: Fungi Imperfecti, Amerosporae.

723
 Number of species known: Three.
Notes: The equatorially disposed pores constitute a characteristic feature of this fungal spore
genus. The fossil spores exhibit remarkable resemblances with the ascospores of Amphisphaerella
included either under Xylariaceae or Amphisphaerellaceae of the pyrenomycetous Ascomycetes
(Eriksson 1966). Palaeoamphisphaerella Ramanujam & Srisailam (1980) appears indistinguishable
from Imprimospora in general and diagnostic morphology. The equatorially disposed pores and
transverse bands constitute the characteristic features of both genera. Imprimospora is therefore
considered a later synonym of Palaeoamphisphaerella.
1.34.1. Species: P. keralensis Ramanujam & Srisailam 1980 (Fig. 8H); Index Fungorum
Registration Identifier: 109135; Location: Palayangadi, Kannur District and Cheruvattur,
Kasaragod District, Kerala, India; Age: Miocene; Notes: The spore wall of this species is
scabrate, unlike the psilate wall of the type species. According to Ramanujam & Srisailam
(1980), Palaeoamphisphaerella keralensis shows significant resemblance to the ascospores of
Amphisphaerella dispersella (Nyl.) O.E. Erikss. 1966 [Current name: Rosellinia dispersella
(Nyl.) P. Karst.].
1.34.2. Species: P. pirozynskii Ramanujam & Srisailam 1980 (Fig. 8I); Index Fungorum
Registration Identifier: 109136; Location: Palayangadi, Kannur District and Cheruvattur,
Kasaragod District, Kerala, India; Age: Miocene.
1.34.3. Species: P. tankensis (G. Norris) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483485; Basionym: Imprimospora tankensis G. Norris 1986; Location: Imperial
Nuktak C–22 Well, Mackenzie Delta Region, District of Mackenzie, North West Territory,
Canada; Age: Eocene; Notes: Imprimospora tankensis, the type of Imprimospora G. Norris,
appears to be similar in general and diagnostic morphology to Palaeoamphisphaerella
pirozynskii described by Ramanujam and Srisailam (1980). The characteristic central part of
the spore is described by Ramanujam and Srisailam (1980) as multiporate with 8–10
equatorial, equidistant pores with prominently thickened margin, whereas Norris (1986)
described it as having 7 or 8 parallel fissures, extending longitudinally between two
transverse, wide ‘shadow bands’. Both the species are regarded as unicellular and aseptate.
Kalgutkar & Jansonius (2000), therefore, treated Imprimospora as a later synonym of
Palaeoamphisphaerella, and transferred its type species to the latter genus.

1.35. Genus: PALAEOPERICONIA C.G. Ibáñez & Zamuner, Mycotaxon 59: 138 (1996) (nom.
inval.) fide Index Fungorum (2021); Index Fungorum Registration Identifier: 27673; Type: P.
fritzschei C.G. Ibáñez & Zamuner 1996.
Original Diagnosis (Combined description): Mycelium immersed, organized in layers (pl. 1,
figs. 1–2); composed of brown hyphae, 4.3–9.6 μm wide, septa being infrequent, walls thin and
delicate with sparse rounded ornamental elements (pl. 1, fig. 6). Conidiophores micronematous,
mononematous. Conidiogenous cells blastic, terminal, with a blistered end. Blastoconidia globose,
brown or dark brown, with verrucose walls (pl. 1, fig. 3), 6.7–14.4 µm, formed singly or in
basipetal chains of up to nine elements (pl. 1, fig. 2–5) from compact apical heads. Spherical
terminal chlamydospore 37.4 × 41.3 μm, with a thick wall (2.4–3.4 μm wide), and with contracted
content, with a supporting hyphal end 8.2 μm in diameter (pl. 1, fig. 6) (Ibañez & Zamuner 1996).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One (not validly published).
Notes: Ibáñez and Zamuner (1996) stated that the studied material was made up of only
asexual structures represented by conidia produced on single conidiophores, not enclosed in
pycnidia or acervuli (conidiomata). This allowed them to assign it within Hyphomycetes -
Moniliales. The presence of conidia and mycelium pigmented, rough-walled and conidial chains,
would relate the sample studied to several Dematiaceae genera. The closely related genera would
be Periconia Tode, Torula Pers., Stachybotrys Corda, Humicola Traaen, Thermomyces Tsikl. and
Chlamydomyces Bainier. The studied material is the first fossil record related to Periconia Tode.
The presence of Palaeopericonia suggests the existence of a warm temperate and very wet habitat.

724
However, the genus was not validly published because its type species was not validly published
and therefore no type was available.
1.35.1. Species: P. fritzschei C.G. Ibáñez & Zamuner 1996 (nom. inval.) fide hoc loco; Index
Fungorum Registration Identifier: 415737; Location: Jaramillo Petrified Forest, Santa Cruz
Province, Argentina; Age: Middle Jurassic; Notes: The species was not validly published
because no holotype was designated.

1.36. Genus: PARAPOTAMOMYCES O’Keefe, Palynology 41(S1): 319 (2017); Index Fungorum
Registration Identifier: 821911; Type: P. maydiformis O’Keefe 2017.
Original Diagnosis: Spores biconical, dark brown, unicellular, with a germ pore at one end.
Thickened verrucae are distributed in three roughly linear bands, one around the equator and one
half-way up each pyramid, or a spiral pattern. Additional verrucae may occur between the upper
bands and the apices (O’Keefe 2017).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
Notes: According to O’Keefe (2017), this genus has many more verrucae than any recorded
species of Potamomyces K.D. Hyde. A similar grain was reported as cf. Gliomastix by Jarzen &
Elsik (1986) from modern fluvial sediments from Zambia, Africa, but is clearly much too big to be
Gliomastix. It is roughly the same size as this grain, but has less robust verrucae. The specimen of
Jarzen & Elsik (1986) looks much more like a Potamomyces with additional tubercles. The generic
name is given because of its similarities to Potamomyces.
1.36.1. Species: P. maydiformis O’Keefe 2017 (Fig. 8J); Index Fungorum Registration Identifier:
821912; Location: Tumbes Province, Peru; Age: Miocene (Heath Formation); Notes: The
species epithet refers to the verrucae looking like corn kernels stuck to the grain; thus,
maydiformis after Zea mays L.

1.37. Genus: PEZIZASPORITES T.C. Huang, Taiwania 26: 47 (1981); Index Fungorum
Registration Identifier: 21234; Type: P. taiwanensis T.C. Huang 1981
Original Diagnosis: Fungal spore with thin reticulate exine
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
1.37.1. Species: P. taiwanensis T.C. Huang 1981 (Fig. 8K); Index Fungorum Registration
Identifier: 107941; Location: Taiwan; Age: Miocene (Peliao Sandstone).

1.38. Genus: POLYPORISPORITES Hammen, Bol. Geol. (Bogota) 2(1): 83 (1954); Index
Fungorum Registration Identifier: 21259; Type: P. elongatus Hammen 1954; Current name:
MONOPORISPORITES Hammen 1954 fide Kalgutkar & Jansonius (2000).
Original Diagnosis: Fungal spores with more than three small pores. (Van der Hammen
1954).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One (the single species has been transferred to Monoporisporites
Hammen 1954).
Notes: Polyporisporites Hammen is a later taxonomic synonym of Monoporisporites
Hammen 1954 sensu Kalgutkar & Jansonius 2000.
1.38.1. Species: P. elongatus Hammen 1954; Index Fungorum Registration Identifier: 337477;
Current name: Monoporisporites elongatus (Hammen) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).

1.39. Genus: PORTALITES Hemer & Nygreen, Micropaleontology 13: 187 (1967); Index
Fungorum Registration Identifier: 25609; Type: P. confertus Hemer & Nygreen 1967.

725
 Original Diagnosis: Amb circular to oval. Thick walled, with a portion of the wall thickened
and containing a simple circular pore, from which a canal extends to the interior (Hemer &
Nygreen 1967).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
Notes: Hemer & Nygreen (1967) knew of no aplanospore of algae containing porate-
canalicular structure of the type exhibited by this form, but did not eliminate possible relationship
with the algae. Elsik (1992) included Portalites and the type species P. confertus in the group
Amerosporae of fossil fungi, and gave an emended diagnosis and description of the one-celled,
monoporate genus.
1.39.1. Species: P. confertus Hemer & Nygreen 1967 (Fig. 8L); Index Fungorum Registration
Identifier: 562009; Location: Borehole ST–8, Arabian American Oil Company’s Stratigraphic
Test No. 8 in Saudi Arabia; Age: Early Carboniferous.

1.40. Genus: PSIAMMOPOMOPIOSPORA Sal.-Cheb. & Locq., C. r. Congr. natn. Socs. sav. Paris,
sect. sci., fasc. 1. (Paleobotanique) 105: 191 (1980); Index Fungorum Registration Identifier:
25597; Type: P. globoides Sal.-Cheb. & Locq. 1980; Current name: MONOPORISPORITES
Hammen 1954 fide Kalgutkar & Jansonius (2000).
Original Diagnosis: Smooth, monoporate, monoapiculate amerospore (Salard-Cheboldaeff &
Locquin 1980).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Three (all the three species, including type species, have been
transferred to other genera).
Notes: With the transfer of type species of Psiammopomopiospora to Monoporisporites
Hammen 1954, the former became a later taxonomic synonym of the latter.
1.40.1. Species: P. ellipsoides Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108420; Current name: Diporisporites ellipsoides (Sal.-Cheb. & Locq.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
1.40.2. Species: P. globoides Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108421; Current name: Monoporisporites globoides (Sal.-Cheb. & Locq.) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.40.3. Species: P. naviculoides Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108422; Current name: Diporisporites naviculoides (Sal.-Cheb. & Locq.) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).

1.41. Genus: PSIAMSPORA Sal.-Cheb. & Locq., C. r. Congr. natn. Socs. sav. Paris, sect. sci., fasc.
1. (Paleobotanique) 105: 191 (1980); Index Fungorum Registration Identifier: 25589; Type: P.
fusiformis Sal.-Cheb. & Locq. 1980.
Original Diagnosis: Smooth, fusiform amerospores (Salard-Cheboldaeff & Locquin 1980).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
Notes: The type species of Psiamspora, P. fusiformis, was described as inaperturate and
fusiform. Inapertisporites Hammen 1954 is only diagnosed as “inaperturate” fungal spores. Hence,
apparently, this genus appears to be similar to Inapertisporites. Kalgutkar & Jansonius (2000),
however, got on loan (from Salard-Cheboldaeff) negative of the photograph of Psiamspora
fusiformis given in the original publication. They observed the negative and found that that there is
no pore or hilum. The spore wall appears to be quite thick, somewhat thinning toward both ends,
but with no more than a possible hint of a flat spot. On the basis of their observation of negative,
they maintained Psiamspora as a genus for inaperturate, rather thick-walled, smooth amerospores
of strictly elongate fusiform shape with narrowly rounded tapered ends.
1.41.1. Species: P. fusiformis Sal.-Cheb. & Locq. 1980 (Fig. 8M); Index Fungorum Registration
Identifier: 107693; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa;

726
Age: Early Eocene-Early Miocene.

1.42. Genus: PSILAINAPERTURITES Pierce, Bull. Minn. geol. Surv., Univ. Minn. 42: 44 (1961);
Index Fungorum Registration Identifier: 581065; Type: P. psilatus Pierce 1961 (gymnosperm
pollen).
Original Diagnosis: Psilate, inaperturate sporomorphs (Pierce 1961).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Two [the type species, P. psilatus Pierce 1961, is a
gymnospermous pollen whereas P. ovalis Mathur 1966 [Current name: Monoporisporites mathurii
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000)] is a fungal spore species].
1.42.1. Species: P. ovalis Mathur 1966; Index Fungorum Registration Identifier: 30413; Current
name: Monoporisporites mathurii Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius
(2000).
1.42.2. Species: P. psilatus Pierce 1961; Botanical affinity: Pinaceae (cf. Laricoipollenites magnus
Potonié 1951); Location: Minnesota, U.S.A.; Age: Early Upper Cretaceous (gymnospermous
pollen).

1.43. Genus: PSILODIPORITES C.P. Varma & Rawat, Grana Palynologica 4(1): 131 (1963);
Index Fungorum Registration Identifier: 25664; Type: P. hammenii C.P. Varma & Rawat 1963
(angiospermous pollen).
Original Diagnosis: Diporate pollen grains with psilate exine (which may sometimes appear
finely scabrate under high power) (Varma & Rawat 1963).
Classification: Angiospermae.
Number of species known: Seven (only four species, which are considered as fungal spores,
have been included in this paper and all of them have been transferred to other genera).
Notes: Varma & Rawat (1963) described their specimens as diporate pollen of angiospermous
plants. Kalgutkar & Jansonius (2000) stated that the type of Psilodiporites, P. hammenii, and also
P. cooksoniae and P. elongatus, named in the same paper, are angiospermous pollen. However,
Varma & Rawat (1963) assigned three species, viz. P. bhardwaji, P. gunniae and P. krempii, to
fungal spores which were later transferred by Kalgutkar & Jansonius (2000) to other genera.
1.43.1. Species: P. bhardwaji C.P. Varma & Rawat 1963; Index Fungorum Registration Identifier:
106421; Current name: Dyadosporites bhardwaji (C.P. Varma & Rawat) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.43.2. Species: P. ellipsoideus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108430; Current name: Dyadosporites ellipsoideus (Sal.-Cheb. & Locq.) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.43.3. Species: P. gunniae C.P. Varma & Rawat 1963; Index Fungorum Registration Identifier:
106426; Current name: Foveodiporites gunniae (C.P. Varma & Rawat 1963) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
1.43.4. Species: P. krempii C.P. Varma & Rawat 1963; Index Fungorum Registration Identifier:
106426; Current name: Biporipsilonites krempii (C.P. Varma & Rawat) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).

1.44. Genus: PUNCTODIPORITES C.P. Varma & Rawat, Grana Palynologica 4(1): 136 (1963);
Index Fungorum Registration Identifier: 28617; Type: P. harrisii C.P. Varma & Rawat 1963;
Current name: FOVEODIPORITES C.P. Varma & Rawat 1963 fide Kalgutkar & Jansonius
(2000).
Original Diagnosis: Diporate grains with punctate exine (Varma & Rawat 1963).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Four (all the species, including type species, have been transferred
to Foveodiporites C.P. Varma & Rawat 1963).

727
 Notes: Punctodiporites C.P. Varma & Rawat 1963 is considered to be a later synonym of
Foveodiporites C.P. Varma & Rawat 1963, as the type species has been transferred to the latter.
1.44.1. Species: P. foedus G. Norris 1997; Index Fungorum Registration Identifier: 483791;
Current name: Foveodiporites foedus (G. Norris) Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
1.44.2. Species: P. granulatus (Rouse) G. Norris 1997; Index Fungorum Registration Identifier:
483790; Current name: Diporisporites pergranulatus (Rouse) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.44.3. Species: P. harrisii C.P. Varma & Rawat 1963; Index Fungorum Registration Identifier:
483373; Current name: Foveodiporites harrisii (C.P. Varma & Rawat) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
1.44.4. Species: P. keshii G. Norris 1997 nom. illeg. fide Index Fungorum (2021).; Index Fungorum
Registration Identifier: 646217; Notes: This is an illegitimate name as the competing
synonym was not validly published.

1.45. Genus: RETICULATASPORITES Ibrahim, Sporenformen des Aegirhorizonts des Ruhr-

Reviers; Dissertation, Technische Hochschule zu Berlin 38 (1933); Type: R. facetus Ibrahim 1933
(Pteridophytic spore).
Original Diagnosis: Spores without trilete mark, and with a measurable reticulate sculpture on
the spore wall; meshes up to 1 μm (Ibrahim 1933).
Classification: Pteridophyta.
Number of species known: Three (the type species, R. facetus Ibrahim 1933, is a fossil
pteridophytic spore and the other two fungal spore species have been transferred to Inapertisporites
Hammen 1954).
1.45.1. Species: R. argentinus R.K. Jain 1968; Index Fungorum Registration Identifier: 486770;
Current name: Inapertisporites argentinus (R.K. Jain) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.45.2. Species: R. cacheutensis R.K. Jain 1968; Index Fungorum Registration Identifier: 646218;
Current name: Inapertisporites cacheutensis (R.K. Jain) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.45.3. Species: R. facetus Ibrahim 1933; Location: Germany; Age: Late Carboniferous
(Pteridophytic spore).

1.46. Genus: RETICULATISPORONITES Elsik, Pollen et Spores 10(2): 274 (1968); Index
Fungorum Registration Identifier: 21279; Type: R. sperryi Elsik 1968; Current name:
MONOPORISPORITES Hammen 1954 fide Kalgutkar & Jansonius (2000).
Original Diagnosis: Elliptical to spatulate, monoporate, reticulate, unicellate fungal spores.
Pore apical (Elsik 1968).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One (the single species has been transferred to Monoporisporites
Hammen 1954).
1.46.1. Species: R. sperryi Elsik 1968; Index Fungorum Registration Identifier: 322416; Current
name: Monoporisporites sperryi (Elsik) Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).

1.47. Genus: RETIDIPORITES C.P. Varma & Rawat, Grana Palynologica 4(1): 137 (1963); Index
Fungorum Registration Identifier: 21280; Type: R. bengalensis C.P. Varma & Rawat 1963.
Original Diagnosis: Diporate grains with reticulate exine (Varma & Rawat 1963).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
1.47.1. Species: R. bengalensis C.P. Varma & Rawat 1963 (Fig. 8N); Index Fungorum Registration
Identifier: 106439; Location: Western and eastern India, including oil exploration areas in

728
 West Bengal and Assam, India; Age: Early-Middle Eocene.

1.48. Genus: SACCISPORONITES Kalgutkar & Janson., AASP Contributions Series (Dallas) 39:
266 (2000); Index Fungorum Registration Identifier: 28624; Type: S. stoughiae (Elsik) Kalgutkar
& Janson. 2000.
Original Diagnosis: One-celled fungal spores, spatulate in overall shape, with a distinct
proximal hilum; spore proper with double wall layer, outer layer thicker than inner one, wall
thickest at distal end; wall turned out at hilum, forming a lip with a characteristic 90° angle at pore;
whole spore loosely enveloped by a filmy perine, that may be lost in preservation or preparation
(Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
Notes: The thin-walled perine is sufficiently persistent to serve as a basis for segregating this
genus from other monocellate spore genera. The development of hilum is also diagnostic.
Kalgutkar & Jansonius (2000) opined that Lacrimasporonites stoughiae Elsik is a misfit in
Lacrimasporonites Elsik and therefore they proposed Saccisporonites to accommodate it. The
genus name was derived from the sac-like membranous perine enveloping the spore.
1.48.1. Species: S. stoughiae (Elsik) Kalgutkar & Janson. 2000 (Fig. 8O); Index Fungorum
Registration Identifier: 483546; Basionym: Lacrimasporonites stoughiae Elsik 1968,
Location: Strip mine approximately 11 km south-west of Rockdale, Milam County, Texas,
U.S.A.; Age: Palaeocene.

1.49. Genus: SCABRADIPORITES Y.K. Mathur, Quarterly Journal of the Geological, Mining &
Metallurgical Society of India 38: 43 (1966); Index Fungorum Registration Identifier: 21292; Type:
S. varias Y.K. Mathur 1966; Current name: DIPORISPORITES Hammen 1954 fide Kalgutkar &
Jansonius (2000).
Original Diagnosis (Combined description): Amb elliptical, 42 × 14 μm in size, diporate,
pores elliptical, larger axis 4 μm. Exine less than 1 μm thick, scabrate, brown (Mathur 1966).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One (the single species has been transferred to Diporisporites
Hammen 1954).
Notes: The specific name is in reference to the variable nature of the spore wall, pore and
size. Kalgutkar & Jansonius (2000) considered this genus to be a later synonym of Diporisporites
Hammen 1954
1.49.1. Species: S. varias Y.K. Mathur 1966; Index Fungorum Registration Identifier: 483818;
Current name: Diporisporites varias (Y.K. Mathur) Kalgutkar & Janson. 2000 fide Kalgutkar
& Jansonius (2000).

1.50. Genus: SENEGALOSPORITES Jardiné & Magloire, Mémoires du Bureau de Recherches

Géologiques et minières, Paris 32: 208 (1965); Index Fungorum Registration Identifier: 21298;
Type: S. costatus Jardiné & Magloire 1965
Original diagnosis: Spores 1-aperturate; lunate, with beak at one end (Jardiné & Magloire
1965).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Two.
1.50.1. Species: S. costatus Jardiné & Magloire 1965; Index Fungorum Registration Identifier:
111026; Location: Senegal; Age: Cretaceous.
1.50.2. Species: S. taiwanensis T.C. Huang 1981; Index Fungorum Registration Identifier: 115817;
Location: Taiwan; Age: Miocene (Talu Shale).

1.51. Genus: SPATULOSPORONITES Z.C. Song in Z.C. Song et al., Fossil Spores and Pollen of
China (Beijing) 1: 825 (1999) (nom. inval.) fide Kalgutkar & Jansonius (2000); Index Fungorum

729
Registration Identifier: 28631; Type: S. minutus Z.C. Song, Qian & Y.H. Zheng in Z.C. Song et al.
1999.
Original Diagnosis: Shape of spore subovate to broadly lanceolate (“spatulo-form”), with at
both bottom and top (at poles of long axis) a finger-like cell projecting outward. Spore composed of
numerous cells, cells arranged both in rows and columns, which produces a regular lattice-like
pattern; all cells of roughly equal size. No pore(s) observed. Overall outline of spore gently
undulate. Colour brown (Song, Qian & Zheng in Song et al. 1999).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One (not validly published).
Notes: The generic name was not validly published for lack of a statement where the type was
deposited.
1.51.1. Species: S. minutus Z.C. Song, Qian & Y.H. Zheng in Z.C. Song 1999 (nom. inval.) fide
Kalgutkar & Jansonius (2000); Index Fungorum Registration Identifier: 483853; Location:
Eastern China, Northern Jiangsu Basin; Age: Early Tertiary (Third Formation of Funing
Group); Notes: Kalgutkar & Jansonius (2000) opined that the overall appearance of this spore
resembles that of Dictyosporites morularis (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000
or that of Dictyosporites symmetricus (V.S. Ediger) Kalgutkar & Janson. 2000.

1.52. Genus: SPIROTREMESPORITES Dueñas, Caldasia 12(60): 539–571 (1979); Index

Fungorum Registration Identifier: 21309; Type: S. simplex Dueñas 1979.
Synonym: Varisulcosporites Rouse & Mustard 1997 fide Kalgutkar & Jansonius (2000),
Index Fungorum Registration Identifier: 28419.
Original Diagnosis: Ellipsoidal to elongate fungal spores possessing one or several spiraling
furrows (Dueñas 1979).
Emended Diagnosis: Psilate, aseptate fungal spores. The aperture is a single furrow at an
angle to the axis of the spore, straight or curved to S-shaped or sigmoidal in outline, or spiral
around the spore axis. The furrow can be short and straight, entirely visible on one face of the
spore; longer and curved; or long and spiral around the outside of the spore. The spore wall is
generally rigid. The spore outline is elongate elliptical to oval, sometimes somewhat reniform in
side view, i.e. with bilateral symmetry. The ends of the spore can be similar or dissimilar; one end
can be truncated by an attachment scar (Elsik 1990a).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: 18 [but we accept only 17 species as legitimate because one
species, viz. S. disciformis (Sheffy & Dilcher) Elsik 1990a, has been transferred here to
Hypoxylonites Elsik 1990a].
Notes: Dueñas (1979) gave the number of apertures as one or several spiraling furrows. Elsik
(1990a) emended the diagnosis to include only spores with one furrow. As contrasted to the
straight, longitudinally parallel furrow of Hypoxylonites, Spirotremesporites has a furrow that is set
at an angle to the axis of the spore, at least for some part of the length of the furrow. This genus has
affinity with Xylariaceae. Varisulcosporites Rouse & Mustard is a later taxonomic synonym of
Spirotremesporites.
1.52.1. Species: S. clinatus Elsik 1990a (Fig. 8P); Index Fungorum Registration Identifier: 130383;
Location: The Gulf Coast, U.S.A.; Age: Neogene; Notes: The species epithet is derived from
Latin clinatus, bent, sloping, slanted.
1.52.2. Species: S. disciformis (Sheffy & Dilcher) Elsik 1990a; Index Fungorum Registration
Identifier: 130347; Current name: Hypoxylonites disciformis (Sheffy & Dilcher) R.K.
Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk comb. nov. This new combination is
described under the section "New species and new combinations"
1.52.3. Species: S. duenasii Elsik 1990a; Index Fungorum Registration Identifier: 130384;
Location: Sabana de Bogota, Colombia, South America; Age: Pleistocene; Notes: The species
epithet is in honour of Dr. Hernando Dueñas-Jimenez.

730
1.52.4. Species: S. ecuatorialis Dueñas 1979; Index Fungorum Registration Identifier: 112635;
Location: Sabana de Bogota, Colombia, South America; Age: Pleistocene.
1.52.5. Species: S. ellipticus Nandi & Shubhra Banerjee in R.K. Saxena 2012 (Fig. 8Q); Index
Fungorum Registration Identifier: 519758; Location: Renkte Kawn-Sherlui Road, Mizoram,
India; Age: Cretaceous-Tertiary (Mahadeo, Langpar, Cherra Sandstone, Siju, Bhuban,
Bokabil, Tipam and Dupitila formations).
1.52.6. Species: S. eminens (Rouse & Mustard) Kalgutkar & Janson. 2000 (Fig. 8R); Index
Fungorum Registration Identifier: 483550; Basionym: Varisulcosporites eminens Rouse &
Mustard 1997; Location: Strait of Georgia, eastern Vancouver Island, the Fraser River
lowlands of southwest British Columbia, Canada and the North-western Washington State,
U.S.A.; Age: Late Eocene-Early Oligocene.
1.52.7. Species: S. longiletus Nandi & Shubhra Banerjee in R.K. Saxena 2012 (Fig. 8S); Index
Fungorum Registration Identifier: 519757; Location: Renkte Kawn-Sherlui Road, Mizoram,
India; Age: Tertiary (Kherapara, Bhuban, Bokabil, Tipam and Dupitila formations).
1.52.8. Species: S. miocenicus Nandi & Shubhra Banerjee in R.K. Saxena 2012 (Fig. 8T); Index
Fungorum Registration Identifier: 519756; Location: Renkte Kawn-Sherlui Road, Mizoram,
India; Age: Tertiary (Kherapara, Bhuban, Bokabil, Tipam, Dupitila and Dihing formations).
1.52.9. Species: S. multiplex Dueñas 1979; Index Fungorum Registration Identifier: 112636;
Location: Sabana de Bogota, Colombia; Age: Pleistocene.
1.52.10. Species: S. neogenicus Elsik 1990a; Index Fungorum Registration Identifier: 130385;
Location: The Gulf Coast, U.S.A.; Age: Neogene.
1.52.11. Species: S. obliquus Elsik 1990a; Index Fungorum Registration Identifier: 130386;
Location: The Green River section, Washington, U.S.A.; Age: Late Eocene.
1.52.12. Species: S. reklawensis Elsik 1990a; Index Fungorum Registration Identifier: 130387;
Location: Texas, U.S.A.; Age: Early Middle Eocene (Reklaw Formation of the Claiborne
Group); Notes: The species epithet is derived from rock strata in which it occurs.
1.52.13. Species: S. reniformis Nandi & Shubhra Banerjee in R.K. Saxena 2012 (Fig. 8U); Index
Fungorum Registration Identifier: 519755; Location: Renkte Kawn-Sherlui Road, Mizoram,
India; Age: Tertiary (Kherapara, Bhuban, Bokabil, Tipam and Dihing formations).
1.52.14. Species: S. shahejiensis Elsik 1990a; Index Fungorum Registration Identifier: 130388;
Location: Leowning Province, China; Age: Eocene-Oligocene (Shahejie Formation); Notes:
The species epithet is derived from rock strata in which it occurs.
1.52.15. Species: S. simplex Dueñas 1979 (Fig. 8V); Index Fungorum Registration Identifier:
112637; Location: Sabana de Bogota, Colombia, South America; Age: Pleistocene.
1.52.16. Species: S. subovoideus Sheffy & Dilcher ex Elsik 1990a; Index Fungorum Registration
Identifier: 130389; Basionym: Inapertisporites subovoideus Sheffy & Dilcher 1971: 38, pl.
15, fig. 7 (only upper one of three specimens illustrated: holotype was designated by Elsik
1990a); Location: Henry County, Tennessee, U.S.A.; Age: Middle Eocene (Claiborne
Group).
1.52.17. Species: S. tertiarus Nandi et al. in R.K. Saxena 2012 (Fig. 8W); Index Fungorum
Registration Identifier: 519754; Location: Renkte Kawn-Sherlui Road, Mizoram, India; Age:
Tertiary (Kherapara, Bhuban, Bokabil, Tipam and Dihing formations).
1.52.18. Species: S. yeguaensis Elsik 1990a; Index Fungorum Registration Identifier: 130390;
Location: Central Texas, U.S.A.; Age: Late-Middle Eocene (Yegua Formation of the
Claiborne Group); Notes: The species epithet is derived from rock strata in which it occurs.

1.53. Genus: STRIADIPORITES C.P. Varma & Rawat, Grana Palynologica 4(1): 137 (1963);
Index Fungorum Registration Identifier: 21324; Type: S. reticulatus C.P. Varma & Rawat 1963.
Synonym: Stridiporosporites P. Ke & Z.Y. Shi 1978 fide Norris (1986); Index Fungorum
Registration Identifier: 21327.
Original Diagnosis: Diporate grains with striated exine (Varma & Rawat 1963).

731
 Emended Diagnosis: Fungal spores of oval to fusiform ambitus and with longitudinally
ribbed to broadly reticulate ornament. Two pores, one at each end of the spore on the long axis.
One cell; no septa, except occasionally a very thin membrane across inner edge of apertures (Elsik
& Jansonius 1974)
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: 15 (but we accept only 14 species as legitimate because one
species, viz. S. anceps G. Norris 1997, has been transferred to Biporipsilonites Kalgutkar & Janson.
2000).
Notes: This genus includes unicellular, diporate fungal spores with striated spore walls. Elsik
& Jansonius (1974) opined that the original diagnosis (“Diporate grains with striated exine”) was
too widely stated and was non-committal regarding the botanical affinity of these grains. The muri
or longitudinal ridges are generally equal to or wider than the exine thickness. This differentiates
the taxon from other diporate fungal spores of similar size and shape with microreticulate, alveolate
or punctate exines. Some morphological similarity exists with the spores of Coelomomyces indicus
Iyengar, as illustrated in Bland & Couch (1973). Striadiporites has been recorded in sediments of
Eocene-Recent, from areas as widely separated as India, Columbia, and Alaska. Norris (1986)
considered that Stridiporosporites P. Ke & Z.Y. Shi is the later synonym of Striadiporites and
formally transferred its type, Stridiporosporites retistriatus, and other species to Striadiporites.
1.53.1. Species: S. anceps G. Norris 1997; Index Fungorum Registration Identifier: 483786;
Current name: Biporipsilonites anceps (G. Norris) Kalgutkar & Janson. 2000 fide Kalgutkar
& Jansonius (2000).
1.53.2. Species: S. asper (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483558; Basionym: Stridiporosporites asper P. Ke & Z.Y. Shi 1978;
Location: Panshan, Liaoning Province; Beidagang, Tianjin Municipality, Coastal region of
Bohai, China; Age: Eocene-Oligocene.
1.53.3. Species: S. bistriatus (P. Ke & Z.Y. Shi) G. Norris 1986; Index Fungorum Registration
Identifier: 126578; Basionym: Stridiporosporites bistriatus P. Ke & Z.Y. Shi 1978; Location:
Panshan, Liaoning Province; Tangjiahe and Beidagang, Tianjin Municipality, Coastal region
of Bohai, China (Ke & Shi 1978), Imperial Nuktak C–22 Well, Mackenzie Delta Region,
District of Mackenzie, Northwest Territories, Canada (Norris 1986); Age: Eocene-Oligocene
(Ke & Shi 1978), Palaeogene (Norris 1986).
1.53.4. Species: S. boletelloides Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108575; Location: Gulf of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene.
1.53.5. Species: S. californicus Elsik & Janson. 1974; Index Fungorum Registration Identifier:
324237; Location: Santa Barbara Channel area, California, U.S.A.; Age: Early Miocene.
1.53.6. Species: S. dolium (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483559; Basionym: Stridiporosporites dolium P. Ke & Z.Y. Shi 1978;
Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
1.53.7. Species: S. inflexus (P. Ke & Z.Y. Shi) G. Norris 1986; Index Fungorum Registration
Identifier: 126579; Basionym: Stridiporosporites inflexus P. Ke & Z.Y. Shi 1978, Location:
Panshan, Liaoning Province; Tangjiahe and Beidagang, Tianjin Municipality, Coastal region
of Bohai, China (Ke & Shi 1978), Imperial Nuktak C–22 Well, Mackenzie Delta Region,
District of Mackenzie, Northwest Territories, Canada (Norris 1986); Age: Eocene-Oligocene
(Ke & Shi 1978), Oligocene (Norris 1986).
1.53.8. Species: S. irregularis Kalgutkar 1993; Index Fungorum Registration Identifier: 483885;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene.
1.53.9. Species: S. minor (Z.C. Song & Z.H. Sun in Z.C. Song et al.) Z.C. Song et al. 1999; Index
Fungorum Registration Identifier: 483846; Basionym: Stridiporosporites minor Z.C. Song &
Z.H. Sun in Z.C. Song et al. 1989; Location: Qingfeng county and Fanxian county of Henan

732
 Province; Shenxian county of Shandong Province, China; Age: Late Eocene-Middle
Oligocene (Shahejie Formation).
1.53.10. Species: S. multistriatus (P. Ke & Z.Y. Shi) G. Norris 1986; Index Fungorum Registration
Identifier: 126580; Basionym: Stridiporosporites multistriatus P. Ke & Z.Y. Shi 1978;
Location: Panshan, Liaoning Province; Coastal region of Bohai, China; Imperial Nuktak C–
22 Well, Mackenzie Delta Region, District of Mackenzie, Northwest Territories, Canada
(Norris 1986); Age: Eocene-Oligocene.
1.53.11. Species: S. reticulatus C.P. Varma & Rawat 1963 (Fig. 8X); Index Fungorum Registration
Identifier: 111055; Location: Western and eastern India, including oil exploration areas in
West Bengal and Assam; Age: Late Oligocene-Early Miocene.
1.53.12. Species: S. retistriatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483560; Basionym: Stridiporosporites retistriatus P. Ke & Z.Y. Shi
1978; Location: Panshan, Liaoning Province; Laoshanggulin and Beidagang, Tianjin
Municipality, Coastal region of Bohai, China; Age: Eocene-Oligocene.
1.53.13. Species: S. sanctaebarbarae Elsik & Janson. 1974; Index Fungorum Registration
Identifier: 324238; Location: Santa Barbara Channel, California, U.S.A.; Age: Late Eocene
and Oligocene.
1.53.14. Species: S. spiralis Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483561; Basionym: Stridiporosporites spiralis Z.C. Song & G.X. Li in Z.C. Song et al. 1989;
Location: Qingfeng county of Henan Province, China; Age: Late Eocene-Early Oligocene
(Shahejie Formation).
1.53.15. Species: S. taiwanensis T.C. Huang 1981; Index Fungorum Registration Identifier:
115822; Location: Taiwan; Age: Miocene (Peliao Sandstone).

1.54. Genus: STRIDIPOROSPORITES P. Ke & Z.Y. Shi, Early Tertiary Spores and Pollen Grains
from the Coastal Region of Bohai (Beijing): 47 (1978); Index Fungorum Registration Identifier:
21327; Type: S. retistriatus P. Ke & Z.Y. Shi 1978; Current name: STRIADIPORITES C.P.
Varma & Rawat 1963 fide Norris (1986).
Original Diagnosis: Spores one-celled and diporate, pores situated at opposite, acute ends of
spore. Pores simple or compound, vestibulate or labiate, etc. Spore surface beset with ridges of
various types, ridges regular or irregular, may anastomose to form reticulum (Ke & Shi 1978).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Eight (Since Stridiporosporites P. Ke & Z.Y. Shi 1978 is a later
synonym of Striadiporites C.P. Varma & Rawat 1963, all its species were transferred to
Striadiporites C.P. Varma & Rawat 1963).
Notes: Song in Song et al. (1999) formulated an emended diagnosis, and simultaneously
proposed the species Stridiporosporites dolium as the new type for the genus. This is contrary to
the rules. In fact, Song in Song et al. (1999) introduced a heterotypic later homonym. Norris (1986)
considered Stridiporosporites to be a later synonym of Striadiporites C.P. Varma & Rawat 1963
and therefore all species of former were transferred to the latter, i.e. Striadiporites.
1.54.1. Species: S. asper P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier: 115674;
Current name: Striadiporites asper (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.54.2. Species: S. bistriatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115673; Current name: Striadiporites bistriatus (P. Ke & Z.Y. Shi) G. Norris 1986 fide
Norris (1986).
1.54.3. Species: S. dolium P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier: 115672;
Current name: Striadiporites dolium (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
1.54.4. Species: S. inflexus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115689; Current name: Striadiporites inflexus (P. Ke & Z.Y. Shi) G. Norris 1986 fide Norris
(1978).

733
1.54.5. Species: S. minor Z.C. Song & Z.H. Sun in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 485248; Current name: Striadiporites minor (Z.C. Song & Z.H. Sun
in Z.C. Song et al.) Z.C. Song et al. 1999 fide Song et al. 1999.
1.54.6. Species: S. multistriatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115638; Current name: Striadiporites multistriatus (P. Ke & Z.Y. Shi) G. Norris 1986 fide
Norris (1986).
1.54.7. Species: S. retistriatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115637; Current name: Striadiporites retistriatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
1.54.8. Species: S. spiralis Z.C. Song & G.X. Li in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 485249; Current name: Striadiporites spiralis (Z.C. Song & G.X. Li
in Z.C. Song et al.) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).

1.55. Genus: TRICHOSPORITES Félix, Zeitschr. Deutsche Geol. Gesell. 46: 273 (1894); Index
Fungorum Registration Identifier: 21341; Type: T. conwentzii Félix 1984 (holotype: in Conwentz
1892).
Original Diagnosis: I designate...those fossil conidia as Trichosporites... that resemble the
spores of the extant Trichosporium to such an extent that a natural affinity of the latter cannot be
excluded (Felix 1894, adopted from Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
Notes: Felix (1894) commented that these conidia greatly resemble those of some
Trichosporium species (e.g. T. fuscum Link). Several species of this genus do indeed live on
decaying wood, such as T. fuscescens and T. splenicum, while T. fuscum is found on rotting fir
branches. These fossil conidia were therefore described as Trichosporites.
1.55.1. Species: T. conwentzii Félix 1984 (Fig. 8Y); Index Fungorum Registration Identifier:
207941; Location: Near Ryedal, Sweden; Age: Late Cretaceous (Holma Sandstone); Notes:
This species was named after H. Conwentz.

1.56. Genus: TRIPORISPORITES Hammen, Bol. Geol. (Bogota) 2(1): 83 (1954); Index Fungorum
Registration Identifier: 21345; Type: T. minutus Hammen 1954; Current name:
INAPERTISPORITES Hammen 1954 fide Kalgutkar & Jansonius (2000).
Original Diagnosis: Fungal spores with three small pores. (Van der Hammen 1954).
Classification: Fungi Imperfecti, Amerosporae
Number of species known: One (the single species has been transferred to Inapertisporites
Hammen 1954).
Notes: Kalgutkar & Jansonius (2000) interpreted the three pores in the type specimen as
being germinal or degradation pores, the number and placement of which have no taxonomic value.
For that reason, they included the genus under Inapertisporites.
1.56.1. Species: T. minutus Hammen 1954; Index Fungorum Registration Identifier: 340461;
Current name: Inapertisporites triporatus Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).

1.57. Genus: TRIPORISPORONITES Sheffy & Dilcher, Palaeontographica Abt. B 133(1-3): 49

(1971); Index Fungorum Registration Identifier: 21346; Type: T. ovalis Sheffy & Dilcher 1971;
Current name: INAPERTISPORITES Hammen 1954 fide Kalgutkar & Jansonius (2000).
Original Diagnosis: Unicellate fungal spore, psilate, aseptate, triporate. Shape variable
(Sheffy & Dilcher 1971).
Classification: Fungi Imperfecti, Amerosporae
Number of species known: Two (both the species have been transferred to other genera).

734
1.57.1. Species: T. ovalis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier: 111989;
Current name: Inapertisporites ovalis Sheffy & Dilcher 1971 fide Kalgutkar & Jansonius
(2000).
1.57.2. Species: T. verus (P. Ke & Z.Y. Shi) Norris 1986; Index Fungorum Registration Identifier:
126581; Current name: Biporipsilonites verus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).

1.58. Genus: UNCINULITES Pampal., Pilzfunde aus der Augsburger Umgebung 8: 125 (1902);
Index Fungorum Registration Identifier: 21350; Type: U. baccarinii Pampal. 1902.
Synonym: Graamspora Sal.-Cheb. & Locq. 1980 fide Kalgutkar & Jansonius (2000), Index
Fungorum Registration Identifier: 25590.
Original Diagnosis (Combined description): Thin walled, dark, astomate, subglobose
perithecia 30–35 µm; with 18–25 simple appendages with hooked tips; appendages nearly equal to
perithecium, undivided, black at their base, dark-coloured at the tip (Pampaloni 1902).
Emended Diagnosis: These are small globose bodies measuring 20–22 μm in diameter,
covered when ripe, on the outside with straight or curved filiform or spine-like projections - the so-
called “appendages” - that attain sometimes to half the diameter of the globose body (Figs. 1–2).
Both the central globose body and the projections are brownish in colour. These globose bodies,
which are termed “perithecia” by Pampaloni (1902) in the as to be rendered opaque (Fig. 1); in a
few cases, however, individuals - evidently in a younger stage of development, as is shown by the
almost complete absence of projections - occur that are more transparent, and in these, under high
power with a strong illumination, it can be seen that each globose body is not a compound mass of
cells, but a single cell (Fig. 2). It seems clear, therefore, that each is to be regarded, not as a
perithecium with appendages, but as a single spore with spinous epispore. On the precise
determination of these spores I am not able at present to throw any light. It may be pointed out
further that neither in the small size of the supposed “perithecium” nor in the nature of the
“appendages” do the present objects bear the slightest resemblance to any of the species of
Uncinula of the Erysiphaceae (Salmon 1903).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: Three.
Notes: Salmon (1903) ruled out slightest resemblance of Uncinulites to any of the species of
Uncinula of the Erysiphaceae. Graamspora Sal.-Cheb. & Locq. 1980 is similar to Uncinulites in
having spinose or echinate spores, but in Uncinulites the spores are spherical whereas in
Graamspora they are oval to elongate in outline (Elsik 1992). Spores in both Graamspora and
Uncinulites are unicellular, aporate and typically spinose. The presence of spines on the spore walls
is a consistent morphological character that makes them similar enough to group them in the same
genus. Hence, Kalgutkar & Jansonius (2000) considered Graamspora to be a later taxonomic
synonym of Uncinulites.
1.58.1. Species: U. artuziae V.S. Ediger & Alisan 1989; Index Fungorum Registration Identifier:
125508; Location: Northern Thrace Basin, Turkey; Age: Middle-Late Eocene to Late
Oligocene, Miocene-Pliocene; Notes: The species epithet honours Professor Samime Artüz.
1.58.2. Species: U. baccarinii Pampal. 1902 (Fig. 8Z); Index Fungorum Registration Identifier:
166879; Location: Italy; Age: Middle Miocene (Disodile beds).
1.58.3. Species: U. pezizaffinis (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483571; Basionym: Graamspora pezizaffinis Sal.-Cheb. & Locq.
1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age: Early
Miocene.

1.59. Genus: VARISULCOSPORITES Rouse & Mustard, Palynology 21: 208 (1997); Index
Fungorum Registration Identifier: 28419; Type: V. eminens Rouse & Mustard 1997; Current name:
SPIROTREMESPORITES Dueñaz 1979 fide Kalgutkar & Jansonius (2000).

735
 Original Diagnosis: Unicellate, elliptical to fusiform fungal spores, isopolar, with thickened
polar caps consisting of polar walls (pl. 11, figs. 5, 6). Apertures single, in form of an elliptical
[oblique] sulcus, or a sulcus plus 2 elongate grooves extending between sulcus and poles (Rouse &
Mustard 1997).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One (the single species has been transferred to Spirotremesporites
Dueñas 1979).
Notes: Spores of the type species, Varisulcosporites eminens, are characterized by the
presence of an oblique furrow or groove. Elsik (1990a) emended Spirotremesporites Dueñas 1979,
restricting it to spores with a single oblique furrow. Varisulcosporites is therefore considered to be
a later taxonomic synonym of Spirotremesporites.
1.59.1. Species: V. eminens Rouse & Mustard 1997; Index Fungorum Registration Identifier:
463994; Current name: Spirotremesporites eminens (Rouse & Mustard) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).

1.60. Genus: XYLARIASPORITES Debi Mukh., International Journal of Geology, Earth and
Environmental Sciences 2(2): 9 (2012); Index Fungorum Registration Identifier: 588470; Type: X.
lanceolatus Debi Mukh. 2012; Current name: HYPOXYLONITES Elsik 1990a fide hoc loco.
Original Diagnosis: Ascospore hyaline, brown in colour, one celled, dark colour with slit-like
germ pore; size ranges 60–70 × 30–40 μm (generally 60 × 40 μm); spores lanceolate, acute, pointed
at the poles; germ pore up to 2 μm wide (Mukherjee 2012).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One (the single species is here transferred to Hypoxylonites Elsik
1990a).
Notes: The genus name is derived from Xylaria (Xylariaceae).
1.60.1. Species: X. lanceolatus Debi Mukh. 2012; Index Fungorum Registration Identifier: 588483;
Current name: Hypoxylonites lanceolatus (Debi Mukh.) R.K. Saxena, Wijayaw., D. Q. Dai,
K. D. Hyde & P. M. Kirk comb. nov. This new combination is described under the section
“New species and new combinations”.

1.61. Genus: XYLOHYPHITES Kalgutkar & Sigler, Mycol. Res. 99(5): 514 (1995); Index
Fungorum Registration Identifier: 13324; Type: X. verrucosus Kalgutkar & Sigler 1995.
Original Diagnosis: Conidia in chains formed in acropetal succession, simple, ellipsoidal or
cylindrical, pale brown, verrucose, aseptate, occasionally 1-septate; conidia tapering at each end but
sometimes with slightly protuberant hilum (Kalgutkar & Sigler 1995).
Classification: Fungi Imperfecti, Amerosporae.
Number of species known: One.
Notes: Kalgutkar & Sigler (1995) suggested affinity of Xylohyphites to dematiaceous
blastoconidial fungus. The genus is named after its affinity to Xylohypha (Fr.) E.W. Mason in
Deighton 1960.
1.61.1. Species: X. verrucosus Kalgutkar & Sigler 1995 (Fig. 8AA); Index Fungorum Registration
Identifier: 414405; Location: Mohgaonkalan, Chhindwara District, Madhya Pradesh, India;
Age: Maastrichtian (Deccan Intertrappean Beds); Notes: The species is named after verrucose
nature of the conidia.

2. Didymosporae

2.1. Genus: AMPULLIFERINITES Kalgutkar & Sigler, Mycol. Res. 99(5): 515 (1995); Index
Fungorum Registration Identifier: 14288; Type: A. axelheibergii Kalgutkar & Sigler 1995.
Original Diagnosis: Fungus filamentous; filaments in short or long, determinate chains of
conidia; filaments traversed by thin and thick-walled septa which alternate along the chains; conidia
in arthroconidial chains separated by dark, thick septa; didymosporous, not or slightly constricted at

736
medium septum and with truncate ends except the terminal conidium which is rounded at the apex
(Kalgutkar & Sigler 1995).
Classification: Fungi Imperfecti, Didymosporae.
Number of species known: One.
Notes: Kalgutkar & Sigler (1995) stated that Ampulliferinites appears to be similar to modern
Ampulliferina B. Sutton 1969 which has didymosporous, catenate conidia that separate by
fragmentation through the thick, dark brown septa along its length of arthroconidial chains.
Ampulliferinites is also characterized by the presence of a basal cell with an attachment scar similar
to one present in Ampulliferina. The genus name indicates its affinity to Ampulliferina B. Sutton.
2.1.1. Species: A. axelheibergi Kalgutkar & Sigler 1995 (Fig. 8AB); Index Fungorum Registration
Identifier: 412403; Location: Yukon Territory, North-western Canada; Age: Late Eocene or
Early Oligocene (Amphitheatre Formation); Notes: The species epithet is after its type
locality on Axel Heiberg Island.

2.2. Genus: CALDESITES Puri, Some plant-micro-fossils from the Cretaceous and Paleocene of
Nigeria; University of Ibadan Botanical Studies 10: 16 (1963); Index Fungorum Registration
Identifier: 92235; Type: C. nigerianus Puri 1963.
Original Diagnosis (combined description): These are two large ascospores of more or less
the same size, lying partly one on the other. The larger measures 42 × nearly 20 μm (in the broadest
part). The smaller spore is only a little smaller in dimensions. Both of these seem to be broken at
the basal portion. They are thin-walled and are divided into more or less equal halves by the
equatorial wall (Puri 1963).
Classification: Ascomycota, Microthyriales.
Number of species known: One.
2.2.1. Species: C. nigerianus Puri 1963; Index Fungorum Registration Identifier: 647741;
Location: Nigeria; Age: Senonian?

2.3. Genus: CLADOSPORITES Félix, Zeitschr. Deutsche Geol. Gesell. 46: 276 (1894); Index
Fungorum Registration Identifier: 21055; Type: C. bipartitus Felix 1894.
Original Diagnosis (Combined description): The conidia are elliptical or pear-shaped, smooth
and of pale-brownish coloration. They are divided by a septum into two halves, one of which is
invariably of roundish, the other of occasionally somewhat rounded three-sided shape. The length
of the conidia is 10.2–11.9 μm, the width 5.1–6.8 μm. Special conidiophores are absent or
rudimentary. In the vicinity of the conidia, brown, septate mycelial filaments occur. They are
ramified, here and there of gnarled appearance. Clamp connections are absent (Félix 1894).
Classification: Fungi Imperfecti, Didymosporae.
Number of species known: Three.
Notes: According to Félix (1894), the conidia greatly resemble those of Cephalothecium
Corda (Current name: Trichothecium Link fide Species Fungorum 2021) and Cladosporium Link.
The former has water-clear conidia but the fossil conidia are partially coloured which may have
been caused by the state of preservation of the organic substance in the conidial wall. The
mycelium of Cephalothecium is very different, and an affiliation of the fossil conidia with the
Cladosporieae is therefore more likely. For this reason, it was named Cladosporites.
2.3.1. Species: C. bipartitus Félix 1894 (Fig. 8AC); Index Fungorum Registration Identifier:
201043; Location: Perekeschkul, near Baku, Azerbaijan; Age: Eocene.
2.3.2. Species: C. fasciculatus E.W. Berry 1916; Index Fungorum Registration Identifier: 483913;
Location: In silicified vessels of Laurinoxylon of Westmorland Bluff, Trinity River, Texas,
U.S.A.; Age: Middle Eocene; Notes: According to Berry (1916), this species is found in
abundance in silicified specimens of lauraceous wood from the Yegua Formation, Claiborne
Group (Middle Eocene) of Texas and is entirely unlike any previously recorded fossil forms.

737
2.3.3. Species: C. oligocaenicus E.W. Berry 1916; Index Fungorum Registration Identifier:
483912; Location: In petrified wood of Palmoxylon cellulosum Knowlton, Bayou Pierre,
Mississippi, U.S.A.; Age: Early Oligocene.

2.4. Genus: DICELLAEPORISPORITES Kalgutkar, Review of Palaeobotany & Palynology

(Amsterdam) 97(1-2): 210 (1997); Index Fungorum Registration Identifier: 27773; Type: D.
poratus Kalgutkar 1997.
Original Diagnosis: Spores dicellate, broadly elliptic-fusiform or ellipsoidal, smooth,
constricted or not at the central septum. Cells uniformly broad with rounded ends or broadest near
the central septum and gradually tapering towards the apices. Apices of each cell thickened or not.
Spores with symmetrically placed germinal pores or lateral germinal slits or furrows. Germinal
pores axial or non-axial, one pore or lateral furrow in each cell (Kalgutkar 1997).
Classification: Fungi Imperfecti, Didymosporae.
Number of species known: Three.
Notes: Dicellaeporisporites can be distinguished from Dicellaesporites Elsik 1968 by the
presence of a germinal pore in each cell.
2.4.1. Species: D. delitschiapites (Kalgutkar & Sigler) Kalgutkar 1997; Index Fungorum
Registration Identifier: 437901; Basionym: Dicellaesporites delitschiapites Kalgutkar &
Sigler 1995; Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories,
Canada; Age: Late Palaeocene-Early Eocene; Notes: Spores of this species are generally
similar to the ascospores of the living loculoascomycetous and coprophilous Delitschia in the
presence of furrows which appear similar to the distinctive germ slits.
2.4.2. Species: D. poratus Kalgutkar 1997 (Fig. 8AD); Index Fungorum Registration Identifier:
437900; Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories, Canada;
Age: Late Palaeocene-Early Eocene; Notes: The species epithet is derived from the porate
cells.
2.4.3. Species: D. siglerae (Kalgutkar) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483291; Basionym: Fusiformisporites siglerae Kalgutkar 1997, Location: Kanguk
Peninsula, Axel Heiberg Island, Northwest Territories, Canada; Age: Late Palaeocene-Early
Eocene; Notes: The species epithet is in honour of Dr. Lynne Sigler.

2.5. Genus: DICELLAESPORITES Elsik, Pollen et Spores 10(2): 269 (1968); Index Fungorum
Registration Identifier: 21074; Type: D. popovii Elsik 1968.
Original Diagnosis: Inaperturate, psilate fungal spores. Two cells, uniseptate. Shape variable
(Elsik 1968).
Emended Diagnoses: Diagnosis of the genus Dicellaesporites was emended by Sheffy &
Dilcher (1971) and Norris (1986), as follows: Inaperturate fungal spores or algal bodies. Two cells,
uniseptate, shape variable. Sculpture psilate to scabrate (Sheffy & Dilcher 1971); Dicellate,
inaperturate, isopolar, equilateral fungal spores. Spore wall laevigate to scabrate (Norris 1986).
Classification: Fungi Imperfecti, Didymosporae.
Number of species known: 68 (but we accept only 58 species as legitimate because eight
species have been transferred to other genera and two species, viz. D. bolharensis Soomro et al.
2010 and D. lingulatus Z.C. Song in Z.C. Song et al. 1999 have not been validly published.
Notes: Both Elsik (1968) and Sheffy & Dilcher (1971) mentioned the shape of
Dicellaesporites as “variable”. Norris (1986) did not accept it and emended the diagnosis to restrict
this genus for dicellate aporate spores with isopolar, equilateral cells. Kalgutkar & Jansonius (2000)
stated that dicellate aporate spores may be isopolar with equilaterally exactly similar cells, but some
have unequal cells as well (even on the same mycelium), e.g. in Dicellaesporites paradoxus P. Ke
& Z.Y. Shi 1978 (pl. 1, figs. 10–11), D. inaequabilis Mart.-Hern. & Tom.-Ort. 1989 (fig. 3c), D.
keralaensis P. Kumar 1990 (pl. 1, fig. 12) and Dicellaesporites sp. 1 (Kalgutkar 1997). They,
therefore, did not accept Norris’ emendation. Elsik (1992) suggested an emended description of the
type as having a pore in one of the cells that is not positioned on the axis of symmetry.

738
2.5.1. Species: D. aculeolatus Sheffy & Dilcher 1971 (Fig. 8AE); Index Fungorum Registration
Identifier: 111404; Location: Puryear clay pit, 800 m south of Puryear, Henry County,
Tennessee, U.S.A.; Age: Middle Eocene (Claiborne Formation); Notes: The species name
refers to the somewhat pointed spore apices.
2.5.2. Species: D. africanus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107909; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene-Early Miocene; Notes: Salard-Cheboldaeff & Locquin (1980) assigned an affinity to
Ascomycota.
2.5.3. Species: D. akoyolii V.S. Ediger & Alisan 1989; Index Fungorum Registration Identifier:
125502; Location: Northern Thrace Basin, Turkey; Age: Middle?-Late Eocene to Late
Oligocene, Miocene-Pliocene; Notes: The species epithet is in honour of Dr. Erol Akyol.
2.5.4. Species: D. antarcticus Z.C. Song & Liu Cao 1994; Index Fungorum Registration Identifier:
483765; Location: King George Island, Antarctica; Age: Late Cretaceous; Notes: The species
epithet is derived from the locality of type specimens.
2.5.5. Species: D. appendiculatus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111405; Current name: Hilidicellites appendiculatus (Sheffy & Dilcher) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
2.5.6. Species: D. arabimarinus Sat. K. Srivast. & Al-Tayyar 2013; Index Fungorum Registration
Identifier: 818887; Location: northern Arabian Gulf; Age: Aptian to late Albian; Notes: The
species epithet is after Arabian Sea, Arabimarinus = of the Arabian Sea.
2.5.7. Species: D. asymmetricus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107910; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene-Early Miocene; Notes: Salard-Cheboldaeff & Locquin (1980) assigned an affinity
to Ascomycota.
2.5.8. Species: D. bellus K.F. Wang & Y.L. Zhang 1986; Index Fungorum Registration Identifier:
646634; Location: China; Notes: Song et al. (1999) cited this name. Since its bibliographic
details were not included in the list of references of Song et al. (1999), it is difficult to verify
whether this species was validly published. Kalgutkar & Jansonius (2000) opined that it is
similar to Dicellaesporites camerounensis Sal.-Cheb. & Locq. 1980, D. inaequabilis Mart.-
Hern. & Tom.-Ort. 1989 and D. scaber Mart.-Hern. & Tom.-Ort. 1989.
2.5.9. Species: D. bigeminatus (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483292; Basionym: Multicellaesporites bigeminatus Sheffy & Dilcher
1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.;
Age: Middle Eocene (Claiborne Formation); Notes: According to Kalgutkar & Jansonius
(2000), the illustrated spores are clearly dicellate and catenate and therefore the species was
assigned to Dicellaesporites Elsik 1968, as emended by Dilcher 1971.
2.5.10. Species: D. bolharensis Soomro et al. 2010 (nom. inval.) fide hoc loco; Index Fungorum
Registration Identifier: 637483; Location: Borehole Core DH No. 18 in the Sonda Coalfield,
Thatta District, Sindh, Pakistan; Age: Palaeocene; Notes: The species name was not validly
published because the author did not specify where the holotype was deposited.
2.5.11. Species: D. camerounensis Sal.-Cheb. & Locq. 1980; Index Fungorum Registration
Identifier: 107911; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa;
Age: Oligocene; Notes: Salard-Cheboldaeff & Locquin (1980) assigned an affinity to
Ascomycota.
2.5.12. Species: D. campanulatus Ambwani 1983 (Fig. 8AF); Index Fungorum Registration
Identifier: 106768; Location: Neyveli Lignite Mine, Cuddalore District, Tamil Nadu, India;
Age: Late Miocene or Pliocene; Notes: The species epithet is derived from the campanulate
upper cell.
2.5.13. Species: D. cartos P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier: 115662;
Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.

739
Figure 8 – A–AA Amerosporae. AB–AI Didymosporae. A Monoporisporites ovaliformis (Anil
Chandra et al.) Kalgutkar & Janson. 2000, Bar = 10 μm. B Monoporisporites psilatus Anil Chandra
et al. 1984, Bar = 20 μm. C Monoporisporites sheffyi Anil Chandra et al. 1984, Bar = 10 μm. D
Monoporisporites singhii A. Gupta 2002, Bar = 10 μm. E Monoporisporites stoveri Elsik 1968, Bar
= 5 μm. F Nailisporites taiwanensis T.C. Huang 1981, Bar = 10 μm. G Nigrosporites neyveliensis
Debi Mukh. 2012, Bar = 12 μm. H Palaeoamphisphaerella keralensis Ramanujam & Srisailam
1980, Bar = 10 μm. I Palaeoamphisphaerella pirozynskii Ramanujam & Srisailam 1980, Bar = 10
μm. J Parapotamomyces maydiformis O’Keefe 2017, Bar = 10 μm. K Pezizasporites taiwanensis
T.C. Huang 1981, Bar = 10 μm. L Portalites confertus Hemer & Nygreen 1967, Bar = 8 μm. M
Psiamspora fusiformis Sal.-Cheb. & Locq. 1980, Bar = 8 μm. N Retidiporites bengalensis C.P.

740
Varma & Rawat 1963, Bar = 15 μm. O Saccisporonites stoughiae (Elsik) Kalgutkar & Janson.
2000, Bar = 10 μm. P Spirotremesporites clinatus Elsik 1990a, Bar = 10 μm. Q Spirotremesporites
ellipticus Nandi & Shubhra Banerjee 2012, Bar = 6 μm. R Spirotremesporites eminens (Rouse &
Mustard) Kalgutkar & Janson. 2000, Bar = 10 μm. S Spirotremesporites longiletus Nandi &
Shubhra Banerjee 2012, Bar = 7 μm. T Spirotremesporites miocenicus Nandi & Shubhra Banerjee
2012, Bar = 5 μm. U Spirotremesporites reniformis Nandi & Shubhra Banerjee 2012, Bar = 6 μm.
V Spirotremesporites simplex Dueñas 1979, Bar = 5 μm. W Spirotremesporites tertiarus Nandi et
al. 2012, Bar = 5 μm. X Striadiporites reticulatus C.P. Varma & Rawat 1963, Bar = 20 μm. Y
Trichosporites conwentzii Félix 1984, Bar = 5 μm. Z Uncinulites baccarinii Pampal. 1902, Bar = 8
μm. AA Xylohyphites verrucosus Kalgutkar & Sigler 1995, Bar = 5 μm. AB Ampulliferinites
axelheibergi Kalgutkar & Sigler 1995, Bar = 10 μm. AC Cladosporites bipartitus Félix 1894, Bar =
7 μm. AD Dicellaeporisporites poratus Kalgutkar 1997, Bar = 10 μm. AE Dicellaesporites
aculeolatus Sheffy & Dilcher 1971, Bar = 5 μm. AF Dicellaesporites campanulatus Ambwani
1983, Bar = 20 μm. AG Dicellaesporites classicus R.K. Saxena & S.K.M. Tripathi 2011, Bar = 25
μm. AH Dicellaesporites constrictus S.C.D. Sah & R.K. Kar 1974, Bar = 20 μm. AI
Dicellaesporites disphaericus Sheffy & Dilcher 1971, Bar = 10 μm.

2.5.14. Species: D. cellaequalis Kalgutkar 1993; Index Fungorum Registration Identifier: 483862;
Location: Peel River, Yukon Territory, Canada; Location: Peel River, Yukon Territory,
Canada; Age: Late Palaeocene-Early Eocene; Notes: The species epithet is derived from the
Latin, cella, cell; aequalis, equal, referring to the spores having both cells equal.
2.5.15. Species: D. classicus R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 8AG); Index Fungorum
Registration Identifier: 561702; Location: Sonapur-Badarpur Road Section, Jaintia Hills,
Meghalaya and Cachar District, Assam, India; Age: Early Miocene (Bhuban Formation)
2.5.16. Species: D. constrictus S.C.D. Sah & R.K. Kar 1974 (Fig. 8AH); Index Fungorum
Registration Identifier: 519767; Location: Palana, Bikaner District, Rajasthan, India; Age:
Early Eocene (Palana lignite).
2.5.17. Species: D. crassiseptus Ramanujam & Srisailam 1980; Index Fungorum Registration
Identifier: 108842; Current name: Hilidicellites dubius (Ramanujam & Srisailam) Kalgutkar
& Janson. 2000 fide Kalgutkar & Jansonius (2000).
2.5.18. Species: D. delitschiapites Kalgutkar & Sigler 1995; Index Fungorum Registration
Identifier: 412946; Current name: Dicellaeporisporites delitschiapites (Kalgutkar & Sigler)
Kalgutkar 1997 fide Kalgutkar (1997).
2.5.19. Species: D. disphaericus Sheffy & Dilcher 1971 (Fig. 8AI); Index Fungorum Registration
Identifier: 111406; Location: Puryear clay pit, 800 m south of Puryear, Henry County,
Tennessee, U.S.A.; Age: Middle Eocene (Claiborne Formation).
2.5.20. Species: D. dolium Z.C. Song 1985; Index Fungorum Registration Identifier: 637484;
Location: Huangshi and Dafengshan, Qaidam Basin, Qinghai Province, China; Age:
Palaeocene-Late Eocene; Early Miocene-Late Pliocene.
2.5.21. Species: D. ellipticus K.P. Jain & R.K. Kar 1979 (Fig. 9A); Index Fungorum Registration
Identifier: 112269; Location: Papanasam and Varkala, Thiruvananthapuram District, Kerala,
India; Age: Miocene.
2.5.22. Species: D. elongatus Ramanujam & K.P. Rao 1978 (Fig. 9B); Index Fungorum
Registration Identifier: 115060; Location: Kannur, Kerala, India; Age: Miocene (Quilon and
Warkalli beds); Notes: The elongate nature of the cells and the considerably thickened
transverse septum are the important features of this species.
2.5.23. Species: D. elongatus P. Kumar 1990; Index Fungorum Registration Identifier: 126551;
Current name: Dicellaesporites perelongatus Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
2.5.24. Species: D. elongatus Z.C. Song 1985; Index Fungorum Registration Identifier: 485267;
Current name: Dicellaesporites largelongatus Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).

741
2.5.25. Species: D. elsikii B. Samant in R.K. Saxena 2009 (Fig. 9C); Index Fungorum Registration
Identifier: 515018; Synonym: Dicellaesporites elsikii B. Samant 2000 (nom. inval.) fide
Saxena (2009); Location: Near Bhavnagar, Cambay Basin, Gujarat, India; Age: Early Eocene
(Kharsalia Clay Formation).
2.5.26. Species: D. foratus Zhong Y. Zhang 1980; Index Fungorum Registration Identifier:
485000; Location: Lunpola Basin, northern Xizang Plateau, Xizang, China; Age: Oligocene
(Niubao Formation).
2.5.27. Species: D. fragilis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111407; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation).
2.5.28. Species: D. fusiformis Sheffy & Dilcher 1971 (Fig. 9D); Index Fungorum Registration
Identifier: 111408; Location: Puryear clay pit, 800 m south of Puryear, Henry County,
Tennessee, U.S.A.; Age: Middle Eocene (Claiborne Formation).
2.5.29. Species: D. granulatus Z.C. Song 1985; Index Fungorum Registration Identifier: 637485;
Location: Dafengshan, Qaidam Basin, Qinghai Province, China; Age: Early Pliocene.
2.5.30. Species: D. granuliformis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111409; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation).
2.5.31. Species: D. guineensis Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107678; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene-Early Miocene; Notes: Salard-Cheboldaeff & Locquin (1980) assigned an affinity
to Ascomycota. Kalgutkar & Jansonius (2000) stated that there is a suggestion in the
photograph of (roughly) longitudinal striations running from one end of the spore to the other.
Although these may represent foreign tissue under- or overlying the specimen, a similar
feature was reported for D. striatus.
2.5.32. Species: D. himachalensis R.K. Saxena & A.P. Bhattach. 1990 (Fig. 9E); Index Fungorum
Registration Identifier: 519768; Location: Manjhi Khad section, Dharmsala, Kangra District,
Himachal Pradesh, India; Age: Oligocene-Early Miocene.
2.5.33. Species: D. himalayaensis A. Gupta 2002 (Fig. 9F); Index Fungorum Registration
Identifier: 540463; Location: Jamtah Road Section, Sirmaur District, Himachal Pradesh,
India; Age: Eocene (Subathu Formation).
2.5.34. Species: D. inaequabilis Mart.-Hern. & Tom.-Ort. 1989; Index Fungorum Registration
Identifier: 483798; Location: Piedras Negras, Coahuila State, Mexico; Age: Maastrichtian;
Notes: The species epithet indicates its cells are of different size.
2.5.35. Species: D. inaequalis (Y.N.R. Varma & R.S. Patil) Kalgutkar & Janson. 2000 (Fig. 9G);
Index Fungorum Registration Identifier: 483293; Basionym: Dyadosporonites inaequalis
Y.N.R. Varma & R.S. Patil 1985; Location: Tonakkal area, Thiruvananthapuram District,
Kerala, India; Age: Miocene; Notes: Dyadosporonites inaequalis is characterized by having
unequal-sized dicellate condition. Kalgutkar & Jansonius (2000) observed that the
photograph of the type does not have axial pores at both ends as suggested by the original
generic assignment. The two “pores” mentioned in the original description apparently refer to
two cracks in the central septum. This form was therefore transferred to Dicellaesporites.
2.5.36. Species: D. indicus A. Gupta 2002 (Fig. 9H); Index Fungorum Registration Identifier:
540464; Location: Jamtah Road Section, Sirmaur District, Himachal Pradesh, India; Age:
Eocene (Subathu Formation).
2.5.37. Species: D. jainii R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 9I); Index Fungorum
Registration Identifier: 519938; Location: Barmer Hill, Barmer District, Rajasthan, India;
Age: Palaeocene (Barmer Sandstone); Notes: The species epithet honours Dr. K.P. Jain,
Birbal Sahni Institute of Palaeosciences, Lucknow, India.
2.5.38. Species: D. keralensis P. Kumar 1990 (Fig. 9J); Index Fungorum Registration Identifier:
126552; Location: Padappakkara, Kollam District, Kerala, India; Age: Early-Middle
Miocene.

742
2.5.39. Species: D. largelongatus Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483294; Basionym: Dicellaesporites elongatus Z.C. Song 1985; Location:
Huatugou and Dafengshan, Qaidam Basin, Qinghai Province, China; Age: Palaeocene - Late
Eocene; Early-Late Miocene; Notes: Dicellaesporites elongatus Z.C. Song 1985 was a later
homonym of Dicellaesporites elongatus Ramanujam & K.P. Rao 1978. Hence, Kalgutkar &
Jansonius (2000) proposed a new name (D. largelongatus) for it. Spores of Dicellaesporites
reniformis Zhong Y. Zhang 1980 are larger, and somewhat reniform.
2.5.40. Species: D. lenghuensis Z.C. Song 1985; Index Fungorum Registration Identifier: 637486;
Location: Eboliang and Dafengshan, Qaidam Basin, Qinghai Province, China; Age:
Palaeocene - Late Eocene; Early Miocene – Early Pliocene.
2.5.41. Species: D. levis Sheffy & Dilcher 1971 (Fig. 9K); Index Fungorum Registration Identifier:
111410; Location: Puryear clay pit, 800 m south of Puryear, Henry, County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation).
2.5.42. Species: D. lingulatus Z.C. Song in Z.C. Song et al. 1999 (nom. inval.) fide Kalgutkar &
Jansonius (2000); Index Fungorum Registration Identifier: 483822; Location: Coastal area of
Bohai Sea, China; Age: Middle-Late Oligocene (Shahejie-Dongying formations); Notes: This
name was not validly published because the author did not specify where the holotype is
deposited, and did not provide a Latin description or its English translation.
2.5.43. Species: D. littoralis Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107912; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene-Early Miocene; Notes: Salard-Cheboldaeff & Locquin (1980) assigned an affinity
to Ascomycota.
2.5.44. Species: D. longus (Trivedi & C.L. Verma) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483295; Basionym: Teleutosporites longus Trivedi & C.L. Verma
1970 (nom. inval.); Location: Near Kuala Lumpur, Malaya; Age: Eocene; Notes: The species
name was not validly published by Trivedi & Verma (1970) because it was not assigned to a
validly published generic name.
2.5.45. Species: D. magnus Doub. & D. Pons 1973; Index Fungorum Registration Identifier:
485252; Current name: Reduviasporonites magnus (Doub. & D. Pons) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
2.5.46. Species: D. major P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier: 115661;
Location: Beidagang, Trianjin Municipality, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
2.5.47. Species: D. megafusiformis Z.C. Song & G.X. Li in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 637487; Location: Qingfeng county of Henan Province, China; Age:
Late Eocene (Shahejie Formation).
2.5.48. Species: D. minutus R.K. Kar & R.K. Saxena 1976 (Fig. 9L); Index Fungorum Registration
Identifier: 112270; Location: Bhuj-Lakhpat Road, Matanomadh Village, District of Kutch,
Gujarat, India; Age: Palaeocene.
2.5.49. Species: D. mollis P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier: 115660;
Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene; Notes: Elsik et al. (1990) considered that Dicellaesporites mollis possibly is an
aberrant form of Anatolinites dongyingensis.
2.5.50. Species: D. nakomanii V.S. Ediger & Alisan 1989; Index Fungorum Registration Identifier:
125504; Location: Northern Thrace Basin, Turkey; Age: Middle?-Late Eocene to Late
Oligocene, Miocene-Pliocene; Notes: The species epithet is in honour of Professor Eran
Nakoman.
2.5.51. Species: D. navicularis K.F. Wang & Y.L. Zhang 1986; Index Fungorum Registration
Identifier: 637487; Location: China; Notes: Song et al. (1999) cited this name. Since its
bibliographic details were not included in the list of references of Song et al. (1999), it is
difficult to verify whether this species was validly published. Kalgutkar & Jansonius (2000)

743
 opined that it is similar to Dicellaesporites obnixus G. Norris 1986 and D. perelongatus
Kalgutkar & Janson. 2000.
2.5.52. Species: D. nodusus V.S. Ediger 1981; Index Fungorum Registration Identifier: 107913;
Location: Thrace Basin, Turkey; Age: Late Eocene-Oligocene, Miocene-Pliocene.
2.5.53. Species: D. oblongatus Z.C. Song & Liu Cao 1994; Index Fungorum Registration
Identifier: 483764; Location: King George Island, Antarctica; Age: Late Cretaceous; Notes:
The species epithet is derived from ellipsoidal shape of spores.
2.5.54. Species: D. obnixus G. Norris 1986; Index Fungorum Registration Identifier: 126571;
Location: Imperial Nuktak C–22 Well, Mackenzie Delta Region, District of Mackenzie,
Northwest Territories, Canada; Age: Eocene; Notes: Kalgutkar & Jansonius (2000) stated that
although the holotype appears to be dicellate, other specimens appear to have (3?-) 4 cells.
The latter also seem to have a curved longitudinal axis.
2.5.55. Species: D. ovatus Z.C. Song & H.C. Luo in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 637489; Location: Shenxian county of Shandong Province, China;
Age: Late Eocene-Middle Oligocene (Shahejie Formation).
2.5.56. Species: D. paradoxus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115659; Current name: Disparidicellites paradoxus (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
2.5.57. Species: D. perelongatus Kalgutkar & Janson. 2000 (Fig. 9M); Index Fungorum
Registration Identifier: 483296; Basionym: Dicellaesporites elongatus P. Kumar 1990;
Location: Padappakkara, Kollam District, Kerala, India; Age: Early-Middle Miocene; Notes:
Dicellaesporites elongatus P. Kumar 1990 was a later homonym of Dicellaesporites
elongatus Ramanujam & K.P. Rao 1978. Hence, Kalgutkar & Jansonius (2000) proposed a
new name (D. perelongatus) for it.
2.5.58. Species: D. popovii Elsik 1968 (Fig. 9N); Index Fungorum Registration Identifier: 312948;
Location: Strip mine approximately 11 km southwest of Rockdale, Milam County, Texas,
U.S.A.; Age: Palaeocene.
2.5.59. Species: D. reniformis Zhong Y. Zhang 1980; Index Fungorum Registration Identifier:
485001; Location: Lunpola Basin, northern Xizang Plateau, Xizang, China; Age: Oligocene
(Niubao Formation).
2.5.60. Species: D. rinconii Doub. & D. Pons 1973; Index Fungorum Registration Identifier:
637490; Location: Cerrejon basin, Colombia, South America; Age: Palaeocene-Eocene; The
species is dedicated to M. Rincon, curator of the “Servicio Geológico Nacional” collection in
Bogota.
2.5.61. Species: D. scaber Mart.-Hern. & Tom.-Ort. 1989; Index Fungorum Registration Identifier:
483797; Location: Piedras Negras, Coahuila State, Mexico; Age: Maastrichtian; Notes: The
species epithet indicates its rough ornamentation.
2.5.62. Species: D. septiconstrictus Kalgutkar 1993; Index Fungorum Registration Identifier:
483863; Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene;
Notes: The species epithet is derived from the Latin, septum, wall; constrictus, contracted,
referring to the spores being constricted at the septum.
2.5.63. Species: D. singhii R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 9O); Index Fungorum
Registration Identifier: 519939; Location: Hoshiarpur-Una Road Section, Hoshiarpur District,
Punjab and Una District, Himachal Pradesh, India; Age: Pliocene (Upper Siwalik); Notes:
The species epithet honours Dr. H.P. Singh, Birbal Sahni Institute of Palaeosciences,
Lucknow, India.
2.5.64. Species: D. striatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115658; Current name: Fusiformisporites striatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
2.5.65. Species: D. subaequatus Zhong Y. Zhang 1980; Index Fungorum Registration Identifier:
484999; Location: Lunpola Basin, northern Xizang Plateau, Xizang, China; Age: Oligocene
(Niubao Formation).

744
2.5.66. Species: D. suborbicularis Z.C. Song & G.X. Li in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 637491; Location: Heze County of Shandong Province, China; Age:
Late Oligocene (Dongying Formation).
2.5.67. Species: D. vermae R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk sp. nov.
(Fig. 9P); This new species is described under the section "New species and new
combinations".
2.5.68. Species: D. volubilis P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115657; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age:
Eocene-Oligocene.

2.6. Genus: DIDYMOPORISPORONITES Sheffy & Dilcher Palaeontographica Abt. B 133(1-3):

42 (1971); Index Fungorum Registration Identifier: 21077; Type: D. psilatus Sheffy & Dilcher
1971.
Original Diagnosis: Spores dicellate, uniseptate, pore at apex of one cell. Sculpture psilate to
punctate, shape variable (Sheffy & Dilcher 1971).
Emended Diagnosis: Generally small to medium dicellate conidia; the proximal cell much
thinner walled and smaller than the distal one, with a distinct pore or hilum (Kalgutkar & Jansonius
2000).
Classification: Fungi Imperfecti, Didymosporae.
Number of species known: 25 (but we accept only ten species as legitimate because fifteen
species have been transferred to other genera).
Notes: Kalgutkar & Jansonius (2000) emended the diagnosis of Didymoporisporonites to
include dicellate spores with dissimilar cells only, of which only the smaller is porate. In the past, a
number of spores (e.g. D. didymus, D. oblongatus) with equal or near-equal cells, of which one is
porate, were also assigned to this genus. Elsik (1992) intended to propose a new “Poridicellites” to
accommodate the latter type. Kalgutkar & Jansonius (2000) considered that the “pore” in those
forms actually is a hilum, and therefore transferred spores of this type to the new Hilidicellites.
2.6.1. Species: D. conicus Kalgutkar 1997; Index Fungorum Registration Identifier: 437904;
Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories, Canada; Age: Late
Palaeocene-Early Eocene; Notes: The species epithet is derived from conical shape of small
cell.
2.6.2. Species: D. crassiseptus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107922; Current name: Hilidicellites crassiseptus (Sal.-Cheb. & Locq.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
2.6.3. Species: D. didymus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107923; Current name: Dyadosporites hilatus Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
2.6.4. Species: D. discitypicus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483309; Basionym: Multicellaesporites discitypicus P. Ke & Z.Y. Shi
1978; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene; Notes: Kalgutkar & Jansonius (2000) stated that the photograph of the type
specimen shows a tear in the equatorial region of the spore, that apparently was interpreted as
a septum; hence the spore was called “three-celled”. They interpreted that the type is two-
celled, one cell much smaller and thinner walled than the other. The small cell has subtle
features suggesting the presence of a terminal pore.
2.6.5. Species: D. discors Kalgutkar 1993; Index Fungorum Registration Identifier: 483866;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes:
Kalgutkar & Jansonius (2000) adjusted the original spelling discordis of the specific epithet
to discors, as grammatically the genitive case of an adjective is not applicable in the context
used. The species epithet is derived from the Latin, discors, different, referring to the unequal
cells.

745
2.6.6. Species: D. elegans P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier: 115654;
Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
2.6.7. Species: D. gigas Kalgutkar & Janson. 2000 (Fig. 9Q); Index Fungorum Registration
Identifier: 483310; Basionym: Lacrimasporonites magnus R.K. Saxena & H.P. Singh 1983;
Location: Hoshiarpur-Una Road section, near Bankhandi, Hoshiarpur District, Punjab, India;
Age: Miocene-Pliocene; Notes: Kalgutkar & Jansonius (2000) transferred it to
Didymoporisporonites because of a septum at one end, as well as the apparent presence of a
pore at the same end.
2.6.8. Species: D. henanensis Z.C. Song & G.X. Li in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 485262; Current name: Hilidicellites henanensis (Z.C. Song & G.X.
Li in Z.C. Song et al.) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
2.6.9. Species: D. inaequalis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111421; Locality: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation); Notes: According to Sheffy & Dilcher
(1971), the spore is similar in shape and dimensions to Shortensis memorabilis [Current
name: Vizella memorabilis (Dilcher) Selkirk 1972] (Dilcher (1965).
2.6.10. Species: D. indicus Anil Chandra et al. 1984; Index Fungorum Registration Identifier:
106776; Current name: Hilidicellites indicus (Anil Chandra et al.) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
2.6.11. Species: D. lacrymosus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107924; Current name: Hilidicellites lacrymosus (Sal.-Cheb. & Locq.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
2.6.12. Species: D. longus (R.K. Kar) Kalgutkar & Janson. 2000 (Fig. 9R); Index Fungorum
Registration Identifier: 483311; Basionym: Lacrimasporonites longus R.K. Kar 1979;
Location: Barkhana nala cutting near Sarangwara, Kutch District, Gujarat, India; Age:
Oligocene.
2.6.13. Species: D. mucronatus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107925; Current name: Dyadosporites mucronatus (Sal.-Cheb. & Locq.) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
2.6.14. Species: D. normalis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111422; Current name: Hilidicellites normalis (Sheffy & Dilcher) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
2.6.15. Species: D. oblongatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115671; Current name: Hilidicellites oblongatus P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
2.6.16. Species: D. obtectus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111423; Current name: Hilidicellites obtectus (Sheffy & Dilcher) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
2.6.17. Species: D. ovatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier: 115653;
Current name: Hilidicellites ovatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
2.6.18. Species: D. oviformis Mart.-Hern. & Tom.-Ort. 1989; Index Fungorum Registration
Identifier: 483799; 261072; Current name: Hilidicellites oviformis (Mart.-Hern. & Tom.-
Ort.) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
2.6.19. Species: D. panshanensis P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115652; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age:
Eocene-Oligocene.
2.6.20. Species: D. psilatus Sheffy & Dilcher 1971 (Fig. 9S); Index Fungorum Registration
Identifier: 111424; Location: Puryear clay pit, 800 m south of Puryear, Henry County,
Tennessee, U.S.A.; Age: Middle Eocene (Claiborne Formation).

746
2.6.21. Species: D. siddiquiei Anil Chandra et al. 1984; Index Fungorum Registration Identifier:
106777; Current name: Hilidicellites siddiquiei (Anil Chandra et al.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
2.6.22. Species: D. strangulatus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107926; Current name: Hilidicellites strangulatus (Sal.-Cheb. & Locq.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
2.6.23. Species: D. teleutosporoides Sal.-Cheb. & Locq. 1980; Index Fungorum Registration
Identifier: 107927; Current name: Hilidicellites teleutosporoides (Sal.-Cheb. & Locq.)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
2.6.24. Species: D. triangulus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115702; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age:
Eocene-Oligocene.
2.6.25. Species: D. varius P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier: 115688;
Current name: Hilidicellites varius (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).

2.7. Genus: DIDYMOSPORONITES Sal.-Cheb. & Locq., C. r. Congr. natn. Socs. sav. Paris, sect.
sci., fasc. 1. (Paleobotanique) 105: 188 (1980); Index Fungorum Registration Identifier: 21079;
Type: D. saccatus Sal.-Cheb. & Locq. 1980.
Original Diagnosis (Combined description): Spore 2-celled, septum thick; biconical, strongly
constricted at the partition, surrounded by a loose smooth sac-like perine; 25 × 17 μm including the
membrane, monohilate (monoporate) (Salard-Cheboldaeff & Locquin 1980).
Classification: Fungi Imperfecti, Didymosporae.
Number of species known: One.
2.7.1. Species: D. saccatus Sal.-Cheb. & Locq. 1980 (Fig. 9T); Index Fungorum Registration
Identifier: 107928; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa;
Age: Oligocene-Early Miocene; Notes: This species was described as “Didymosporonites”
saccatus which is probably an unintentional orthographical error for Didymoporisporonites.
Kalgutkar & Jansonius (2000) accepted the generic name in its original orthography as
validly published with a combined description, because the unusual two-layered wall
construction warrants the species to be recognized in a separate genus.

2.8. Genus: DIPLODITES Teterevn-Babajan & Tasl. ex Kalgutkar et al., Review of Palaeobotany
and Palynology (Amsterdam) 77(1-2): 111 (1993); Index Fungorum Registration Identifier:
532862; Type: D. sweetii Kalgutkar et al. 1993.
Synonym: Palaeodiplodites Kyoto Watan. et al. 1999, Index Fungorum Registration
Identifier: 28342.
Original Diagnosis: Hyphae intercellular, in the pericarp, septate, branched, smooth, thick-
walled. Pycnidia superficial or immersed, with no definite orientation; shape and size variable,
globose to subglobose, ovate-oblong or pyriform; dark, thick-walled, wall tissue
pseudoparenchymatous. Pycnidia generally ostiolate, astomous when immersed; solitary or
aggregated in small groups; a subicle or stroma present. Stroma typically dark brown or black,
composed of thick-walled cells, uniloculate. Immature pycnidia filled with thin-walled cells.
Conidia 1-septate or aseptate, both kinds occurring in the same pycnidium, size variable,
ellipsoidal-oblong to ovate, light to dark brown, septa twice as thick as spore walls; globose conidia
1-celled, lightly coloured; two-celled conidia brown, with striations occasionally present.
Conidiophores or their remnants not present (Kalgutkar et al. 1993).
Classification: Fungi Imperfecti, Sphaeropsidales.
Number of species known: Five.
Notes: Kalgutkar (1993) validated the name Diplodites to encompass fossil taxa that are
morphologically similar to the extant fungi Diplodia, Botryodiplodia, and other related genera such
as Dothiorella and Macrophoma.

747
2.8.1. Species: D. mohgaoensis (Barlinge & Paradkar) Kalgutkar et al. 1993; Index Fungorum
Registration Identifier: 532825; Basionym: Botryodiplodia mohgaoensis Barlinge & Paradkar
1982; Location: Mohgaonkalan, Chhindwara District, Madhya Pradesh, India; Age: Late
Cretaceous (Maastrichtian).
2.8.2. Species: D. rodei (Mahab.) Kalgutkar et al. 1993; Index Fungorum Registration Identifier:
532806; Basionym: Diplodia rodei Mahab. 1969; Location: Mohgaonkalan locality in
Chhindwara District, Madhya Pradesh, India; Age: Late Cretaceous (Maastrichtian).
2.8.3. Species: D. sahnii (Singhai) Kalgutkar et al. 1993; Index Fungorum Registration Identifier:
532805; Basionym: Diplodia sahnii Singhai 1974; Location: Mohgaonkalan locality in
Chhindwara District, Madhya Pradesh, India; Age: Late Cretaceous (Maastrichtian).
2.8.4. Species: D. sweetii Kalgutkar et al. 1993; Index Fungorum Registration Identifier: 533027;
Location: Mohgaonkalan locality in Chhindwara District, Madhya Pradesh, India; Age: Late
Cretaceous (Maastrichtian).
2.8.5. Species: D. yezoensis (Kyoto Watan. et al.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483312; Basionym: Palaeodiplodites yezoensis Kyoto Watan. et al.
1999; Location: On bisexual cone of Cycadeoidella japonica Ogura, recovered from shales in
the Middle Yezo Group, Kami-kinenbetsu River, Hokkaido, Japan; Age: Late Cretaceous -
Middle Turonian; Notes: The species epithet is derived from the ancient Japanese name for
Hokkaido (Yezo).

2.9. Genus: DIPLONEUROSPORA K.P. Jain & R.C. Gupta, Palaeobotanist 18(2): 180 (1970);
Index Fungorum Registration Identifier: 21081; Type: D. tewarii K.P. Jain & R.C. Gupta 1970.
Original Diagnosis: Ascospores two-celled, uniseriate, elliptical, margin uneven, cells
unequal; upper cell prominent, dark brown, thick-walled, wall sculptured with longitudinal ribs.
Lower cell hyaline, appendage-like, small in size, rib sculpture faint (Jain & Gupta 1970).
Classification: Fungi Imperfecti, Didymosporae.
Number of species known: One.
Notes: Jain & Gupta (1970) named the present two-celled fossil ascospore genus as
Diploneurospora for its similarity to the single celled ascospores of extant Neurospora.
2.9.1. Species: D. tewarii K.P. Jain & R.C. Gupta 1970 (Fig. 9U); Index Fungorum Registration
Identifier: 313231; Location: Padappakkara, Kollam District, Kerala, India; Age: Miocene;
Notes: The species epithet honours Dr. J.P. Tewari.

2.10. Genus: DISPARIDICELLITES Kalgutkar & Janson., AASP Contributions Series (Dallas)
39: 94 (2000); Index Fungorum Registration Identifier: 28614; Type: D. paradoxus (P. Ke & Z.Y.
Shi) Kalgutkar & Janson. 2000.
Original Diagnosis: Small to medium-sized, dicellate, inaperturate fungal spores; cells
distinctly unequal, the proximal cell much smaller and thinner-walled than the distal cell; septum
may show a perforation and/or septal folds (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Didymosporae.
Number of species known: One.
Notes: In Didymoporisporites the smaller cell has a distinct terminal pore. Dicellaesporites
has spores with two equal cells. The name of the genus is derived from Latin disparatus, dissimilar,
and the dicellate structure of the spore.
2.10.1. Species: D. paradoxus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000 (Fig. 9V); Index
Fungorum Registration Identifier: 483327; Basionym: Dicellaesporites paradoxus P. Ke &
Z.Y. Shi 1978, Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age:
Eocene-Oligocene.

2.11. Genus: DYADOSPORITES R.T. Clarke, Mountain Geologist 2(2): 90, pl. 1, fig. 13–14.
(1965); Index Fungorum Registration Identifier: 21091; Type: D. ellipsus R.T. Clarke 1965.

748
 Synonyms: Dyadosporites Hammen 1954 (nom. inval.) fide Kalgutkar & Jansonius (2000),
Index Fungorum Registration Identifier: 21091; Dyadosporonites Elsik 1968, Index Fungorum
Registration Identifier: 21092; Psidimobipiospora Sal.-Cheb. & Locq. 1980, Index Fungorum
Registration Identifier: 25598.
Original Diagnosis: Fungal spores bilocular (didymosporous), elliptical, central septum
simple, cell wall psilate to finely punctate, pore at apex of each cell (Clarke 1965).
Classification: Fungi Imperfecti, Didymosporae.
Number of species known: 43 (but we accept only 42 species as legitimate because one
species, viz. D. umirensis Hammen 1955, was not validly published).
Notes: Van der Hammen (1955) did not validly publish Dyadosporites. He gave generic
diagnosis and the name of the type species but the latter was never described or illustrated
(Jansonius & Hills 1976). Clarke (1965) treated Dyadosporites as having been validly published by
Van der Hammen (1955). Kalgutkar & Jansonius (2000) considered that Clarke (1965) was the first
to validly publish the generic name and was the first to assign a species Dyadosporites ellipsus R.T.
Clarke 1965 to it which became the type species through the principle of monotypy.
Dyadosporonites Elsik 1968 and Psidimobipiospora Sal.-Cheb. & Locq. 1980 are later taxonomic
synonyms of Dyadosporites.
2.11.1. Species: D. acutus (Rouse & Mustard) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483328; Basionym: Diporicellaesporites acutus Rouse & Mustard
1997; Location: Strait of Georgia, eastern Vancouver Island, the Fraser River lowlands of
southwest British Columbia, Canada and the North-western Washington State, U.S.A.; Age:
Late mid-Eocene-Early late Eocene.
2.11.2. Species: D. annulatus (E.J. Romero & M.T. Castro) Bianchin., Alej. Martínez & R.K.
Saxena in Alej. Martínez et al. 2016; Index Fungorum Registration Identifier: 812336;
Basionym: Dyadosporonites annulatus E.J. Romero & M.T. Castro 1986; Notes: These
spores resemble Trichodelitschia (Phaeotrichaceae, Pleosporales) ascospores as illustrated in
Cugny et al. (2010).
2.11.3. Species: D. antarcticus Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483330; Basionym: Diporicellaesporites stenosus Z.C. Song & Liu Cao 1994; Location:
King George Island, Antarctica; Age: Late Cretaceous.
2.11.4. Species: D. bhardwaji (C.P. Varma & Rawat) Kalgutkar & Janson. 2000 (Fig. 9W); Index
Fungorum Registration Identifier: 483331; Basionym: Psilodiporites bhardwaji C.P. Varma
& Rawat 1963; Location: Western and eastern India, including oil exploration areas in West
Bengal and Assam; Age: Eocene-Miocene (Varma & Rawat 1963).
2.11.5. Species: D. cannanorensis (Ramanujam & K.P. Rao) Kalgutkar & Janson. 2000 (Fig. 9X);
Index Fungorum Registration Identifier: 483332; Basionym: Dyadosporonites cannanorensis
Ramanujam & K.P. Rao 1978; Location: Kannur, Kerala, India; Age: Miocene (Quilon and
Warkalli beds).
2.11.6. Species: D. clarkii (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483333; Basionym: Psidimobipiospora clarkii Sal.-Cheb. & Locq.
1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene.
2.11.7. Species: D. denticulatus (Ramanujam & K.P. Rao) Kalgutkar & Janson. 2000 (Fig. 9Y);
Index Fungorum Registration Identifier: 483334; Basionym: Dyadosporonites denticulatus
Ramanujam & K.P. Rao 1978; Location: Alleppey, Alappuzha District, Kerala, India; Age:
Miocene (Quilon and Warkalli beds); Notes: Prominent pores with thickened rims and teeth-
like or wedge-shaped thickenings on one side of the septa are the diagnostic features of this
species.
2.11.8. Species: D. didymus (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483335; Basionym: Dyadosporonites didymus Sheffy & Dilcher 1971;
Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.; Age:
Middle Eocene (Claiborne Formation).

749
2.11.9. Species: D. dubius P. Kumar 1990 (Fig. 9Z); Index Fungorum Registration Identifier:
126556; Location: Padappakkara, Kollam District, Kerala, India; Age: Early-Middle
Miocene.
2.11.10. Species: D. dyadosporus (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483336; Basionym: Psidimobipiospora dyadospora Sal.-
Cheb. & Locq. 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon,
Africa; Age: Oligocene.
2.11.11. Species: D. ellipsoideus (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483337; Basionym: Psilodiporites ellipsoideus Sal.-Cheb. & Locq.
1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene; Notes: Kalgutkar & Jansonius (2000) stated that although the original description
makes no mention of a septum, and although the authors grouped this species with the
amerospores, there is no mistaking the central septum.
2.11.12. Species: D. ellipsus R.T. Clarke 1965 (Fig. 9AA); Index Fungorum Registration Identifier:
330252; Location: Canon City coal field, Fremont County, Colorado, U.S.A.; Age: Late
Cretaceous; Notes: The species epithet indicates elliptical shape of the spore.
2.11.13. Species: D. elsikii (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483338; Basionym: Psidimobipiospora elsikii Sal.-Cheb. & Locq.
1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age: Late
Eocene-Oligocene.
2.11.14. Species: D. grandiporus (H.P. Singh et al. 1986) Kalgutkar & Janson. 2000 (Fig. 9AB);
Index Fungorum Registration Identifier: 483339; Basionym: Dyadosporonites grandiporus
H.P. Singh et al. 1986; Location: Surma group, Sonapur-Badarpur Road section, Jaintia Hills,
Meghalaya and Cachar, Assam; Age: Early Miocene.
2.11.15. Species: D. hilatus Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483340; Basionym: Didymoporisporonites didymus Sal.-Cheb. & Locq. 1980; Location:
Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age: Oligocene.
2.11.16. Species: D. inaequalis Kalgutkar 1993; Index Fungorum Registration Identifier: 483876;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes:
The specific epithet is derived from the Latin, inaequalis, unequal, referring to the unequal
size of the cells.
2.11.17. Species: D. incisus Kalgutkar 1993; Index Fungorum Registration Identifier: 483877;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes:
The specific epithet is derived from the Latin, incisus, cut, referring to the pores pulling apart.
2.11.18. Species: D. inornatus (Mart.-Hern. & Tom.-Ort.) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483341; Basionym: Dyadosporonites inornatus Mart.-Hern
& Tom.-Ort. 1989, Location: Piedras Negras, Coahuila State, Mexico; Age: Campanian;
Notes: The specific epithet inornatus indicates lack of ornamentation.
2.11.19. Species: D. megaporus (Z.C. Song) Z.C. Song in Z.C. Song et al. 1999; Index Fungorum
Registration Identifier: 483847; Basionym: Dyadosporonites megaporus Z.C. Song 1985;
Location: Youshashan and Dafengshan, Qaidam Basin, Qinghai Province, China; Age:
Palaeocene-Late Eocene; Middle Miocene-Late Miocene; Late Pliocene.
2.11.20. Species: D. minor Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107943; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age: Late
Eocene-Oligocene; Notes: Salard-Cheboldaeff & Locquin (1980) assigned an affinity to
Ascomycota.
2.11.21. Species: D. mucronatus (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483342; Basionym: Didymoporisporonites mucronatus Sal.-Cheb. &
Locq 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene.
2.11.22. Species: D. novus P. Kumar 1990 (Fig. 9AC); Index Fungorum Registration Identifier:
126557; Location: Padappakkara, Kollam District, Kerala, India; Age: Early-Middle

750
Miocene.
2.11.23. Species: D. oblongatus (P. Ke & Z.Y. Shi) G. Norris 1986; Index Fungorum Registration
Identifier: 126572; Basionym: Dyadosporonites oblongatus P. Ke & Z.Y. Shi 1978;
Location: Panshan, Liaoning Province, Coastal region of Bohai, China; (Norris): Imperial
Nuktak C–22 Well, Mackenzie Delta Region, District of Mackenzie, Northwest Territories,
Canada; Age: Eocene-Oligocene; (Norris): Eocene-Oligocene.
2.11.24. Species: D. obscurus Z.C. Song & Liu Cao 1994; Index Fungorum Registration Identifier:
483766; Location: King George Island, Antarctica; Age: Late Cretaceous; Notes: The specific
epithet obscurus is derived from the dark colour of spores.
2.11.25. Species: D. okayi (V.S. Ediger & Alisan) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483343; Basionym: Dyadosporonites okayi V.S. Ediger & Alisan
1989; Location: Northern Thrace Basin, Turkey; Age: Middle?-Late Eocene to Late
Oligocene, Miocene-Pliocene, Notes: The specific epithet is in honour of Professor Hayrettin
B. Okay.
2.11.26. Species: D. puryearensis (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483344; Basionym: Diporicellaesporites puryearensis Sheffy &
Dilcher 1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation); Notes: The specific epithet is after the
name of the pit from where the material was collected.
2.11.27. Species: D. quadratus (Rouse & Mustard) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483345; Basionym: Diporicellaesporites quadratus Rouse & Mustard
1997; Location: Strait of Georgia, eastern Vancouver Island, the Fraser River lowlands of
southwest British Columbia, Canada, and North-western Washington State, U.S.A.; Age:
Late mid-Eocene-Early Late Eocene.
2.11.28. Species: D. reticulatus (Ramanujam & K.P. Rao) Kalgutkar & Janson. 2000 (Fig. 9AD);
Index Fungorum Registration Identifier: 483346; Basionym: Dyadosporonites reticulatus
Ramanujam & K.P. Rao 1978; Location: Kannur, Kerala, India; Age: Miocene (Quilon and
Warkalli beds).
2.11.29. Species: D. sahnii (C.P. Varma & Rawat) Kalgutkar & Janson. 2000 (Fig. 9AE); Index
Fungorum Registration Identifier: 483347; Basionym: Granodiporites sahnii C.P. Varma &
Rawat 1963; Location: Western and eastern India, including oil exploration areas in West
Bengal and Assam; Age: Eocene-Miocene; The species epithet is in honour of Professor
Birbal Sahni.
2.11.30. Species: D. scabratus (P. Kumar) Kalgutkar & Janson. 2000 (Fig. 9AF); Index Fungorum
Registration Identifier: 483348; Basionym: Psidimobipiospora scabrata Kumar 1990;
Location: Padappakkara, Kollam District, Kerala, India; Age: Early-Middle Miocene.
2.11.31. Species: D. schwabii (Elsik) Kalgutkar & Janson. 2000 (Fig. 9AG); Index Fungorum
Registration Identifier: 126573; Basionym: Dyadosporonites schwabii Elsik 1968, Location:
Strip mine approximately 7 miles southwest of Rockdale, Milam County, Texas, U.S.A.;
Age: Palaeocene.
2.11.32. Species: D. singhii R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk sp. nov.
(Fig. 9AH); This new species is described under the section "New species and new
combinations".
2.11.33. Species: D. solidus (P. Ke & Z.Y. Shi) Z.C. Song in Z.C. Song et al. 1999; Index
Fungorum Registration Identifier: 483848; Basionym: Dyadosporonites solidus P. Ke & Z.Y.
Shi 1978, p. 49, pl. 5, fig. 10; Location: Kenli, Shandong Province, Coastal region of Bohai,
China; Age: Eocene-Oligocene.
2.11.34. Species: D. stenosus (Z.C. Song & G.X. Li in Z.C. Song et al.) Z.C. Song in Z.C. Song et
al. 1999; Index Fungorum Registration Identifier: 483849; Basionym: Dyadosporonites
stenosus Z.C. Song & G.X. Li in Z.C. Song et al. 1989; Location: Shenxian county of
Shandong Province, China; Age: Late Eocene-Middle Oligocene (Shahejie Formation).

751
Figure 9 – A–AI Didymosporae. A Dicellaesporites ellipticus K.P. Jain & R.K. Kar 1979, Bar = 20
μm. B Dicellaesporites elongatus Ramanujam & K.P. Rao 1978, Bar = 10 μm. C Dicellaesporites
elsikii B. Samant in R.K. Saxena 2009, Bar = 10 μm. D Dicellaesporites fusiformis Sheffy &
Dilcher 1971, Bar = 10 μm. E Dicellaesporites himachalensis R.K. Saxena & A.P. Bhattach. 1990,
Bar = 20 μm. F Dicellaesporites himalayaensis A. Gupta 2002, Bar = 7 μm. G Dicellaesporites
inaequalis (Y.N.R. Varma & R.S. Patil) Kalgutkar & Janson. 2000, Bar = 20 μm. H
Dicellaesporites indicus A. Gupta 2002, Bar = 5 μm. I Dicellaesporites jainii R.K. Saxena &
S.K.M. Tripathi 2011, Bar = 10 μm. J Dicellaesporites keralensis P. Kumar 1990, Bar = 10 μm. K
Dicellaesporites levis Sheffy & Dilcher 1971, Bar = 10 μm. L Dicellaesporites minutus R.K. Kar &
R.K. Saxena 1976, Bar = 10 μm. M Dicellaesporites perelongatus Kalgutkar & Janson. 2000, Bar =

752
10 μm. N Dicellaesporites popovii Elsik 1968, Bar = 12 μm. O Dicellaesporites singhii R.K.
Saxena & S.K.M. Tripathi 2011, Bar = 20 μm. P Dicellaesporites vermae R.K. Saxena, Wijayaw.,
D.Q. Dai, K.D. Hyde & P.M. Kirk sp. nov., Bar = 17 μm. Q Didymoporisporonites gigas Kalgutkar
& Janson. 2000, Bar = 10 μm. R Didymoporisporonites longus (R.K. Kar) Kalgutkar & Janson.
2000, Bar = 20 μm. S Didymoporisporonites psilatus Sheffy & Dilcher 1971, Bar = 7 μm. T
Didymosporonites saccatus Sal.-Cheb. & Locq. 1980, Bar = 10 μm. U Diploneurospora tewarii
K.P. Jain & R.C. Gupta 1970, Bar = 8 μm. V Disparidicellites paradoxus (P. Ke & Z.Y. Shi)
Kalgutkar & Janson. 2000, Bar = 10 μm. W Dyadosporites bhardwaji (C.P. Varma & Rawat)
Kalgutkar & Janson. 2000, Bar = 20 μm. X Dyadosporites cannanorensis (Ramanujam & K.P.
Rao) Kalgutkar & Janson. 2000, Bar = 10 μm. Y Dyadosporites denticulatus (Ramanujam & K.P.
Rao) Kalgutkar & Janson. 2000, Bar = 10 μm. Z Dyadosporites dubius P. Kumar 1990, Bar = 10
μm. AA Dyadosporites ellipsus R.T. Clarke 1965, Bar = 15 μm. AB Dyadosporites grandiporus
(H.P. Singh et al. 1986) Kalgutkar & Janson. 2000, Bar = 10 μm. AC Dyadosporites novus P.
Kumar 1990, Bar = 10 μm. AD Dyadosporites reticulatus (Ramanujam & K.P. Rao) Kalgutkar &
Janson. 2000, Bar = 10 μm. AE Dyadosporites sahnii (C.P. Varma & Rawat) Kalgutkar & Janson.
2000, Bar = 10 μm. AF Dyadosporites scabratus (P. Kumar) Kalgutkar & Janson. 2000, Bar = 10
μm. AG Dyadosporites schwabii (Elsik) Kalgutkar & Janson. 2000, Bar = 5 μm. AH Dyadosporites
singhii R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk sp. nov., Bar = 20 μm. AI
Dyadosporites udarii (A. Gupta) Kalgutkar & Janson. 2000, Bar = 10 μm.

2.11.35. Species: D. subovalis (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483349; Basionym: Dyadosporonites subovalis Sheffy & Dilcher
1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.;
Age: Middle Eocene (Claiborne Formation).
2.11.36. Species: D. substrangulatus (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483350; Basionym: Psidimobipiospora substrangulata
Sal.-Cheb. & Locq. 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon,
Africa; Age: Late Eocene-Oligocene.
2.11.37. Species: D. taiwanensis T.C. Huang 1981; Index Fungorum Registration Identifier:
115746; Location: Taiwan; Age: Miocene.
2.11.38. Species: D. udarii (A. Gupta) Kalgutkar & Janson. 2000 (Fig. 9AI); Index Fungorum
Registration Identifier: 483351; Basionym: Dyadosporonites udarii A. Gupta 1984, Location:
Barkhana nala cutting, near the village Sarangwara, District of Kutch, western India; Age:
Oligocene Notes: The species epithet is in honour of Professor Ram Udar.
2.11.39. Species: D. umirensis Hammen 1955 (nom. inval.) fide Kalgutkar & Jansonius (2000);
Index Fungorum Registration Identifier: 509580.
2.11.40. Species: D. urniformis Kalgutkar 1993; Index Fungorum Registration Identifier: 483878;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes:
The specific epithet is derived from the Latin, urniformis, urn-shaped, referring to the shape
of the cells.
2.11.41. Species: D. verrucatus (Ramanujam & Srisailam) Kalgutkar & Janson. 2000 (Fig. 10A);
Index Fungorum Registration Identifier: 483352; Basionym: Dyadosporonites verrucatus
Ramanujam & Srisailam 1980; Location: Kannur Beach area, Palayangadi and Cheruvattur
(southern side of Karingottu River), Kerala, India; Location: Palayangadi; Age: Miocene.
2.11.42. Species: D. wilkinsonii (R.K. Saxena & N.K. Misra) Kalgutkar & Janson. 2000 (Fig.
10B); Index Fungorum Registration Identifier: 483353; Basionym: Diporicellaesporites
wilkinsonii R.K. Saxena & N.K. Misra 1990; Location: Ratnagiri beds, Amberiwadi section,
Sindhudurg District, Maharashtra, India; Age: Neogene.

2.12. Genus: DYADOSPORONITES Elsik, Pollen Spores 10(2): 278 (1968); Index Fungorum
Registration Identifier: 21092; Type: D. schwabii Elsik 1968; Current name: DYADOSPORITES
R.T. Clarke 1965 fide Kalgutkar & Jansonius (2000).

753
 Original Diagnosis: Diporate, uniseptate fungal spores. Shape and ornamentation variable.
Single pore at each end of spore. Pores may be modified, i.e., atrium, annulus, or pore chamber
formed by thin septum across end of spore (Elsik 1968).
Classification: Fungi Imperfecti, Didymosporae.
Number of species known: 22 (all the species have been transferred to Dyadosporites Clarke
1965 or to some other appropriate genera).
Notes: Kalgutkar & Jansonius (2000) considered Dyadosporonites Elsik 1968 to be a later
taxonomic synonym of Dyadosporites R.T. Clarke 1965.
2.12.1. Species: D. annulatus E.J. Romero & M.T. Castro 1986; Index Fungorum Registration
Identifier: 812335; Current name: Dyadosporites annulatus (E.J. Romero & M.T. Castro)
Bianchin., Alej. Martínez & R.K. Saxena in Alej. Martínez et al. 2016 fide Martinez et al.
(2016).
2.12.2. Species: D. bhardwaji (C.P. Varma & Rawat) Elsik 1968 (nom. inval.); Index Fungorum
Registration Identifier: 313433; Current name: Dyadosporites bhardwaji (C.P. Varma &
Rawat) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
2.12.3. Species: D. cannanorensis Ramanujam & K.P. Rao 1978; Index Fungorum Registration
Identifier: 124439, Current name: Dyadosporites cannanorensis (Ramanujam & K.P. Rao)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
2.12.4. Species: D. constrictus Y.K. Mathur & K. Mathur 1969; Index Fungorum Registration
Identifier: 483857; Current name: Hilidicellites constrictus (Y.K. Mathur & K. Mathur)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
2.12.5. Species: D. constrictus R.K. Kar 1979 = Dyadosporonites udarii A. Gupta 1984 (nom. nov.
pro Dyadosporonites constrictus R.K. Kar); Index Fungorum Registration Identifier: 112282;
Current name: Dyadosporites udarii (A. Gupta) Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
2.12.6. Species: D. denticulatus Ramanujam & K.P. Rao 1978; Index Fungorum Registration
Identifier: 115064; Current name: Dyadosporites denticulatus (Ramanujam & K.P. Rao)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
2.12.7. Species: D. didymus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111446; Current name: Dyadosporites didymus (Sheffy & Dilcher) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
2.12.8. Species: D. ellipsus (R.T. Clarke) Elsik 1968 (nom. inval.); Index Fungorum Registration
Identifier: 313434; Current name: Dyadosporites ellipsus Clarke 1965 fide Kalgutkar &
Jansonius (2000).
2.12.9. Species: D. grandiporus H.P. Singh et al. 1986; Index Fungorum Registration Identifier:
131931; Current name: Dyadosporites grandiporus (H.P. Singh et al.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
2.12.10. Species: D. inaequalis Y.N.R. Varma & R.S. Patil 1985; Index Fungorum Registration
Identifier: 133494; Current name: Dicellaesporites inaequalis (Y.N.R. Varma & R.S. Patil)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
2.12.11. Species: D. inornatus Mart.-Hern. & Tom.-Ort. 1989; Index Fungorum Registration
Identifier: 483802; Current name: Dyadosporites inornatus (Mart.-Hern. & Tom.-Ort.)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
2.12.12. Species: D. megaporus Z.C. Song 1985; Index Fungorum Registration Identifier: 485263;
Current name: Dyadosporites megaporus (Z.C. Song) Z.C. Song in Z.C. Song et al. 1999 fide
Kalgutkar & Jansonius (2000).
2.12.13. Species: D. oblongatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115636; Current name: Dyadosporites oblongatus (P. Ke & Z.Y. Shi) G. Norris 1986 fide
Norris (1986).
2.12.14. Species: D. okayi V.S. Ediger & Alisan 1989; Index Fungorum Registration Identifier:
125505; Current name: Dyadosporites okayi (V.S. Ediger & Alisan) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).

754
2.12.15. Species: D. reticulatus Ramanujam & K.P. Rao 1978, p. 295, pl. 1, fig. 12. Index
Fungorum Registration Identifier: Current name: Dyadosporites reticulatus (Ramanujam &
Rao) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
2.12.16. Species: D. sahnii (C.P. Varma & Rawat) Elsik 1968 (nom. inval.) (= Granodiporites
sahnii C.P. Varma & Rawat 1963); Index Fungorum Registration Identifier: 115065; Current
name: Dyadosporites sahnii (C.P. Varma & Rawat) Kalgutkar & Janson. 2000 fide Kalgutkar
& Jansonius (2000).
2.12.17. Species: D. schwabii Elsik 1968; Index Fungorum Registration Identifier: 313436; Current
name: Dyadosporites schwabii (Elsik) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius
(2000).
2.12.18. Species: D. solidus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115635; Current name: Dyadosporites solidus (P. Ke & Z.Y. Shi) Z.C. Song in Z.C. Song et
al. 1999 fide Kalgutkar & Jansonius (2000).
2.12.19. Species: D. stenosus Z.C. Song & G.X. Li in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 485264; Current name: Dyadosporites stenosus (Z.C. Song & G.X. Li
in Z.C. Song et al.) Z.C. Song in Z.C. Song et al. 1999 fide Kalgutkar & Jansonius (2000).
2.12.20. Species: D. subovalis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111447; Current name: Dyadosporites subovalis (Sheffy & Dilcher) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
2.12.21. Species: D. udarii A. Gupta 1984; Index Fungorum Registration Identifier: 483858;
Current name: Dyadosporites udarii (A. Gupta) Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
2.12.22. Species: D. verrucatus Ramanujam & Srisailam 1980; Index Fungorum Registration
Identifier: 108852; Current name: Dyadosporites verrucatus (Ramanujam & Srisailam)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).

2.13. Genus: FELIXITES Elsik ex Janson. & Hills, Pollen et Spores 31(1-2): 155 (1990b); Index
Fungorum Registration Identifier: 25443; Type: F. pollenisimilis (Horst) Elsik 1990b.
Original Diagnosis: Psilate, aporate, dicellate fungal spores with an elliptical outline. The
spore outline occasionally can be slightly indented at the septum; individual dispersed cells can
have an ovate outline. The septal area is generally very dark due to its excessive development; the
septum can be several times the thickness of the cell wall. The septum typically is thick and darkly
translucent to practically opaque. The septum is of two main layers, one continuing into the wall of
each cell; in extreme cases the cells are found isolated as a dome-shaped cell truncated by the dark
septal layer. The septal pore is distinct or not discernible. The spore wall is rigid at the septum, less
rigid to folded away from the septum. Rarely a third cell is present; in those specimens the spore is
oriented with a trilobate outline; the dark central septal area serves as a common junction for all
three cells, but the exact nature of attachment and the septum of the third cell is generally obscured
by the opacity of the attachment area (Elsik 1990b).
Classification: Fungi Imperfecti, Didymosporae.
Number of species known: Two.
Notes: Elsik (1990b) stated that Felixites was described for late Palaeozoic fungal
didymospores that are aporate and characterized by a very thick medium septum. Chaetosphaerites
Felix 1894 is tetracellate and the two end cells are more lightly pigmented than the two central
cells. The genus is named in honour of Dr. Johannes Felix.
2.13.1. Species: F. playfordii Elsik 1990b; Index Fungorum Registration Identifier: 126549;
Location: Spitsbergen; Age: Early Carboniferous; Notes: Felixites playfordii is characterized
by its bicampanulate outline. The species epithet is in honour of Professor Geoffrey Playford.
2.13.2. Species: F. pollenisimilis (Horst) Elsik 1990b (Fig. 10C); Index Fungorum Registration
Identifier: 126548; Basionym: Sporonites pollenisimilis Horst 1955; Synonym:
Chaetosphaerites pollenisimilis (Horst) M.A. Butterworth & R.W. Williams 1958 fide Elsik
1990a; Location: Concordia mine, Adit Andreas IV, Upper Silesia, S. Poland (Horst 1955),

755
 Lothians, Central and West Fife coalfields, Scotland (M.A. Butterworth & R.W. Williams
1958), Billefjorden Sandstones, Spitsbergen, Norway (Playford 1962); Age: Carboniferous
(Horst 1955, Playford 1962), Early Carboniferous (Butterworth & Williams 1958).

2.14. Genus: FUSIFORMISPORITES Rouse, Micropaleontology 8(2): 210 (1962); Index

Fungorum Registration Identifier: 21113; Type: F. crabbii Rouse 1962.
Synonym: Striadyadosporites Dueñas 1979 fide Kalgutkar & Jansonius (2000), Index
Fungorum Registration Identifier: 21325.
Original Diagnosis: Spores? very distinctly fusiform in outline. The unit is split into two
equal halves by an equatorial wall that appears to be continuous, thus completely dividing the unit.
Longitudinal grooves spread out along the wall from either pole like a spindle; some reach the
equator, others stop short of it. Only occasionally is a groove continuous across the dividing wall.
The wall is moderately thick, about 3 µm. Ornamentation laevigate. Size-range 20–100 µm (Rouse
1962).
Emended Diagnosis: Inaperturate, dicellate fungal spores bearing characteristic elongate
striae, ribs, ridges or costae oriented parallel to the long axis of the spore. Wall of one or more
layers. Inner surface of wall psilate to punctate or scabrate. Equatorial septum of two layers.
Equatorial constriction of wall may or may not be present (Elsik 1968).
Classification: Fungi Imperfecti, Didymosporae.
Number of species known: 22 (but we accept only 20 species as legitimate because two
species have been transferred to other genera).
Notes: Elsik (1968) emended the generic diagnosis to include forms with less obvious parallel
ornamentational elements. Spores of Fusiformisporites have a close resemblance to the ascospores
of the modern Cookeina (Wolf & Cavaliere 1966). Wolf (1970) also reported non-petrified spores
from Pleistocene and Eocene sediments that are similar in appearance to Cookeina. Although
Fusiformisporites is generally known from the late Palaeocene to Recent (Elsik 1992), Martínez-
Hernández & Tomasini-Ortiz (1989) reported F. striaoctoformis from Maastrichtian strata.
2.14.1. Species: F. acutus P. Kumar 1990 (Fig. 10D); Index Fungorum Registration Identifier:
126560; Location: Padappakkara, Kollam District, Kerala, India; Age: Early-Middle
Miocene.
2.14.2. Species: F. annafrancescae G. Norris 1997; Index Fungorum Registration Identifier:
483789; Location: Imperial ADGO F–28 Well, Mackenzie River delta, Canada; Age:
Palaeocene-Eocene; Notes: Norris (1997) named this species in honour of his wife Mrs. Anne
Frances Norris.
2.14.3. Species: F. barmerensis R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 10E); Index Fungorum
Registration Identifier: 519942; Location: Barmer Hills, Barmer District, Rajasthan, India;
Age: Palaeocene (Barmer Sandstone).
2.14.4. Species: F. crabbii Rouse 1962 (Fig. 10F); Index Fungorum Registration Identifier:
109769; Location: Terminal Dock, Vancouver, British Columbia, Canada; Age: Late
Cretaceous-Middle Eocene (Burrard Formation).
2.14.5. Species: F. duenasii Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483379; Basionym: Striadyadosporites elongatus Dueñas 1979; Location: Tarragona, Sabana
de Bogota, Colombia, South America; Age: Pleistocene.
2.14.6. Species: F. elongatus Ramanujam & K.P. Rao 1978 (Fig. 10G); Index Fungorum
Registration Identifier: 115067; Location: Kannur, Kerala, India; Age: Miocene (Quilon and
Warkalli beds).
2.14.7. Species: F. foedus S.K. Salujha et al. 1974 (Fig. 10H); Index Fungorum Registration
Identifier: 519807; Location: Bali-Chara Nadi traverse, Khasi-Jaintia Hills, India; Age:
Palaeogene (Disang Formation); Notes: A comparable specimen is illustrated by Baksi (1962)
under Fungus striatus.

756
2.14.8. Species: F. keralensis Ramanujam & K.P. Rao 1978 (Fig. 10I); Index Fungorum
Registration Identifier: 115068; Location: Kannur, Kerala, India; Age: Miocene (Quilon and
Warkalli beds).
2.14.9. Species: F. lineatus Rouse & Mustard 1997; Index Fungorum Registration Identifier:
463997; Location: Strait of Georgia, eastern Vancouver Island, the Fraser River lowlands of
southwest British Columbia, Canada, and the North-western Washington State, U.S.A.; Age:
Late Eocene-Early Oligocene; Notes: According to Rouse & Mustard (1997), this species is a
good index fossil for Late Eocene-Early Oligocene age of western coastal deposits in North
America.
2.14.10. Species: F. lineolatus Sheffy & Dilcher 1971 (Fig. 10J); Index Fungorum Registration
Identifier: 111493; Location: Puryear clay pit, 800 m south of Puryear, Henry County,
Tennessee, U.S.A.; Age: Middle Eocene (Claiborne Formation); Notes: The species epithet
lineolatus refers to the continuous longitudinal lines.
2.14.11. Species: F. mackenziei M.G. Parsons & G. Norris 1999; Index Fungorum Registration
Identifier: 483920; Location: Caribou Hills, Mackenzie River delta, Canada; Age: Early
Eocene; Notes: The species epithet is derived from the Mackenzie Delta.
2.14.12. Species: F. marii Elsik 1968; Index Fungorum Registration Identifier: 314247; Location:
Strip mine approximately 7 miles southwest of Rockdale, Milam County, Texas, U.S.A.;
Age: Palaeocene.
2.14.13. Species: F. microstriatus Hopkins 1969; Index Fungorum Registration Identifier: 637492;
Location: Kitsilano outcrops and excavations of the Highbury Tunnel, southwestern British
Columbia, Canada; Age: Late Eocene-Early Oligocene.
2.14.14. Species: F. paucistriatus Rouse & Mustard 1997; Index Fungorum Registration Identifier:
463996; Location: Strait of Georgia, eastern Vancouver Island, the Fraser River lowlands of
southwest British Columbia, Canada, and the North-western Washington State, U.S.A.; Age:
Late Palaeocene; Notes: Rouse & Mustard (1997) stated that this species is distinguishable
from others by generally low number of weak, thin, and often short striae concentrated in the
mid-section of each hemisphere.
2.14.15. Species: F. pseudocrabbii Elsik 1968 (Fig. 10K); Index Fungorum Registration Identifier:
314248; Location: Strip mine approximately 7 miles southwest of Rockdale, Milam County,
Texas, U.S.A.; Age: Palaeocene.
2.14.16. Species: F. rugosus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111494; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation); Notes: The species epithet refers to the
irregular creases in the cell wall.
2.14.17. Species: F. sahii R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk sp. nov. (Fig.
10L); This new species is described under the section "New species and new combinations".
2.14.18. Species: F siglerae Kalgutkar 1997; Index Fungorum Registration Identifier: 437392;
Current name: Dicellaeporisporites siglerae (Kalgutkar) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
2.14.19. Species: F. striaoctoformis Mart.-Hern. & Tom.-Ort. 1989; Index Fungorum Registration
Identifier: 263516; Location: Piedras Negras, Coahuila State, Mexico; Age: Maastrichtian;
Notes: The species epithet is derived from its figure-eight shape and its striation.
2.14.20. Species: F. striatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483380; Basionym: Dicellaesporites striatus P. Ke & Z.Y. Shi 1978;
Location: Tangjiahe, Tianjin Municipality, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
2.14.21. Species: F. taiwanensis T.C. Huang 1981; Index Fungorum Registration Identifier:
115777; Location: Taiwan; Age: Miocene.
2.14.22. Species: F. tonakkalensis Y.N.R. Varma & R.S. Patil 1985; Index Fungorum Registration
Identifier: 133495; Current name: Varmasporites tonakkalensis (Y.N.R. Varma & R.S. Patil)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).

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2.15. Genus: HILIDICELLITES Kalgutkar & Janson., AASP Contributions Series (Dallas) 39:
133 (2000); Index Fungorum Registration Identifier: 28618; Type: H. appendiculatus (Sheffy &
Dilcher) Kalgutkar & Janson. 2000.
Original Diagnosis: Small to medium-sized dicellate fungal spores, with the proximal end
flattened or truncate, due to the presence of a hilum or pore-like structure; the two cells generally of
comparable size; spore wall thin or of medium thickness, smooth or with subdued sculpture,
generally thinner than the septal base (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Didymosporae.
Number of species known: 18.
Notes: In Didymoporisporonites, the proximal cell is much smaller than the distal cell.
Dicellaesporites lacks a hilum or pore. The genus name is derived from Latin hilum, scar, and the
dicellate structure of these spores.
2.15.1. Species: H. appendiculatus (Sheffy & Dilcher) Kalgutkar & Janson. 2000 (Fig. 10M);
Index Fungorum Registration Identifier: 483381; Basionym: Dicellaesporites appendiculatus
Sheffy & Dilcher 1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County,
Tennessee, U.S.A.; Age: Middle Eocene (Claiborne Formation).
2.15.2. Species: H. constrictus (Y.K. Mathur & K. Mathur) Kalgutkar & Janson. 2000 (Fig. 10N);
Index Fungorum Registration Identifier: 519810; Basionym: Dyadosporonites constrictus
Y.K. Mathur & K. Mathur 1969; Location: Naera and Baraia area of Kutch, Gujarat, India;
Age: Pliocene; Notes: Mathur & Mathur (1969) incorrectly cited the generic name as
“Dyadosporonites Hammen 1954”. Kalgutkar & Jansonius (2000) stated that this error in
bibliographic citation does not invalidate the publication of the species name. The specific
epithet is derived from the strong constriction at the septum.
2.15.3. Species: H. crassiseptus (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483383; Basionym: Didymoporisporonites crassiseptus Sal.-Cheb. &
Locq. 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene.
2.15.4. Species: H. dubius Kalgutkar & Janson. 2000 (Fig. 10O); Index Fungorum Registration
Identifier: 483384; Basionym: Dicellaesporites crassiseptus Ramanujam & Srisailam 1980;
Location: Kannur Beach area, Palayangadi and Cheruvattur (southern side of Karingottu
River), Kerala, India; Age: Miocene.
2.15.5. Species: H. henanensis (Z.C. Song & G.X. Li in Z.C. Song et al.) Kalgutkar & Janson.
2000; Index Fungorum Registration Identifier: 483385; Basionym: Didymoporisporonites
henanensis Z.C. Song & G.X. Li in Z.C. Song et al. 1989, Location: Qingteng county of
Henan Province, China; Age: Late Eocene-Early Oligocene (Shahejie Formation); Notes:
This species can be distinguished from other species of Didymoporisporonites by its large
size, and having folds on the surface.
2.15.6. Species: H. indicus (Anil Chandra et al.) Kalgutkar & Janson. 2000 (Fig. 10P); Index
Fungorum Registration Identifier: 483386; Basionym: Didymoporisporonites indicus Anil
Chandra et al. 1984; Location: Cores from the Arabian Sea, type locality Core no. 1; Age:
Late Quaternary.
2.15.7. Species: H. lacrymosus (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483387; Basionym: Didymoporisporonites lacrymosus Sal.-Cheb. &
Locq. 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene.
2.15.8. Species: H. major (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483388; Basionym: Lacrimasporonites major P. Ke & Z.Y. Shi 1978;
Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
2.15.9. Species: H. normalis (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483389; Basionym: Didymoporisporonites normalis Sheffy & Dilcher

758
 1971, Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.;
Age: Middle Eocene (Claiborne Formation).
2.15.10. Species: H. oblongatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483390; Basionym: Didymoporisporonites oblongatus P. Ke & Z.Y.
Shi 1978, Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age:
Eocene-Oligocene.
2.15.11. Species: H. obtectus (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483391; Basionym: Didymoporisporonites obtectus Sheffy & Dilcher
1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.;
Age: Middle Eocene (Claiborne Formation); Notes: The species epithet refers to the presence
of an external psilate sheath.
2.15.12. Species: H. ovatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483392; Basionym: Didymoporisporonites ovatus P. Ke & Z.Y. Shi
1978; Location: Tanggu, Tianjin Municipality; Kenli, Shandong Province, Coastal region of
Bohai, China; Age: Eocene-Oligocene.
2.15.13. Species: H. oviformis (Mart.-Hern. & Tom.-Ort.) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483393; Basionym: Didymoporisporonites oviformis
Mart.-Hern. & Tom.-Ort. 1989; Location: Piedras Negras, Coahuila State, Mexico; Age:
Maastrichtian; Notes: The genus name was incorrectly spelled “Dydimoporisporites Sheffy &
Dilcher” in Martínez-Hernández & Tomasini-Ortiz. (1989). The epithet is derived from its
oval shape.
2.15.14. Species: H. siddiquiei (Anil Chandra et al.) Kalgutkar & Janson. 2000 (Fig. 10Q); Index
Fungorum Registration Identifier: 483394; Basionym: Didymoporisporonites siddiquiei Anil
Chandra et al. 1984; Location: Cores from the Arabian Sea, type locality Core no. 2; Age:
Late Quaternary; Notes: The present species epithet is in honour of Dr. H.N. Siddiquie,
National Institute of Oceanography, Dona Paula, Goa, India.
2.15.15. Species: H. strangulatus (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483395; Basionym: Didymoporisporonites strangulatus
Sal.-Cheb. & Locq. 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon,
Africa; Age: Oligocene.
2.15.16. Species: H. teleutosporoides (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483396; Basionym: Didymoporisporonites
teleutosporoides Sal.-Cheb. & Locq. 1980, Location: Coast of Equatorial Africa, Gulf of
Guinea, Cameroon, Africa; Age: Oligocene-Early Miocene.
2.15.17. Species: H. trivedii Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483397; Basionym: Teleutosporites ovatus Trivedi & C.L. Verma 1970 (nom. inval.) fide
Kalgutkar & Jansonius (2000); Location: Near Kuala Lumpur, Malaya; Age: Eocene; Notes:
Trivedi & Verma (1970) stated that these spores come closest to the teleutospores of rusts.
The specific epithet is derived from the oval shape of the spores. Kalgutkar & Jansonius
(2000) stated that the species name T. ovatus was not validly published by Trivedi & Verma
(1970) because it was not assigned to a validly published generic name. They validated the
species name and transferred it to Hilidicellites. The name Hilidicellites ovatus was
preoccupied, necessitating a new specific epithet. The species epithet is in honour of
Professor B.S. Trivedi, Department of Botany, Lucknow University, Lucknow, India.
2.15.18. Species: H. varius (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483398; Basionym: Didymoporisporonites varius P. Ke & Z.Y. Shi
1978; Location: Nanjiao, Tianjin Municipality, Coastal region of Bohai, China; Age: Eocene-
Oligocene.

2.16. Genus: PALAEODIPLODITES Kyoto Watan. et al., Int. J. Plant Sci. 160(2): 440 (1999);
Index Fungorum Registration Identifier: 28342, Type: P. yezoensis Kyoto Watan. et al. 1999;
Current name: DIPLODITES Kalgutkar et al. 1993.

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 Original Diagnosis: Fossil pycnidia and acervuli; semi-immersed, globose, composed of
thick-walled cells, textura angularis, and producing numerous conidia; conidia dark, ellipsoidal
with dark medial septum and truncate base; conidial wall ornamented with angularly reticulate
ridges (Watanabe et al. 1999).
Classification: Fungi Imperfecti, Sphaeropsidales.
Number of species known: One (the single species has been transferred to Diplodites
Kalgutkar et al. 1993).
Notes: The genus name is derived from Greek Palaeo and modern genus Diplodia.
2.16.1. Species: P. yezoensis Kyoto Watan. et al. 1999; Fungorum Registration Identifier: 460927;
Current name: Diplodites yezoensis (Kyoto Watan. et al.) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).

2.17. Genus: PSIDIMOBIPIOSPORA Sal.-Cheb. & Locquin, C. r. Congr. natn. Socs. sav. Paris,
sect. sci., fasc. 1. (Paleobotanique) 105: 191 (1980); Index Fungorum Registration Identifier:
25598; Type: P. dyadospora Sal.-Cheb. & Locq. 1980; Current name: DYADOSPORITES Clarke
1965 fide Kalgutkar & Jansonius (2000).
Original Diagnosis: Smooth dihilate didymospores (Salard-Cheboldaeff & Locquin 1980).
Classification: Fungi Imperfecti, Didymosporae.
Number of species known: Five (all the species have been transferred to Dyadosporites
Clarke 1965).
2.17.1. Species: P. clarkii Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108423; Current name: Dyadosporites clarkii (Sal.-Cheb. & Locq.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
2.17.2. Species: P. dyadospora Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108424; Current name: Dyadosporites dyadosporus (Sal.-Cheb. & Locq.) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
2.17.3. Species: P. elsikii Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108425; Current name: Dyadosporites elsikii (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
2.17.4. Species: P. scabrata P. Kumar 1990; Index Fungorum Registration Identifier: 126568;
Current name: Dyadosporites scabratus (P. Kumar) Kalgutkar & Janson. 2000 fide Kalgutkar
& Jansonius (2000).
2.17.5. Species: P. substrangulata Sal.-Cheb. & Locq. 1980; Index Fungorum Registration
Identifier: 108426; Current name: Dyadosporites substrangulatus (Sal.-Cheb. & Locq.)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).

2.18. Genus: PUCCINIASPORONITES Ramanujam & Ramachar, Records of the Geological

Survey of India 113(5): 82 (1980); Index Fungorum Registration Identifier: 28628; Type: P.
arcotensis Ramanujam & Ramachar 1980.
Original Diagnosis: Teliospores borne singly on pedicels, two-celled by prominent horizontal
septum; wall thick, pigmented; one germ pore in each cell, more or less terminal in upper [distal]
cell, and lateral in lower [proximal] cell (Ramanujam & Ramachar 1980).
Classification: Basidiomycota, Pucciniomycotina, Pucciniomycetes, Pucciniales,
Pucciniaceae.
Number of species known: One.
2.18.1. Species: P. arcotensis Ramanujam & Ramachar 1980 (Fig. 10R); Index Fungorum
Registration Identifier: 483757; Location: Neyveli Lignite Mine, Cuddalore District, Tamil
Nadu, India; Age: Miocene. Notes: According to Ramanujam & Ramachar (1980), the fossil
spores are quite similar to spores of modern Puccinia, which parasitizes members of the
family Poaceae.

760
2.19. Genus: STRIADYADOSPORITES Dueñas, Caldasia 12(60): 564 (1979); Index Fungorum
Registration Identifier: 21325; Type: S. elongatus Dueñas 1979; Current name:
FUSIFORMISPORITES Rouse 1962 fide Kalgutkar & Jansonius (2000).
Original Diagnosis: Elongate two-celled fungal spores with striate surface. [Jansonius & Hills
1980, card no. 3770.]
Classification: Fungi Imperfecti, Didymosporae.
Number of species known: One (the single species has been transferred to Fusiformisporites
Rouse 1962).
2.19.1. Species: S. elongatus Dueñas 1979; Index Fungorum Registration Identifier: 112651;
Current name: Fusiformisporites duenasii Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).

2.20. Genus: VALSARITES Puri, Some Plant-micro-fossils from the Cretaceous and Palaeocene of
Nigeria; University of Ibadan Botanical Studies 10: 14 (1963); Index Fungorum Registration
Identifier: 646215; Type: V. senonianus Puri 1963.
Original Diagnosis (Combined description): This is an ascospore of some Ascomycota,
measuring 21 × 12 μm in the middle of the broader segment. It is divided into two halves by an
equatorial wall that is continuous. The lower half is narrower, about 10 μm or so. There seems to be
some sort of irregular and faint reticulation. The wall of the spore is not thickened.
Classification: Ascomycota, Sphaeriales.
Number of species known: One.
Notes: According to Puri (1963), spores of this genus resemble ascospores of Endothia Fr.,
Didymosphaeria Fuckel, and Valsaria Ces. & De Not. Spores of Valsaria insitiva (Tode) Ces. &
De Not. are closest in size.
2.20.1. Species: V. senonianus Puri 1963 (Fig. 10S); Index Fungorum Registration Identifier:
IF646216; Location: Nigeria; Age: Senonian.

2.21. Genus: VERRUDISPORONITES O’Keefe, Palynology 41(S1): 322 (2017); Index Fungorum
Registration Identifier: 821913; Type: V. elsikianus O’Keefe 2017.
Original Diagnosis: Dicellate, diporate fungal spores with broadly fusiform to oval outlines
which may or may not be indented at the median septum. Median septum annulate. Spore axis is
straight or nearly so. Typically, 18–24 µm wide × 25–33 µm long. Pores may or may not be
surrounded by protruding collars and are typically annulate, the annulus being formed by a
thickening of the wall. Pores may be modified by a basal septum into pore chambers, which may be
obscured in some species by very thick, dark spore walls. Surface has scattered to abundant
verrucae (O’Keefe 2017).
Classification: Fungi Imperfecti, Didymosporae.
Number of species known: One.
Notes: The name of the genus is derived from the verrucate spore wall and two cells of this
spore.
2.21.1. Species: V. elsikianus O’Keefe 2017 (Fig. 10T); Index Fungorum Registration Identifier:
821914; Location: Tumbes Province, Peru; Age: Miocene (Heath Formation); Notes: The
specific epithet was chosen to honour Dr. William C. Elsik.

3. Phragmosporae

3.1. Genus: ALLEPPEYSPORONITES Ramanujam & K.P. Rao, Proc. IV International

Palynology Conference, Lucknow 1976–1977 1: 299 (1978); Index Fungorum Registration
Identifier: 21008; Type: A. scabratus Ramanujam & K.P. Rao 1978.
Original Diagnosis: Spores branched, brownish in colour, multicellular, nonaperturate, septa
only transverse, branches one or two per spore, gently curved. Basal and terminal cells each with a
conspicuous appendage. Spore wall psilate to scabrate (Ramanujam & Rao 1978).

761
 Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: One.
Notes: Ramanujam & Rao (1978) commented that the branched nature and the presence of
appendages are the important features of this spore type and that the fossil taxon shows striking
similarity to the dematiaceous fungus Grallomyces F. Stevens (Barnett 1956, Ellis 1971,
Subramanian 1971). The spore wall in Grallomyces is minutely verrucate whereas in the fossil
spores it is essentially scabrate. Grallomyces is common in moist tropics. Ramanujam and Rao
(1978) illustrated two specimens of this species. Of these, the holotype (pl. 3, fig. 40) complies with
the specific diagnosis whereas the other specimen (pl. 3, fig. 41) appears to be different.
3.1.1. Species: A. scabratus Ramanujam & K.P. Rao 1978 (Fig. 10U); Index Fungorum
Registration Identifier: 115028; Location: Alleppey, Alappuzha District, Kerala, India; Age:
Miocene (Warkalli Beds).

3.2. Genus: ANATOLINITES Elsik et al., Palynology 14: 92 (1990); Index Fungorum Registration
Identifier: 21010; Type: A. dongyingensis (P. Ke & Z.Y. Shi) Elsik et al. 1990.
Synonym: Cupulisporonites Z.C. Song & Liu Cao 1994 fide Kalgutkar & Jansonius (2000),
Index Fungorum Registration Identifier: 28632.
Original Diagnosis: Diporate, tricellate to tetracellate, generally psilate to vaguely sculptured
fungal spores with a straight to practically straight axis. The pores are centered on the ends of the
spore axis. The pore at the distal (wider) end of the spore is simple, with the spore wall thinning
from the inside of the cell into the pore; or slightly thickened, sometimes with a faint suggestion of
an attachment scar. The pore at the proximal (smaller) end of the spore is in a relatively thinner
wall, and is simple, with the wall thickness unchanged or annulate. The overall spore outline is
broadly to narrowly obovate. The spore outline is straight to slightly indented over the ends of the
septa. The cells are arranged in a graded series of distally increasing size. The ends of the spore are
generally rounded, as the pores are simple; if not, the former presence and subsequent loss of a cell
can be inferred. The spore wall is generally of uniform to variable thickness, and is predominantly
psilate, although the sculpture can be scabrate to pitted at optimum magnifications. A subdued
infrasculpture is also possible. Variable spore wall thickness can be suggested by the variable
development of the brown pigmentation; the cell(s) at the distal end of the spore is/are generally of
darker colour. The septa are of variable thickness. The septal pore is generally distinct (Elsik et al.
1990).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: 14.
Notes: Anatolinites Elsik et al. 1990 and Brachysporisporites R.T. Lange & P.H. Sm. 1971
have similar shapes and spore outlines. Anatolinites has two simple pores, upon which the genus
can be differentiated from Brachysporisporites and similar brachysporid fungal spores that have a
single pore. In addition, the pore of Brachysporisporites can be a compound pore chamber.
3.2.1. Species: A. alaskaensis Elsik et al. 1990; Index Fungorum Registration Identifier: 412404;
Location: Coal Bay, Port Moller Quadrangle, Alaska Peninsula; Age: ?Eocene
(undifferentiated); Notes: The species epithet is after its occurrence in Alaska.
3.2.2. Species: A. alternarioides Elsik et al. 1990; Index Fungorum Registration Identifier: 412405;
Location: Kerala, India; Age: Early to Middle Miocene (Quilon and Warkalli beds); Notes:
The species epithet is after its similarity to extant Alternaria.
3.2.3. Species: A. antarcticus (Z.C. Song & Liu Cao) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483246; Basionym: Pluricellaesporites antarcticus Z.C. Song & Liu
Cao 1994; Location: King George Island, Antarctica; Age: Late Cretaceous; Notes: The
species epithet is derived from the locality of the type specimens.
3.2.4. Species: A. chubutensis Elsik et al. 1990; Index Fungorum Registration Identifier: 412406;
Location: Northern Patagonian Cordillera, Chubut Province, Argentina; Age: Late
Palaeocene (Andean, Andesitic Series); Notes: The species epithet is derived from Chubut
Province, the place of its occurrence.

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3.2.5. Species: A. claibornensis Elsik et al. 1990; Index Fungorum Registration Identifier: 412407;
Location: Central Texas, U.S.A.; Age: Middle Eocene (Stone City Formation of the
Claiborne Group); Notes: The species epithet indicates its occurrence in the Claiborne Group.
3.2.6. Species: A. cupuliformis (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483247; Basionym: Monoporisporites cupuliformis Sheffy & Dilcher
1971; Synonym: Lacrimasporonites cupuliformis (Sheffy & Dilcher) D.L.E. Glass et al. 1986
fide Kalgutkar & Jansonius (2000); Location: East and south-central Texas, U.S.A.; Age:
Late Eocene (Manning Formation); Notes: The species epithet refers to the cup shape of the
spore.
3.2.7. Species: A. dongyingensis (P. Ke & Z.Y. Shi) Elsik et al. 1990 (Fig. 10V); Index Fungorum
Registration Identifier: 412408; Basionym: Multicellaesporites dongyingensis P. Ke & Z.Y.
Shi 1978; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age:
Oligocene; Notes: Elsik et al. (1990) stated that Anatolinites dongyingensis appears to be
endemic to some Turkish Tertiary basins, such as the Thrace Basin in North-western Turkey
and the Tekman Basin in eastern Turkey. The species occurs from the Late Eocene to the
Early? Miocene in the northern Thrace Basin, where it can be considered as an index fossil
for the Late Oligocene, based on its abundance in those strata (Ediger 1981, Ediger & Alisan
1989). The overall range of Anatolinites dongyingensis apparently is Late Palaeocene to
Early? Miocene.
3.2.8. Species: A. holocenicus Elsik et al. ex Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 412409; Location: Alblasserwaard, province of Zuid-Holland, The
Netherlands; Age: Holocene; Notes: Anatolinites holocenicus was not validly published by
Elsik et al. (1990) because the author did not specify where the holotype was deposited.
Kalgutkar & Jansonius (2000) validated the species name by citing information on where the
type is deposited, i.e. Hugo de Vries Laboratorium, University of Amsterdam. The species
epithet indicates its occurrence in the Holocene sediments.
3.2.9. Species: A. megaporus (Z.C. Song & Liu Cao) Janson. et al. 1998, card no. 5071; Index
Fungorum Registration Identifier: 637493; Basionym: Cupulisporonites megaporus Z.C.
Song & Liu Cao 1994; Location: King George Island, Antarctica; Age: Late Cretaceous;
Notes: The species epithet refers to the large size of the pore.
3.2.10. Species: A. reklawensis Elsik et al. 1990; Index Fungorum Registration Identifier: 412410;
Location: Central Texas, U.S.A.; Age: Early Middle Eocene (Reklaw Formation, lower
Claiborne Group); Notes: The species epithet indicates its occurrence in the Reklaw
Formation.
3.2.11. Species: A. spinatus M.G. Parsons & G. Norris 1999; Index Fungorum Registration
Identifier: 483921; Location: Caribou Hills, Mackenzie Delta, northern Canada; Age: Late
Palaeocene-Early to Middle Eocene.
3.2.12. Species: A. subcapsilaris (Sheffy & Dilcher) Elsik et al. 1990; Index Fungorum
Registration Identifier: 412411; Basionym: Pluricellaesporites subcapsilaris Sheffy &
Dilcher 1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation); Notes: The holotype was selected by
Kalgutkar & Jansonius (2000).
3.2.13. Species: A. tenuis (Z.C. Song & Liu Cao) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483248; Basionym: Lacrimasporonites tenuis Z.C. Song & Liu Cao
1994; Location: King George Island, Antarctica; Age: Late Cretaceous; Notes: The species
epithet indicates thinning of spore wall.
3.2.14. Species: A. thraceus (V.S. Ediger) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483249; Basionym: Brachysporisporites thraceus V.S. Ediger 1981,
Orth. corr. pro Brachysporisporites thraceous; Location: Thrace Basin, Turkey; Age: Late
Eocene-Oligocene, Miocene-Pliocene; Notes: Kalgutkar & Jansonius (2000) assigned
Brachysporisporites thraceus to Anatolinites. It may be a taxonomic synonym of Anatolinites
dongyingensis (V.S. Ediger) Elsik et al. 1990 (Kalgutkar & Jansonius 2000). The specific

763
 epithet originally was spelled “thraceous”, an obvious typographic or orthographic error.
Kalgutkar and Jansonius (2000) corrected it as “thraceus” (= from Thrace).

3.3. Genus: ASPERGILLITES Trivedi & C.L. Verma, J. Palynology 5(2): 68 (1970) ex Janson. et
al. 1998; Index Fungorum Registration Identifier: 21023; Type: A. torulosus Trivedi & C.L. Verma
ex Janson. et al., 1998; Current name: CERCOSPORITES Salmon 1903.
Synonym: Aspergillites Trivedi & C.L. Verma 1970 (nom. inval.).
Original Diagnosis: (combined description): Fungal spores in chains consisting of five to
many conidial spores, spore outline almost circular, spores without connective wall, smooth, dark
brown, [wall] about 1 μm thick, basal part of the chain where conidia are formed from
conidiophore clearly seen. Length of individual chains varies, depending on the number of conidia
present in it; terminal spore 13 × 13 μm, middle spore 17 × 17 μm (Trivedi & Verma 1970).
Classification: Ascomycota, Eurotiales.
Number of species known: One (the single species has been transferred to Cercosporites
Salmon 1903).
3.3.1. Species: A. torulosus Trivedi & C.L. Verma ex Janson. et al. 1998; Index Fungorum
Registration Identifier: 309200; Current name: Cercosporites torulosus (Janson. et al.)
Kalgutkar & Janson. 2000.

3.4. Genus: AXISPORONITES Kalgutkar & Janson., AASP Contributions Series (Dallas) 39: 36
(2000); Index Fungorum Registration Identifier: 28611; Type: A. indicus (Kumar) Kalgutkar &
Janson. 2000.
Original Diagnosis: Medium sized, inaperturate, tricellate fungal spores; overall shape more
or less elliptical; two polar cells smaller, triangular, with dark pigmentation and thicker wall than
large, hyaline central cell; septa thicker than wall of central cell (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: One.
Notes: The genus name is derived from the Latin word axis = (geographic) pole.
3.4.1. Species: A. indicus (P. Kumar) Kalgutkar & Janson. 2000 (Fig. 10W); Index Fungorum
Registration Identifier: 483266; Basionym: Multicellaesporites indicus Kumar 1990;
Location: Clay mine section near Kanjantheria House, Padappakkara, Kollam District, Kerala,
India; Age: Early-Middle Miocene (Quilon Beds); Notes: The species epithet is after its
occurrence in India.

3.5. Genus: BRACHYSPORISPORITES R.T. Lange & P.H. Sm., Neues Jb. für Geologie und
Paläontologie 11: 677 (1971); Index Fungorum Registration Identifier: 21037; Type: B. pyriformis
R.T. Lange & P.H. Sm. 1971.
Synonym: Granatisporites Elsik & Janson. 1974 fide Kalgutkar & Jansonius (2000), Index
Fungorum Registration Identifier: 21121.
Original Diagnosis: Obovate, turbinate or pyriform, phaeophragmospores of several cells, the
cells much broader than long in a sharply graded series of diminishing size from a large domed
apical cell to a small hyaline attachment cell, with extremely dark thick bands of septa, similarly
graded towards the attachment cell (Lange & Smith 1971).
Emended Diagnosis: Fungal spores of three or more cells and two or more septa, symmetrical
along the long axis; clavate, obovate, pyriform or ovoid shape; cells are arranged in a graded series
of diminishing size from a large domed apical cell to a relatively small hyaline or so attachment
cell; one or two cells at aporate end usually bigger [and making up the bulk of the spore];
monoporate, pore is situated on the long axis at the narrower end of the spore; exine thickest on the
biggest cell, psilate or scabrate; there may be some pits especially on the biggest cell; thickness of
septa variable, there may be at least one opening on septa (Ediger 1981).
Classification: Fungi Imperfecti, Phragmosporae.

764
 Number of species known: 24 (but we accept only fifteen species as legitimate because six
species have been transferred to other genera and three species, viz. B. communis Z.C. Song in Z.C.
Song et al. 1999, B. elongatus Z.C. Song in Z.C. Song et al. 1999 and B. lageniformis Z.C. Song in
Z.C. Song et al. 1999, have not been validly published).
Notes: Spores of Brachysporisporites are usually compared to the conidia of the modern
Brachysporium Sacc.
3.5.1. Species: B. antarcticus Z.C. Song & Liu Cao 1994; Index Fungorum Registration Identifier:
483767; Location: King George Island, Antarctica; Age: Late Cretaceous; Notes: The species
is derived from the provenance of the type specimens.
3.5.2. Species: B. atratus Kalgutkar 1993; Index Fungorum Registration Identifier: 483860;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes:
The species epithet refers to the dark colour of the spores, Latin, atratus, dressed in black.
3.5.3. Species: B. bullatus Kalgutkar 1993; Index Fungorum Registration Identifier: 483861;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes:
The species epithet is derived from the Latin, bullatus, inflated, referring to the bulged cell.
3.5.4. Species: B. catinus (Elsik & Janson.) Kalgutkar & Janson. 2000 (Fig. 10X); Index Fungorum
Registration Identifier: 483795; Basionym: Granatisporites catinus Elsik & Janson. 1974;
Location: Mackenzie River Delta, Northwest Territories, Canada; Age: Palaeogene.
3.5.5. Species: B. communis Z.C. Song in Z.C. Song et al. 1999 (nom. inval.) fide Kalgutkar &
Jansonius (2000); Index Fungorum Registration Identifier: 483834; Notes: Song (1999) did
not validly publish the name, as he did not cite where the type is stored and did not provide a
Latin description or its English translation.
3.5.6. Species: B. conicus (P. Ke & Z.Y. Shi) Norris 1997, Current name: Paragranatisporites
conicus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.5.7. Species: B. cotalis (Elsik & Janson.) Norris 1986; Index Fungorum Registration Identifier:
126569; Basionym: Granatisporites cotalis Elsik & Janson. 1974; Location: Mackenzie
Delta, Northwest Territories, Canada (Elsik & Jansonius 1974), Imperial Nuktak C–22 Well,
Mackenzie Delta Region, District of Mackenzie, Northwest Territories, Canada (Norris
1986); Age: Palaeogene (Elsik & Jansonius 1974), Eocene (Norris 1986); Notes: The species
epithet is derived from the Greek kotalis, pestle.
3.5.8. Species: B. elongatus Z.C. Song in Z.C. Song et al. 1999 (nom. inval.) fide Kalgutkar &
Jansonius (2000); Index Fungorum Registration Identifier: 483835; Notes: Song (1999) did
not validly publish the name, as he did not cite where the holotype is deposited and did not
provide a Latin description or its English translation.
3.5.9. Species: B. endophragmia Kalgutkar & Sigler 1995; Index Fungorum Registration Identifier:
412616; Current name: Jansoniisporites endophragmia (Kalgutkar & Sigler) Kalgutkar 1997
fide Kalgutkar (1997).
3.5.10. Species: B. fustitudinus G. Norris 1997; Index Fungorum Registration Identifier: 483793;
Location: Imperial ADGO F–28 Well, Mackenzie River delta, Canada; Age: Palaeocene-
Eocene; Notes: The species epithet is derived from the Latin fustitudinus: cudgel-walloping.
3.5.11. Species: B. grandus Z.C. Song & Liu Cao 1994; Index Fungorum Registration Identifier:
483768; Orthogr. corr. pro Brachysporisporites grandus Z.C. Song & Liu Cao 1994;
Location: King George Island, Antarctica; Age: Late Cretaceous; Notes: The species epithet
is derived from the from large bulk of spore.
3.5.12. Species: B. incurvus (P. Ke & Z.Y. Shi) Z.C. Song in Z.C. Song et al. 1999; Index
Fungorum Registration Identifier: 483836; Current name: Paragranatisporites incurvus (P.
Ke & Z.Y. Shi) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.5.13. Species: B. infacetus Kalgutkar 1997; Index Fungorum Registration Identifier: 437390;
Current name: Pluricellaesporites infacetus (Kalgutkar) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
3.5.14. Species: B. inuvikensis M.G. Parsons & G. Norris 1999; Index Fungorum Registration
Identifier: 483922; Location: Caribou Hills, Mackenzie River delta, northern Canada; Age:

765
 Late Palaeocene - Early to Middle Eocene; The species epithet is derived from the
community of Inuvik.
3.5.15. Species: B. lageniformis Z.C. Song in Z.C. Song et al. 1999 (nom. inval.) fide Kalgutkar &
Jansonius (2000); Index Fungorum Registration Identifier: 483837; Notes: The name of the
species was not validly published by Song (1999) because the author did not specify where
the holotype is stored, and did not provide a Latin description or its English translation.
3.5.16. Species: B. longovatus Z.C. Song & Liu Cao 1994 (Fig. 10Y); Index Fungorum
Registration Identifier: 483770; Location: King George Island, Antarctica; Age: Late
Cretaceous; Notes: The species epithet refers to elongated egg-shaped spore.
3.5.17. Species: B. lunpolaensis (Zhong Y. Zhang) Z.C. Song in Z.C. Song et al. 1999; Index
Fungorum Registration Identifier: 483914; Current name: Paragranatisporites lunpolaensis
Zhong Y. Zhang 1980 fide Kalgutkar & Jansonius (2000).
3.5.18. Species: B. magnus B. Samant in R.K. Saxena 2009 (Fig. 10Z); Index Fungorum
Registration Identifier: 515015; Synonym: Brachysporisporites magnus B. Samant 2000,
(nom. inval.) fide Kalgutkar & Jansonius (2000); Location: Near Bhavnagar, Cambay Basin,
Gujarat, India; Age: Early Eocene (Kharsalia Clay Formation).
3.5.19. Species: B. maximus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483282; Basionym: Multicellaesporites maximus P. Ke & Z.Y. Shi
1978; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
3.5.20. Species: B. opimus (Elsik & Janson.) G. Norris 1986; Index Fungorum Registration
Identifier: 126570; Basionym: Granatisporites opimus Elsik & Janson. 1974; Location:
Mackenzie Delta Region, District of Mackenzie, Northwest Territories, Canada; Age:
Palaeogene (Elsik & Jansonius 1974), Eocene (Norris 1986); Notes: The species epithet is
derived from the Latin opimus, fat.
3.5.21. Species: B. ovoidus Z.C. Song & Liu Cao 1994; Index Fungorum Registration Identifier:
483769; Location: King George Island. Antarctica; Age: Late Cretaceous; Notes: The species
epithet indicates its almost ovate shape.
3.5.22. Species: B. pyriformis R.T. Lange & P.H. Sm. 1971 (Fig. 10AA); Index Fungorum
Registration Identifier: 309928; Location: Maslin Bay, South Australia; Age: Early-Middle
Eocene; Notes: Spore Type B described by Ramanujam & Rao (1978) from Quilon and
Warkalli beds (Miocene) of Kerala resembles Brachysporisporites pyriformis in possessing
similar shape and nature of cells, septa and spore wall.
3.5.23. Species: B. tenuis P. Kumar 1990 (Fig. 10AB); Index Fungorum Registration Identifier:
126550; Location: Padappakkara, Kollam District, Kerala, India; Age: Early-Middle Miocene
(Quilon Beds); Notes: Didymoporisporonites sp. described by Varma & Patil (1985) from the
Miocene sediments of Tonakkal area in Thiruvanthapuram District, Kerala resembles
Brachysporisporites tenuis Kumar (1990) in all morphological features, hence merged to it.
3.5.24. Species: B. thraceous V.S. Ediger 1981; Index Fungorum Registration Identifier: 107761;
Current name: Anatolinites thraceus (V.S. Ediger) Kalgutkar & Janson. 2000 fide Kalgutkar
& Jansonius (2000).

3.6. Genus: CANNANOROSPORONITES Ramanujam & K.P. Rao 1978; Index Fungorum
Registration Identifier: 21044; Type: C. raoi Ramanujam & K.P. Rao 1978; Current name:
CHAETOSPHAERITES Felix 1894 fide Kalgutkar & Jansonius (2000).
Original Diagnosis: Spores melanin-coloured, tetracellate, barell-shaped, cells unequal in
size, basal and terminal cells smaller than central cells. Cells exhibit differential pigmentation, two
central cells dark coloured, basal and terminal cells sub-hyaline to hyaline. Central cells with
buldging lateral walls. Septa considerably thickened. Spore wall psilate. Apical cell conidiogenous,
bearing a large simple pore (Ramanujam & Rao 1978).
Classification: Fungi Imperfecti, Phragmosporae.

766
 Number of species known: One (the single species has been transferred to Chaetosphaerites
Felix 1894).
3.6.1. Species: C. raoi Ramanujam & K.P. Rao 1978; Index Fungorum Registration Identifier:
115037; Current name: Chaetosphaerites raoi (Ramanujam & K.P. Rao) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).

3.7. Genus: CERATOHIRUDISPORA R. Kar et al., Review of Palaeobotany & Palynology

(Amsterdam) 158(3-4): 246 (2010); Index Fungorum Registration Identifier: 541644; Type: C.
miocenica R. Kar et al. 2010.
Original 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 (Kar et al.
2010).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: Two.
Notes: Ceratosporium Schwein. and Hirudinaria Ces. 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). The
genus name is after the genera Ceratosporium and Hirudinaria of the Hyphomycetes.
3.7.1. Species: C. miocenica R. Kar et al. 2010 (Fig. 10AC); Index Fungorum Registration
Identifier: 542233; Location: Tlangsam, Mizoram, India; Age: Miocene (Bhuban Formation);
Notes: The species epithet is after its occurrence in Miocene sediments.
3.7.2. Species: C. triradiata R. Kar et al. 2010 (Fig. 10AD); Index Fungorum Registration
Identifier: 542234; Notes: The species epithet is after its three armed conidia.

3.8. Genus: CERCOSPORITES E.S. Salmon, J. Bot., Lond. 41: 127 (1903); Index Fungorum
Registration Identifier: 21048; Type: C. salmonii Kalgutkar 1997.
Synonym: Aspergillites Trivedi & C.L. Verma ex Janson. et al. 1998 fide Kalgutkar &
Jansonius (2000), Index Fungorum Registration Identifier: 21023.
Original Diagnosis: Mycelium consisting of pale brown, septate, filamentous hyphae that
grow individually and have a diameter of 5–8 μm; at more or less irregular intervals the hyphae
may suddenly inflate into larger, more or less globose cells of 15–23 μm diameter which, when
mature, are opaque dark brown and aligned into chains of 3 to 6 cells or rarely into biseriate
aggregates, and which probably function as sclerotia (Salmon 1903).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: Three.
3.8.1. Species: C. catenatus (Ramanujam & K.P. Rao) Kalgutkar & Janson. 2000 (Fig. 10AE);
Index Fungorum Registration Identifier: 483287; Basionym: Pluricellaesporites catenatus
Ramanujam & K.P. Rao; Location: Varkala, Thiruvananthapuram District, Kerala, India;
Age: Miocene (Quilon and Warkalli beds).
3.8.2. Species: C. salmonii Kalgutkar 1997 (Fig. 10AF); Index Fungorum Registration Identifier:
437391; Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories, Canada;
Age: Late Palaeocene-Early Eocene (Iceberg Bay Formation); Notes: The species epithet is in
honour of Dr. Ernest Salmon, author of the generic name.
3.8.3. Species: C. torulosus (Trivedi & C.L. Verma) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483288; Basionym: Aspergillites torulosus Trivedi & C.L. Verma ex
Janson. et al. 1998; Argillites torulosus Trivedi & C.L. Verma 1970 (nom. inval.); Location:
Near Kuala Lumpur, Malaya; Age: Eocene; Notes: The name Aspergillites torulosus was not
validly published by Trivedi & Verma (1970) because two different specimens, on two
different slides, were listed under “Holotype Preparation”.

767
3.9. Genus: CHAETOSPHAERITES Félix, Zeitschr. Deutsche Geol. Gesell. 46: 272 (1894); Index
Fungorum Registration Identifier: 21053; Type: C. bilychnis Felix 1894.
Synonym: Cannanorosporonites Ramanujam & K.P. Rao 1978 fide Kalgutkar & Jansonius
(2000), Index Fungorum Registration Identifier: 21044.
Original Diagnosis: Some of the silicified spores are characterized particularly by the two
middle cells being dark brown, the two end cells pale brown. Since the boundaries of the respective
colours coincide exactly with the sharp delimitations of the individual segments, it is probably not
justified to assume that the differences in shade only resulted from the state of preservation. It is
unlikely, particularly in view of the minute size of the objects, for differences to occur in the state
of preservation of the individual parts; in addition, within the genera Chaetosphaeria, Lophiostoma,
Massaria and Melanomma there occur many species in which only the middle cells are of darker
colour – generally dark brown or blackish-than the end cells which often appear almost colourless.
The shape of the sporidia is strongly obtuse spindle-shaped, almost like that of a cylinder with
rounded ends. They probably consist of 4 segments, but the midmost septum is not clearly visible
on account of the dark coloration of this area. The length is 0.0238 mm [23.8 μm], the width 0.0085
mm [8.5 μm]. The two median, dark coloured cells are larger than the two others; their combined
length is 0.0148 mm [14.8 μm] (Felix 1894).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: Five (but we accept only three species as legitimate because two
species have been transferred to other genera).
Notes: This genus is characterized by spores having two middle cells being dark brown and
two end cells pale brown. The shape of the sporidia is strongly obtuse spindle-shaped, almost like
that of a cylinder with rounded ends. Cannanorosporonites Ramanujam & K.P. Rao is a later
taxonomic synonym of Chaetosphaerites.
3.9.1. Species: C. bilychnis Félix 1894 (Fig. 10AG); Index Fungorum Registration Identifier:
246518; Location: Perekeschkul, near Baku, Azerbaijan; Age: Eocene.
3.9.2. Species: C. camerounensis Sal.-Cheb. & Locq. 1980; Index Fungorum Registration
Identifier: 107812; Current name: Multicellites camerounensis (Sal.-Cheb. & Locq.)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.9.3. Species: C. obscurus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483289; Basionym: Multicellaesporites obscurus P. Ke & Z.Y. Shi
1978; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
3.9.4. Species: C. pollenisimilis (Horst) M.A. Butterworth & R.W. Williams 1958; Index Fungorum
Registration Identifier: 637472; Current name: Felixites pollenisimilis (Horst) Elsik 1990b
fide Elsik (1990b).
3.9.5. Species: C. raoi (Ramanujam & K.P. Rao) Kalgutkar & Janson. 2000 (Fig. 10AH); Index
Fungorum Registration Identifier: 483286; Basionym: Cannanorosporonites raoi Ramanujam
& K.P. Rao 1978; Location: Kannur, Kerala, India; Age: Miocene (Quilon and Warkalli
beds); Notes: The species epithet is in honour of Professor A.R. Rao, one of the pioneer
workers in the field of palaeomycology in India.

3.10. Genus: CHORDECYSTIA C.B. Foster, Jitsuyo Sakumotsu Byorigaku 372: 109 (1979); Index
Fungorum Registration Identifier: 27204; Type: C. chalasta C.B. Foster 1979.
Original Diagnosis: Microfossils occurring in chains of two or more individuals, or as discrete,
disarticulated cells. Median members of chain originally cylindrical, terminal members ellipsoidal
or club-shaped. Termini of cells intact; invaginated or convex, depending upon position in chain;
often constricted and slightly thicker (folded?) at points of interconnection. Wall apparently one-
layered, without obvious structure. One or two narrow clefts or grooves may extend ±diagonally
between the ends of the cell, or from one end to approximately midway along opposite wall of cell.
When chain is intact, grooves appear to spiral along its length. The grooves may delimit dehiscence
area (Foster 1979).

768
 Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: One.
3.10.1. Species: C. chalasta C.B. Foster 1979 (Fig. 10AI); Index Fungorum Registration Identifier:
560974; Location: Australia; Age: Late? Permian; Notes: Elsik (1992) treated the name of
this genus as a later synonym of Reduviasporonites. Kalgutkar & Jansonius (2000) did not
accept the synonymy proposed by Elsik (1999).

Figure 10 – A–S Didymosporae. T–AI Phragmosporae. A Dyadosporites verrucatus (Ramanujam

& Srisailam) Kalgutkar & Janson. 2000, Bar = 10 μm. B Dyadosporites wilkinsonii (R.K. Saxena &
N.K. Misra) Kalgutkar & Janson. 2000, Bar = 10 μm. C Felixites pollenisimilis (Horst) Elsik

769
1990b, Bar = 20 μm. D Fusiformisporites acutus P. Kumar 1990, Bar = 10 μm. E Fusiformisporites
barmerensis R.K. Saxena & S.K.M. Tripathi 2011, Bar = 20 μm. F Fusiformisporites crabbii Rouse
1962, Bar = 12 μm. G Fusiformisporites elongatus Ramanujam & K.P. Rao 1978, Bar = 10 μm. H
Fusiformisporites foedus S.K. Salujha et al. 1974, Bar = 10 μm. I Fusiformisporites keralensis
Ramanujam & K.P. Rao 1978, Bar = 10 μm. J Fusiformisporites lineolatus Sheffy & Dilcher 1971,
Bar = 10 μm. K Fusiformisporites pseudocrabbii Elsik 1968, Bar = 10 μm. L Fusiformisporites
sahii R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk sp. nov., Bar = 15 μm.
M Hilidicellites appendiculatus (Sheffy & Dilcher) Kalgutkar & Janson. 2000, Bar = 20 μm. N
Hilidicellites constrictus (Y.K. Mathur & K. Mathur) Kalgutkar & Janson. 2000, Bar = 10 μm. O
Hilidicellites dubius Kalgutkar & Janson. 2000, Bar = 20 μm. P Hilidicellites indicus (Anil Chandra
et al.) Kalgutkar & Janson. 2000, Bar = 10 μm. Q Hilidicellites siddiquiei (Anil Chandra et al.)
Kalgutkar & Janson. 2000, Bar = 10 μm. R Pucciniasporonites arcotensis Ramanujam & Ramachar
2000, Bar = 10 μm. S Valsarites senonianus Puri 1963, Bar = 10 μm. T Verrudisporonites
elsikianus O’Keefe 2017, Bar = 10 μm. U Alleppeysporonites scabratus Ramanujam & K.P. Rao
1978, Bar = 20 μm. V Anatolinites dongyingensis (P. Ke & Z.Y. Shi) Elsik et al. 1990, Bar = 10
μm. W Axisporonites indicus (P. Kumar) Kalgutkar & Janson. 2000, Bar = 10 μm. X
Brachysporisporites catinus (Elsik & Janson.) Kalgutkar & Janson. 2000, Bar = 20 μm. Y
Brachysporisporites longovatus Z.C. Song & Liu Cao 1994, Bar = 20 μm. Z Brachysporisporites
magnus B. Samant in R.K. Saxena 2009, Bar = 30 μm. AA Brachysporisporites pyriformis R.T.
Lange & P.H. Sm. 1971, Bar = 10 μm. AB Brachysporisporites tenuis P. Kumar 1990, Bar = 20
μm. AC Ceratohirudispora miocenica R. Kar et al. 2010, Bar = 10 μm. AD Ceratohirudispora
triradiata R. Kar et al. 2010, Bar = 10 μm. AE Cercosporites catenatus (Ramanujam & K.P. Rao)
Kalgutkar & Janson. 2000, Bar = 10 μm. AF Cercosporites salmonii Kalgutkar 1997, Bar = 10 μm.
AG Chaetosphaerites bilychnis Félix 1894, Bar = 5 μm. AH Chaetosphaerites raoi (Ramanujam &
K.P. Rao) Kalgutkar & Janson. 2000, Bar = 12 μm. AI Chordecystia chalasta C.B. Foster 1979,
Bar = 10 μm. AJ Circinoconites arthrus R. Kar et al. 2010, Bar = 8 μm.

3.11. Genus: CIRCINOCONITES R. Kar et al., Review of Palaeobotany & Palynology

(Amsterdam) 158(3-4): 246 (2010); Index Fungorum Registration Identifier: 541647; Type: C.
arthrus R. Kar et al. 2010.
Original Diagnosis: Fungal conidia, conidia acrogenous, strongly spiraled, 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
(Kar et al. 2010).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: One.
Notes: Kar et al. (2010) stated that the coiled conidia generally occur amongst the aquatic fungi.
Besides, there are also aeroaquatic or helicosporous fungi which also produce helicoid conidia (van
Beverwijk 1953). The name of the genus is derived from Greek arthron (Gr.) = joint.
3.11.1. Species: C. arthrus R. Kar et al. 2010 (Fig. 10AJ); Index Fungorum Registration Identifier:
542238; Location: Tlangsam, Mizoram, India; Age: Miocene (Bhuban Formation).

3.12. Genus: CLADOSPORIUMSPORINITES Debi Mukh., International Journal of Geology,

Earth and Environmental Sciences 2(2): 8 (2012); Index Fungorum Registration Identifier: 588468;
Type: C. cylindricus Debi Mukh. 2012.
Original Diagnosis: Conidia cylindrical, oblong, rounded at both the ends, 3-4 septate,
fuliginous (dark soot colour); size 80–100 × 15–30 μm; septa slightly constricted (Mukherjee
2012).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: One.

770
 Notes: Mukherjee (2012) opined that these conidia are borne on the apical part of the
conidiophores but are found detached in dispersed condition. They resemble to the conidia of extant
Cladosporium (growing on leaves of Dianthus barbatus and other Caryophyllaceae plants).
3.12.1. Species: C. cylindricus Debi Mukh. 2012 (Fig. 11A); Index Fungorum Registration
Identifier: 588481; Location: Neyveli Lignite Mine-I, Cuddalore District, Tamil Nadu, India;
Age: Miocene (Neyveli Lignite); Notes: The specific epithet refers to the cylindrical shape.

3.13. Genus: DIPORICELLAESPORITES Elsik, Mikol. Fitopatol. 7(3): 181 (1968); Index
Fungorum Registration Identifier: 21083; Type: D. stacyi Elsik 1968.
Original Diagnosis: Elongate, diporate, multicellate fungal or algal spores. One pore at each
end of the spore. Shape and ornamentation variable except never coiled. Two or more septae (Elsik
1968).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: 68 (but we accept only 50 species as legitimate because 18 species
have been transferred to other genera).
3.13.1. Species: D. acuminatus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111432; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation); Notes: The specific epithet indicates
pointed spore apex.
3.13.2. Species: D. acutus Rouse & Mustard 1997; Index Fungorum Registration Identifier:
464001; Current name: Dyadosporites acutus (Rouse & Mustard) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.13.3. Species: D. aequalibus Kalgutkar 1993; Index Fungorum Registration Identifier: 483867;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes:
The specific epithet is derived from the Latin, aequabilis, equal, referring to spores with
symmetrical cells.
3.13.4. Species: D. anisosporus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107929; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age: Late
Eocene-Oligocene.
3.13.5. Species: D. antarcticus Z.C. Song & Liu Cao 1994; Index Fungorum Registration
Identifier: 483773; Location: King George Island, Antarctica; Age: Late Cretaceous; Notes:
The species epithet is after the locality of type specimens.
3.13.6. Species: D. attenuatus Ramanujam & Srisailam 1980; Index Fungorum Registration
Identifier: 108846; Current name: Quilonia attenuata (Ramanujam & Srisailam) Kalgutkar
& Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.13.7. Species: D. belluloides Z.C. Song 1985; Index Fungorum Registration Identifier: 374794;
Current name: Biporipsilonites belluloides (Z.C. Song) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
3.13.8. Species: D. bellulus P. Ke & Z.Y. Shi (nom. inval.); Index Fungorum Registration
Identifier: 115648, Current name: Biporipsilonites bellulus ex Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
3.13.9. Species: D. chitaleyae Kalgutkar 1993; Index Fungorum Registration Identifier: 483868;
Location: Peel River, Yukon Territory, Canada; AGE: Late Palaeocene-Early Eocene; Notes:
The species epithet is in honour of Dr. Shya Chitaley.
3.13.10. Species: D. concavus P. Kumar 1990 (Fig. 11B); Index Fungorum Registration Identifier:
126553; Location: Padappakkara, Kollam District, Kerala, India; Age: Early-Middle Miocene
(Quilon Beds).
3.13.11. Species: D. dilcheri (Anil Chandra et al.) Kalgutkar & Janson. 2000 (Fig. 11C); Index
Fungorum Registration Identifier: 483313; Basionym: Inapertisporites dilcheri Anil Chandra
1984; Location: Sediment core no. 1 (Lat. 17°57.9′N: Long. 70°46.0′E), Arabian Sea; Age:
Late Quaternary; Notes: The species epithet is in honour of Professor D.L. Dilcher.

771
3.13.12. Species: D. doliiformis Kalgutkar 1997; Index Fungorum Registration Identifier: 437905;
Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories, Canada; Age: Late
Palaeocene-Early Eocene; Notes: The species is named for barrel-shape of spores.
3.13.13. Species: D. dolium P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115647; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age:
Eocene-Oligocene.
3.13.14. Species: D. elegans Z.C. Song & H.C. Luo in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 637494; Location: Shenxian county of Shandong Province, China;
Age: Late Eocene-Middle Oligocene (Shahejie Formation); Notes: According to Kalgutkar &
Jansonius (2000), the single figure illustrated appears to have some longitudinal septa,
subdividing the central three cells (that themselves are separated by two transverse septa).
3.13.15. Species: D. ellipticus Zhong Y. Zhang 1980; Index Fungorum Registration Identifier:
485003; Location: Lunpola Basin, northern Xizang Plateau, Xizang, China; Age: Oligocene
(Niubao Formation).
3.13.16. Species: D. elongatus (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483314; Basionym: Multicellaesporites elongatus Sheffy & Dilcher
1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.;
Age: Middle Eocene (Claiborne Formation); Notes: According to Sheffy & Dilcher (1971),
this spore resembles the shape of teliospores of Xenodochus Schlect, in Ramanujam &
Ramachar (1963).
3.13.17. Species: D. elsikii Mart.-Hern. & Tom.-Ort. 1989; Index Fungorum Registration Identifier:
483800; Location: Piedras Negras, Coahuila State, Mexico; Age: Maastrichtian; Notes: The
species epithet is in honour of Dr. William C. Elsik.
3.13.18. Species: D. elsikii B. Samant & Tapaswi 2000; Index Fungorum Registration Identifier:
515004; Current name: Diporicellaesporites samantiae R.K. Saxena 2009 fide Saxena
(2009); Notes: Diporicellaesporites elsikii B. Samant & Tapaswi 2000 is illegitimate, being
later homonym of Diporicellaesporites elsikii Mart.-Hern. & Tom.-Ort. 1989. For this reason,
Saxena (2009) replaced it with a new name.
3.13.19. Species: D. endogranulosus Kemp 1978; Index Fungorum Registration Identifier: 111433;
Current name: Foveodiporites endogranulosus (Kemp) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
3.13.20. Species: D. extensus Rouse & Mustard 1997; Index Fungorum Registration Identifier:
464000; Location: Strait of Georgia, eastern Vancouver Island, the Fraser River lowlands of
southwest British Columbia, Canada, and northwestern Washington State, U.S.A.; Age: Late
Paleocene.
3.13.21. Species: D. fusiformis Ramanujam & Srisailam 1980 (Fig. 11D); Index Fungorum
Registration Identifier: 108847; Location: Kannur Beach area, Palayangadi and Cheruvattur
(southern side of Karingottu River), Kerala, India; Age: Miocene (Warkalli Beds).
3.13.22. Species: D. fusiformis Z.C. Song & H.C. Luo in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 485265; Current name: Diporicellaesporites minifusiformis
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.13.23 Species: D. fusoides Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107930; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age: Early
Miocene.
3.13.24. Species: D. giganteus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115646; Location: Huanghua, Hebei Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
3.13.25. Species: D. hillsii Kalgutkar 1993; Index Fungorum Registration Identifier: 483869,
Current name: Quilonia hillsii (Kalgutkar) Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
3.13.26. Species: D. icebergi Kalgutkar & Sigler 1995; Index Fungorum Registration Identifier:
412959; Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories, Canada;

772
 Age: Late Palaeocene or Early Eocene (Iceberg Bay Formation); Notes: The species epithet is
after its occurrence in Iceberg Bay Formation.
3.13.27. Species: D. incurviusculus Kalgutkar 1997; Index Fungorum Registration Identifier:
437906; Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories, Canada;
Age: Late Palaeocene-Early Eocene; Notes: The species epithet is after the incurved pores
found the spores of this species.
3.13.28. Species: D. intrastriatus H.P. Fan in X.T. Guan et al. 1989; Index Fungorum Registration
Identifier: 637495; Location: Southern Bohai Sea region, China; Age: Neogene (Guantao
Formation).
3.13.29. Species: D. jansonii Kalgutkar 1993; Index Fungorum Registration Identifier: 483870;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes:
Kalgutkar (1993) opined that the tendency of these spores to form in succession and their
morphological similarity are comparable to conidia encountered in species of modern
Annellophora, a dematiaceous hyphomycetous fungus. The species epithet is honour of Dr.
Jan Jansonius.
3.13.30. Species: D. laevigataeformis P. Ke & Z.Y. Shi 1978; Index Fungorum Registration
Identifier: 261342; Location: Panshan, Liaoning Province, Coastal region of Bohai, China;
Age: Eocene-Oligocene.
3.13.31. Species: D. lageniformis Zhong Y. Zhang 1980; Index Fungorum Registration Identifier:
485005; Current name: Quilonia lageniformis (Zhong Y. Zhang) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.13.32. Species: D. lanceolatus Z.C. Song 1985; Index Fungorum Registration Identifier: 485257;
Current name: Multicellaesporites songii Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
3.13.33. Species: D. liaoningensis P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115622; Location: Panshan, Liaoning Province; Kenli, Shandong Province, Coastal region of
Bohai, China; Age: Eocene-Oligocene.
3.13.34. Species: D. macellus Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483315; Basionym: Multicellaesporites attenuatus Sheffy & Dilcher 1971; Location: Puryear
clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.; Age: Middle Eocene
(Claiborne Formation); Notes: The species epithet is derived from Latin macellus = rather
slender, thin.
3.13.35. Species: D. mediocoloratus Kalgutkar 1993; Index Fungorum Registration Identifier:
483871; Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene;
Notes: The species epithet is derived from the Latin, medius, middle; coloratus, coloured,
referring to the middle cell pigment.
3.13.36. Species: D. minifusiformis Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483316; Basionym: Diporicellaesporites fusiformis Z.C. Song & H.C. Luo in Z.C.
Song et al. 1989; Location: Shenxian county of Shandong Province, China; Age: Late
Eocene-Middle Oligocene (Shahejie Formation); Notes: Diporicellaesporites minifusiformis
Kalgutkar & Janson. 2000 is a replacement name of Diporicellaesporites fusiformis Z.C.
Song & H.C. Luo in Z.C. Song et al. 1989.
3.13.37. Species: D. minusculus (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483317; Basionym: Pluricellaesporites minusculus Sheffy & Dilcher
1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.;
Age: Middle Eocene (Claiborne Formation).
3.13.38. Species: D. minutigranulatus (Hammen) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483318; Basionym: Pluricellaesporites minutigranulatus Hammen
1954, p. 104, Location: Magdalena Valley, Eastern Cordellera, Colombia, South America;
Age: Maastrichtian.

773
3.13.39. Species: D. minutus Zhong Y. Zhang 1980; Index Fungorum Registration Identifier:
485004; Location: Lunpola Basin, northern Xizang Plateau, Xizang, China; Age: Oligocene
(Niubao Formation).
3.13.40. Species: D. multicellatus R.K. Saxena & S. Khare 1992; Index Fungorum Registration
Identifier: 483893; Current name: Quilonia multicellata (R.K. Saxena & S. Khare) Kalgutkar
& Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.13.41. Species: D. navicularis Kalgutkar 1993; Index Fungorum Registration Identifier: 483872;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes:
The species epithet is derived from the Latin, navicularis, boat-shaped, referring to the shape
of the spores.
3.13.42. Species: D. oblongatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115644; Location: Coastal region of Bohai, China; Age: Early Tertiary.
3.13.43. Species: D. oculinus M.G. Parsons & G. Norris 1999; Index Fungorum Registration
Identifier: 483923; Location: Caribou Hills, Mackenzie Delta, northern Canada; Age: Late
Palaeocene-Early to Middle Eocene; Notes: The species epithet is derived from Latin oculus,
eye, and -inus, like, referring to the eye-like shape of this spore.
3.13.44. Species: D. ordinatus (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483319; Basionym: Fractisporonites ordinatus Sheffy & Dilcher
1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.;
AGE: Middle Eocene (Claiborne Formation).
3.13.45. Species: D. padappakkarensis P. Kumar 1990; Index Fungorum Registration Identifier:
126554; Current name: Biporipsilonites padappakkarensis (P. Kumar) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.13.46. Species: D. papillatus Kalgutkar 1997; Index Fungorum Registration Identifier: 437907;
Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories, Canada; Age: Late
Palaeocene-Early Eocene; Notes: The species epithet is after papillate spores.
3.13.47. Species: D. pluricellus R.K. Kar & R.K. Saxena 1976 (Fig. 11E); Index Fungorum
Registration Identifier: 112280; Location: Matanomadh, Bhuj-Lakhpat Road, Kutch District,
Gujarat, India; Age: Palaeocene (Matanomadh Formation).
3.13.48. Species: D. prakashii Anil Chandra et al. 1984; Index Fungorum Registration Identifier:
106780; Current name: Quilonia prakashii (Anil Chandra et al.) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.13.49. Species: D. psilatus Elsik & Dilcher 1974; Index Fungorum Registration Identifier:
637496; Location: Lawrence Clay Pit, Henry County, Tennessee, U.S.A.; Age: Middle
Eocene.
3.13.50. Species: D. puryearensis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111434; Current name: Dyadosporites puryearensis (Sheffy & Dilcher) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.13.51. Species: D. quadratus Rouse & Mustard 1997; Index Fungorum Registration Identifier:
464002; Current name: Dyadosporites quadratus (Rouse & Mustard) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.13.52. Species: D. quaternarius Kalgutkar 1993; Index Fungorum Registration Identifier:
483873; Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene;
Notes: The species epithet is derived from the Latin, quaternarius, consisting of four,
referring to the spores being typically four-celled.
3.13.53. Species: D. ramanujamii Kalgutkar 1997; Index Fungorum Registration Identifier:
437908; Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories, Canada;
Age: Late Palaeocene-Early Eocene; Notes: The species epithet is in honour of Professor
C.G.K. Ramanujam.
3.13.54. Species: D. reticulatus Elsik & Dilcher 1974; Index Fungorum Registration Identifier:
637497; Location: Lawrence Clay Pit, Henry County, Tennessee; Age: Middle Eocene.

774
3.13.55. Species: D. samantiae R.K. Saxena 2009 (Fig. 11F); Index Fungorum Registration
Identifier: 515005; Basionym: Diporicellaesporites elsikii B. Samant & Tapaswi 2000;
Location: Near Bhavnagar, Cambay Basin, Gujarat, India; Age: Early Eocene (Kharsalia Clay
Formation); Notes: Diporicellaesporites samantiae R.K. Saxena 2009 is a replacement name
of Diporicellaesporites elsikii B. Samant & Tapaswi 2000.
3.13.56. Species: D. scalaris Kalgutkar 1993; Index Fungorum Registration Identifier: 483874,
Current name: Scolecosporites scalaris (Kalgutkar 1993) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
3.13.57. Species: D. segmentatus Rouse & Mustard 1997; Index Fungorum Registration Identifier:
464003; Location: Strait of Georgia, eastern Vancouver Island, the Fraser River lowlands of
southwest British Columbia, Canada, and North-western Washington State, U.S.A.; Age:
Late Eocene-Early Oligocene.
3.13.58. Species: D. serratulus Traverse & Ash 1994; Index Fungorum Registration Identifier:
313249; Location: Wallowa terrane in Hells Canyon, Idaho, U.S.A.; Age: Early Jurassic to
early Middle Jurassic; Notes: The species epithet is derived from the Latin for toothed.
3.13.59. Species: D. stacyi Elsik 1968 (Fig. 11G); Index Fungorum Registration Identifier: 313249;
Location: Strip mine approximately 11 km south-west of Rockdale, Milam County, Texas,
U.S.A.; Age: Palaeocene (Rockdale lignite).
3.13.60. Species: D. stenosus Z.C. Song & Liu Cao 1994; Index Fungorum Registration Identifier:
483774, Current name: Dyadosporites antarcticus Kalgutkar & Janson. 2000 fide Kalgutkar
& Jansonius (2000).
3.13.61. Species: D. suboblongatus (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483320; Basionym: Pluricellaesporites suboblongatus
Sheffy & Dilcher 1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County,
Tennessee, U.S.A.; Age: Middle Eocene (Claiborne Formation).
3.13.62. Species: D. taeniolelloides Kalgutkar 1997; Index Fungorum Registration Identifier:
437902; Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories, Canada;
Age: Late Palaeocene-Early Eocene; Notes: According to Kalgutkar (1997), the present
species resembles the catenate conidia of species of the modern Taeniolella S. Hughes,
particularly Taeniolella rudis (Sacc.) S. Hughes. In species of Taeniolella, the conidia are
generally smooth whereas in Diporicellaesporites taeniolelloides, the spores are larger and
broader, and generally coarsely rough. The species epithet is derived from the presumed
affinity to Taeniolella.
3.13.63. Species: D. taiwanensis T.C. Huang 1981; Index Fungorum Registration Identifier:
115742; Location: Taiwan; Age: Miocene.
3.13.64. Species: D. tetralocularis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111435; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation).
3.13.65. Species: D. tiruchirappalliensis R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M.
Kirk sp. nov. (Fig. 11H); This new species is described under the section "New species and
new combinations".
3.13.66. Species: D. vermiculatus Kalgutkar 1993; Index Fungorum Registration Identifier:
483875; Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene;
Notes: The species epithet is derived from the Latin, vermiculus, little worm (grub), referring
to the worm-like shape of the spores.
3.13.67. Species: D. verrucatus H.P. Singh et al. 1986 (Fig. 11I); Index Fungorum Registration
Identifier: 131930; Location: Sonapur-Badarpur road Section, Jaintia Hills, Meghalaya and
Cachar, Assam, India; Age: Early Miocene (Bhuban Formation, Surma group); Notes: The
species epithet refers to the verrucate spore wall.
3.13.68. Species: D. wilkinsonii R.K. Saxena & N.K. Misra 1990; Index Fungorum Registration
Identifier: 483360; Current name: Dyadosporites wilkinsonii (R.K. Saxena & N.K. Misra)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).

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3.14. Genus: DIPORIPOLLIS S.K. Dutta & S.C.D. Sah, Palaeontographica Abt. B 131: 45 (1970);
Index Fungorum Registration Identifier: 519817; Type: D. assamicus S.K. Dutta & S.C.D. Sah
1970.
Original Diagnosis: Diporate; amb more or less globular to sub-spheroidal; small to medium
in size; pores placed one over the other, circular in shape, and encircled by one or more thickened
rims; exine thin, surface sculpture psilate to scabrate or finely granulate (Dutta & Sah 1970).
Emended Diagnosis: Small to medium sized, tri- to tetracellate conidia, of which the distal
cell is very much larger than the other cells. Although the basic structure can be compared to that of
Brachysporisporites, here the distal cell is so inflated, that it is spherical, or its width even larger
than its length, commonly causing the spores to be (proximo-distally) compressed in the equatorial
plane. This in turn causes the two or three proximalmost cells to collapse into each other. Spore
wall thin to very thin, and prone to concentric folding, in the distal cell; septa thin, with distinct
rimmed pores, anchored by thick, opaque septal bases, that in proximo-distal compression form
concentric rings. When the compression is somewhat oblique, the proximal cells show that they are
forming a short, more or less tapering tube, the end of which has a small, imperceptible pore,
indicating that it is functionally a hilum (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: Two.
Notes: Dutta & Sah (1970), though proposed Diporipollis as a pollen genus, compared it with
fungal spore genera, viz. Diporisporites Hammen (1954), Diporites Hammen (1954). Kalgutkar &
Jansonius (2000), on the basis of personal communication with William C. Elsik, recognized its
fungal affinity and considered it as ‘Fungi Imperfecti, Phragmosporae’.
3.14.1. Species: D. assamicus S.K. Dutta & S.C.D. Sah 1970 (Fig. 11J); Index Fungorum
Registration Identifier: 519812; Location: Umstew, South Shillong Plateau, Meghalaya, India;
Age: Early Eocene (Cherra Formation); Notes: Dutta & Sah (1970) mentioned that “pollen
grains of Diporipollis assamicus are not comparable to any of the fossil species known so far.
Although the present pollen grains from Assam show a striking resemblance to pollen within
the family Gramineae, their affinity with the family Gramineae remains questionable owing to
number, position and nature of the apertures”. According to Kalgutkar & Jansonius (2000),
William C. Elsik (personal communication with Jan Jansonius) recognized this grain as a fungal
spore: the “thickened rims” mark the septal boundaries, the largest of which located where the
proximal cylinder of cells is attached to the distal-most cell (Jansonius & Hills 1976).
3.14.2. Species: D. melanii (Elsik) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483321; Basionym: Pluricellaesporites melanii Elsik 1968; Synonym:
Granatisporites melanii (Elsik) Elsik & Janson. 1974 fide Kalgutkar & Jansonius (2000);
Location: Strip mine approximately 7 miles southwest of Rockdale, Milam County, Texas,
U.S.A.; Age: Palaeocene.

3.15. Genus: DWAYABEEJAESPORONITES Debi Mukh., International Journal of Geology,

Earth and Environmental Sciences 2(2): 4 (2012); Index Fungorum Registration Identifier: 588462;
Type: D. undulatus Debi Mukh. 2012.
Original Diagnosis: Conidiophores with 14–15 cells arranged in acropetal order, thick walled,
enclosed in an undulating conidiophores wall; conidiophores up to 200 μm size, apical cell mostly
smaller; scoliospores range 12–25 × 25–32 μm (Mukherjee 2012).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: One.
Notes: The name of the genus is after extant Dwayabeeja Subramanian 1971.
3.15.1. Species: D. undulatus Debi Mukh. 2012 (Fig. 11K); Index Fungorum Registration
Identifier: 588475; Location: Neyveli Lignite Mine-I, Cuddalore District, Tamil Nadu, India;
Age: Miocene (Neyveli Lignite); Notes: The specific epithet refers to its undulate nature.

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3.16. Genus: EDMUNDMASONAESPORITES Debi Mukh., International Journal of Geology,
Earth and Environmental Sciences 2(2): 4 (2012); Index Fungorum Registration Identifier: 588463;
Type: E. globulatus Debi Mukh. 2012.
Generic Diagnosis: Spore multicelled, septate, dark in colour; 5 celled, apical cell enlarged,
globular, vacuolated; spores 100–120 × 18–30μm, apical cell large measures 33μm, basal one 18
μm; spore wall laevigate (Mukherjee 2012).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: One.
Notes: The present fossil spore shows resemblance with the extant Edmundmasonia Subram.
under Hyphomycetes.
3.16.1. Species: E. globulatus Debi Mukh. 2012 (Fig. 11L); Index Fungorum Registration
Identifier: 588476; Location: Neyveli Lignite Mine-I, Cuddalore District, Tamil Nadu, India;
Age: Miocene (Neyveli Lignite); Notes: The specific epithet refers to globular apical cell.

3.17. Genus: FOVEOLETISPORONITES Ramanujam & K.P. Rao, Proc. IV International

Palynology Conference, Lucknow 1976–1977 1: 299 (1978); Index Fungorum Registration
Identifier: 21102; Type: F. miocenicus Ramanujam & K.P. Rao 1978.
Original Diagnosis: Spores simple, light to dark brown, multicellate, inaperturate. Spore wall
two-layered, inner layer forming transverse septa, surface conspicuously foveolate (Ramanujam &
Rao 1978).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: Three.
Notes: The diagnostic features of this genus are the simple, multiseptate nature and the
conspicuously foveolate sculpturing of the spore wall.
3.17.1. Species: F. indicus Ramanujam & Srisailam 1980 (Fig. 11M); Index Fungorum
Registration Identifier: 108883; Location: Kannur Beach area, Palayangadi and Cheruvattur
(southern side of Karingottu River), Kerala, India; Age: Miocene (Warkalli Beds).
3.17.2. Species: F. keralensis R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 11N); Index Fungorum
Registration Identifier: 519940; Location: Around Kollam and Varkala, Kerala, India; Age:
Miocene.
3.17.3. Species: F. miocenicus Ramanujam & K.P. Rao 1978 (Fig. 11O); Index Fungorum
Registration Identifier: 115066; Location: Varkala, Kerala, India; Age: Miocene (Quilon and
Warkalli Beds).

3.18. Genus: FRACTISPORONITES R.T. Clarke, Mountain Geologist 2(2): 91 (1965); Index
Fungorum Registration Identifier: 21106, Type: F. canalis R.T. Clarke 1965.
Original Diagnosis: Fungal spores uniseriate, fragments consist of four to many rectangular to
square cells, sides generally parallel (Clarke 1965).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: Nine (but we accept only eight species as legitimate because F.
ordinatus Sheffy & Dilcher 1971 has been transferred to Diporicellaesporites Elsik 1968).
Notes: The prefix of the generic name is derived from the Latin word fractus meaning
broken. Fractisporonites is for fragments of very long fungal spores that generally has breaks at
both ends, which differentiates it from all other psilate scalariform species.
3.18.1. Species: F. canalis R.T. Clarke 1965 (Fig. 11P); Index Fungorum Registration Identifier:
330975; Location: Canon City coal field, Fremont County, Colorado, U.S.A.; Age: Late
Cretaceous; Notes: The species epithet is derived from Latin canalis for channel, or canal,
and is given in reference to the tubular structure extending the length of the spore.
3.18.2. Species: F. doliiformis Kalgutkar 1993; Index Fungorum Registration Identifier: 483880;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes:
The species epithet is derived from Latin, doliiformis, vessel-shaped, referring to the cells
being barrel-shaped.

777
3.18.3. Species: F. elongatus (Sat. K. Srivast.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483376; Basionym: Pluricellaesporites elongatus Sat. K. Srivast.
1968, Location: East Coulee locality, sec. 27, twp. 27, rge. 18, W. 4th mer., Alberta, Canada;
Age: Maastrichtian; Notes: The species epithet is derived from the elongated cells of the
species.
3.18.4. Species: F. filiformis (Hammen) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483377; Basionym: Pluricellaesporites filiformis Hammen 1954; Location:
Magdalena Valley, Eastern Cordellera, Colombia, South America; Age: Maastrichtian.
3.18.5. Species: F. lucibiliporus Sat. K. Srivast. & H. Al. -Tayyar 2013; Index Fungorum
Registration Identifier: 818888; Location: Northern Arabian Gulf; Age: Late middle Albian;
Notes: Fusiform conidia of Scolicosporium macrosporium (Berk.) Sutton of Coelomycetes
has a pore on the wall of each cell (Ellis & Ellis 1985). The species epithet is derived from
the Latin lucibilis = bright.
3.18.6. Species: F. moniliformis R.T. Clarke 1965; Index Fungorum Registration Identifier:
330976; Location: Canon City coal field, Fremont County, Colorado; Age: Late Cretaceous;
Notes: The species epithet is derived from the regularly constricted intervals at the septa
(Latin monile is beaded necklace). Fractisporonites moniliformis differs from F. canalis by
its septal constrictions.
3.18.7. Species: F. nodosus (Sat. K. Srivast.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483378; Basionym: Pluricellaesporites nodosus Sat. K. Srivast. 1968;
Location: Morrin Bridge locality, sec. 16, twp. 31, rge. 21, W. 4th mer., Alberta, Canada;
Age: Maastrichtian; Notes: The species epithet is derived from the Latin nodosus = having
knots.
3.18.8. Species: F. ordinatus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111492; Current name: Diporicellaesporites ordinatus (Sheffy & Dilcher) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.18.9. Species: F. pittsburgensis Traverse & Ash 1994; Index Fungorum Registration Identifier:
363118; Location: Wallowa terrane in Hells Canyon, Idaho, U.S.A.; Age: Early Jurassic to
early Middle Jurassic.

3.19. Genus: GRANATISPORITES Elsik & Janson., Can. J. Bot. 52(5): 953 (1974); Index
Fungorum Registration Identifier: 21121; Type: G. cotalis Elsik & Janson. 1974; Current name:
BRACHYSPORISPORITES R.T. Lange & P.H. Sm. 1971 fide Kalgutkar & Jansonius (2000).
Original Diagnosis: Fungal spores of three or more cells and two or more septa, symmetrical
or nearly so around the long axis. Ambitus oval to clavate; widest towards aporate end. One
aperture, pore, hilum or exitus, generally situated on the long axis at the narrower end of the spore.
Exine psilate, or has ornament of low relief. One or two (rarely three) cells at aporate end always
much larger and constituting the bulk of the spore (Elsik & Jansonius 1974).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: Five (all the species have been transferred to other genera).
Notes: Lange & Smith (1975) stated that the type species of Granatisporites should be
assigned to Brachysporisporites R.T. Lange & P.H. Sm. 1971, the correct name for this form-
generic concept. This makes Granatisporites a later synonym of Brachysporisporites. Therefore, G.
cotalis Elsik & Janson. 1974 (type species) and two other species, described by the same authors,
have been transferred to Brachysporisporites R.T. Lange & P.H. Sm. 1971. However, the
remaining two species have been transferred to other genera.
3.19.1. Species: G. catinus Elsik & Janson. 1974; Index Fungorum Registration Identifier: 314702;
Current name: Brachysporisporites catinus (Elsik & Janson.) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
3.19.2. Species: G. coahuilensis Mart-Hern. & Tom.-Ort. 1989; Index Fungorum Registration
Identifier: 483804; Current name: Pluricellaesporites coahuilensis (Mart-Hern. & Tom.-Ort.)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).

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3.19.3. Species: G. cotalis Elsik & Janson. 1974; Index Fungorum Registration Identifier: 314703;
Current name: Brachysporisporites cotalis (Elsik & Janson.) G. Norris 1986 fide Norris
(1986).
3.19.4. Species: G. melanii (Elsik) Elsik & Janson. 1974; Index Fungorum Registration Identifier:
314704; Current name: Diporipollis melanii (Elsik) Kalgutkar & Janson. 2000 fide Kalgutkar
& Jansonius (2000).
3.19.5. Species: G. opimus Elsik & Janson. 1974; Index Fungorum Registration Identifier: 314705;
Current name: Brachysporisporites opimus (Elsik & Janson.) G. Norris 1986 fide Norris
(1986).

3.20. Genus: HAPALOPHRAGMITES Ramanujam & Ramachar, Records of the geological

Survey of India 113(5): 82 (1980); Index Fungorum Registration Identifier: 28629; Type: H.
cumminsii Ramanujam & Ramachar 1980, p. 82, pl. 1, fig. 7.
Original Diagnosis: Teliospores triquetrously three-celled, pedicellate, odd cell terminal, the
two basal cells borne on a common stalk; wall cinnamon-brown; one germ pore in each cell
(Ramanujam & Ramachar 1980).
Classification: Basidiomycota, Uredinales.
Number of species known: One.
3.20.1. Species: H. cumminsii Ramanujam & Ramachar 1980 (Fig. 11Q); Index Fungorum
Registration Identifier: 483758; Location: Neyveli Lignite Mine, Cuddalore District, Tamil
Nadu, India; Age: Miocene (Neyveli lignite); Notes: Named for G.B. Cummins, famed
uredinologist of Arizona University, U.S.A.

3.21. Genus: HETEROCYSTINELLA Cookson & Eisenack, Neues Jahrbuch für Geologie und
Paläontologie, Monatshefte 2: 79 (1979); Index Fungorum Registration Identifier: 637498; Type:
H. bulbosa Cookson & Eisenack 1979.
Original Diagnosis: The coenobium consists of several approximately spherical cells of
widely different size, arranged in a straight row in order of decreasing size (Cookson & Eisenack
1979).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: One.
3.21.1. Species: H. bulbosa Cookson & Eisenack 1979 (Fig. 11R); Index Fungorum Registration
Identifier: 637499; Location: Eucla basin, Western Australia; Age: Middle Cretaceous;
Notes: Eisenack (1979) suggested affinity to the Algae. Kalgutkar & Jansonius (2000)
considered this form a possible multicelled fungal spore (Jansonius & Hills 1982).

3.22. Genus: JANSONIISPORITES Kalgutkar, Review of Palaeobotany & Palynology

(Amsterdam) 97(1-2): 216 (1997); Index Fungorum Registration Identifier: 27774; Type: J.
endophragmia (Kalgutkar & Sigler) Kalgutkar 1997.
Original Diagnosis: Spores obovoid to clavate, smooth, light brown to brown, not dark, 2–4
septate; septa slightly notched. Cells often unequally coloured. Spores truncated at the base. Spore
wall consisting of two layers that commonly are not appressed, and that may be somewhat
separated between the septa; characterized by distinct presence of folds along the internal cell wall
and encircling the septa (Kalgutkar 1997).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: One.
3.22.1. Species: J. endophragmia (Kalgutkar & Sigler) Kalgutkar 1997 (Fig. 11S); Index
Fungorum Registration Identifier: 437914; Location: Kanguk Peninsula, Axel Heiberg Island,
Northwest Territories, Canada; Age: Late Palaeocene-Early Eocene.

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3.23. Genus: KUMARISPORITES Kalgutkar & Janson., AASP Contributions Series (Dallas) 39:
157 (2000); Index Fungorum Registration Identifier: 28619; Type: K. ramanujamii (P. Kumar.)
Kalgutkar & Janson. 2000.
Original Diagnosis: Small to medium-sized tricellate, inaperturate fungal spores; central cell
may be larger than the tapering terminal cells; septa (or septal bases) thicker than spore wall; spore
wall ornamented by longitudinal ribs running full length of the spore, tapering towards the poles
(Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: One.
Notes: Imprimospora ramanujamii P. Kumar 1990 is a misfit in Imprimospora and therefore
Kalgutkar & Jansonius (2000) proposed Kumarisporites to accommodate it.
3.23.1. Species: K. ramanujamii (P. Kumar) Kalgutkar & Janson. 2000 (Fig. 11T); Index
Fungorum Registration Identifier: 483411; Basionym: Imprimospora ramanujamii P. Kumar
1990; Location: Clay mine section near Kanjantheria House, Padappakkara, Kollam District,
Kerala, India; Age: Early-Middle Miocene (Quilon Beds).

3.24. Genus: MATHURISPORITES Kalgutkar & Janson., AASP Contributions Series (Dallas) 39:
166 (2000); Index Fungorum Registration Identifier: 28620; Type: M. ellipticus (Y.K. Mathur & K.
Mathur) Kalgutkar & Janson. 2000.
Original Diagnosis: Medium-sized hilate spores, generally consisting of a darker central part
with 2–4(–6) cells, and proximal and distal parts of a single to few hyaline cells. No distal pore.
Septa distinct, as thick as, or thicker than, the spore wall. Differs from Pluricellaesporites in the
swollen dark central cells (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: Two.
Notes: Kalgutkar & Jansonius (2000) opined that Pluricellaesporites ellipticus Y.K. Mathur
& K. Mathur is a misfit in Pluricellaesporites Hammen and therefore they proposed
Mathurisporites to accommodate it.
3.24.1. Species: M. ellipticus (Y.K. Mathur & K. Mathur) Kalgutkar & Janson. 2000 (Fig. 11U);
Index Fungorum Registration Identifier: 483417; Basionym: Pluricellaesporites ellipticus
Y.K. Mathur & K. Mathur 1969; Location: Naera and Baraia, Kutch District, Gujarat, India;
Age: Pliocene.
3.24.2. Species: M. glomeratus (Sat. K. Srivast.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483418; Basionym: Pluricellaesporites glomeratus Sat. K. Srivast.
1968; Location: East Coulee, Alberta, Canada; Age: Maastrichtian (Horseshoe Canyon
Formation).

3.25. Genus: MONILITES Pampal., Pilzfunde aus der Augsburger Umgebung 8: 128 (1902); Index
Fungorum Registration Identifier: 21171; Type: M. albidus Pampal. 1902.
Original Diagnosis: Hyphae septate, hyaline, loosely but copiously branching. Conidia
globose, elliptical, hyaline, smooth, 18–21 μm, obtuse on both sides, in short chains that
occasionally branch. (Pampaloni 1902).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: One.
3.25.1. Species: M. albidus Pampal. 1902 (Fig. 11V); Index Fungorum Registration Identifier:
141284; Location: “Disodile” beds, Italy; Age: Middle Miocene.

3.26. Genus: MULTICELLAESPORITES Elsik, Pollen et Spores 10(2): 269 (1968); Index
Fungorum Registration Identifier: 21181; Type: M. nortonii Elsik 1968.
Synonym: Warkallisporonites Ramanujam & K.P. Rao 1978 fide Kalgutkar & Jansonius
(2000), Index Fungorum Registration Identifier: 21361.

780
 Original Diagnosis: Inaperturate, psilate fungal spores of three or more cells; two or more
septa. Shape variable around a long axis (Elsik 1968).
Emended Diagnoses: Diagnosis of Multicellaesporites was emended by Sheffy & Dilcher
(1971) and Kumar (1990), as follows: Inaperturate, psilate to scabrate fungal spores or algal bodies
of three or more cells; two or more septa. Shape variable around a long axis (Sheffy & Dilcher
1971); Fungal spores multicellate, elongate. A longitudinal slit or furrow present. Spore wall
smooth or ornamented or differentially coloured or thickened (Kumar 1990).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: 91 (but we accept only thirteen species as legitimate because 71
species have been transferred to other genera by various authors and seven species, viz. M.
biconicus Z.C. Song in Z.C. Song et al. 1999, M. leguminosus Z.C. Song in Z.C. Song et al. 1999,
M. longiovatus Z.C. Song, Qian & Y.H. Zheng in Z.C. Song et al. 1999, M. margarodes Z.C. Song
in Z.C. Song et al. 1999, M. megafusus Z.C. Song, Qian & Y.H. Zheng in Z.C. Song et al. 1999, M.
spatulatus Z.C. Song, Qian & Y.H. Zheng in Z.C. Song et al. 1999 and M. spindlus Z.C. Song,
Qian & Y.H. Zheng in Z.C. Song et al. 1999 have not been validly published).
Notes: Sheffy & Dilcher (1971) emended the generic diagnosis to include a wider range of
ornamentation than found in the type species. Kumar (1990) also emended the generic diagnosis.
Kalgutkar & Jansonius (2000) adopted the emended diagnosis of Kumar (1990) and paraphrased
the same as follows: “Fungal spores, generally of three to five cells; overall shape fusiform to
elliptical, commonly with a slight curvature in the long axis; generally, the equatorial section is a
plane of symmetry; commonly with a subtle, but distinct, longitudinal furrow, crease, thinning or
rupture of the wall on the concave side of the spore, that may be expressed in only the terminal
cells, or along the whole spore; spore wall smooth or with minute sculpturing; septa distinct or thin,
commonly with a perforation or septal folds.” Warkallisporonites Ramanujam & K.P. Rao 1978 is
the later taxonomic synonym of Multicellaesporites Elsik 1968.
3.26.1. Species: M. acuminatus Rouse & Mustard 1997; Index Fungorum Registration Identifier:
463998; Synonym: Piriurella acuminata (Rouse & Mustard) M.G. Parsons & G. Norris 1999
fide Kalgutkar & Jansonius (2000); Current name: Polycellaesporonites acuminatus (Rouse
& Mustard) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.2. Species: M. allomorphus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111662; Current name: Multicellites allomorphus (Sheffy & Dilcher) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.3. Species: M. articulatus Sat. K. Srivast. & Al-Tayyar 2013; Index Fungorum Registration
Identifier: 818889; Location: northern Arabian Gulf; Age: Aptian; Notes: Srivastava and Al-
Tayyar (2013) stated that similar conidiophores occur in Lobatopedis foliicola P.M. Kirk of
Hyphomycetes (Ellis & Ellis 1985). The species epithet is derived from Latin articulus = joint
3.26.4. Species: M. attenuatus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111663; Current name: Diporicellaesporites macellus Kalgutkar & Janson. 2000, fide
Kalgutkar & Jansonius (2000).
3.26.5. Species: M. bellus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier: 115677;
Current name: Multicellites bellus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
3.26.6. Species: M. biconicus Z.C. Song in Z.C. Song et al. 1999 (nom. inval.) fide Kalgutkar &
Jansonius (2000); Index Fungorum Registration Identifier: 483823; Notes: This species name
was not validly published because the author did not specify where the holotype is stored, and
did not provide a Latin description or its English translation.
3.26.7. Species: M. bigeminatus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111664; Current name: Dicellaesporites bigeminatus (Sheffy & Dilcher) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.8. Species: M. bilobus Rouse & Mustard 1997; Index Fungorum Registration Identifier:
463999; Location: Strait of Georgia, eastern Vancouver Island, the Fraser River lowlands of

781
 southwest British Columbia, Canada and the North-western Washington State, U.S.A.; Age:
Late Palaeocene.
3.26.9. Species: M. capsularis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111665; Current name: Pluricellaesporites capsularis (Sheffy & Dilcher) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.10. Species: M. cerrejonensis Doub. & D. Pons 1973; Index Fungorum Registration Identifier:
485250; Current name: Reduviasporonites cerrejonensis (Doub. & D. Pons) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.11. Species: M. cingulatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115691; Current name: Multicellites cingulatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.12. Species: M. circularis B. Samant & Tapaswi 2000; Index Fungorum Registration
Identifier: 519787; Current name: Multicellites circularis (B. Samant & Tapaswi) R.K.
Saxena & S.K.M. Tripathi 2011 fide Saxena & Tripathi (2011); Notes: This species does not
possess a longitudinal furrow, an essential character of Multicellaesporites (sensu Kalgutkar
& Jansonius 2000) and therefore Saxena & Tripathi (2011) transferred it to Multicellites
Kalgutkar & Janson. 2000.
3.26.13. Species: M. compactilis P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115690; Current name: Multicellites compactilis (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.14. Species: M. confusus Anil Chandra et al. 1984; Index Fungorum Registration Identifier:
107038; Current name: Multicellites confusus (Anil Chandra et al.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.15. Species: M. conicus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115656; Current name: Paragranatisporites conicus (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.16. Species: M. conspicuus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115655; Current name: Multicellites conspicuus (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.17. Species: M. crassisporus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration
Identifier: 108235; Current name: Multicellites crassisporus (Sal.-Cheb. & Locq.) Kalgutkar
& Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.18. Species: M. curvatus Ambwani 1983; Index Fungorum Registration Identifier: 107039;
Current name: Reduviasporonites curvatus (Ambwani) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
3.26.19. Species: M. dawaensis Wang & Leng 1982; Index Fungorum Registration Identifier:
637500; Location: China; Notes: This species was cited in Song et al. (1999). Since its
bibliographic details were not included in the list of references of Song et al. (1999), it is
difficult to verify whether this species was validly published. Kalgutkar & Jansonius (2000)
opined that it has close similarity with Dyadosporites clarkii (Sal.-Cheb. & Locq.) Kalgutkar
& Janson. 2000.
3.26.20. Species: M. denticulatus (Ramanujam & K.P. Rao) Kalgutkar & Janson. 2000 (Fig. 11W);
Index Fungorum Registration Identifier: 483445; Basionym: Warkallisporonites denticulatus
Ramanujam & K.P. Rao 1978; Location: Kannur, Kerala, India; Age: Miocene (Quilon and
Warkalli beds).
3.26.21. Species: M. desmodes P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115687; Current name: Multicellites desmodes (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.26.22. Species: M. didymus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111666; Current name: Multicellites didymus (Sheffy & Dilcher) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).

782
3.26.23. Species: M. differentialis Ramanujam & Srisailam 1980; Index Fungorum Registration
Identifier: 109089; Current name: Ramasricellites differentialis (Ramanujam & Srisailam)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000)
3.26.24. Species: M. dilcheri B. Samant in R.K. Saxena 2009 (Fig. 11X); Index Fungorum
Registration Identifier: 515016; Synonym: Multicellaesporites dilcheri B. Samant 2000 (nom.
inval.); Location: Bhavnagar, Cambay Basin, Gujarat, India; Age: Early Eocene (Kharsalia
Clay Formation); Notes: Samant (2000) described the new species “Multicellaesporites
dilcheri” but did not validly publish the name, as she did not cite where the type was
deposited. The species was validated by Saxena (2009) by providing holotype location.
3.26.25. Species: M. discitypicus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115686; Current name: Didymoporisporonites discitypicus (P. Ke & Z.Y. Shi) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.26. Species: M. dongyingensis P. Ke & Z.Y. Shi 1978; Index Fungorum Registration
Identifier: 115670; Current name: Anatolinites dongyingensis (P. Ke & Z.Y. Shi) Elsik et al.
1990 fide Elsik et al. (1990).
3.26.27. Species: M. doualae Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108236; Current name: Multicellites doualae (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.26.28. Species: M. ellipticus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111667; Current name: Multicellites ellipticus (Sheffy & Dilcher) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.26.29. Species: M. elongatus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111668; Current name: Diporicellaesporites elongatus (Sheffy & Dilcher) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.30. Species: M. elongatus B. Samant 2000; Index Fungorum Registration Identifier: 529875;
Current name: Multicellites psilatus (R.K. Saxena) R.K. Saxena & S.K.M. Tripathi 2011 fide
Saxena & Tripathi (2011); Notes: Samant (2000) described the new species
“Multicellaesporites elongatus” but did not validly publish the name, as she did not cite
where the type was deposited. The species was validated by Saxena (2009) by providing
holotype location, obtained from personal communication with Dr. Bandana Samant. Further,
the epithet “elongatus” cannot be used because of the existence of Multicellaesporites
elongatus Sheffy & Dilcher 1971. Saxena (2009) therefore proposed a new name
Multicellaesporites psilatus Saxena 2009. Kalgutkar & Jansonius (2000), however, redefined
Multicellaesporites and restricted this genus only for spores having a distinct longitudinal
furrow. They transferred all the species of this genus, which lack longitudinal furrow, under
Multicellites. This species also was therefore transferred to Multicellites Kalgutkar & Janson.
2000 by Saxena & Tripathi (2011).
3.26.31. Species: M. elsikii R.K. Kar & R.K. Saxena 1976; Index Fungorum Registration Identifier:
112453; Current name: Multicellites elsikii (R.K. Kar & R.K. Saxena) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.32. Species: M. elsikii V.S. Ediger & Alisan 1989; Index Fungorum Registration Identifier:
125503; Current name: Pluricellaesporites edigeri Kalgutkar & Janson. 2000 fide Kalgutkar
& Jansonius (2000).
3.26.33. Species: M. elsikii (Ramanujam & Srisailam) P. Kumar 1990; Index Fungorum
Registration Identifier: 126566; Current name: Staphlosporonites elsikii Ramanujam &
Srisailam 1980 fide Kalgutkar & Jansonius (2000).
3.26.34. Species: M. evidens P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115684; Current name: Multicellites evidens (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.26.35. Species: M. evidens Zhong Y. Zhang 1980 (=Multicellaesporites lunpolaensis Z.C. Song
in Z.C. Song et al. 1999); Index Fungorum Registration Identifier: 484271; Current name:

783
 Ramasricellites lunpolaensis (Z.C. Song in Z.C. Song et al. 1999) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.26.36. Species: M. fusiformis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111669; Current name: Pluricellaesporites malevisus Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
3.26.37. Species: M. fusus V.S. Ediger 1981; Index Fungorum Registration Identifier: 108237;
Current name: Multicellites fusus (V.S. Ediger) Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
3.26.38. Species: M. grandiusculus Sheffy & Dilcher 1971; Index Fungorum Registration
Identifier: 111670; Current name: Multicellites grandiusculus (Sheffy & Dilcher) Kalgutkar
& Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.39. Species: M. granulatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115701; Current name: Multicellites granulatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.40. Species: M. himalayaensis A. Gupta 2002; Index Fungorum Registration Identifier:
540672; Current name: Multicellites himalayaensis (A. Gupta) R.K. Saxena & S.K.M.
Tripathi 2011 fide Saxena & Tripathi (2011); Notes: This species does not possess a
longitudinal furrow, an essential character of Multicellaesporites (sensu Kalgutkar &
Jansonius 2000) and therefore has been transferred to Multicellites Kalgutkar & Janson. 2000
by Saxena & Tripathi (2011).
3.26.41. Species: M. incurvus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115649; Current name: Paragranatisporites incurvus (P. Ke & Z.Y. Shi) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.42. Species: M. indicus P. Kumar 1990; Index Fungorum Registration Identifier: 126567;
Current name: Axisporonites indicus (Kumar) Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
3.26.43. Species: M. irregularis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111671; Current name: Pluricellaesporites irregularis (Sheffy & Dilcher) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.44. Species: M. jainii A. Gupta 2002; Index Fungorum Registration Identifier: 540673;
Current name: Multicellites jainii (A. Gupta) R.K. Saxena & S.K.M. Tripathi 2011 fide
Saxena & Tripathi (2011); Notes: This species does not possess a longitudinal furrow, an
essential character of Multicellaesporites [sensu Kalgutkar & Jansonius 2000) and therefore
Saxena & Tripathi (2011) transferred it to Multicellites Kalgutkar & Janson. 2000].
3.26.45. Species: M. karii Anil Chandra et al. 1984; Index Fungorum Registration Identifier:
106573; Current name: Biporipsilonites karii (Anil Chandra et al.) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.26.46. Species: M. kumarii R.K. Saxena 2009; Index Fungorum Registration Identifier: 515008;
Basionym: Multicellaesporites elsikii (Ramanujam & Srisailam) P. Kumar 1990; Notes:
Multicellaesporites kumarii R.K. Saxena 2009 is a replacement name of Multicellaesporites
elsikii (Ramanujam & Srisailam) Kumar 1990.
3.26.47. Species: M. laevigataeformis Wang & Leng, 1982; Index Fungorum Registration
Identifier: 637501; Location: China; Notes: This species was cited in Song et al. (1999).
Since its bibliographic details were not included in the list of references of Song et al. (1999),
it is difficult to verify whether this species was validly published. Kalgutkar & Jansonius
(2000) opined that it resembles Dyadosporites verrucatus (Ramanujam & Srisailam)
Kalgutkar & Janson. 2000.
3.26.48. Species: M. lanceolatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
483467; Current name: Multicellites lanceolatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.49. Species: M. leguminosus Z.C. Song in Z.C. Song et al. Z.C. Song 1999 (nom. inval.) fide
Kalgutkar & Jansonius (2000); Index Fungorum Registration Identifier: 483824; Notes: Song

784
 (1999) did not validly publish the name, as he did not cite where the type was deposited and
did not provide a Latin description or its English translation.
3.26.50. Species: M. leptaleus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115666; Current name: Multicellites leptaleus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.26.51. Species: M. littoralis Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108238; Current name: Multicellites littoralis (Sal.-Cheb. & Locq.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.52. Species: M. longiovatus Z.C. Song, Qian & Y.H. Zheng in Z.C. Song et al. 1999 (nom.
inval.) fide Kalgutkar & Jansonius (2000); Index Fungorum Registration Identifier: 483825;
Location: China; Age: Late Cretaceous/Tertiary; Notes: This name was not validly published
because the author did not specify where the holotype is stored, and did not provide a Latin
description or its English translation.
3.26.53. Species: M. lunpolaensis Z.C. Song in Z.C. Song et al. 1999; Index Fungorum Registration
Identifier: 483826; Current name: Multicellites evidens P. Ke & Z.Y. Shi 1978 fide Kalgutkar
& Jansonius (2000).
3.26.54. Species: M. margaritus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115665; Current name: Multicellites margaritus (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.55. Species: M. margarodes Z.C. Song in Z.C. Song et al. 1999 (nom. inval.) fide Kalgutkar &
Jansonius (2000); Index Fungorum Registration Identifier: 483827; Notes: This name was not
validly published because the author did not specify where the holotype was deposited, and
did not provide a Latin description or its English translation.
3.26.56. Species: M. maximus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115664; Current name: Brachysporisporites maximus (P. Ke & Z.Y. Shi) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.57. Species: M. megafusus Z.C. Song, Qian & Y.H. Zheng in Z.C. Song et al. 1999 (nom.
inval.) fide Kalgutkar & Jansonius (2000); Index Fungorum Registration Identifier: 483828;
Notes: This name was not validly published because the author did not specify where the
holotype was deposited, and did not provide a Latin description or its English translation.
3.26.58. Species: M. megareniformis (Zhong Y. Zhang) Z.C. Song in Z.C. Song et al. 1999; Index
Fungorum Registration Identifier: 483829; Current name: Dicellaesporites reniformis Zhong
Y. Zhang 1980 fide Kalgutkar & Jansonius (2000).
3.26.59 Species: M. mollis P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115680; Current name: Biporipsilonites mollis (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.26.60. Species: M. mustus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115694; Current name: Biporipsilonites mustus (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.61. Species: M. nortonii Elsik 1968 (Fig. 11Y); Index Fungorum Registration Identifier:
317946; Location: Strip mine approximately 7 miles southwest of Rockdale, Milam County,
Texas, U.S.A.; Age: Palaeocene (Rockdale Lignite).
3.26.62. Species: M. obscurus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115693; Current name: Chaetosphaerites obscurus (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.63. Species: M. oculeus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115698; Current name: Multicellites oculeus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.26.64. Species: M. ornatus Wang & Leng 1982; Index Fungorum Registration Identifier: 637502;
Location: China; Notes: This species was cited in Song et al. (1999). Since its bibliographic
details were not included in the list of references of Song et al. (1999), it is difficult to verify
whether this species was validly published. Kalgutkar & Jansonius (2000) opined that it is

785
 similar to Diporicellaesporites antarcticus Z.C. Song & Liu Cao 1994 or possibly
Hilidicellites indicus (Anil Chandra et al.) Kalgutkar & Janson. 2000.
3.26.65. Species: M. ovatoides Z.C. Song & G.X. Li in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 485256; Current name: Mulicellites ovatoides (Z.C. Song & G.X. Li
in Z.C. Song et al.) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.66. Species: M. ovatus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111672; Current name: Multicellites ovatus (Sheffy & Dilcher) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.26.67. Species: M. pachydermus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115697; Current name: Multicellites pachydermus (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.68. Species: M. pachysporus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration
Identifier: 108239; Current name: Multicellites pachysporus (Sal.-Cheb. & Locq.) Kalgutkar
& Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.69. Species: M. pandus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111673; Current name: Multicellites pandus (Sheffy & Dilcher) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.26.70. Species: M. prakashii Ambwani 1982; Index Fungorum Registration Identifier: 483901;
Current name: Reduviasporonites prakashii (Ambwani) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
3.26.71. Species: M. psilatus R.K. Saxena 2009; Index Fungorum Registration Identifier: 515014;
Basionym: Multicellaesporites elongatus Samant 2000 (nom. inval.) fide Saxena 2009;
Current name: Multicellites psilatus (R.K. Saxena) R.K. Saxena & S.K.M. Tripathi 2011 fide
Saxena & Tripathi (2011).
3.26.72. Species: M. pulvinus Debi Mukh. 2012; Index Fungorum Registration Identifier: 565665;
Location: Neyveli Lignite Mine-I, Cuddalore District, Tamil Nadu, India; Age: Miocene
(Neyveli Lignite).
3.26.73. Species: M. qingfengensis Z.C. Song & H.C. Luo in Z.C. Song et al. 1989; Index
Fungorum Registration Identifier: 485251; Current name: Reduviasporonites qingfengensis
(Z.C. Song & H.C. Luo in Z.C. Song et al.) Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
3.26.74. Species: M. quattorodecimcellus Mart.-Hern. & Tom.-Ortiz; Index Fungorum Registration
Identifier: 271930; Current name: Multicellites quattorodecimcellus (Mart.-Hern. & Tom.-
Ortiz) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.75. Species: M. ramanujamii A. Gupta 2002; Index Fungorum Registration Identifier: 540674;
Current name: Multicellites ramanujamii (A. Gupta 2002) R.K. Saxena & S.K.M. Tripathi
2011 fide Saxena & Tripathi (2011); Notes: This species does not possess a longitudinal
furrow, an essential character of Multicellaesporites (sensu Kalgutkar & Jansonius 2000) and
therefore was transferred to Multicellites Kalgutkar & Janson. 2000.
3.26.76. Species: M. reniformis P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115704; Current name: Multicellites reniformis (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.77. Species: M. reticulatus B. Samant & Tapaswi 2000; Index Fungorum Registration
Identifier: 519788; Current name: Multicellites reticulatus (B. Samant & Tapaswi) R.K.
Saxena & S.K.M. Tripathi 2011 fide Saxena & Tripathi (2011); Notes: This species does not
possess a longitudinal furrow, an essential character of Multicellaesporites (sensu Kalgutkar
& Jansonius 2000) and therefore Saxena & Tripathi (2011) transferred this species to
Multicellites Kalgutkar & Janson. 2000.
3.26.78. Species: M. sacciformis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111674; Current name: Pluricellaesporites sacciformis (Sheffy & Dilcher) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).

786
3.26.79. Species: M. serpentinus Debi Mukh. 2012; Index Fungorum Registration Identifier:
565666; Location: Neyveli Lignite Mine-I, Cuddalore District, Tamil Nadu, India; Age:
Miocene (Neyveli Lignite).
3.26.80. Species: M. simplicissimus Sheffy & Dilcher 1971; Index Fungorum Registration
Identifier: 111675; Current name: Multicellites simplicissimus (Sheffy & Dilcher) Kalgutkar
& Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.81. Species: M. songii Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483446; Basionym: Diporicellaesporites lanceolatus Z.C. Song 1985; Location:
Jiandingshan, Qaidam Basin, Qinghai Province, China; Age: Early-Middle Miocene; Notes:
Song (1985) assigned this form to Diporicellaesporites. Kalgutkar & Jansonius (2000) could
not observe pore at any of the terminal poles. They, rather, saw a small incision at the lower
end that lines up with the longitudinal furrow that is well expressed near the left margin. They
therefore placed this species under Multicellaesporites (with a question mark). We hereby
confirm, with certainty, the transfer of this species to Multicellaesporites.
3.26.82. Species: M. spatulatus Z.C. Song, Qian & Y.H. Zheng in Z.C. Song et al. 1999 (nom.
inval.) fide Kalgutkar & Jansonius (2000); Index Fungorum Registration Identifier: 483830;
Notes: This name was not validly published because the author did not specify where the
holotype was deposited, and did not provide a Latin description or its English translation.
3.26.83. Species: M. spindlus Z.C. Song, Qian & Y.H. Zheng in Z.C. Song et al. 1999 (nom. inval.)
fide Kalgutkar & Jansonius (2000); Index Fungorum Registration Identifier: 483831; Notes:
This name was not validly published because the author did not specify where the holotype is
stored, and did not provide a Latin description or its English translation.
3.26.84. Species: M. squamosus Debi Mukh. 2012; Index Fungorum Registration Identifier:
565667; Location: Neyveli Lignite Mine-I, Cuddalore District, Tamil Nadu, India; Age:
Miocene (Neyveli Lignite); Notes: Mukherjee (2012) placed this species in Ascomycota.
3.26.85. Species: M. subglobosus Zhong Y. Zhang 1980; Index Fungorum Registration Identifier:
485002; Current name: Pluricellaesporites subglobosus (Zhong Y. Zhang) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.86. Species: M. tener P. Ke & Z.Y. Shi 1978, Orth. corr. pro Multicellaesporites tenerus P. Ke
& Z.Y. Shi 1978; Current name: Multicellites tener (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.87. Species: M. tricellatus Anil Chandra et al. 1984; Index Fungorum Registration Identifier:
107040; Current name: Pluricellaesporites tricellatus (Anil Chandra et al.) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.26.88. Species: M. tricyclosus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115692; Current name: Multicellites tricyclosus (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.89. Species: M. vermiculatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115703; Current name: Multicellites vermiculatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.26.90. Species: M. verus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115669; Current name: Biporipsilonites verus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.26.91. Species: M. volubilis P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115633; Current name: Multicellites volubilis (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).

3.27. Genus: MULTICELLITES Kalgutkar & Janson. 2000, AASP Contributions Series (Dallas)
39: 189 (2000); Index Fungorum Registration Identifier: 28622; Type: M. tener (P. Ke & Z.Y. Shi)
Kalgutkar & Janson. 2000.
Original Diagnosis: Multicellate, uniserial, inaperturate fungal spores; number of cells three
to many, terminal cells usually rounded; spore wall usually smooth, of medium thickness, usually

787
thinner than the septa (or septal bases); septa generally perforate, or with septal folds (Kalgutkar &
Jansonius 2000).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: 46.
Notes: The generic name is derived from the main diagnostic character, which is the presence
of many (Latin multus) cells in each linear spore.
3.27.1. Species: M. allomorphus (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483447; Basionym: Multicellaesporites allomorphus Sheffy &
Dilcher 1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation); Notes: The specific epithet allomorphus
refers to the strange shape of the spore.
3.27.2. Species: M. bellus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483448; Basionym: Multicellaesporites bellus P. Ke & Z.Y. Shi 1978;
Location: Kenli, Shandong Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
3.27.3. Species: M. camerounensis (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483449; Basionym: Chaetosphaerites camerounensis Sal.-
Cheb. & Locq. 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon,
Africa; Age: Oligocene; Notes: Salard-Cheboldaeff & Locquin (1980) suggested affinity of
this species Ascomycota.
3.27.4. Species: M. chandrae R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 11Z); Index Fungorum
Registration Identifier: 519947; Location: Sediment core no. 2 (Lat. 18°35.2'N: Long.
69°17.2'E), Arabian Sea; Age: Late Quaternary; Notes: The species epithet is in honour of Dr.
Anil Chandra, Birbal Sahni Institute of Palaeosciences, Lucknow, India.
3.27.5. Species: M. cingulatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483450; Basionym: Multicellaesporites cingulatus P. Ke & Z.Y. Shi
1978; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
3.27.6. Species: M. circularis (B. Samant & Tapaswi) R.K. Saxena & S.K.M. Tripathi 2011 (Fig.
11AA); Index Fungorum Registration Identifier: 519935; Basionym: Multicellaesporites
circularis B. Samant & Tapaswi 2000; Location: Cambay Basin Gujarat, India; Age: Early
Eocene (Cambay Shale).
3.27.7. Species: M. clarkei (Sat. K. Srivast.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483451; Basionym: Pluricellaesporites clarkei Sat. K. Srivast. 1968;
Location: East Coulee locality, sec. 27, twp. 27, rge. 18, W. 4th mer., Alberta, Canada; Age:
Maastrichtian; Notes: The specific epithet is given in honour of Professor Robert T. Clarke.
3.27.8. Species: M. compactilis (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483452; Basionym: Multicellaesporites compactilis P. Ke & Z.Y. Shi
1978; Location: Guangrao, Shandong Province, Coastal region of Bohai, China; Age:
Eocene-Oligocene.
3.27.9. Species: M. confusus (Anil Chandra et al.) Kalgutkar & Janson. 2000 (Fig. 11AB); Index
Fungorum Registration Identifier: 483453; Basionym: Multicellaesporites confusus Anil
Chandra et al. 1984; Location: Sediment core no. 4 (Lat. 21°10.0′N: Long. 70°26.9′E),
Arabian Sea; Age: Late Quaternary.
3.27.10. Species: M. conspicuus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483454; Basionym: Multicellaesporites conspicuus P. Ke & Z.Y. Shi
1978; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
3.27.11. Species: M. crassisporus (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000 (Fig. 11AC);
Index Fungorum Registration Identifier: 483455; Basionym: Multicellaesporites crassisporus
Sal.-Cheb. & Locq. 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon,

788
 Africa; Age: Late Eocene-Oligocene; Notes: Salard-Cheboldaeff & Locquin (1980) suggested
affinity of this species with Ascomycota.
3.27.12. Species: M. desmodes (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483456; Basionym: Multicellaesporites desmodes P. Ke & Z.Y. Shi
1978; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
3.27.13. Species: M. didymus (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483457; Basionym: Multicellaesporites didymus Sheffy & Dilcher
1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.;
Age: Middle Eocene (Claiborne Formation); Notes: The specific epithet didymus means twin
or similar pairs, referring to the two pairs of cells separated by the equatorial constriction.
3.27.14. Species: M. doualae (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483458; Basionym: Multicellaesporites doualae Sal.-Cheb. & Locq.
1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age: Early
Miocene; Notes: Salard-Cheboldaeff and Locquin (1980) suggested affinity of this species
with Ascomycota.
3.27.15. Species: M. ellipticus (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483459; Basionym: Multicellaesporites ellipticus Sheffy & Dilcher
1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.;
Age: Middle Eocene (Claiborne Formation).
3.27.16. Species: M. elsikii (R.K. Kar & R.K. Saxena) Kalgutkar & Janson. 2000 (Fig. 11AD);
Index Fungorum Registration Identifier: 483460; Basionym: Multicellaesporites elsikii R.K.
Kar & R.K. Saxena 1976; Location: Bhuj-Lakhpat Road, Matanomadh, Kutch District,
Gujarat, India; Age: Palaeocene (Matanomadh Formation).
3.27.17. Species: M. erdtmanii (Hammen) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483461; Basionym: Pluricellaesporites erdtmanii Hammen 1954;
Location: Magdalena Valley, Eastern Cordellera, Colombia, South America; Age:
Maastrichtian.
3.27.18. Species: M. evidens (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483462; Basionym: Multicellaesporites evidens P. Ke & Z.Y. Shi
1978; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
3.27.19. Species: M. fusus (V.S. Ediger) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483463; Basionym: Multicellaesporites fusus V.S. Ediger 1981; Location: Thrace
Basin, Turkey; Age: Upper Eocene-Oligocene, Miocene-Pliocene.
3.27.20. Species: M. grandiusculus (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483464; Basionym: Multicellaesporites grandiusculus
Sheffy & Dilcher 1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County,
Tennessee, U.S.A.; Age: Middle Eocene (Claiborne Formation); Notes: The specific epithet
indicates that the spore is somewhat large in size.
3.27.21. Species: M. granulatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483465; Basionym: Multicellaesporites granulatus P. Ke & Z.Y. Shi
1978; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
3.27.22. Species: M. himalayaensis (A. Gupta) R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 11AE);
Index Fungorum Registration Identifier: 519900; Basionym: Multicellaesporites
himalayaensis A. Gupta 2002; Location: Dadahu Road Section, Sirmaur District, Himachal
Pradesh, India; Age: Late Palaeocene to Late Eocene (Subathu Formation).
3.27.23. Species: M. jainii (A. Gupta) R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 11AF); Index
Fungorum Registration Identifier: 519901; Basionym: Multicellaesporites jainii A. Gupta
2002; Location: Dadahu Road Section, Sirmaur District, Himachal Pradesh, India; Age: Late

789
 Palaeocene to Late Eocene (Subathu Formation); Notes: The species epithet honours Dr. K.P.
Jain, Birbal Sahni Institute of Palaeosciences, Lucknow, India.
3.27.24. Species: M. krauselii (Hammen) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483466; Basionym: Pluricellaesporites krauselii Hammen 1954; Location:
Magdalena Valley, Eastern Cordellera, Colombia, South America; Age: Maastrichtian.
3.27.25. Species: M. lanceolatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483467; Basionym: Multicellaesporites lanceolatus P. Ke & Z.Y. Shi
1978; Location: Huangxian, Shandong Province, Coastal region of Bohai, China; Age:
Eocene-Oligocene.
3.27.26. Species: M. leptaleus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483468; Basionym: Multicellaesporites leptaleus Ke &Shi 1978;
Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
3.27.27. Species: M. ligeae (Félix) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483469; Basionym: Leptosphaerites ligeae Felix 1894; Location: Perekeschkul,
near Baku, Azerbaijan; Age: Eocene.
3.27.28. Species: M. littoralis (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483470; Basionym: Multicellaesporites littoralis Sal.-Cheb. & Locq.
1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene; Notes: Salard-Cheboldaeff & Locquin (1980) suggested affinity of this species to
Ascomycota.
3.27.29. Species: M. margaritus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483471; Basionym: Multicellaesporites margaritus P. Ke & Z.Y. Shi
1978; Location: Panshan, Liaoning Province; Beidagang, Tianjin Municipality, Coastal
region of Bohai, China; Age: Eocene-Oligocene.
3.27.30. Species: M. oculeus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483472; Basionym: Multicellaesporites oculeus P. Ke & Z.Y. Shi
1978; Location: Kenli, Shandong Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
3.27.31. Species: M. ovatoides (Z.C. Song & G.X. Li in Z.C. Song et al.) Kalgutkar & Janson.
2000; Index Fungorum Registration Identifier: 483473; Basionym: Multicellaesporites
ovatoides Z.C. Song & G.X. Li in Z.C. Song et al. 1989; Location: Qingfeng county of Henan
Province, China; Age: Late Oligocene (Dongying Formation).
3.27.32. Species: M. ovatus (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483474; Basionym: Multicellaesporites ovatus Sheffy & Dilcher
1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.;
Age: Middle Eocene (Claiborne Formation).
3.27.33. Species: M. pachydermus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483475; Basionym: Multicellaesporites pachydermus P. Ke & Z.Y.
Shi 1978; Location: Kenli, Shandong Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene; Notes: The species epithet refers to the indicates thicker spore wall.
3.27.34. Species: M. pachysporus (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483476; Basionym: Multicellaesporites pachysporus Sal.-
Cheb. & Locq. 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon,
Africa; Age: Late Eocene-Oligocene; Notes: Salard-Cheboldaeff & Locquin (1980) suggested
affinity of this species Ascomycota.
3.27.35. Species: M. pandus (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483477; Basionym: Multicellaesporites pandus Sheffy & Dilcher
1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.;
Age: Middle Eocene (Claiborne Formation); Notes: The specific name pandus means bent,
crooked, or curved, which describes the shape of the spore.

790
Figure 11 – A–AI Phragmosporae. A Cladosporiumsporinites cylindricus Debi Mukh. 2012, Bar =
15 μm. B Diporicellaesporites concavus P. Kumar 1990, Bar = 10 μm. C Diporicellaesporites
dilcheri (Anil Chandra et al.) Kalgutkar & Janson. 2000, Bar = 20 μm. D Diporicellaesporites
fusiformis Ramanujam & Srisailam 1980, Bar = 10 μm. E Diporicellaesporites pluricellus R.K. Kar
& R.K. Saxena 1976, Bar = 20 μm. F Diporicellaesporites samantiae R.K. Saxena 2009, Bar = 10
μm. G Diporicellaesporites stacyi Elsik 1968, Bar = 10 μm. H Diporicellaesporites
tiruchirappalliensis R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk sp. nov., Bar = 8
μm. I Diporicellaesporites verrucatus H.P. Singh et al. 1986, Bar = 50 μm. J Diporipollis

791
assamicus S.K. Dutta & S.C.D. Sah 1970, Bar = 10 μm. K Dwayabeejaesporonites undulatus Debi
Mukh. 2012, Bar = 8 μm. L Edmundmasonaesporites globulatus Debi Mukh. 2012, Bar = 12 μm.
M Foveoletisporonites indicus Ramanujam & Srisailam 1980, Bar = 20 μm. N Foveoletisporonites
keralensis R.K. Saxena & S.K.M. Tripathi 2011, Bar = 30 μm. O Foveoletisporonites miocenicus
Ramanujam & K.P. Rao 1978, Bar = 12 μm. P Fractisporonites canalis R.T. Clarke 1965, Bar = 15
μm. Q Hapalophragmites cumminsii Ramanujam & Ramachar 1980, Bar = 20 μm. R
Heterocystinella bulbosa Cookson & Eisenack 1979, Bar = 10 μm. S Jansoniisporites
endophragmia (Kalgutkar & Sigler) Kalgutkar 1997, Bar = 4 μm. T Kumarisporites ramanujamii
(P. Kumar) Kalgutkar & Janson. 2000, Bar = 12 μm. U Mathurisporites ellipticus (Y.K. Mathur &
K. Mathur) Kalgutkar & Janson. 2000, Bar = 10 μm. V Monilites albidus Pampal. 1902, Bar = 12
μm. W Multicellaesporites denticulatus (Ramanujam & K.P. Rao) Kalgutkar & Janson. 2000, Bar
= 20 μm. X Multicellaesporites dilcheri B. Samant in R.K. Saxena 2009, Bar = 5 μm. Y
Multicellaesporites nortonii Elsik 1968, Bar = 12 μm. Z Multicellites chandrae R.K. Saxena &
S.K.M. Tripathi 2011, Bar = 10 μm. AA Multicellites circularis (B. Samant & Tapaswi) R.K.
Saxena & S.K.M. Tripathi 2011, Bar = 10 μm. AB Multicellites confusus (Anil Chandra et al.)
Kalgutkar & Janson. 2000, Bar = 20 μm. AC Multicellites crassisporus (Sal.-Cheb. & Locq.)
Kalgutkar & Janson. 2000, Bar = 20 μm. AD Multicellites elsikii (R.K. Kar & R.K. Saxena)
Kalgutkar & Janson. 2000, Bar = 15 μm. AE Multicellites himalayaensis (A. Gupta) R.K. Saxena &
S.K.M. Tripathi 2011, Bar = 10 μm. AF Multicellites jainii (A. Gupta) R.K. Saxena & S.K.M.
Tripathi 2011, Bar = 5 μm. AG Multicellites psilatus (R.K. Saxena) R.K. Saxena & S.K.M. Tripathi
2011, Bar = 5 μm. AH Multicellites ramanujamii (A. Gupta) R.K. Saxena & S.K.M. Tripathi 2011,
Bar = 10 μm. AI Multicellites reticulatus (B. Samant & Tapaswi) R.K. Saxena & S.K.M. Tripathi
2011, Bar = 5 μm. AJ Multicellites tamilensis R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde &
P.M. Kirk sp. nov., Bar = 20 μm.

3.27.36. Species: M. psilatus (R.K. Saxena) R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 11AG);
Index Fungorum Registration Identifier: 519903; Basionym: Multicellaesporites psilatus
R.K. Saxena 2009 (= Multicellaesporites elongatus B. Samant 2000, nom. inval. fide Saxena
2009); Location: Bhavnagar, Cambay Basin, Gujarat, India; Age: Early Eocene (Kharsalia
Clay Formation).
3.27.37. Species: M. quattorodecimcellus (Mart.-Hern. & Tom.-Ort.) Kalgutkar & Janson. 2000;
Index Fungorum Registration Identifier: 271940; Basionym: Multicellaesporites
quattorodecimcellus Mart.-Hern. & Tom.-Ort. 1989; Location: Piedras Negras, Coahuila
State, Mexico; Age: Maastrichtian; Notes: The species epithet quattorodecimcellus refers to
the spore having 14 cells.
3.27.38. Species: M. ramanujamii (A. Gupta) R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 11AH);
Index Fungorum Registration Identifier: 519902; Basionym: Multicellaesporites ramanujamii
A. Gupta 2002; Location: Jamath Road Section, Sirmaur District, Himachal Pradesh, India;
Age: Late Palaeocene to Late Eocene (Subathu Formation).
3.27.39. Species: M. reniformis (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483479; Basionym: Multicellaesporites reniformis P. Ke & Z.Y. Shi
1978; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
3.27.40. Species: M. reticulatus (B. Samant & Tapaswi) R.K. Saxena & S.K.M. Tripathi 2011 (Fig.
11AI); Index Fungorum Registration Identifier: 519936; Basionym: Multicellaesporites B.
Samant & Tapaswi 2000; Location: Surat District, Gujarat, India; Age: Early Eocene.
3.27.41. Species: M. simplicissimus (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483480; Basionym: Multicellaesporites simplicissimus
Sheffy & Dilcher 1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County,
Tennessee, U.S.A.; Age: Middle Eocene (Claiborne Formation).

792
3.27.42. Species: M. tamilensis R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk sp. nov.
(Fig. 11AJ); This new species is described under the section "New species and new
combinations".
3.27.43. Species: M. tener (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000 (Fig. 12A); Index
Fungorum Registration Identifier: 483481; Basionym: Multicellaesporites tener P. Ke & Z.Y.
Shi 1978; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age:
Eocene-Oligocene; Notes: The species epithet refers to the comparative thinness and fragility
of the spore wall.
3.27.44. Species: M. tricyclosus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483482; Basionym: Multicellaesporites tricyclosus P. Ke & Z.Y. Shi
1978; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
3.27.45 Species: M. vermiculatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483483; Basionym: Multicellaesporites vermiculatus P. Ke & Z.Y.
Shi 1978; Location: Huangxian, Shandong Province, Coastal region of Bohai, China; Age:
Eocene-Oligocene.
3.27.46. Species: M. volubilis (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483484; Basionym: Multicellaesporites volubilis P. Ke & Z.Y. Shi
1978; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.

3.28. Genus: ORNASPORONITES Proc. IV International Palynology Conference, Lucknow

1976–1977 1: 298 (1978); Index Fungorum Registration Identifier: 21200; Type: O. inaequalis
Ramanujam & K.P. Rao 1978.
Original Diagnosis: Spores brownish yellow to pale yellow, 4-celled, fusiform, diporate, cells
unequal in size, basal and apical cells much smaller than two central cells; transverse septa three,
central septum straight, other two septa curved. One simple pore at each end of spore along its long
axis. Spore wall rugulate-reticuloid (Ramanujam & Rao 1978).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: One.
Notes: This monotypic genus differs from Fusiformisporites because the latter possesses only
a single septum and is longitudinally ribbed and inaperturate.
3.28.1. Species: O. inaequalis Ramanujam & K.P. Rao 1978 (Fig. 12B); Index Fungorum
Registration Identifier: 115078; Location: Alleppey, Alappuzha District, Kerala, India; Age:
Miocene (Quilon and Warkalli beds).

3.29. Genus: PALAEOCURVULARIA Dörfelt & A.R. Schmidt in A.R. Schmidt et al.,
Palaeontographica Abt. B 283(4-6): 161 (2010); Index Fungorum Registration Identifier: 622339;
Type: P. variabilis Dörfelt & A.R. Schmidt in A.R. Schmidt et al. 2010.
Original Diagnosis: Conidiogenous fungus having characters of extant genera
Helminthosporium, Drechslera, Curvularia, Bipolaris and Exserohilum. Conidia pigmented, one to
eight celled, 1–7 transverse septa present, rarely unicellular. Rarely occurring single-celled spores
possess a basal hilar appendix, while the multicellular spores possess a basal hilar cell (Schmidt et
al. 2010).
Classification: Pleosporaceae, Pleosporales, Pleosporomycetidae, Dothideomycetes,
Pezizomycotina, Ascomycota.
Number of species known: One.
3.29.1. Species: P. variabilis Dörfelt & A.R. Schmidt in A.R. Schmidt et al. 2010 (Fig. 12C); Index
Fungorum Registration Identifier: 622340; Location: Ethiopia; Age: Cretaceous; Notes: The
fungus was recorded from the faecal pellets of insect embedded in amber, collected from the
Cretaceous sediments of Ethiopia.

793
3.30. Genus: PARAGRANATISPORITES Zhong Y. Zhang, Acta palaeont. sin. 19(4): 298 (1980);
Index Fungorum Registration Identifier: 28685; Type: P. lunpolaensis Zhong Y. Zhang 1980.
Original Diagnosis: Fungal spores consisting of three or more cells, with a single germinal
aperture, and noticeably asymmetrical around their long axis. One or two cells at the inaperturate
end of the spore are commonly quite large, giving the spore a swollen appearance in that area. The
germinal aperture at the narrow end of the spore, offset from its long axis. Spore wall psilate to
scabrate. Two or more transverse septa are observed (Zhang 1980).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: Five.
Notes: Zhang (1980) distinguished Paragranatisporites from Granatisporites Elsik & Janson.
1974 by the fact that its spores are conspicuously asymmetrical about their long axis, whereas those
of Granatisporites are symmetrical or nearly symmetrical around the long axis. Kalgutkar &
Jansonius (2000) considered that Granatisporites Elsik & Janson. 1974 is a later synonym of
Brachysporisporites R.T. Lange & P.H. Sm. 1971. In Palaeocirrenalia Ramanujam & Srisailam
1980, the long axis of the spore is more curved to helicoidal; also, it has a more than hemispherical,
darkly pigmented distal cell.
3.30.1. Species: P. conicus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483279; Basionym: Multicellaesporites conicus P. Ke & Z.Y. Shi
1978; Synonym: Brachysporisporites conicus (P. Ke & Z.Y. Shi) Norris 1997 fide Kalgutkar
& Jansonius (2000); Location: Panshan, Liaoning Province, Coastal region of Bohai, China;
Age: Eocene-Oligocene.
3.30.2. Species: P. incurvus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483280; Basionym: Multicellaesporites incurvus P. Ke & Z.Y. Shi
1978; Location: Kenli, Shandong Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
3.30.3. Species: P. lunpolaensis Zhong Y. Zhang 1980 (Fig. 12D); Index Fungorum Registration
Identifier: 484998; Location: Lunpola Basin, northern Xizang Plateau, Xizang, China; Age:
Oligocene (Niubao Formation).
3.30.4. Species: P. putus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483508; Basionym: Involutisporonites putus P. Ke & Z.Y. Shi 1978;
Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
3.30.5. Species: P. vermiculus (V.S. Ediger) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483509; Basionym: Pluricellaesporites vermiculus V.S. Ediger 1981;
Location: Thrace Basin, Turkey; Age: Late Eocene-Oligocene, Miocene-Pliocene.

3.31. Genus: PHIALOPHORONITES Debi Mukh., International Journal of Geology, Earth and
Environmental Sciences 2(2): 5 (2012); Index Fungorum Registration Identifier: 588465; Type: P.
magnus Debi Mukh. 2012.
Original Diagnosis: Phialophores in a vesicle (head-like structure), generally solitary, colour
light brown; circular to subcircular shape; size 600–700 × 350–400 μm; 30–40 microspores per
phialophore; size of the microspores 40–80 μm, spore exine laevigate (Mukherjee 2012).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: One.
Notes: Similar spore bearing bodies of Phialophora, a hyphomycetous taxon, are known to
occur generally on humic matters (Subramanian 1971). The name of the genus is based on extant
Phialophora.
3.31.1. Species: P. magnus Debi Mukh. 2012 (Fig. 12E); Index Fungorum Registration Identifier:
588478; Location: Neyveli Lignite Mine-I, Cuddalore District, Tamil Nadu, India; Age:
Miocene (Neyveli Lignite); Notes: The species epithet refers to the large size of the spores.

794
3.32. Genus: PIRIURELLA Cookson & Eisenack, Neues Jb. für Geologie und Paläontologie (2): 78
(1979); Index Fungorum Registration Identifier: 21250; Type: P. elongata Cookson & Eisenack
1979; Current name: PLURICELLAESPORITES Hammen 1954 fide Kalgutkar & Jansonius
(2000).
Original Diagnosis: Coenobium elongate pear-shaped, flat, in its broader parts with paired
cells, in the narrower parts with a row of single cells. In immature forms with only a few cells lined
up in a row (Jansonius & Hills 1982).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: Three (all the species have been transferred to other genera).
Notes: Smith & Chaloner (1979) opined that Piriurella is a conidium of an Alternaria-like
taxon. Elsik (1992) accepted similarity of Piriurella to Alternaria and commented that the name
has priority only if a form genus was created, i.e. if it is a fossil form and not a modern
contaminant. As Piriurella is considered to include fossil fungal spores with an affinity to modern
Alternaria, Kalgutkar & Sigler (1995) described their Alternaria-type conidia under Piriurella.
Kalgutkar & Jansonius (2000) transferred the type species, P. elongatus, to Pluricellaesporites (as
P. cooksoniae Kalgutkar & Janson. 2000) making Piriurella a later synonym of Pluricellaesporites
Hammen 1954.
3.32.1. Species: P. acuminata (Rouse & Mustard) Parsons & G. Norris 1999; Index Fungorum
Registration Identifier: 483924; Current name: Polycellaesporonites acuminatus (Rouse &
Mustard) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.32.2. Species: P. alternariata Kalgutkar & Sigler 1995; Index Fungorum Registration Identifier:
413840; Current name: Polycellaesporonites alternariatus (Kalgutkar & Sigler) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.32.3. Species: P. elongata Cookson & Eisenack 1979; Index Fungorum Registration Identifier:
483892; Current name: Pluricellaesporites cooksoniae Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).

3.33. Genus: PLURICELLAESPORITES Hammen, Bol. Geol. (Bogota) 2(1): 83 (1954); Index
Fungorum Registration Identifier: 21255; Type: P. typicus Hammen 1954 (designated by Van der
Hammen 1955: 14).
Synonym: Piriurella Cookson & Eisenack 1979 fide Kalgutkar & Jansonius (2000), Index
Fungorum Registration Identifier: 21250.
Original Diagnosis: Fungal spores composed of several [grains or] cells aligned along a
single axis (Van der Hammen 1954).
Restated diagnosis: Fungal spores uniseriate, individuals consisting of five to many cells,
cells flattened at common boundary, convex on sides, each cell connected by a slit-like opening
through the septa (Clarke 1965).
Emended Diagnoses: Diagnosis of Pluricellaesporites was emended by Elsik (1968), Sheffy
& Dilcher (1971) and Elsik & Jansonius (1974), as follows: Monoporate, psilate fungal or algal
spores of three or more cells. Symmetrical or nearly symmetrical around one long axis. Two or
more septa (Elsik 1968); Monoporate, psilate to scabrate fungal or algal spores of three or more
cells; two or more septa. Cells linear along one long axis (Sheffy & Dilcher 1971); Fungal spores of
three or more cells, two or more septa, symmetrical or very nearly so around the long axis. There is
a single aperture, pore, hilum or exitus, at one end. Septa may be entire, perforate or split. Cells are
short to long in relation to overall spore length. Spore outline is lenticular, oval or cylindrical. One
or two cells at aporate end never constitute the bulk of the spore. Exine is psilate to variously
ornamented; if ornament is present it is subdued, i.e. of low relief (Elsik & Jansonius 1974).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: 84 (but we accept only 56 species as legitimate because 26 species
have been transferred to other genera and two species, viz. P. collaris Z.C. Song, Qian & Y.H.
Zheng in Z.C. Song et al. 1999 and P. lagenosus Z.C. Song, Qian & Y.H. Zheng in Z.C. Song et
al., have not been validly published).

795
 Notes: Lange & Smith (1971) considered the generic circumscription of Pluricellaesporites,
as provided by Van der Hammen (1954), unacceptably broad. Later, Lange & Smith (1975), in their
paper on Ctenosporites and other Palaeogene fungal spores, stated that “Pluricellaesporites is at
risk of becoming a terminological catchall for few-celled linear phragmospores, unless an effective
revision of this group is accomplished soon”. However, in the mean time, Elsik & Jansonius (1974)
emended the diagnosis of Pluricellaesporites to include monoporate/ monohilate, symmetrical or
nearly symmetrical fungal spores of three or more cells showing different characteristics of shape,
size, ornamentation and septation, and made its description more complete and generally
acceptable. Kalgutkar & Jansonius (2000) followed the generic diagnosis, as emended by Elsik &
Jansonius (1974).
3.33.1. Species: P. alleppeyensis Ramanujam & K.P. Rao 1978; Index Fungorum Registration
Identifier: 115085; Current name: Quilonia alleppeyensis (Ramanujam & K.P. Rao)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.33.2. Species: P. annulatus Anil Chandra et al. 1984 (Fig. 12F); Index Fungorum Registration
Identifier: 107177; Location: Sediment core no. 1 (Lat. 17°57.9′N: Long. 70°46.0′E), Arabian
Sea; Age: Late Quaternary; Notes: Chandra et al. (1984) suggested a possible affinity with
Curvularia sp.
3.33.3. Species: P. antarcticus Z.C. Song & Liu Cao 1994; Index Fungorum Registration Identifier:
483771; Current name: Anatolinites antarcticus (Z.C. Song & Liu Cao) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.33.4. Species: P. apiculatus Kalgutkar 1993; Index Fungorum Registration Identifier: 483883;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes:
According to Kalgutkar (1993), this species shows a close affinity to the conidia of the extant
dematiaceous fungus Phaeodactylium alpiniae in general morphology. The species epithet is
derived From the Latin apiculatus, with a point, referring to the spores being pointed at their
base.
3.33.5. Species: P. beaufortensis M.G. Parsons & G. Norris 1999; Index Fungorum Registration
Identifier: 483925; Location: Caribou Hills, Mackenzie River delta, northern Canada; Age:
Late Palaeocene - Early to Middle Eocene; Notes: The species epithet is after the Beaufort
Sea.
3.33.6. Species: P. capsularis (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483515; Basionym: Multicellaesporites capsularis Sheffy & Dilcher
1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.;
Age: Middle Eocene (Claiborne Formation).
3.33.7. Species: P. catenatus Ramanujam & K.P. Rao 1978; Index Fungorum Registration
Identifier: 115086; Current name: Cercosporites catenatus (Ramanujam & K.P. Rao)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.33.8. Species: P. clarkei Sat. K. Srivast. 1968; Index Fungorum Registration Identifier: 337277;
Current name: Multicellites clarkei (Sat. K. Srivast.) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
3.33.9. Species: P. clavellatus Z.C. Song & G.X. Li in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 485254; Current name: Polycellaesporonites clavellatus (Z.C. Song
& G.X. Li in Z.C. Song et al.) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.33.10. Species: P. coahuilensis (Mart.-Hern. & Tom.-Ort.) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483516; Basionym: Granatisporites coahuilensis Mart.-
Hern. & Tom.-Ort. 1989; Location: Piedras Negras, Coahuila State, Mexico; Age:
Maastrichtian; Notes: The species epithet indicates its occurrence in the state of Coahuila.
3.33.11. Species: P. collaris Z.C. Song, Qian & Y.H. Zheng in Z.C. Song et al. 1999 (nom. inval.)
fide Kalgutkar & Jansonius (2000); Index Fungorum Registration Identifier: 483844; Notes:
This name was not validly published because the author did not specify where the holotype is
stored, and did not provide a Latin description or its English translation.

796
3.33.12. Species: P. conspicuus (P. Ke & Z.Y. Shi) G. Norris 1997; Index Fungorum Registration
Identifier: 483792; Basionym: Multicellaesporites conspicuus P. Ke & Z.Y. Shi 1978;
Current name: Multicellites conspicuus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
3.33.13. Species: P. cooksoniae Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483517; Basionym: Piriurella elongata Cookson & Eisenack 1979; Location:
Eucla basin, Western Australia; Age: Middle Cretaceous.
3.33.14. Species: P. delicatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115678; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age:
Eocene-Oligocene.
3.33.15. Species: P. dentatus Trivedi & C.L. Verma 1970 ex Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 320739; Basionym: Pluricellaesporites dentatus Trivedi &
C.L. Verma 1970 (nom. inval.), lectotype was designated by Kalgutkar and Jansonius 2000;
Location: Near Kuala Lumpur, Malaya; Age: Eocene; Notes: Trivedi & Verma (1970) did not
validly publish the name of species because they included two, entirely different, specimens
under the heading “Holotype”. Kalgutkar & Jansonius (2000) validated the species name by
designating the lectotype.
3.33.16. Species: P. edigeri Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483518; Basionym: Multicellaesporites elsikii V.S. Ediger & Alisan 1989; Location:
Northern Thrace Basin, Turkey; Age: Middle?-Late Eocene to Late Oligocene, Miocene-
Pliocene; Notes: The species epithet of the basionym honours Dr. William C. Elsik and the
new name honours Dr. V.S. Ediger, author of the homonym.
3.33.17. Species: P. ellipticus Y.K. Mathur & K. Mathur 1969; Index Fungorum Registration
Identifier: 483856; Current name: Mathurisporites ellipticus (Y.K. Mathur & K. Mathur)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.33.18. Species: P. ellipticus Mart.-Hern. & Tom.-Ort. 1989; Index Fungorum Registration
Identifier: 483808; Current name: Pluricellaesporites mexicanus Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.33.19. Species: P. elongatus Sat. K. Srivast. 1968; Index Fungorum Registration Identifier:
337278; Current name: Fractisporonites elongatus (Sat. K. Srivast.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.33.20. Species: P. elsikii B. Samant & Tapaswi 2000; Index Fungorum Registration Identifier:
515011; Current name: Pluricellaesporites suratensis R.K. Saxena 2009 fide Saxena (2009);
Notes: Pluricellaesporites elsikii B. Samant & Tapaswi 2000 is illegitimate, being later
homonym of Pluricellaesporites elsikii Kalgutkar 1997. Hence, Saxena (2009) replaced it
with a new name Pluricellaesporites suratensis R.K. Saxena 2009.
3.33.21. Species: P. elsikii Kalgutkar 1997; Index Fungorum Registration Identifier: 437932;
Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Teritories, Canada; Age: Late
Palaeocene-Early Eocene; Notes: According to Kalgutkar (1997), this species strikingly
resembles the conidia of modern dematiaceous hyphomycetous fungus Clasterosporium
cocoicola M.B. Ellis (Shaw & Ellis 1959). The species epithet honours Dr. William C. Elsik.
3.33.22. Species: P. eocenicus B. Samant & Tapaswi 2000 (Fig. 12G); Index Fungorum
Registration Identifier: 519789; Location: Surat District, Gujarat, India; Age: Early Eocene
(Cambay Shale).
3.33.23. Species: P. erdtmanii Hammen 1954; Index Fungorum Registration Identifier: 337279;
Current name: Multicellites erdtmanii (Hammen) Kalgutkar & Janson. 2000 fide Kalgutkar
& Jansonius (2000).
3.33.24. Species: P. excipularis Kalgutkar & Sigler 1995; Index Fungorum Registration Identifier:
413859; Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories, Canada;
Age: Late Palaeocene or Early Eocene (Iceberg Bay Formation); Notes: According to
Kalgutkar & Sigler (1995), the conidia of Pluricellaesporites excipularis morphologically
resemble modern Excipularia Sacc. 1884 (Ascomycota genera incertae sedis fide

797
 Wijayawardene et al. 2020a). The species epithet is derived from its apparent affinity to
Excipularia Sacc.
3.33.25. Species: P. filiformis Hammen 1954; Index Fungorum Registration Identifier: 337280;
Current name: Fractisporonites filiformis (Hammen) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
3.33.26. Species: P. fusiformis P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115679; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age:
Eocene-Oligocene.
3.33.27. Species: P. globatus B. Samant in R.K. Saxena 2009 (Fig. 12H); Index Fungorum
Registration Identifier: 515002; Basionym: Pluricellaesporites globatus B. Samant 2000
(nom. inval.); Location: Bhavnagar, Cambay Basin, Gujarat, India; Age: Early Eocene
(Kharsalia Clay Formation).
3.33.28. Species: P. glomeratus Sat. K. Srivast. 1968; Index Fungorum Registration Identifier:
337281; Current name: Mathurisporites glomeratus (Sat. K. Srivast.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.33.29. Species: P. grahamii Mart.-Hern. & Tom.-Ort. 1989; Index Fungorum Registration
Identifier: 483809; Location: Piedras Negras, Coahuila State, Mexico; Age: Maastrichtian;
Notes: The species epithet honours Dr. A. Graham.
3.33.30. Species: P. guptae R.K. Saxena 2009 (Fig. 12I); Index Fungorum Registration Identifier:
515010; Basionym: Pluricellaesporites minutus A. Gupta 2002; Location: Dadahu Road
Section, Sirmaur District, Himachal Pradesh, India; Age: Late Palaeocene to Late Eocene
(Subathu Formation); Notes: Pluricellaesporites guptae R.K. Saxena 2009 is a replacement
name of Pluricellaesporites minutus A. Gupta 2002.
3.33.31. Species: P. heterosporus (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483519; Basionym: Dictyosporites heterosporus Sal.-
Cheb. & Locq. 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon,
Africa; Age: Oligocene.
3.33.32. Species: P. hillsii Elsik 1968 (Fig. 12J); Index Fungorum Registration Identifier: 519795;
Location: Strip mine approximately 7 miles southwest of Rockdale, Milam County, Texas,
U.S.A.; Age: Palaeocene.
3.33.33. Species: P. himachalensis R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 12K); Index
Fungorum Registration Identifier: 519948; Location: Bhakra-Nangal Section, Bilaspur
District, Himachal Pradesh, India; Age: Early Pliocene (Middle Siwalik).
3.33.34. Species: P. idahoensis Traverse & Ash 1994; Index Fungorum Registration Identifier:
363119; Location: Wallowa terrane in Hells Canyon, Idaho, U.S.A.; Age: Early Jurassic to
early Middle Jurassic; Notes: The species epithet idahoensis refers to the state of Idaho.
3.33.35. Species: P. infacetus (Kalgutkar) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483281; Basionym: Brachysporisporites infacetus Kalgutkar 1997; Location:
Kanguk Peninsula, Axel Heiberg Island, Northwest Territories, Canada; Age: Late
Palaeocene-Early Eocene; Notes: The spores of Pluricellaesporites infacetus are similar to
the conidia of Brachysporium bloxami (Cooke) Sacc. 1886 in general appearance, although
the latter are smooth and distally narrow (Kalgutkar 1997). The species epithet indicates
coarse spore surface.
3.33.36. Species: P. informis P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115696; Current name: Quilonia informis (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
3.33.37. Species: P. irregularis (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483520; Basionym: Multicellaesporites irregularis Sheffy & Dilcher
1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.;
Age: Middle Eocene (Claiborne Formation).
3.33.38. Species: P. karii A. Gupta 2002 (Fig. 12L); Index Fungorum Registration Identifier:
540757; Location: Jamtah Road Section, Sirmaur District, Himachal Pradesh, India; Age:

798
 Eocene (Subathu Formation); Notes: The species epithet honours Dr. R.K. Kar, Birbal Sahni
Institute of Palaeosciences, Lucknow, India.
3.33.39. Species: P. keralensis R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 12M); Index Fungorum
Registration Identifier: 519950; Location: Around Kollam and Varkala, Kerala, India; Age:
Miocene.
3.33.40. Species: P. krauselii Hammen 1954; Index Fungorum Registration Identifier: 337282;
Current name: Multicellites krauselii (Hammen) Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
3.33.41. Species: P. lagenosus Z.C. Song, Qian & Y.H. Zheng in Z.C. Song et al. 1999 (nom.
inval.) fide Kalgutkar & Jansonius (2000); Index Fungorum Registration Identifier: 483845;
Notes: Song et al. (1999) did not validly publish this name because they did not specify
where the holotype was deposited, and did not provide a Latin description or its English
translation.
3.33.42. Species: P. lanceolatus Z.C. Song & G.X. Li in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 485247; Current name: Quilonia lanceolata (Z.C. Song & G.X. Li in
Z.C. Song et al.) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.33.43. Species: P. longicollus Sheffy & Dilcher 1971 (Fig. 12N); Index Fungorum Registration
Identifier: 111833; Location: Puryear clay pit, 800 m south of Puryear, Henry County,
Tennessee, U.S.A.; Age: Middle Eocene (Claiborne Formation); Notes: The specific epithet
refers to the presence of a long terminal neck.
3.33.44. Species: P. magnus Rouse & Mustard 1997; Index Fungorum Registration Identifier:
464004; Location: Strait of Georgia, eastern Vancouver Island, the Fraser River lowlands of
southwest British Columbia, Canada, and North-western Washington State, U.S.A.; Age:
Late Eocene-Early Oligocene.
3.33.45. Species: P. malayensis (Trivedi & C.L. Verma) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483240.; Basionym: Alternaria malayensis Trivedi & C.L.
Verma 1970; Location: Near Kuala Lumpur, Malaya; Age: Eocene; Notes: According to
Trivedi & Verma (1970), spores described in this species resemble the conidia of Alternaria
Nees (Pleosporaceae, Pleosporales, Dothideomycetes fide Wijayawardene et al. 2020a),
where conidia divide transversely into two or more by transverse septa, forming two- or more
celled conidial spores. Fossil spore resembles the living spore of Alternaria in all respect
except that they are smaller in size.
3.33.46. Species: P. malevisus Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483521; Basionym: Multicellaesporites fusiformis Sheffy & Dilcher 1971; Location: Puryear
clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.; Age: Middle Eocene
(Claiborne Formation); Notes: According to Sheffy & Dilcher (1971), this spore resembles
the material which Bradley (1931) identified as Leptosphaeria Ces. & De Not.
(Leptosphaeriaceae, Pleosporales, Dothideomycetes fide Wijayawardene et al. 2020a). The
new species epithet is from Latin male = poorly, and visus = observed.
3.33.47. Species: P. maximus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115695; Current name: Quilonia maximus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
3.33.48. Species: P. mehrotrae R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 12O); Index Fungorum
Registration Identifier: 519951; Location: Sonapur-Badarpur Road Section, Jaintia Hills,
Meghalaya and Cachar District, Assam, India; Age: Early Miocene (Bhuban Formation).
3.33.49. Species: P. melanii Elsik 1968; Index Fungorum Registration Identifier: 320740;
Synonym: Granatisporites melanii (Elsik) Elsik & Janson. 1974 fide Kalgutkar & Jansonius
(2000); Current name: Diporipollis melanii (Elsik) Kalgutkar & Janson. 2000 fide Kalgutkar
& Jansonius (2000).
3.33.50. Species: P. mexicanus Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483522; Basionym: Pluricellaesporites ellipticus Mart.-Hern. & Tom.-Ortiz 1989; Location:
Piedras Negras, Coahuila State, Mexico; Age: Maastrichtian; Notes: Pluricellaesporites

799
 mexicanus Kalgutkar & Janson. 2000 is a replacement name of Pluricellaesporites ellipticus
Mart.-Hern. & Tom.-Ortiz 1989.
3.33.51. Species: P. minusculus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111834; Current name: Diporicellaesporites minusculus (Sheffy & Dilcher) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.33.52. Species: P. minutigranulatus Hammen 1954; Index Fungorum Registration Identifier:
337283; Current name: Diporicellaesporites minutigranulatus (Hammen) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.33.53. Species: P. minutus (Trivedi & Verma) ex Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483523; Basionym: Brachysporium minutum Trivedi & C.L. Verma
1970 (nom. inval.), lectotype was designated by Kalgutkar & Jansonius 2000); Location:
Near Kuala Lumpur, Malaya; Age: Eocene; Notes: Trivedi & Verma (1970) did not validly
publish Brachysporium minutum because they included two elements under the designation
“Holotype”. Kalgutkar & Jansonius (2000) designated Fig. 4 of Trivedi & Verma (1970) as
lectotype, and assigned the species to Pluricellaesporites.
3.33.54. Species: P. minutus A. Gupta 2002; Index Fungorum Registration Identifier: 515009;
Current name: Pluricellaesporites guptae R.K. Saxena 2009 fide Saxena (2009); Notes:
Pluricellaesporites minutus A. Gupta 2002 is illegitimate, being later homonym of
Pluricellaesporites minutus Kalgutkar & Janson. 2000. Hence, Saxena (2009) replaced it with
a new name, Pluricellaesporites guptae R.K. Saxena 2009.
3.33.55. Species: P. mirus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115700; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age:
Eocene-Oligocene.
3.33.56. Species: P. misrae Anil Chandra et al. 1984 (Fig. 12P); Index Fungorum Registration
Identifier: 107178; Location: Sediment core no. 4 (Lat. 21°10.0′N: Long. 70°26.9′E), Arabian
Sea; Age: Late Quaternary; Notes: The species epithet honours Professor Ramesh C. Misra.
3.33.57. Species: P. nodosus Sat. K. Srivast. 1968; Index Fungorum Registration Identifier:
337284; Current name: Fractisporonites nodosus (Sat. K. Srivast.) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
3.33.58. Species: P. ocellatus Z.C. Song & Liu Cao 1994; Index Fungorum Registration Identifier:
483772; Location: King George Island, Antarctica; Age: Late Cretaceous; Notes: The species
epithet refers to the shape of small pore.
3.33.59. Species: P. ovatus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111835; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation).
3.33.60. Species: P. patagonicus Bianchin., Alej. Martınez & R.K. Saxena in Alej. Martınez et al.
2016; Index Fungorum Registration Identifier: 812334; Location: Rio Foyel section, Nirihuau
Basin, Argentina; Age: Palaeogene (El Foyel Group); Notes: These spores resemble those of
Bactrodesmium Cooke. The species epithet refers to the zone from which the material was
collected.
3.33.61. Species: P. planus Trivedi & C.L. Verma ex Kalgutkar & Janson. 2000 (Fig. 12Q); Index
Fungorum Registration Identifier: 320741; Basionym: Pluricellaesporites planus Trivedi &
C.L. Verma 1970 (nom. inval.), lectotype was designated by Kalgutkar & Jansonius (2000);
Location: Near Kuala Lumpur, Malaya; Age: Eocene; Notes: Trivedi & Verma (1970) did not
validly publish the name of species as they did not designate the holotype. Kalgutkar &
Jansonius (2000) validated the species name by designating the lectotype.
3.33.62. Species: P. psilatus R.T. Clarke 1965 (Fig. 12R); Index Fungorum Registration Identifier:
337285; Location: Canon City Coalfield, Fremont County, Colorado, U.S.A.; Age: Late
Cretaceous; Notes: The species epithet psilatus indicates smooth cell walls of the spore.
3.33.63. Species: P. rectangulatus Y.K. Mathur & K. Mathur 1969 (Fig. 12S); Index Fungorum
Registration Identifier: 483855; Location: Naera and Baraia area of Kutch District, Gujarat,
India; Age: Pliocene; Notes: According to Mathur & Mathur (1969), the spores of this species

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 are teliospore of Uredinales. The specific epithet is in reference to the rectangular shape of
the cells.
3.33.64. Species: P. regularis J.T.F. Guim. et al. 2013; Index Fungorum Registration Identifier:
637503; Location: BOP2 outcrop, central and coastal Amazon Region, North Brazil; Age:
Miocene (Barreiras Formation); Notes: Guimarães et al. (2013) stated taxonomic affinity of
this species with Curvularia, Pleosporaceae. The species epithet indicates regular size of the
cells.
3.33.65. Species: P. sacciformis (Sheffy & Dilcher) M.G. Parsons & G. Norris 1999; Index
Fungorum Registration Identifier: 483926; Basionym: Multicellaesporites sacciformis Sheffy
& Dilcher 1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County,
Tennessee, U.S.A.; Age: Middle Eocene (Claiborne Formation).
3.33.66. Species: P. scabiosus R.T. Clarke 1965; Index Fungorum Registration Identifier: 337286;
Location: Canon City coal field, Fremont County, Colorado, U.S.A.; Age: Late Cretaceous;
Notes: The specific epithet is Latin for rough and is given in reference to the appearance of
the cell wall.
3.33.67. Species: P. scalaris Kalgutkar 1997; Index Fungorum Registration Identifier: 437933;
Current name: Scolecosporites modicus Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
3.33.68. Species: P. semicircularis Debi Mukh. 2012; Index Fungorum Registration Identifier:
565672; Location: Neyveli Lignite Mine-I, Cuddalore District, Tamil Nadu, India; Age:
Miocene (Neyveli Lignite).
3.33.69. Species: P. serratus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111836; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation); Notes: The specific epithet serratus
means jagged, and refers to the appearance of the septa.
3.33.70. Species: P. sheffyi Mart.-Hern. & Tom.-Ort. 1989; Index Fungorum Registration
Identifier: 483810; Location: Piedras Negras, Coahuila State, Mexico; Age: Maastrichtian;
Notes: The species epithet is in honour of Dr. M.V. Sheffy.
3.33.71. Species: P. simplicissimus Sheffy & Dilcher 1971; Index Fungorum Registration
Identifier: 111837; Location: Puryear clay pit, 800 m south of Puryear, Henry County,
Tennessee, U.S.A.; Age: Middle Eocene (Claiborne Formation).
3.33.72. Species: P. sirmaurensis A. Gupta 2002 (Fig. 12T); Index Fungorum Registration
Identifier: 540758; Location: Dadahu Road Section, Sirmaur District, Himachal Pradesh,
India; Age: Eocene (Subathu Formation).
3.33.73. Species: P. subcapsilaris Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111838; Current name: Anatolinites subcapsilaris (Sheffy & Dilcher) Elsik et al. 1990 fide
Elsik et al. (1990); Notes: Elsik et al. (1990) transferred the species to Anatolinites. They
emended the description of the species, and drew attention to the tendency of the individual
cells of the spore to break loose by splitting along the septa.
3.33.74. Species: P. subglobosus (Zhong Y. Zhang) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483524; Basionym: Multicellaesporites subglobosus Zhong Y. Zhang
1980; Location: Lunpola Basin, northern Xizang Plateau, Xizang, China; Age: Oligocene
(Niubao Formation); Notes: Kalgutkar & Jansonius (2000) transferred the species to
Pluricellaesporites on the basis of the presence of a flat spot at one end of the type specimen
which they considered a hilum.
3.33.75. Species: P. suboblongatus Sheffy & Dilcher 1971; Index Fungorum Registration
Identifier: 111839; Current name: Diporicellaesporites suboblongatus (Sheffy & Dilcher)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
3.33.76. Species: P. suratensis R.K. Saxena 2009 (Fig. 12U); Index Fungorum Registration
Identifier: 515012; Basionym: Pluricellaesporites elsikii B. Samant & Tapaswi 2000;
Location: Surat District, Gujarat, India; Age: Early Eocene (Surat lignite, Cambay Shale);

801
 Notes: Pluricellaesporites suratensis R.K. Saxena 2009 is a replacement name of
Pluricellaesporites elsikii B. Samant & Tapaswi 2000.
3.33.77. Species: P. tamilensis R.K. Saxena & S. Khare 1992 (Fig. 12V); Index Fungorum
Registration Identifier: 483894; Location: Borehole – JC–12, Jayamkondacholapuram, 45 km
south of Neyveli, Tiruchirappalli District, Tamil Nadu, India; Age: Miocene.
3.33.78. Species: P. tenuis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111840; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation); Notes: The species epithet indicates the
long slender shape of the spore.
3.33.79. Species: P. tricellatus (Anil Chandra et al.) Kalgutkar & Janson. 2000 (Fig. 12W); Index
Fungorum Registration Identifier: 483525; Basionym: Multicellaesporites tricellatus Anil
Chandra et al. 1984; Location: Sediment core no. 5 (Lat. 24°04.5′N: Long. 69°26.0′E),
Arabian Sea; Age: Late Quaternary.
3.33.80. Species: P. trichocladites Kalgutkar 1997; Index Fungorum Registration Identifier:
437934; Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Teritories, Canada;
Age: Late Palaeocene-Early Eocene. Etymology: Similarity to dematiaceous fungus
Trichocladium Harz.; Notes: According to Kalgutkar (1997), Pluricellaesporites
trichocladites shows resemblance to conidia of extant dematiaceous fungus Trichocladium
opacum (Corda) S. Hughes in general morphology and in having spores ranging from 2 to
many cells. Conidia of extant Pithomyces griminicolus Roy & Rai described by Matsushima
(1989) also appear similar to spores of Pluricellaesporites trichocladites, but are verrucose.
3.33.81. Species: P. typicus Hammen 1954 (Fig. 12X); Index Fungorum Registration Identifier:
337287; Location: Magdalena Valley, Eastern Cordillera, Colombia, South America; Age:
Maastrichtian.
3.33.82. Species: P. vermiculus V.S. Ediger 1981; Index Fungorum Registration Identifier: 108390;
Current name: Paragranatisporonites vermiculus (V.S. Ediger 1981) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
3.33.83. Species: P. verrucatus H.P. Singh et al. 1986 (Fig. 12Y); Index Fungorum Registration
Identifier: 131934; Location: Sonapur-Badarpur Road section, Jaintia Hills, Meghalaya and
Cachar, Assam, India; Age: Early Miocene (Bhuban Formation).
3.33.84. Species: P. woodianus O'Keefe 2017; Index Fungorum Registration Identifier: 821915;
Location: Tumbes Province, Peru; Age: Miocene (Heath Formation); Notes: The species
epithet honours Dr. Gordon Wood.

3.34. Genus: QUILONIA K.P. Jain & R.C. Gupta, Palaeobotanist 18(2): 180 (1970); Index
Fungorum Registration Identifier: 21272; Type: Q. typica K.P. Jain & R.C. Gupta 1970.
Original Diagnosis: Body multicellular, filamentous. Exine thick, margin undulated. Apical
and basal portions narrow, central [section] wide. Basal stalk prominent with one or two
rectangular thick-walled cells; apical cell mostly incomplete, curved, central portion broad,
elongate with irregularly shaped furrow-like suture, inside the filament at different places occur one
to four small circular, ostiolate bodies (Jain & Gupta 1970).
Emended Diagnosis: Pluricellate hilate fungal spores, with an oval to elongate obpyriform
pigmented central section, the greatest width of which tends to be near the proximal end; spore
distally extended into a (very) elongated multiseptate narrow stalk that terminates in a closed cell,
although the tip of the stalk is commonly lacking; proximally, there is a short tapering stalk with a
hilate scar. Both stalks tend to be thin-walled or hyaline (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: Eleven.
3.34.1. Species: Q. alleppeyensis (Ramanujam & K.P. Rao) Kalgutkar & Janson. 2000 (Fig. 12Z);
Index Fungorum Registration Identifier: 483529; Basionym: Pluricellaesporites
alleppeyensis Ramanujam & K.P. Rao 1978; Location: Alleppey, Alappuzha District, Kerala,
India; Age: Miocene (Quilon and Warkalli beds); Notes: Ramanujam & Rao (1978) stated

802
 that this species shows significant similarity to conidia of the dematiaceous hyphomycetous
taxon Sporidesmium tropicale (Ellis 1974). Sporidesmium is commonly encountered in moist
tropical regions.
3.34.2. Species: Q. attenuata (Ramanujam & Srisailam) Kalgutkar & Janson. 2000 (Fig. 12AA);
Index Fungorum Registration Identifier: 483530; Basionym: Diporicellaesporites attenuatus
Ramanujam & Srisailam 1980; Location: Kannur Beach area, Palayangadi and Cheruvattur,
Kerala, India; Age: Miocene (Warkalli Beds).
3.34.3. Species: Q. hillsii (Kalgutkar) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483531; Basionym: Diporicellaesporites hillsii Kalgutkar 1993; Location: Peel
River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes: The species
epithet is in honour of Dr. L.V. Hills.
3.34.4. Species: Q. informis (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483532; Basionym: Pluricellaesporites informis P. Ke & Z.Y. Shi
1978; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age: Eocene-
Oligocene.
3.34.5. Species: Q. lageniformis (Zhong Y. Zhang) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483533; Basionym: Diporicellaesporites lageniformis Zhong Y.
Zhang 1980; Location: Lunpola Basin, northern Xizang Plateau, Xizang, China; Age:
Oligocene (Niubao Formation).
3.34.6. Species: Q. lanceolata (Z.C. Song & G.X. Li in Z.C. Song et al.) Kalgutkar & Janson.
2000; Index Fungorum Registration Identifier: 483534; Basionym: Pluricellaesporites
lanceolatus Z.C. Song & G.X. Li in Z.C. Song et al. 1989; Location: Qingfeng county of
Henan Province, China; Age: Late Eocene-Early Oligocene (Shahejie Formation).
3.34.7. Species: Q. maxima (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483535; Basionym: Pluricellaesporites maximus P. Ke & Z.Y. Shi
1978; Location: Laoshanggulin, Tianjin Municipality, Coastal region of Bohai, China; Age:
Eocene-Oligocene.
3.34.8. Species: Q. miocenica (H.P. Singh et al.) Kalgutkar & Janson. 2000 (Fig. 12AB); Index
Fungorum Registration Identifier: 483536; Basionym: Inapertisporites miocenicus H.P.
Singh et al. 1986; Location: 173 km stone, Sonapur-Badarpur Road Section, Jaintia Hills,
Meghalaya, India; Age: Early Miocene (Bhuban Formation).
3.34.9. Species: Q. multicellata (R.K. Saxena & S. Khare) Kalgutkar & Janson. 2000 (Fig. 12AC);
Index Fungorum Registration Identifier: 483537; Basionym: Diporicellaesporites
multicellatus R.K. Saxena & S. Khare 1992; Location: Jayamkondacholapurum Well–12, 45
km south of Neyveli, Tiruchirappalli District, Tamil Nadu, India; Age: Eocene (Neyveli
Formation).
3.34.10. Species: Q. prakashii (Anil Chandra et al.) Kalgutkar & Janson. 2000 (Fig. 12AD); Index
Fungorum Registration Identifier: 483538; Basionym: Diporicellaesporites prakashii Anil
Chandra et al. 1984; Location: Sediment core no. 1 (Lat. 17°57.9′N: Long. 70°46.0′E),
Arabian Sea; Age: Late Quaternary; Notes: The species epithet is in honour of Dr. Uttam
Prakash, Birbal Sahni Institute of Palaeosciences, Lucknow, India.
3.34.11. Species: Q. typica K.P. Jain & R.C. Gupta 1970 (Fig. 12AE); Index Fungorum
Registration Identifier: 322212; Location: Padappakkara, Kollam District, Kerala, India; Age:
Miocene; Notes: According to Jain & Gupta (1970), this species most closely resembles the
extant Annellophora (S. Hughes) Ellis 1958.

3.35. Genus: RAMASRICELLITES Kalgutkar & Janson., AASP Contributions Series (Dallas) 39:
259 (2000); Index Fungorum Registration Identifier: 28623; Type: R. differentialis (Ramanujam &
Srisailam) Kalgutkar & Janson 2000.
Original Diagnosis: Fungal spores inaperturate, tetracellate, ellipsoidal, with central cells
broader, thicker walled and more pigmented than the terminal cells; terminal cells thin-walled to

803
hyaline, with rounded ends; septa (or septal bases) thick and dark, evenly spaced (Kalgutkar &
Jansonius 2000).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: Two.
Notes: Kalgutkar & Jansonius (2000) opined that Multicellaesporites differentialis
Ramanujam & Srisailam is a misfit in Multicellaesporites and therefore they proposed
Ramasricellites to accommodate it. The sharp differentiation between the dark, broad central cells
and the narrower, elongate hyaline terminal cells, as well as the lack of constriction at the median
septum, differentiate this form from species in Multicellites. The name of the genus is a composite
of the beginnings of the surnames Ramanujam and Srisailam, the authors of the type species.
3.35.1. Species: R. differentialis (Ramanujam & Srisailam) Kalgutkar & Janson. 2000 (Fig. 12AF);
Index Fungorum Registration Identifier: 483539; Basionym: Multicellaesporites differentialis
Ramanujam & Srisailam 1980; Location: Kannur Beach area, Palayangadi and Cheruvattur
(southern side of Karingottu River), Kerala, India; Age: Miocene (Warkalli Beds); Notes: The
differences in the pigmentation, shape and wall thickness of the terminal two cells from the
central two cells, constitute the diagnostic features of this species. The fossil spore shows
striking resemblances with the conidia of some species of the modern dematiaceous genus
Spiropes Cif., viz. S. clavatus (Ellis & G. Martin) M.B. Ellis and S. effusus (Pat.) M.B. Ellis.
3.35.2. Species: R. lunpolaensis (Z.C. Song in Z.C. Song et al. 1999) Kalgutkar & Janson. 2000;
Index Fungorum Registration Identifier: 483540; Basionym: Multicellaesporites lunpolaensis
Z.C. Song in Z.C. Song et al. 1999; Synonym: Multicellaesporites evidens Zhong Y. Zhang
1980 fide Kalgutkar & Jansonius (2000); Location: Lunpola Basin, Xizang, China; Age:
Middle-Late Oligocene.

3.36. Genus: REDUVIASPORONITES L.R. Wilson, Okla. Geol. Notes 22(4): 91–92 (1962);
Index Fungorum Registration Identifier: 21278; Type: R. catenulatus L.R. Wilson 1962.
Original Diagnosis: Conidia-like spores occurring in uniseriate chains (phragmospores?) of
several or more individuals; subspherical, slightly flattened at the contacts with adjacent spores, all
approximately same diameter, walls 1–2 μm thick, uniform, smooth or slightly rough, yellow or
brown, translucent (Wilson 1962).
Classification: Fungi imperfecti, Phragmosporae.
Number of species known: Nine.
Notes: Wilson (1962) stated that these fossils differ from Penicillites curtipes Berk. 1848
(Eocene), as illustrated in Hirmer (1927), in having spores of subspherical rather than elliptical
shape and walls that are relatively thinner than those in Penicillites. According to him, another
fossil fungus with which Reduviasporonites should be compared is the Eocene Torulites
moniliformis Menge (Casp. 1907). This form is illustrated as having spores that are progressively
smaller towards the apex (Hirmer 1927). The shape of the spores and mycelium of Torulites
resembles more closely the modern Hormiscum Kunze ex Wallr. than with Torula Persoon ex Fr.
The first part of the generic name Reduviasporonites has been derived from the Latin reduvia
(fragment) and suffix -sporonites indicates that the specimens are fossil fungus spores not
organically attached to other structures. Sporonites is used in preference to -sporites to indicate that
the spore is of fungal affinity.
3.36.1. Species: R. anangus G. Norris 1986; Index Fungorum Registration Identifier: 126575;
Location: Imperial Nuktak C–22 Well, Mackenzie Delta Region, District of Mackenzie, N.
W. T., Canada; Age: Oligocene.
3.36.2. Species: R. catenarius (G. Playford) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483541; Basionym: Synsphaeridium catenarium G. Playford in
Playford & Dring 1981; Location: Carnarvon Basin, Western Australia; Age: Late Devonian
(Late? Frasnian).
3.36.3. Species: R. catenulatus L.R. Wilson 1962 (Fig. 12AG); Index Fungorum Registration
Identifier: 338248; Location: North bank of Salt Fork of the Red River, Greer County,

804
 Oklahoma, U.S.A.; Age: Late Permian; Notes: Wilson (1962) observed several hundred
specimens of this species during an examination of 150 microscope-slide preparations of the
Flowerpot Shale. This abundance may indicate that the fungus was of a marine type and
thrived in the shallow waters of the sea; however, no account of modern marine fungus
comparable to Reduviasporonites catenulatus has been found during a search of the literature.
The specific epithet is derived from the Latin catena (chain) and is given in reference to the
occurrence of the spores in chain-like units.
3.36.4. Species: R. cerrejonensis (Doub. & D. Pons) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483542; Basionym: Multicellaesporites cerrejonensis Doub. & D.
Pons 1973; Location: Cerrejon basin, Colombia, South America; Age: Palaeocene-Eocene.
3.36.5. Species: R. curvatus (Ambwani) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483543; Basionym: Multicellaesporites curvatus Ambwani 1983; Location:
Neyveli Lignite Mine, Cuddalore District, Tamil Nadu, India; Age: Late Miocene or Pliocene
(Neyveli Lignite).
3.36.6. Species: R. magnus (Doub. & D. Pons) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483544; Basionym: Dicellaesporites magnus Doub. & D. Pons 1973;
Location: Cerrejon basin, Colombia, South America; Age: Palaeocene-Eocene.
3.36.7. Species: R. prakashii (Ambwani) Kalgutkar & Janson. 2000 (Fig. 12AH); Index Fungorum
Registration Identifier: 483545; Basionym: Multicellaesporites prakashii Ambwani 1982;
Location: Kotta-Bommuru Village, Rajamahendravaram, East Godavari District, Andhra
Pradesh, India; Age: Early Eocene; Notes: The species epithet is in honour of Dr. Uttam
Prakash, Birbal Sahni Institute of Palaeosciences, Lucknow, India.
3.36.8. Species: R. qingfengensis (Z.C. Song & H.C. Luo in Z.C. Song et al.) Z.C. Song in Z.C.
Song et al. 1999; Index Fungorum Registration Identifier: 483832; Basionym:
Multicellaesporites qingfengensis Z.C. Song & H.C. Luo in Z.C. Song et al. 1989; Location:
Qingfeng county of Henan Province, China; Age: Late Eocene-Middle Oligocene (Shahejie
Formation).
3.36.9. Species: R. ramosus Kalgutkar 1993; Index Fungorum Registration Identifier: 483884;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes:
The species epithet is derived from the Latin, ramosus, branched.

3.37. Genus: RETICELLITES D.L.E. Glass et al., Pollen et Spores 28: 414 (1986); Index
Fungorum Registration Identifier: 11439; Type: R. houstonii D.L.E. Glass et al. 1986.
Original Diagnosis: Fungal spores of 3 or more cells, rarely 2, generally of unequal size, with
sculpture of indistinct to distinct reticulum. Spore outline lanceolate, spatulate or ovate, indented or
not across the septa. Axis appears to be straight, but when a pore is present the axis is occasionally
curved through the smaller cell to the pore. Pore present or not, often very obscure. Spore wall of
one or two layers; layers tightly appressed or, less commonly, separated. Spores with melanin
pigment; the outer wall layer over the larger cells and both layers over the smaller cell can be
yellow with practically no brownish component. Septa of two layers; layering not always evident.
Septal pores are obscure in the type species (Glass et al. 1986).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: One.
Notes: The combination of overall shape and the reticulate spore wall is unique for these
spores. The name of the genus denotes sculpture and the multiple number of cells.
3.37.1. Species: R. houstonii D.L.E. Glass et al. 1986 (Fig. 12AI); Index Fungorum Registration
Identifier: 360321; Location: South-central Texas, U.S.A.; Age: Late Eocene (Manning
Formation); Notes: The species epithet is in memory of Sam Houston, on the occasion of the
Texas sesquicentennial.

3.38. Genus: SCOLECOSPORITES R.T. Lange & P.H. Sm., Neues Jb. für Geologie und
Paläontologie 11: 674 (1971); Index Fungorum Registration Identifier: 21296; Type:

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S. maslinensis R.T. Lange & P.H. Sm. 1971.
Original Diagnosis: Scoleco-phragmospores of lengths 15-30 times of the breadth, the outline
of walls and septa ladder-like (Lange & Smith 1971).
Emended Diagnosis: Long to very long, linear filamentous phragmospores, hilate, with or
without distal pore; length many times width of spore. Spores scalariform, commonly broken and
lacking proximal and/ or distal portions; wall and septa commonly thin; septa often with septal
folds. Not or barely indented at septa (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: Five [but we accept only four species as legitimate because one
species, viz. S. vermiculatus (P. Ke & Z.Y. Shi) Z.C. Song 1999, has been transferred to
Multicellites Kalgutkar & Janson. 2000].
Notes: According to Lange & Smith (1971) this species has phragmospores typically of more
than 10 cells and typically with length greater than 4 times breadth. Kalgutkar & Jansonius (2000)
emended the generic diagnosis.
3.38.1. Species: S. longus Z.C. Song & Z.H. Sun in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 637504; Location: Qingfeng county of Henan Province, China; Age:
Late Eocene-Middle Oligocene (Shahejie Formation).
3.38.2. Species: S. maslinensis R.T. Lange & P.H. Sm. 1971 (Fig. 12AJ); Index Fungorum
Registration Identifier: 323295; Location: Maslin Bay, South Australia; Age: Early-Middle
Eocene.
3.38.3. Species: S. modicus Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483547; Basionym: Pluricellaesporites scalaris Kalgutkar 1997; Location: Kanguk
Peninsula, Axel Heiberg Island, Northwest Territories, Canada; Age: Late Palaeocene-Early
Eocene; Notes: The species epithet is derived from Latin modicus = modest, moderate, for its
modest length compared to other species in this genus.
3.38.4. Species: S. scalaris (Kalgutkar) Kalgutkar & Janson. 2000 (Fig. 12AK); Index Fungorum
Registration Identifier: 483548; Basionym: Diporicellaesporites scalaris Kalgutkar 1993;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes:
The species epithet is derived from the Latin, scalaris, scalariform, referring to the ladder-like
appearance of the spores.
3.38.5. Species: S. vermiculatus (P. Ke & Z.Y. Shi) Z.C. Song in Z.C. Song et al. 1999; Index
Fungorum Registration Identifier: 483833; Basionym: Multicellaesporites vermiculatus P. Ke
& Z.Y. Shi 1978; Current name: Multicellites vermiculatus (P. Ke & Z.Y. Shi) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).

3.39. Genus: TRIPITHONITES Sat. K. Srivast. & Al-Tayyar, Savitriana 3: 180 (2013); Index
Fungorum Registration Identifier: 817993; Type: T. amoenus Sat. K. Srivast. & Al-Tayyar 2013.
Original Diagnosis: Tricellate fungal spore, middle cell dark coloured, end-cells hyaline;
middle cell oval to elliptical, middle cell generally with a longitudinal furrow; exine surface
smooth, middle cell with two wall-layers, inner wall layer dark, smooth to striate (Srivastava & Al-
Tayyar 2013).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: Two.
Notes: Tripithonites is distinct from Dicellaesporites Elsik 1968 and Tricellaesporonites
Sheffy & Dilcher 1971, in having two hyaline cells attached at the extremities of a large black cell.
The name of the genus is derived from the Greek tri - = three; pithon = cellar.
3.39.1. Species: T. amoenus Sat. K. Srivast. & Al-Tayyar 2013 (Fig. 12AL); Index Fungorum
Registration Identifier: 818890; Location: Northern Arabian Gulf; Age: Late middle Albian;
Notes: Tripithonites amoenus consists of tricellate fungal spores having a large dark elliptical
cells and two hyaline end-cells. The species epithet is derived from the Latin amoenus =
delightful, lovely.

806
3.39.2. Species: T. argoperatus Sat. K. Srivast. & Al-Tayyar 2013; Index Fungorum Registration
Identifier: 818891; Location: Northern Arabian Gulf; Age: Late middle Albian; Notes: The
species epithet is derived from the Greek argos = white; peratos = end, extremity.

Figure 12 – A–AM Phragmosporae. AN Dictyosporae. A Multicellites tener (P. Ke & Z.Y. Shi)
Kalgutkar & Janson. 2000, Bar = 8 μm. B Ornasporonites inaequalis Ramanujam & K.P. Rao
1978, Bar = 15 μm. C Palaeocurvularia variabilis Dörfelt & A.R. Schmidt 2010, Bar = 10 μm. D

807
Paragranatisporites lunpolaensis Zhong Y. Zhang 1980, Bar = 8 μm. E Phialophoronites magnus
Debi Mukh. 2012, Bar = 120 μm. F Pluricellaesporites annulatus Anil Chandra et al. 1984, Bar =
10 μm. G Pluricellaesporites eocenicus B. Samant & Tapaswi 2000, Bar = 10 μm. H
Pluricellaesporites globatus B. Samant in R.K. Saxena 2009, Bar = 40 μm. I Pluricellaesporites
guptae R.K. Saxena 2009, Bar = 8 μm. J Pluricellaesporites hillsii Elsik 1968, Bar = 10 μm. K
Pluricellaesporites himachalensis R.K. Saxena & S.K.M. Tripathi 2011, Bar = 15 μm. L
Pluricellaesporites karii A. Gupta 2002, Bar = 10 μm. M Pluricellaesporites keralensis R.K.
Saxena & S.K.M. Tripathi 2011, Bar = 30 μm. N Pluricellaesporites longicollus Sheffy & Dilcher
1971, Bar = 5 μm. O Pluricellaesporites mehrotrae R.K. Saxena & S.K.M. Tripathi 2011, Bar = 20
μm. P Pluricellaesporites misrae Anil Chandra et al. 1984, Bar = 10 μm. Q Pluricellaesporites
planus Trivedi & C.L. Verma ex Kalgutkar & Janson. 2000, Bar = 10 μm. R Pluricellaesporites
psilatus R.T. Clarke 1965, Bar = 20 μm. S Pluricellaesporites rectangulatus Y.K. Mathur & K.
Mathur 1969, Bar = 10 μm. T Pluricellaesporites sirmaurensis A. Gupta 2002, Bar = 10 μm. U
Pluricellaesporites suratensis R.K. Saxena 2009, Bar = 10 μm. V Pluricellaesporites tamilensis
R.K. Saxena & S. Khare 1992, Bar = 20 μm. W Pluricellaesporites tricellatus (Anil Chandra et al.)
Kalgutkar & Janson. 2000, Bar = 10 μm. X Pluricellaesporites typicus Hammen 1954, Bar = 20
μm. Y Pluricellaesporites verrucatus H.P. Singh et al. 1986, Bar = 20 μm. Z Quilonia
alleppeyensis (Ramanujam & K.P. Rao) Kalgutkar & Janson. 2000, Bar = 20 μm. AA Quilonia
attenuata (Ramanujam & Srisailam) Kalgutkar & Janson. 2000, Bar = 5 μm. AB Quilonia
miocenica (H.P. Singh et al.) Kalgutkar & Janson. 2000, Bar = 10 μm. AC Quilonia multicellata
(R.K. Saxena & S. Khare) Kalgutkar & Janson. 2000, Bar = 10 μm. AD Quilonia prakashii (Anil
Chandra et al.) Kalgutkar & Janson. 2000, Bar = 10 μm. AE Quilonia typica K.P. Jain & R.C.
Gupta 1970, Bar = 20 μm. AF Ramasricellites differentialis (Ramanujam & Srisailam) Kalgutkar &
Janson. 2000, Bar = 7 μm. AG Reduviasporonites catenulatus L.R. Wilson 1962, Bar = 8 μm. AH
Reduviasporonites prakashii (Ambwani) Kalgutkar & Janson. 2000, Bar = 8 μm. AI Reticellites
houstonii D.L.E. Glass et al. 1986, Bar = 6 μm. AJ Scolecosporites maslinensis R.T. Lange & P.H.
Sm. 1971, Bar = 8 μm. AK Scolecosporites scalaris (Kalgutkar) Kalgutkar & Janson. 2000, Bar =
20 μm. AL Tripithonites amoenus Sat. K. Srivast. & Al-Tayyar 2013, Bar = 6 μm. AM
Varmasporites tonakkalensis (Y.N.R. Varma & R.S. Patil) Kalgutkar & Janson. 2000, Bar = 10 μm.
AN Centonites symmetricus Peppers 1964, Bar = 12 μm.

3.40. Genus: VARMASPORITES Kalgutkar & Janson., AASP Contributions Series (Dallas) 39:
309 (2000); Index Fungorum Registration Identifier: 28626; Type: V. tonakkalensis (Y.N.R.
Varma & R.S. Patil) Kalgutkar & Janson. 2000.
Original Diagnosis: Fusiform, four-celled, inaperturate fungal spores, with a pronounced
constriction at the thick median septum, and with a distinct ribbed or striate sculpture parallel to the
long axis. The two centrifugal septa may be less strongly developed (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: One.
Notes: Kalgutkar & Jansonius (2000) opined that Fusiformisporites tonakkalensis Y.N.R.
Varma & R.S. Patil is a misfit in Fusiformisporites Rouse and therefore they proposed
Varmasporites to accommodate it.
3.40.1. Species: V. tonakkalensis (Y.N.R. Varma & R.S. Patil) Kalgutkar & Janson. 2000 (Fig.
12AM); Index Fungorum Registration Identifier: 483572; Basionym: Fusiformisporites
tonakkalensis Y.N.R. Varma & R.S. Patil 1985; Location: Tonakkal area,
Thiruvananthapuram District, Kerala, India; Age: Miocene.

3.41. Genus: WARKALLISPORONITES Ramanujam & K.P. Rao, Proc. IV International Palynology
Conference, Lucknow 1976–1977 1: 298 (1978); Index Fungorum Registration Identifier: 21361;
Type: W. denticulatus Ramanujam & K.P. Rao 1978; Current name: MULTICELLAESPORITES
Elsik 1968 fide Kalgutkar & Jansonius (2000).

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 Original Diagnosis: Spores simple, melanin-coloured, multicellular, uniseriate, fusiform,
multiporate. Spores distinctly constricted in the central region. Denticulate or wedge-shaped
thickenings on some septa, pore slit-like in each septum. Spore wall granular to scabrate
(Ramanujam & Rao 1978).
Classification: Fungi Imperfecti, Phragmosporae.
Number of species known: One (the single species has been transferred to Multicellaesporites
Elsik 1968).
Notes: Since Kalgutkar & Jansonius (2000) transferred the type species to Multicellaesporites
Elsik 1968, Warkallisporonites Ramanujam & K.P. Rao 1978 became a later taxonomic synonym
of Multicellaesporites Elsik 1968.
3.41.1. Species: W. denticulatus Ramanujam & K.P. Rao 1978; Index Fungorum Registration
Identifier: 115097; Current name: Multicellaesporites denticulatus (Ramanujam & K.P. Rao)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).

4. Dictyosporae

4.1. Genus: ARBUSCULITES Paradkar, Journal of Palynology 10(2): 120 (1976); Index Fungorum
Registration Identifier: 21013; Type: A. dicotylophylli Paradkar 1976; Current name:
DICTYOSPORITES Felix 1894 fide Kalgutkar & Jansonius (2000).
Original Diagnosis: Saprophytic [saprobic] fungus, septate mycelium, dictyospores formed
on a conidial head (Paradkar 1976).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: One (the single species has been transferred to Dictyosporites
Felix 1894).
4.1.1. Species: A. dicotylophylli Paradkar 1976; Index Fungorum Registration Identifier: 483814;
Current name: Dictyosporites dicotylophylli (Paradkar) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).

4.2. Genus: CENTONITES Peppers, Bull. Ill. St. geol. Surv. 90: 47 (1964); Index Fungorum
Registration Identifier: 21032; Type: C. symmetricus Peppers 1964.
Original Diagnosis: The microfossils are composed of 5 to approximately 15 polygonal
segments joined to form radially or bilaterally symmetrical bodies, their symmetry depending upon
their number and arrangement. Most of the microfossils are flatly compressed, but many are folded.
Straight grooves are present on one side of the body where the segments are joined. Elevated,
flattened ridges or folds occur adjacent to and along both sides of the grooves but are absent along
the smooth outer margin of the microfossils. The microfossils usually are made up of an odd
number of segments. The most common variation consists of five segments arranged in a boat-
shaped pattern with two small, pointed segments at one end, two larger segments in the middle, and
one pointed segment at the opposite end. In the largest specimens two to four segments in the
center are almost completely bordered by other segments. Along the margins of the largest
specimens three small indentations may occur, almost equal distances apart, where segments are
absent. These indentations either have smooth edges and look almost like pores or they have
uneven edges as if segments had been torn out. The microfossils are revealed under oil immersion
objective as laevigate. They are 1 to 2 µm thick. The known size range is 50.2 to 123.2 µm in
maximum diameter (Peppers 1964).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: One.
Notes: Elsik (1992) indicated that the spore wall of Centonites appears to be not exclusively
fungal; thus, an algal affinity cannot be ruled out.
4.2.1. Species: C. symmetricus Peppers 1964 (Fig. 12AN); Index Fungorum Registration Identifier:
560969; Location: Illinois, U.S.A.; Age: Late Pennsylvanian; Notes: The species epithet is in
reference to the symmetrical arrangement of the segments.

809
4.3. Genus: CTENOSPORITES Elsik & Janson., Can. J. Bot. 52(5): 956 (1974); Index Fungorum
Registration Identifier: 21066; Type: C. eskerensis Elsik & Janson. 1974.
Original Diagnosis: Multicellular structures of fungal origin; one main stem of a few to
several (commonly seven-nine) cells and lateral or secondary septate branches (cf. filaments) along
one side of the main stem. Main stem and lateral branches are straight to slightly curved; apex of
the main stem may be curved towards the side of the lateral branches; lateral branches are curved
concave to the apex of the main stem. Basal cell of the stem is generally torn and may be thinner-
walled than that portion of the stem bearing branches. Branches are of a few to several (commonly
five-seven) cells. Branches may have incomplete septa but in all cases the apical branch has fewer
septa (i.e., in most cases none) than the basal branches; the progression (with occasional exception)
is one more septum with each additional branch counting from the apex to the base of the stem.
Apical cell(s) of the stem may mimic the most apical branch or may be lacking altogether (Elsik &
Jansonius 1974).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: Three (but we accept only two species as legitimate because one
species, C. wolfei Elsik & Janson. 1974, has been considered to be a synonym of Ctenosporites
eskerensis Elsik & Janson. 1974).
Notes: The species epithet is derived from Greek kteis, ktenos = comb.
4.3.1. Species: C. eskerensis Elsik & Janson. 1974 (Fig. 13A); Index Fungorum Registration
Identifier: 312369; Synonym: Ctenosporites wolfei Elsik & Janson. 1974 (named in honour of
Dr. J.A. Wolfe); Location Gulf of Alaska, Alaska (Elsik & Jansonius 1974), Lower Headon
deposits, Hordle Cliff, Hampshire, England (Smith 1978); Age: Late Eocene-Early
Oligocene; Notes: Smith (1978) considered differences in Ctenosporites eskerensis and C.
wolfei as morphological variations produced by the same mycelium and placed Ctenosporites
wolfei in synonymy with Ctenosporites eskerensis.
4.3.2. Species: C. sherwoodiae Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483290; Location: Fremount County, Colorado, U.S.A.; Age: Late Cretaceous (Vermejo
Formation coal beds); Notes: Kalgutkar & Jansonius (2000) formally named Fungal Spore sp.
A (in Clarke 1965) after Martha Sherwood, in recognition of her important contributions to
the knowledge of fossil fungal spores.
4.3.3. Species: C. wolfei Elsik & Janson. 1974; Index Fungorum Registration Identifier: 312370;
Current name: Ctenosporites eskerensis Elsik & Janson. 1974 fide Kalgutkar & Jansonius
(2000).

4.4. Genus: DACTYLOSPORITES Paradkar 1976; Index Fungorum Registration Identifier: 21070;
Type: D. dicotylophylli Paradkar 1976; Current name: DICTYOSPORITES Felix 1894 fide
Kalgutkar & Jansonius (2000).
Original Diagnosis: Fungus saprophytic [saprobic], mycelium septate, phragmospores smooth
(Paradkar 1976).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: One (the single species has been transferred to Dictyosporites
Felix 1894).
Notes: Dactylosporites Paradkar (1976) is a later synonym of Dictyosporites Felix (1894).
4.4.1. Species: D. dicotylophylli Paradkar 1976; Index Fungorum Registration Identifier: 483815;
Current name: Dictyosporites paradkariae Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).

4.5. Genus: DICTYOSPORITES Félix 1894, Zeitschr. Deutsche Geol. Gesell. 46: 277 (1894);
Index Fungorum Registration Identifier: 21075; Type: D. loculatus Felix 1894 (type selected by
Jansonius & Hills 1976).
Synonyms: Arbusculites Paradkar 1976, Index Fungorum Registration Identifier: 21013;
Dactylosporites Paradkar 1976, Index Fungorum Registration Identifier: 21070; Pleosporonites

810
R.T. Lange & P.H. Sm. 1971, Index Fungorum Registration Identifier: 21253; Ravenelites
Ramanujam & Ramachar 1980, Index Fungorum Registration Identifier: 28630.
Original Diagnosis: The so-called wall-shaped conidia become multicellular by repeated
transverse and longitudinal divisions. In addition to large conidia, whose growth can probably be
regarded as complete, uni- and bicellular conidia representing the initial developmental stages also
occur. They are all of brownish coloration. Their outlines are rather variable, depending on the
position of the conidium to the plane of section. Viewed from the top or bottom, they often appear
spherical with flatly indented outlines; longitudinal sections are of rather irregular shape; elliptical,
pear-shaped or resembling short, corpulent snails (e.g. Turbo). The maximum length is 0.0204 mm
[20.4 μm], the maximum diameter 0.0153 mm [15.3 μm]; the respective dimensions of an only
bicellular conidium are 0.0102 and 0.0085 mm [10.2 and 8.5 μm] (Felix 1894).
Emended Diagnosis: Inaperturate, multicellate (apparently by internal septation, of irregular
pattern), muriform fungal spores, cells rounded to rounded polygonal. Overall shape rounded,
oval/ovoid to elongate; indentations may occur where septa intersect the amb. A hilum cannot be
discerned. Staphlosporonites differs in showing a distinct hilum, or proximal hilar cell.
Papulosporonites consists of spore clusters or aggregates, in which there is no suggestion of linear
or planar symmetry (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: 20 (but we accept only 15 species as legitimate because five
species have been transferred to other genera).
Notes: According to Kalgutkar & Jansonius (2000), different species of Dictyosporites are
comparable to the conidia of some modern genera like Alternaria, Dictyosporium, Septosporium
and Stemphylium, all belonging to dematiaceous hyphomycetes, and the ascospores of Pleospora.
4.5.1. Species: D. dicotylophylli (Paradkar) Kalgutkar & Janson. 2000 (Fig. 13B); Index Fungorum
Registration Identifier: 483297; Basionym: Arbusculites dicotylophylli Paradkar 1976;
Location: Mohgaonkalan, Chhindwara District, Madhya Pradesh, India; Age: Late Cretaceous
(Maastrichtian); Notes: According to Paradkar (1976), this species can be compared with the
Arbuscula Bat. & Peres (Batista & Peres 1965) (Current name: Neoarbuscula B. Sutton 1983
fide Species Fungorum 2021). Arbuscula is recorded on fallen Eugenia leaves. The specific
name is after its occurrence on a dicotyledonous leaf.
4.5.2. Species: D. dictyosus (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483298; Basionym: Pleosporonites dictyosus Sal.-Cheb. & Locq.
1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene.
4.5.3. Species: D. eccentricus Kalgutkar 1993; Index Fungorum Registration Identifier: 483865;
Current name: Kutchiathyrites canadensis Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
4.5.4. Species: D. elsikii Kalgutkar 1993; Index Fungorum Registration Identifier: 483864; Current
name: Staphlosporonites billelsikii Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius
(2000).
4.5.5. Species: D. felixii R.T. Lange & P.H. Sm. 1971; Index Fungorum Registration Identifier:
312993; Current name: Staphlosporonites felixii (R.T. Lange & P.H. Sm.) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
4.5.6. Species: D. firbasii (Hammen) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483299; Basionym: Polyadosporites firbasii Hammen 1954; Location: Magdalena
Valley, Eastern Cordellera, Colombia, South America; Age: Maastrichtian.
4.5.7. Species: D. garciabarrigae (Hammen) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483300; Basionym: Polyadosporites garciabarrigae Hammen 1954;
Location: Magdalena Valley, Eastern Cordellera, Colombia, South America; Age:
Maastrichtian.
4.5.8. Species: D. globimuriformis Kalgutkar 1997; Index Fungorum Registration Identifier:
437903; Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories, Canada;

811
 Age: Late Palaeocene-Early Eocene; Notes: The species epithet refers to the rounded and
muriform nature of spores.
4.5.9. Species: D. heterosporus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107915; Current name: Pluricellaesporites heterosporus (Sal.-Cheb. & Locq.) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
4.5.10. Species: D. hyalinus (R.T. Lange & P.H. Sm. 1971) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 483301; Basionym: Pleosporonites hyalinus R.T. Lange &
P.H. Sm. 1971; Location: Maslin Bay, South Australia; Age: Early-Middle Eocene; Notes:
The transfer of this species, the type of Pleosporonites, makes Pleosporonites a later
taxonomic synonym of Dictyosporites.
4.5.11. Species: D. loculatus Félix 1894 (Fig. 13C); Index Fungorum Registration Identifier:
189946; Location: Perekeschkul, near Baku, Azerbaijan; Age: Eocene; Notes: Félix (1894)
stated that as compared with the conidia of recent forms, the fossils show the closest
resemblance to those of the genera Macrosporium Bon., Septosporium Zopf., Stemphylium
Wallr. and Stigmella Lév.
4.5.12. Species: D. morularis (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483302; Basionym: Pleosporonites morularis Sal.-Cheb. & Locq.
1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age:
Oligocene.
4.5.13. Species: D. moruloides (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483303; Basionym: Pleosporonites moruloides Sal.-Cheb. & Locq.
1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age: Early
Miocene.
4.5.14. Species: D. ovalis (Sheffy & Dilcher) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483304; Basionym: Staphlosporonites ovalis Sheffy & Dilcher 1971;
Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.; Age:
Middle Eocene (Claiborne Formation).
4.5.15. Species: D. ovoideus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
107916; Basionym: Pleosporonites ovoideus Sal.-Cheb. & Locq. 1980; Location: Coast of
Equatorial Africa, Gulf of Guinea, Cameroon, Africa; Age: Oligocene.
4.5.16. Species: D. pachycellularis Sal.-Cheb. & Locq. 1980; Index Fungorum Registration
Identifier: 107917; Current name: Staphlosporonites pachycellularis (Sal.-Cheb. & Locq.)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius 2000.
4.5.17. Species: D. paradkariae Kalgutkar & Janson. 2000 (Fig. 13D); Index Fungorum
Registration Identifier: 483305; Basionym: Dactylosporites dicotylophylli Paradkar 1976;
Location: Mohgaon Kalan, Chhindwara District, Madhya Pradesh, India; Age: Late
Cretaceous (Maastrichtian); Notes: Paradkar (1976) stated that such spores are found in the
genera Alternaria and Dactylosporium (Barnett 1965). More resemblance is seen, however,
with Dactylosporium in the shape, size and number of cells in the compound spores, than
with Alternaria. Kalgutkar & Jansonius (2000) transferred this species to Dictyosporites Felix
1894 with a new name (Dictyosporites paradkariae Kalgutkar & Janson. 2000, ‘paradkarii’)
because the name Dictyosporites dicotylophylli (Paradkar) Kalgutkar & Janson. 2000 was
preoccupied, The species epithet is in honour of Professor B.S. Trivedi, Department of
Botany, Lucknow University, Lucknow, India.
4.5.18. Species: D. symmetricus (V.S. Ediger) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483306; Basionym: Transeptaesporites symmetricus V.S. Ediger
1981; Location: Thrace Basin, Turkey; Age: Late Eocene-Oligocene, Miocene-Pliocene;
Notes: Ediger (1981) opined that this species has probable affinity with Alternaria.
4.5.19. Species: D. tirumalacharii (Ramanujam & Ramachar) Kalgutkar & Janson. 2000 (Fig.
13E); Index Fungorum Registration Identifier: 483307; Basionym: Ravenelites tirumalacharii
Ramanujam & Ramachar 1980; Location: Neyveli Lignite Mine, Cuddalore District, Tamil
Nadu, India; Age: Miocene (Neyveli lignite).

812
4.5.20. Species: D. tristratosus (Sheffy & Dilcher) Kalgutkar & Janson. 2000 (Fig. 13F); Index
Fungorum Registration Identifier: 483308; Basionym: Staphlosporonites tristratosus Sheffy
& Dilcher 1971; Location: Puryear clay pit, 800 m south of Puryear, Henry County,
Tennessee, U.S.A.; Age: Middle Eocene (Claiborne Formation); Notes: The specific name
refers to the arrangement of the cells into three layers.

4.6. Genus: DICTYOSPORIUMINITES Debi Mukh., International Journal of Geology, Earth and
Environmental Sciences 2(2): 5 (2012); Index Fungorum Registration Identifier: 588466; Type: D.
intermedius Debi Mukh. 2012.
Original Diagnosis: Conidiophore compact in sporodochium-like structure, clustered, dark in
colour; size mediumly large, 400–450 × 125–150 μm; conidia produced singly, acrogenous;
branches multiseptate, 5–8 septa observed; 4–5 branches present (Mukherjee 2012).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: One.
Notes: The genus is named after extant Dictyosporium (Hyphomycetes).
4.6.1. Species: D. intermedius Debi Mukh. 2012 (Fig. 13G); Index Fungorum Registration
Identifier: 588479; Location: Neyveli Lignite Mine-I, Cuddalore District, Tamil Nadu, India;
Age: Miocene (Neyveli Lignite); Notes: Dictyosporiminites intermedium can be compared
with the extant Dictyosporium in having sporodochium-like structure with multiseptate
conidiophores (united together).

4.7. Genus: DICTYOSTROMA R. Kar et al., Review of Palaeobotany & Palynology (Amsterdam)
158(3-4): 247 (2010); Index Fungorum Registration Identifier: 622180; Type: D. perfectum R. Kar
et al. 2010; Current name: KUTCHIATHYRITES R.K. Kar 1979 fide Saxena & Tripathi (2011).
Original 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 (Kar et al.
2010).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: One (the single species has been transferred to Kutchiathyrites
R.K. Kar 1979).
Notes: Kar et al. (2010) proposed Dictyostroma (“Dictyostromata”), which is identical to
Kutchiathyrites R.K. Kar 1979 and therefore the former is considered here as later synonym of the
latter.
4.7.1. Species: D. perfectum R. Kar et al. 2010; Index Fungorum Registration Identifier: 622214;
Current name: Kutchiathyrites perfectus (R. Kar et al.) R.K. Saxena & S.K.M. Tripathi 2011
fide Saxena & Tripathi (2011).

4.8. Genus: KUTCHIATHYRITES R.K. Kar, Palaeobotanist 26(1): 32 (1979); Index Fungorum
Registration Identifier: 21145; Type: K. eccentricus R.K. Kar 1979.
Synonym: Dictyostroma R. Kar et al. 2010 fide Saxena & Tripathi 2011, Index Fungorum
Registration Identifier: 622180.
Original Diagnosis: Microthyriaceous asco-stromata eccentric in development, no free
hyphae present, dimidiate, nonostiolate, radially arranged hyphae thick, dark, diverging from one
another, transverse hyphae comparatively thinner, ± translucent, interconnecting radial ones to form
squarish, pseudoparenchymatous cells without any pore; Description: Microthyriaceous
ascostromata of approximately semicircular shape in most specimens, in others they look like fish
scales, size range 64–110 × 41–73 μm. Upper surface of ascostromata darker than inner one; radial
hyphae also well pronounced in former. Radial hyphae look like dark strands; transverse hyphae ill-
developed, sometimes hardly discernable at places (Kar 1979).
Emended Diagnosis: Hilate conidia, fan-shaped, formed by numerous linear filaments
radiating out from the hilum; conidia may be flattened (i.e. two-) or three dimensional; filaments

813
may be joined to their neighbours, or partially free, and may branch towards the periphery; hilum
may or may not show the stipe from which it developed (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: Five.
Notes: Kalgutkar & Jansonius (2000) opined that Kutchiathyrites is not a microthyriaceous
fructification, as stated by Kar (1979), but a multicellular spore/conidium showing a clear
attachment area, scar or pore. Kutchiathyrites eccentricus, as described by Kar (1979),
demonstrates a close similarity to the conidia of the hyphomycetous fungus Mycoenterolobium
platysporum Goos 1970. This similarity was also pointed out by Jain & Kar (1979), who referred to
Kendrick & Carmichael (1973), where such eccentric structures are shown as conidia of the
hyphomycetous Mycoenterolobium platysporum.
4.8.1. Species: K. canadensis Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483412; Basionym: Dictyosporites eccentricus Kalgutkar 1993; Location: Peel River, Yukon
Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes: The species epithet refers to
the place of its occurrence in Canada.
4.8.2. Species: K. eccentricus R.K. Kar 1979 (Fig. 13H); Index Fungorum Registration Identifier:
112385; Location: Barkhana nala cutting, near Sarangwara village, Kutch District, Gujarat,
India; Age: Oligocene (Maniyara Fort Formation); Notes: The species epithet refers to fan
like shape of the conidia.
4.8.3. Species: K. mehrotrae R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 13I); Index Fungorum
Registration Identifier: 519945; Basionym: Kutchiathyrites sp. in Singh et al. 1986; Location:
Sonapur-Badarpur Road Section, Jaintia Hills, Meghalaya and Cachar District, Assam, India;
Age: Early Miocene (Bhuban Formation).
4.8.4. Species: K. palmatus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483414; Basionym: Microthyriacites palmatus P. Ke & Z.Y. Shi
1978; Location: Kenlixian and Bingxian, Shandong Province, Coastal region of Bohai,
China; Age: Eocene-Oligocene.
4.8.5. Species: K. perfectus (R. Kar et al.) R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 13J); Index
Fungorum Registration Identifier: 519897; Basionym: Dictyostromata perfecta R. Kar et al.
2010; Location: Tlangsam, Mizoram, India; Age: Miocene (Bhuban Formation).

4.9. Genus: LIRASPORIS R. Potonié & S.C.D. Sah, Palaeobotanist 7(2): 132 (1960); Index
Fungorum Registration Identifier: 21154; Type: L. intergranifer R. Potonié & S.C.D. Sah 1960.
Original Diagnosis: Size varies from 69 × 103 μm to 116 × 134 μm; outline oval, longitudinal
ends of oval broadly rounded or somewhat tapering, sometimes showing irregular protuberances
which form a jumbled mass; extrema lineamenta somewhat smooth except the longitudinal ends
which are always nearly notched; following the longer axis exist perhaps 20–30 parallel but narrow
ribs showing between them spaced grana (Potonié & Sah 1960).
Emended Diagnosis: Fungal bodies oval-elliptical with equal or unequal, broad, generally
notched ends. Mycelia, long, septate, ± parallel to one another, extending from one end to other;
wall generally laevigate, sometimes granulose (Jain & Kar 1979).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: Three (but we accept only two species as legitimate because L.
dongpuensis is not validly published).
Notes: Jain & Kar (1979) opined that the parallel ribs seem to be septate fungal mycelia
arranged in longitudinal direction and transferred from Polyplicates to Fungi. Kalgutkar &
Jansonius (2000) had reservations about the fungal nature these forms.
4.9.1. Species: L. dongpuensis Z.C. Song in Z.C. Song et al. 1999 (nom. inval.) fide Kalgutkar &
Jansonius (2000); Index Fungorum Registration Identifier: 483854; Notes: Lirasporis
dongpuensis was not validly published by Song (1999) because the author did not specify
where the holotype is deposited and also did not provide a Latin or English translation.

814
4.9.2. Species: L. elongatus R.K. Kar in R.K. Saxena 2012 (Fig. 13K); Index Fungorum
Registration Identifier: 519798; Location: Rokhia borehole; Tripura, north-east India; Age:
Early-Middle Miocene; Notes: Kar (1990) did not validly publish Lirasporis elongatus as he
did not cite information on where its type is stored. Saxena (2012) validated the name of this
species and ascribed it to Kar, because its description and illustrations published by Kar
(1990) are the validating ones. The holotype of this species name is that designated by Kar
(1990).
4.9.3. Species: L. intergranifer R. Potonié & S.C.D. Sah 1960 (Fig. 13L); Index Fungorum
Registration Identifier: 519797; Location: European Club, Kannur Beach, (Potonié & Sah
1960); Padappakkara, Edvai and Varkala, Kerala, India (Jain & Kar 1979); Age: Late
Miocene or Pliocene (Potonié & Sah 1960); Early Miocene-Early Pliocene (Jain & Kar
1979).

4.10. Genus: OCTOSPORITES Sal.-Cheb. & Locq., C. r. Congr. natn. Socs. sav. Paris, sect. sci.,
fasc. 1. (Paleobotanique) 105: 191 (1980); Index Fungorum Registration Identifier: 25608; Type:
O. stauroides Sal.-Cheb. & Locq. 1980.
Original Diagnosis: Monohilate, octocellate murospores, globular, the middle partitions
forming a cross; 15 µm (Salard-Cheboldaeff & Locquin 1980).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: One.
4.10.1. Species: O. stauroides Sal.-Cheb. & Locq. 1980 (Fig. 13M); Index Fungorum Registration
Identifier: 108279; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon, Africa;
Age: Early Miocene.

4.11. Genus: PALAMBAGES Wetzel, Micropaleontology 7(3): 338 (1961); Index Fungorum
Registration Identifier: 21217; Type: P. morulosa Wetzel 1961.
Original Diagnosis: Spheroidal bodies, composed of many (8–18?) oval membranous cells,
sometimes with a flat peripheral portion (with aperture?) (Wetzel 1961).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: Three.
Notes: Kalgutkar & Jansonius (2000) opined that Palambages is characterized by cells or
spores arranged in clusters of varying sizes, similar to groups of spores that occur in
Polyadosporites. In Palambages, the colonies are membranous and consist of thin-walled, hyaline
cells; in Polyadosporites, the spores in colonies are thicker-walled and pigmented. Colonies of
Palambages are similar in appearance to coenobial of colonial algae. The genus name is derived
from Latin palaeo = old; ambages = going round, winding, ambiguous.
4.11.1. Species: P. canadiana Sat. K. Srivast. 1968; Index Fungorum Registration Identifier:
335542; Location: Drumheller locality, sec. 30, twp. 29, rge. 20, W. 4th mer., Alberta,
Canada; Age: Maastrichtian;
4.11.2. Species: P. colonicus Trivedi & C.L. Verma 1970 ex Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 319134; Basionym: Palambages colonica Trivedi & C.L.
Verma 1970 (nom. inval.), holotype designated by Kalgutkar & Jansonius 2000; Location:
Near Kuala Lumpur, Malaysia (‘Malaya’); Age: Eocene; Notes: Trivedi & Verma (1970) did
not validly publish the name of this species because two different specimens were included
under the designation “Holotype”. Kalgutkar & Jansonius (2000) selected fig. 5 as holotype
and validated the species name.
4.11.3. Species: P. morulosa Wetzel 1961 (Fig. 13N); Index Fungorum Registration Identifier:
319135; Location: Baltic Cretaceous flintstones; Age: Cretaceous; Notes: The species epithet
is derived from Latin morula = diminutive of morum, a mulberry.

815
4.12. Genus: PAPULOSPORONITES Schmied. & A. J. Schwab, Deutsche Akademie der
Wissenschaften zu Berlin, 6: 686 (1964); Index Fungorum Registration Identifier: 21220; Type: P.
sphaeromorphus Schmied. & A. J. Schwab 1964.
Original Diagnosis: Fungal remains of globular to elongate shape, consisting of numerous
more or less polygonal cells that are firmly fused into mulberry-shaped aggregates. Cells without
any regular order, or concentrically arranged. No differentiation of an outer wall layer; however,
one to three of the innermost cells commonly much larger. Occasionally individual aggregates
fused together (Schmiedeknecht & Schwab 1964).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: Eight (but we accept only 7 species as legitimate because one
species, viz. P. mohgaoensis (Chitaley & Yawale) Kalgutkar & Janson. 2000, is here transferred to
Thecaphora Fingerh 1836).
Notes: Kalgutkar & Jansonius (2000) opined that Palambages appears to represent thinner-
walled vesicles, which are less tightly aggregated into spherical units.
4.12.1. Species: P. enormis (V.S. Ediger) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483501; Basionym: Polyadosporites enormis V.S. Ediger 1981; Location: Thrace
Basin, Turkey; Age: Late Eocene-Oligocene, Miocene-Pliocene.
4.12.2. Species: P. hammenii (Haseld.) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483502; Basionym: Polyadosporites hammenii Haseld. 1973; Location: Arén,
Ribagorzana Valley, Pyrenees, Spain; Age: Early Palaeocene-Late Eocene.
4.12.3. Species: P. mohgaoensis (Chitaley & Yawale) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483503; Synonym: Sorosporium mohgaoense Chitaley & Yawale
1978 fide Kalgutkar & Jansonius (2000), Index Fungorum Registration Identifier: 111035;
Current name: Thecaphora mohgaoensis (Chitaley & Yawale) R.K. Saxena, Wijayaw., D.Q.
Dai, K.D. Hyde & P.M. Kirk comb. nov. This new combination is described under the section
"New species and new combinations"
4.12.4. Species: P. multicellatus (R.K. Saxena & H.P. Singh) Kalgutkar & Janson. 2000 (Fig.
13O); Index Fungorum Registration Identifier: 483504; Basionym: Staphlosporonites
multicellatus R.K. Saxena & H.P. Singh 1983; Location: Hoshiarpur-Una Road section, near
Bankhandi, Hoshiarpur District, Punjab, India; Age: Miocene-Pliocene.
4.12.5. Species: P. orbis (V.S. Ediger) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483505; Basionym: Polyadosporites orbis V.S. Ediger 1981; Location: Thrace
Basin, Turkey; Age: Late Eocene-Oligocene, Miocene-Pliocene.
4.12.6. Species: P. siwalikus (R.K. Saxena & A.P. Bhattach.) Kalgutkar & Janson. 2000 (Fig. 13P);
Index Fungorum Registration Identifier: 483506; Basionym: Staphlosporonites siwalikus
R.K. Saxena & A.P. Bhattach. 1987, Location: Kala Amb-Nahan area, Sirmaur District,
Himachal Pradesh, India; Age: Miocene.
4.12.7. Species: P. sphaeromorphus Schmied. & A.J. Schwab 1964 (Fig. 13Q); Index Fungorum
Registration Identifier: 637505; Location: Zwischenflöz, Tagebau des Braunkohlenwerkes
Nachterstedt, Germany; Age: Middle Eocene.
4.12.8. Species: P. subcircularis (Anil Chandra et al.) Kalgutkar & Janson. 2000 (Fig. 13R); Index
Fungorum Registration Identifier: 483507; Basionym: Staphlosporonites subcircularis Anil
Chandra et al. 1984; Location: Sediment core no. 2 (Lat. 18°35.2′N: Long. 69°17.2′E),
Arabian Sea; Age: Late Quaternary.

4.13. Genus: PLEOSPORONITES R.T. Lange & P.H. Sm., Neues Jb. für Geologie und
Paläontologie 11: 672 (1971); Index Fungorum Registration Identifier: 21253; Type: P. hyalinus
R.T. Lange & P.H. Sm. 1971; Current name: DICTYOSPORITES Felix 1894 fide Kalgutkar &
Jansonius (2000).
Original Diagnosis: Hyaline muriform fungal spores, ellipsoid to ovoid in outline, and many-
celled, apparently by internal septation of irregular pattern (Lange & Smith 1971).
Classification: Fungi Imperfecti, Dictyosporae.

816
 Number of species known: Five (all the species have been transferred to Dictyosporites Felix
1894).
Notes: According to Kalgutkar & Jansonius (2000), Pleosporonites appears to be closely
related to Dictyosporites Felix 1894. Different species of Dictyosporites are comparable to modern
genera, such as the conidia of Alternaria Nees, Dictyosporium Corda, Septosporium Corda and
Stemphylium Wallr. and the ascospores of Pleospora Rabenh. ex Ces. & De Not.
4.13.1. Species: P. dictyosus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108376; Current name: Dictyosporites dictyosus (Sal.-Cheb. & Locq.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
4.13.2. Species: P. hyalinus R.T. Lange & P.H. Sm. 1971; Index Fungorum Registration Identifier:
320665; Current name: Dictyosporites hyalinus (R.T. Lange & P.H. Sm.) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
4.13.3. Species: P. morularis Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108377; Current name: Dictyosporites morularis (Sal.-Cheb. & Locq.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
4.13.4. Species: P. moruloides Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108378; Current name: Dictyosporites moruloides (Sal.-Cheb. & Locq.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).
4.13.5. Species: P. ovoideus Sal.-Cheb. & Locq. 1980; Index Fungorum Registration Identifier:
108379; Current name: Dictyosporites ovoideus (Sal.-Cheb. & Locq.) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).

4.14. Genus: POLYADOSPORITES Hammen 1954, Boletín Geológico (Bogota) 2(1): 82 (1954);
Index Fungorum Registration Identifier: 21256; Type: P. suescae Hammen 1954.
Original Diagnosis: Fungal spores composed of several grains or cells that are united along
several axes or in a more or less irregular manner (Van der Hammen 1954).
Emended Diagnosis: Spores (sub) spherical, loosely aggregated in clusters, with individual
cells not connected to others by shared walls; clusters (colonies?) more or less regularly spherical to
subspherical (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: Nine (but we accept only three species as legitimate because six
species have been transferred to other genera).
4.14.1. Species: P. conoideus (Sheffy & Dilcher) V.S. Ediger 1981; Index Fungorum Registration
Identifier: 108394; Current name: Staphlosporonites conoideus Sheffy & Dilcher 1971 fide
Kalgutkar & Jansonius (2000).
4.14.2. Species: P. enormis V.S. Ediger 1981; Index Fungorum Registration Identifier: 108395;
Current name: Papulosporonites enormis (V.S. Ediger) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
4.14.3. Species: P. firbasii Hammen 1954; Index Fungorum Registration Identifier: 337454;
Current name: Dictyosporites firbasii (Hammen) Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).
4.14.4. Species: P. garciabarrigae Hammen 1954; Index Fungorum Registration Identifier:
337455; Current name: Dictyosporites garciabarrigae (Hammen) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
4.14.5. Species: P. hammenii Haseld. 1973; Index Fungorum Registration Identifier: 483817;
Current name: Papulosporonites hammenii (Haseld.) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
4.14.6. Species: P. nadahensis M.R. Rao & R. Patnaik 2001 (Fig. 13S); Index Fungorum
Registration Identifier: 519784; Location: Nadah, Panchkula, Haryana, India; Age: Late
Pliocene (Pinjor Formation).

817
4.14.7. Species: P. orbis V.S. Ediger 1981; Index Fungorum Registration Identifier: 108396;
Current name: Papulosporonites orbis (V.S. Ediger) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
4.14.8. Species: P. siwalikus M.R. Rao & R. Patnaik 2001 (Fig. 13T); Index Fungorum
Registration Identifier: 519785; Location: Nadah, Panchkula, Haryana, India; Age: Late
Pliocene (Pinjor Formation).
4.14.9. Species: P. suescae Hammen 1954 (Fig. 13U); Index Fungorum Registration Identifier:
337456; Location: Magdalena Valley, Eastern Cordillera, Colombia, South America; Age:
Maastrichtian.

4.15. Genus: POLYCELLAESPORONITES Anil Chandra et al. 1984, Biovigyanam 10: 49 (1984);
Index Fungorum Registration Identifier: 25604; Type: P. bellus Anil Chandra et al. 1984.
Original Diagnosis: Capsular fungal spores; inaperturate; one end of the spore is rounded
while the other gives rise to a tube-like projection; multicellate; cells arranged in clusters, and not
in a row or along a single axis; spore wall laevigate (Chandra et al. 1984).
Emended Diagnoses: The diagnosis of Polycellaesporonites was emended by Kalgutkar &
Jansonius (2000) and Gupta (2002), as follows: Muriform spores with a hilum, and distally with an
elongated, knob-like or beaked, extension; overall structure as that in the modern Alternaria
(Kalgutkar & Jansonius 2000); Capsular spore, one end of the spore gives rise to tube like
projection, multicellate, inaperturate, cells arranged in clusters and not in a row or along a single
axis, spore wall laevigate to ornamented (Gupta 2002).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: Seven.
Notes: Kalgutkar & Jansonius (2000) opined that the type resembles to Staphlosporonites
Sheffy & Dilcher 1971, but differs in that the elongate projection is distal, rather than being a
(proximal) hold-fast or hilum.
4.15.1. Species: P. acuminatus (Rouse & Mustard) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483526; Basionym: Multicellaesporites acuminatus Rouse & Mustard
1997; Synonym: Piriurella acuminata (Rouse & Mustard) M.G. Parsons & G. Norris 1999
fide Kalgutkar & Jansonius (2000); Location: Strait of Georgia, eastern Vancouver Island, the
Fraser River lowlands of southwest British Columbia, Canada and the North-western
Washington State, U.S.A.; Age: Late Palaeocene.
4.15.2. Species: P. alternariatus (Kalgutkar & Sigler) Kalgutkar & Janson. 2000 (Fig. 13V); Index
Fungorum Registration Identifier: 483527; Basionym: Piriurella alternariata Kalgutkar &
Sigler 1995; Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories,
Canada; Age: Late Palaeocene or Early Eocene (Iceberg Bay Formation).
4.15.3. Species: P. bellus Anil Chandra et al. 1984 (Fig. 13W); Index Fungorum Registration
Identifier: 107183; Location: Sediment core no. 1 (Lat. 17°57.9′N: Long. 70°46.0′E), Arabian
Sea; Age: Late Quaternary; Notes: Chandra et al. (1984) suggested possible affinity of this
species to Alternaria sp.
4.15.4. Species: P. clavellatus (Z.C. Song & G.X. Li in Z.C. Song et al.) Kalgutkar & Janson.
2000; Index Fungorum Registration Identifier: 483528; Basionym: Pluricellaesporites
clavellatus Z.C. Song & G.X. Li in Z.C. Song et al. 1989; Location: Heze County and
Shenxian County of Shandong Province, China; Age: Middle-Late Oligocene (Shahejie and
Dongying formations).
4.15.5. Species: P. psilatus A. Gupta 2002 (Fig. 13X); Index Fungorum Registration Identifier:
540760; Location: Dadahu Road Section, Sirmaur District, Himachal Pradesh, India; Age:
Eocene (Subathu Formation).
4.15.6. Species: P. saxenae A. Gupta 2002 (Fig. 13Y); Index Fungorum Registration Identifier:
540761; Location: Jamtah Road Section, Sirmaur District, Himachal Pradesh, India; Age:
Eocene (Subathu Formation); Notes: The species epithet honours Dr. Ramesh K. Saxena,
Birbal Sahni Institute of Palaeosciences, Lucknow, India.

818
4.15.7. Species: P. sirmaurensis A. Gupta 2002 (Fig. 13Z); Index Fungorum Registration
Identifier: 540762; Location: Dadahu Road Section, Sirmaur District, Himachal Pradesh,
India; Age: Eocene (Subathu Formation).

4.16. Genus: RAVENELITES Ramanujam & Ramachar, Records of the Geological Survey of India
113(5): 83 (1980); Index Fungorum Registration Identifier: 28630; Type: R. tirumalacharii
Ramanujam & Ramachar 1980; Current name: DICTYOSPORITES Felix 1894 fide Kalgutkar &
Jansonius (2000).
Original Diagnosis: Teliospores one-celled but strongly adherent, forming discoid heads.
Number of spores in each telial head variable; outer walls of peripheral cells (spores) of each head
smooth or ornamented; wall pigmented; one germ pore in each cell (spore) of discoid head
(Ramanujam & Ramachar 1980).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: One (the single species has been transferred to Dictyosporites
Felix 1894).
Notes: Kalgutkar & Jansonius (2000) considered this genus as a later synonym of
Dictyosporites Felix 1894.
4.16.1. Species: R. tirumalacharii Ramanujam & Ramachar 1980; Index Fungorum Registration
Identifier: 483759; Current name: Dictyosporites tirumalacharii (Ramanujam & Ramachar)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).

4.17. Genus: STAPHLOSPORONITES Sheffy & Dilcher 1971, Palaeontographica Abt. B 133 (1-
3): 48 (1971); Index Fungorum Registration Identifier: 21316; Type: S. conoideus Sheffy &
Dilcher 1971.
Synonym: Transeptaesporites V.S. Ediger 1981 fide Kalgutkar & Jansonius (2000), Index
Fungorum Registration Identifier: 25599.
Original Diagnosis: Inaperturate, psilate to punctuate fungal or algal bodies of four or more
irregular cells. Cells in clusters, shape variable along more than one axis (Sheffy & Dilcher 1971).
Emended Diagnosis: Inaperturate multicellate fungal spores, with muriform architecture
(cells internally dividing without a regular pattern), lacking a plane or axis of symmetry. Cells
rounded or rounded polygonal, septa may be depressed where they intersect the amb. Overall shape
generally more or less elongate; sometimes oval to ellipsoidal, rarely subspherical. Always with a
distinct proximal hold-fast cell and/or a hilar scar (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: 23 (but we accept only 18 species as legitimate because five
species have been transferred to other genera).
Notes: Presence of a hilate scar differentiates this genus from Dictyosporites.
4.17.1. Species: S. allomorphus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111945; Location: Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee,
U.S.A.; Age: Middle Eocene (Claiborne Formation); Notes: The specific name refers to the
strange shape of the spore. According to Kalgutkar & Jansonius (2000), the narrow end
appears to be hilate, possibly with a hyphal fragment attached.
4.17.2. Species: S. billelsikii Kalgutkar & Janson. 2000; Index Fungorum Registration Identifier:
483554; Basionym: Dictyosporites elsikii Kalgutkar 1993; Location: Peel River, Yukon
Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes: The species epithet honours
Dr. William C. Elsik.
4.17.3. Species: S. chandrae A. Gupta 2002 (Fig. 13AA); Index Fungorum Registration Identifier:
540809; Location: Jamtah Road Section, Sirmaur District, Himachal Pradesh, India; Age:
Eocene (Subathu Formation); Notes: The species epithet is in honours of Dr. Anil Chandra,
Birbal Sahni Institute of Palaeosciences, Lucknow, India.

819
4.17.4. Species: S. conoideus Sheffy & Dilcher 1971 (Fig. 13AB); Index Fungorum Registration
Identifier: 111946; Location: Puryear clay pit, 800 m south of Puryear, Henry County,
Tennessee, U.S.A.; Age: Middle Eocene (Claiborne Formation).
4.17.5. Species: S. delumbus Norris 1986; Index Fungorum Registration Identifier: 126577;
Location: Imperial Nuktak C–22 Well. Mackenzie Delta Region, District of Mackenzie,
Northwest Territories, Canada; Age: Eocene to Oligocene.
4.17.6. Species: S. dichotomus A. Gupta 2002 (Fig. 13AC); Index Fungorum Registration
Identifier: 540810; Location: Jamtah Road Section, Sirmaur District, Himachal Pradesh,
India; Age: Eocene (Subathu Formation).
4.17.7. Species: S. discitypicus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115634; Location: Panshan, Liaoning Province, Coastal region of Bohai, China; Age:
Eocene-Oligocene.
4.17.8. Species: S. elongatus A. Gupta 2002 (Fig. 13AD); Index Fungorum Registration Identifier:
540811; Location: Jamtah Road Section, Sirmaur District, Himachal Pradesh, India; Age:
Eocene (Subathu Formation).
4.17.9. Species: S. elsikii Ramanujam & Srisailam 1980 (Fig. 13AE); Index Fungorum Registration
Identifier: 109551; Location: Kannur Beach area, Palayangadi and Cheruvattur (southern side
of Karingottu River), Kerala, India; Age: Miocene; Notes: Kumar (1990) transferred this
species to Multicellaesporites, as Multicellaesporites elsikii (Ramanujam & Srisailam) Kumar
1990. However, Kalgutkar & Jansonius (2000) did not accept this transfer.
4.17.10. Species: S. felixii (R.T. Lange & P.H. Sm.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 282643; Basionym: Dictyosporites felixii R.T. Lange & P.H. Sm.
1971; Location: Maslin Bay, South Australia; Age: Early-Middle Eocene.
4.17.11. Species: S. irregularis (V.S. Ediger) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483556; Basionym: Transeptaesporites irregularis V.S. Ediger 1981;
Location: Thrace Basin, Turkey; Age: Upper Eocene-Oligocene, Miocene-Pliocene; Notes:
Ediger (1981) suggested possible affinity of this species to Alternaria sp.
4.17.12. Species: S. laetevirens K.F. Wang & Y.L. Zhang 1986; Index Fungorum Registration
Identifier: 637506; Location: China; Notes: This species was cited in Song et al. (1999).
Since its bibliographic details were not included in the list of references of Song et al. (1999),
it is difficult to verify whether this species was validly published. Kalgutkar & Jansonius
(2000) opined that it is similar to Staphlosporonites irregularis (Ediger) Kalgutkar & Janson.
2000.
4.17.13. Species: S. lanceolatus K.F. Wang & Y.L. Zhang 1986; Index Fungorum Registration
Identifier: 637507; Location: China; Notes: This species was cited in Song et al. (1999).
Since its bibliographic details were not included in the list of references of Song et al. (1999),
it is difficult to verify whether this species was validly published. Kalgutkar & Jansonius
(2000) opined that it is similar to Staphlosporonites allomorphus Sheffy & Dilcher 1971.
4.17.14. Species: S. multicellatus R.K. Saxena & H.P. Singh 1983; Index Fungorum Registration
Identifier: 485255; Current name: Papulosporonites multicellatus (R.K. Saxena & H.P.
Singh) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
4.17.15. Species: S. neyveliensis Ambwani 1983 (Fig. 13AF); Index Fungorum Registration
Identifier: 107315; Location: Neyveli Lignite Mine, Cuddalore District, Tamil Nadu, India;
Age: Late Miocene or Pliocene; Notes: According to Ambwani (1983), this species appears
similar to the spore type illustrated by Graham (1962) as Alternaria but differs in having
larger number of cells and considerably bigger size.
4.17.16. Species: S. ovalis Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111947; Current name: Dictyosporites ovalis (Sheffy & Dilcher) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
4.17.17. Species: S. pachycellularis (Sal.-Cheb. & Locq.) Kalgutkar & Janson. 2000; Index
Fungorum Registration Identifier: 637508; Basionym: Dictyosporites pachycellularis Sal.-

820
 Cheb. & Locq. 1980; Location: Coast of Equatorial Africa, Gulf of Guinea, Cameroon,
Africa; Age: Oligocene.

Figure 13 – A–AH Dictyosporae.A Ctenosporites eskerensis Elsik & Janson. 1974, Bar = 5 μm. B
Dictyosporites dicotylophylli (Paradkar) Kalgutkar & Janson. 2000, Bar = 20 μm. C Dictyosporites
loculatus Félix 1894, Bar = 5 μm. D Dictyosporites paradkariae Kalgutkar & Janson. 2000, Bar =
20 μm. E Dictyosporites tirumalacharii (Ramanujam & Ramachar) Kalgutkar & Janson. 2000, Bar
= 20 μm. F Dictyosporites tristratosus (Sheffy & Dilcher) Kalgutkar & Janson. 2000, Bar = 5 μm.
G Dictyosporiuminites intermedius Debi Mukh. 2012, Bar = 20 μm. H Kutchiathyrites eccentricus
R.K. Kar 1979, Bar = 10 μm. I Kutchiathyrites mehrotrae R.K. Saxena & S.K.M. Tripathi 2011,

821
Bar = 40 μm. J Kutchiathyrites perfectus (R. Kar et al.) R.K. Saxena & S.K.M. Tripathi 2011, Bar =
10 μm. K Lirasporis elongatus R.K. Kar 2012, Bar = 10 μm. L Lirasporis intergranifer R. Potonié
& S.C.D. Sah 1960, Bar = 12 μm. M Octosporites stauroides Sal.-Cheb. & Locq. 1980, Bar = 12
μm. N Palambages morulosa Wetzel 1961, Bar = 12 μm. O Papulosporonites multicellatus (R.K.
Saxena & H.P. Singh) Kalgutkar & Janson. 2000, Bar = 20 μm. P Papulosporonites siwalikus (R.K.
Saxena & A.P. Bhattach.) Kalgutkar & Janson. 2000, Bar = 20 μm. Q Papulosporonites
sphaeromorphus Schmied. & A. J. Schwab 1964, Bar = 10 μm. R Papulosporonites subcircularis
(Anil Chandra et al.) Kalgutkar & Janson. 2000, Bar = 10 μm. S Polyadosporites nadahensis M.R.
Rao & R. Patnaik 2001, Bar = 40 μm. T Polyadosporites siwalikus M.R. Rao & R. Patnaik 2001,
Bar = 10 μm. U Polyadosporites suescae Hammen 1954, Bar = 10 μm. V Polycellaesporonites
alternariatus (Kalgutkar & Sigler) Kalgutkar & Janson. 2000, Bar = 15 μm. W
Polycellaesporonites bellus Anil Chandra et al. 1984, Bar = 10 μm. X Polycellaesporonites psilatus
A. Gupta 2002, Bar = 10 μm. Y Polycellaesporonites saxenae A. Gupta 2002, Bar = 5 μm. Z
Polycellaesporonites sirmaurensis A. Gupta 2002, Bar = 15 μm. AA Staphlosporonites chandrae
A. Gupta 2002, Bar = 20 μm. AB Staphlosporonites conoideus Sheffy & Dilcher 1971, Bar = 8 μm.
AC Staphlosporonites dichotomus A. Gupta 2002, Bar = 20 μm. AD Staphlosporonites elongatus
A. Gupta 2002, Bar = 8 μm. AE Staphlosporonites elsikii Ramanujam & Srisailam 1980, Bar = 10
μm. AF Staphlosporonites neyveliensis Ambwani 1983, Bar = 20 μm. AG Staphlosporonites raoi
A. Gupta 2002, Bar = 10 μm. AH Staphlosporonites settyi A. Gupta 2002, Bar = 10 μm.

4.17.18. Species: S. raoi A. Gupta 2002 (Fig. 13AG); Index Fungorum Registration Identifier:
540812; Location: Jamtah Road Section, Sirmaur District, Himachal Pradesh, India; Age:
?Eocene (Dagshai Formation).
4.17.19. Species: S. settyi A. Gupta 2002 (Fig. 13AH); Index Fungorum Registration Identifier:
540813; Location: Jamtah Road Section, Sirmaur District, Himachal Pradesh, India; Age:
Eocene (Subathu Formation).
4.17.20. Species: S. siwalikus R.K. Saxena & A.P. Bhattach. 1987; Index Fungorum Registration
Identifier: 483890; Current name: Papulosporonites siwalikus (R.K. Saxena & A.P.
Bhattach.) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
4.17.21. Species: S. subcircularis Anil Chandra et al. 1984; Index Fungorum Registration
Identifier: 107316; Current name: Papulosporonites subcircularis (Anil Chandra et al.)
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
4.17.22. Species: S. tetracellatus A. Gupta 2002 (Fig. 14A); Index Fungorum Registration
Identifier: 540814; Location: Jamtah Road Section, Sirmaur District, Himachal Pradesh,
India; Age: Eocene (Subathu Formation).
4.17.23. Species: S. tristratosus Sheffy & Dilcher 1971; Index Fungorum Registration Identifier:
111948; Current name: Dictyosporites tristratosus (Sheffy & Dilcher) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).

4.18. Genus: TRANSEPTAESPORITES V.S. Ediger 1981; Index Fungorum Registration Identifier:
25599; Type: T. irregularis V.S. Ediger 1981; Current name: STAPHLOSPORONITES Sheffy &
Dilcher 1971 fide Kalgutkar & Jansonius (2000).
Original Diagnosis: Fungal spores of five or more cells, four or more septa; shape is oval to
irregularly elongated; septa thicker than exine, imperforate, at least one of them transverse to the
others; individual cells variable in shape and size, usually rounded, lighter, a weak area is always
present at the middle of the cells, cells are light coloured to hyaline (Ediger 1981).
Classification: Fungi Imperfecti, Dictyosporae.
Number of species known: Five (all the species have been transferred to other genera).
Notes: According to Ediger (1981), these fossil fungal spores resemble the conidia of the
modern Alternaria, however, the new genus is proposed because of inadequate knowledge about
the relationship between the fossil and modern genera. Kalgutkar & Jansonius (2000) opined that
the illustrated specimens assigned to Transeptaesporites irregularis by Ediger (1981) appear so

822
variable that they may not all be conspecific. Ediger (1981) cited Staphlosporonites as a synonym
of the new Transeptaesporites, but transferred its type species S. conoideus to Polyadosporites.
Jansonius & Hills (1982) argued that Ediger (1981) did not provide bibliographic reference of
basionyms in sufficient detail to validly publish the new combinations. Three other species of
Staphlosporonites, named by Sheffy & Dilcher (1971), were also invalidly transferred to
Transeptaesporites by Ediger (1981).
4.18.1. Species: T. allomorphus (Sheffy & Dilcher) V.S. Ediger 1981; Index Fungorum
Registration Identifier: 107699; Current name: Staphlosporonites allomorphus Sheffy &
Dilcher 1971 fide Kalgutkar & Jansonius (2000).
4.18.2. Species: T. irregularis V.S. Ediger 1981; Index Fungorum Registration Identifier: 108586;
Current name: Staphlosporonites irregularis (V.S. Ediger) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
4.18.3. Species: T. ovalis (Sheffy & Dilcher) V.S. Ediger 1981; Index Fungorum Registration
Identifier: 108587; Current name: Dictyosporites ovalis (Sheffy & Dilcher) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
4.18.4. Species: T. symmetricus V.S. Ediger 1981; Index Fungorum Registration Identifier: 108588;
Current name: Dictyosporites symmetricus (V.S. Ediger) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
4.18.5. Species: T. tristratosus (Sheffy & Dilcher) V.S. Ediger 1981; Index Fungorum Registration
Identifier: 108589; Current name: Dictyosporites tristratosus (Sheffy & Dilcher) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).

5. Helicosporae

5.1. Genus: COLLIGERITES K.P. Jain & R.K. Kar, Palaeobotanist 26(2): 110 (1979); Index
Fungorum Registration Identifier: 21058; Type: C. kutchensis (R.K. Kar & R.K. Saxena) K.P. Jain
& R.K. Kar 1979.
Original Diagnosis: Spores multicellular, coiled, cells generally smaller, rounded in central
region and bigger, rectangular in outer region. Spore wall mostly laevigate, sometimes granulose.
Pore may be present or absent in each cell (Jain & Kar 1979).
Classification: Fungi Imperfecti, Helicosporae.
Number of species known: Three (but we accept only two species as legitimate because one
species, C. chowdhryi K.P. Jain & R.K. Kar 1979, has been transferred to Involutisporonites R.T.
Clarke 1965).
5.1.1. Species: C. chowdhryi K.P. Jain & R.K. Kar 1979; Index Fungorum Registration Identifier:
112034; Current name: Involutisporonites chowdhryi (K.P. Jain & R.K. Kar) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
5.1.2. Species: C. kutchensis (R.K. Kar & R.K. Saxena) K.P. Jain & R.K. Kar 1979 (Fig. 14B);
Index Fungorum Registration Identifier: 112205; Basionym: Involutisporonites kutchensis
R.K. Kar & R.K. Saxena 1976; Location: Bhuj-Lakhpat Road, Matanomadh Village, Kutch
District, Gujarat, India (Kar & Saxena 1976); Papanasam and Varkala, Kerala, India (Jain &
Kar 1979); Age: Palaeocene (Kar & Saxena 1976); Miocene (Jain & Kar 1979).
5.1.3. Species: C. trochus B. Samant in R.K. Saxena 2009 (Fig. 14C); Index Fungorum
Registration Identifier: 515017; Synonym: Colligerites trochus B. Samant 2000 (nom. inval.)
fide Saxena (2009).; Location: Near Bhavnagar, Cambay Basin, Gujarat, India; Age: Early
Eocene (Kharsalia Clay Formation).

5.2. Genus: ELSIKISPORONITES P. Kumar, Review of Palaeobotany & Palynology

(Amsterdam) 63(1-2): 18 (1990); Index Fungorum Registration Identifier: 25441; Type: E.
tubulatus Kumar 1990.
Original Diagnosis: Fungal spores monoporate, aseptate, tubular and coiled. Pore at outer
end, nozzle-like. Spore wall smooth and pale (Kumar 1990).

823
 Classification: Fungi Imperfecti, Helicosporae.
Number of species known: One.
Notes: The generic name honours Dr. William C. Elsik.
5.2.1. Species: E. tubulatus P. Kumar 1990 (Fig. 14D); Index Fungorum Registration Identifier:
126558; Location: Padappakkara, Kollam District, Kerala, India; Age: Early-Middle
Miocene.

5.3. Genus: HELICOMINITES Barlinge & Paradkar, Botanique, Nagpur 10(1-4): 167 (1982);
Index Fungorum Registration Identifier: 21126; Type: H. salvinites Barlinge & Paradkar 1982.
Original Diagnosis: Fungus saprophytic [saprobic]; mycelium septate, branched, hyphae faint
in colour; pycnidium and acervulus absent; conidia coiled in loose spirals and narrow at both ends
(Barlinge & Paradkar 1982).
Classification: Fungi Imperfecti, Helicosporae.
Number of species known: One.
5.3.1. Species: H. salvinites Barlinge & Paradkar 1982 (Fig. 14E); Index Fungorum Registration
Identifier: 108905; Location: Mohgaon Kalan, Chhindwara District, Madhya Pradesh, India;
Age: Cretaceous – Maastrichtian (Deccan Intertrappean Series).

5.4. Genus: HELICOONITES Kalgutkar & Sigler (as 'Helicoönites'), Mycol. Res. 99(5): 519
(1995); Index Fungorum Registration Identifier: 622357; Type: H. goosii Kalgutkar & Sigler 1995
Original Diagnosis: Conidia simple, tightly coiled or twisted in three planes to form an ovoid,
ellipsoidal (doliiform) to cylindrical or beehive to barrel-shaped spiral; spirals made up of variable
numbers of ascending coils or gyres, with each successive gyre usually of smaller diameter;
filaments multiseptate, fuscous; cells rectangular (Kalgutkar & Sigler 1995).
Classification: Fungi Imperfecti, Helicosporae.
Number of species known: One.
5.4.1. Species: H. goosii Kalgutkar & Sigler 1995 (Fig. 14F); Index Fungorum Registration
Identifier: 627571; Location: Strand Fiord, Axel Heiberg Island, Northwest Territories,
Canada; Age: Early Eocene (Iceberg Bay Formation); Notes: According to Kalgutkar &
Sigler (1995), Helicoonites goosii appears more closely allied to species of Helicoon (Goos et
al. 1986) such as H. richonis than to species of Helicodendron (Goos et al. 1985). The species
epithet honours Dr. R.D. Goos.

5.5. Genus: HELICOSPORIATES Kalgutkar & Sigler, Mycol. Res. 99(5): 520 (1995); Index
Fungorum Registration Identifier: 15012; Type: H. pirozynskii Kalgutkar & Sigler 1995.
Original Diagnosis: Conidia simple, pale brown to brown, helicoid; spirals of loose to tightly
coiled filaments; filaments slender, multicellular. Conidia usually helically coiled in one plane or
somewhat cochleate (Kalgutkar & Sigler 1995).
Classification: Fungi Imperfecti, Helicosporae.
Number of species known: One.
5.5.1. Species: H. pirozynskii Kalgutkar & Sigler 1995 (Fig. 14G); Index Fungorum Registration
Identifier: 413166; Location: Strand Fiord, Axel Heiberg Island, Northwest Territories,
Canada; Age: Early Eocene (Iceberg Bay Formation); Notes: The species epithet honours Dr.
Kris A. Pirozynski.

5.6. Genus: INVOLUTISPORONITES R.T. Clarke, Mountain Geologist 2(2): 90 (1965); Index
Fungorum Registration Identifier: 21142; Type: I. foraminus R.T. Clarke 1965.
Original Diagnosis: Fungal spores planispiral, individual cells lobate, septa simple with an
opening through each septum (Clarke 1965).
Emended Diagnosis: Monoporate, psilate, multiseptate, coiled fungal spores (Elsik 1968).
Classification: Fungi Imperfecti, Helicosporae.

824
 Number of species known: Eight (but we accept only four species as legitimate because four
species have been transferred to other genera).
Notes: Clarke (1965) named the genus in reference to the involute arrangement of the cells.
5.6.1. Species: I. chowdhryi (K.P. Jain & R.K. Kar) Kalgutkar & Janson. 2000 (Fig. 14H); Index
Fungorum Registration Identifier: 483410; Basionym: Colligerites chowdhryi K.P. Jain &
R.K. Kar 1979; Location: Papanasam and Varkala, Kerala, India; Age: Miocene; Notes: Jain
& Kar (1979) opined that Colligerites chowdhryi resembles closely the spores of
Helicominopsis fici (illustrated by Subramanian 1971). The spores of Helicomina caperoniae
are also helicoid but the coilings are not as prominent as in the present species. The species
epithet honours Dr. Harsh Chowdhry.
5.6.2. Species: I. crassus Z.C. Song & Liu Cao 1994; Index Fungorum Registration Identifier:
483775; Location: Antarctica; Age: Late Cretaceous.
5.6.3. Species: I. foraminus R.T. Clarke 1965 (Fig. 14I); Index Fungorum Registration Identifier:
332602; Location: Canon City coal field, Fremont County, Colorado, U.S.A.; Age: Late
Cretaceous; Notes: The species epithet refers to the resemblance of the species to planispiral
foraminifera.
5.6.4. Species: I. kutchensis R.K. Kar & R.K. Saxena 1976; Index Fungorum Registration
Identifier: 112379; Current name: Colligerites kutchensis (R.K. Kar & R.K. Saxena) Jain &
Kar 1979 fide Jain & Kar (1979).
5.6.5. Species: I. minutus Rouse & Mustard 1997; Index Fungorum Registration Identifier: 463995;
Current name: Palaeoslimacomyces minutus (Rouse & Mustard) Kalgutkar & Janson. 2000
fide Kalgutkar & Jansonius (2000).
5.6.6. Species: I. putus P. Ke & Z.Y. 1978; Index Fungorum Registration Identifier: 115668;
Current name: Paragranatisporites putus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
5.6.7. Species: I. trapezoides Kalgutkar 1993; Index Fungorum Registration Identifier: 483881;
Location: Peel River, Yukon Territory, Canada; Age: Late Palaeocene-Early Eocene; Notes:
The species epithet is derived from the Latin, trapezoideus, irregularly four-sided.
5.6.8. Species: I. wilcoxii Elsik 1968; Index Fungorum Registration Identifier: 315918; Current
name: Paleoslimacomyces wilcoxii (Elsik) Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).

5.7. Genus: PALAEOCIRRENALIA Ramanujam & Srisailam, Botanique, Nagpur 9(1-4): 124
(1980); Index Fungorum Registration Identifier: 21205; Type: P. elegans Ramanujam & Srisailam
1980.
Original Diagnosis: Spores light brown to reddish brown, inaperturate, helicoid, 1 to 1 1/4
times loosely coiled, multicellular, 2- to 6-septate, septa transverse, prominent, as thick and dark
bands, cells of unequal size, terminal cell dome-shaped and broader, basal cell usually cuneate,
pale-coloured, surface psilate (Ramanujam & Srisailam 1980).
Classification: Fungi Imperfecti, Helicosporae.
Number of species known: Two
Notes: Ramanujam & Srisailam (1980) stated that “In their loosely coiled 2- to 6-septate
nature with the terminal cell dome-shaped, these spores are remarkably similar to the conidia of the
modern dematiaceous hyphomycete Cirrenalia (Ellis 1976). The species of Cirrenalia are
characteristic of brackish to marine habitats and hence are environmentally significant. They are
generally found on driftwood. No information is available with regard to the nature of the
conidiophores of the fossil specimens, i.e. whether they are simple or branched.” Kalgutkar &
Jansonius (2000) opined that the characteristics of a more than hemispherical to globular dark distal
cell, the curved longitudinal axis, and an indistinct proximal hilum, define this genus.
5.7.1. Species: P. elegans Ramanujam & Srisailam 1980 (Fig. 14J); Index Fungorum Registration
Identifier: 109520; Location: Kannur Beach area, Palayangadi and Cheruvattur (southern side
of Karingottu River), Kerala, India; Age: Miocene; Notes: According to Ramanujam &

825
 Srisailam (1980), these spores exhibit remarkable similarity with the conidia of Cirrenalia
macrocephala (Kohlm.) Meyers & R.T. Moore 1960 (Ellis 1976).
5.7.2. Species: P. oligoseptata Ramanujam & Srisailam 1980 (Fig. 14K); Index Fungorum
Registration Identifier: 109521; Location: Kannur Beach area, Palayangadi and Cheruvattur
(southern side of Karingottu River), Kerala, India; Age: Miocene.

5.8. Genus: PALEOSLIMACOMYCES Kalgutkar & Sigler, Mycol. Res. 99(5): 521 (1995); Index
Fungorum Registration Identifier: 27617; Type: P. canadensis Kalgutkar & Sigler 1995.
Original Diagnosis: Conidia simple, solitary, helicoid, curved to hemi-circinate, brown to
fuscous, smooth. Conidia 2–3-septate; septa dark, often thick; conidial filaments short, made up of
3–4 broadly curved cells; cells, except the apical cell, darkly-pigmented; apical cell hyaline to pale
brown (Kalgutkar & Sigler 1995).
Classification: Fungi Imperfecti, Helicosporae.
Number of species known: Three.
Notes: According to Kalgutkar & Sigler (1995), conidia of Paleoslimacomyces show some
similarity to the conidia of extant Slimacomyces monospora (W.B. Kendr.) Minter, which was
originally described by Kendrick (1958) in Helicoma Corda.
5.8.1. Species: P. canadensis Kalgutkar & Sigler 1995 (Fig. 14L); Index Fungorum Registration
Identifier: 413657; Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories,
Canada; Age: Late Palaeocene or Early Eocene (Iceberg Bay Formation); Notes: The species
epithet is derived from the place of its occurrence in Canada.
5.8.2. Species: P. minutus (Rouse & Mustard) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483499; Basionym: Involutisporonites minutus Rouse & Mustard
1997; Location: Strait of Georgia, eastern Vancouver Island, the Fraser River lowlands of
southwest British Columbia, Canada, and the North-western Washington State, U.S.A.; Age:
Late Palaeocene.
5.8.3. Species: P. wilcoxii (Elsik) Kalgutkar & Janson. 2000 (Fig. 14M); Index Fungorum
Registration Identifier: 483500; Basionym: Involutisporonites wilcoxii Elsik 1968; Location:
Strip mine approximately 11 km southwest of Rockdale, Milam County, Texas, U.S.A.; Age:
Palaeocene.

5.9. Genus: RETIHELICOSPORONITES Ramanujam & K.P. Rao, Proc. IV International

Palynology Conference, Lucknow 1976–1977 1: 3 (1978); Index Fungorum Registration Identifier:
21281; Type: R. elsikii Ramanujam & K.P. Rao 1978.
Original Diagnosis: Spores simple, uniseriate, multicellular, inaperturate, basal cell cuneate,
other cells rectangular; apical part of spore helical. Spore wall reticulate (Ramanujam & Rao 1978).
Classification: Fungi Imperfecti, Helicosporae.
Number of species known: One.
Notes: Ramanujam & Rao (1978) opined that Involutisporonites Elsik 1968, although
multiseptate and helical, is distinguishable by its monoporate and psilate nature. Helical spores
(conidia) are found in various genera of Hyphomycetes, viz. Helicoma, Helicomina, Helicoon,
Helicodendron, Hiospira and Xenosporella (Barnett 1956, Ellis 1971, Ainsworth et al. 1973). In
possessing a reticulate cell wall, the fossil spores show remarkable resemblance to the conidia of
the Hiospira state of Brooksia tropicalis. Hiospira is commonly encountered in moist tropical
regions. Elsik (1992) considered spores of Retihelicosporonites to be monoporate and differentiated
them from other multicellate monoporate form genera on their combination of reticulate sculpture
and their tendency to be loosely coiled.
5.9.1. Species: R. elsikii Ramanujam & K.P. Rao 1979 (Fig. 14N); Index Fungorum Registration
Identifier: 115089; Location: Alleppey, Alappuzha District, Kerala, India, locality: Age:
Miocene (Quilon and Warkalli beds); Notes: The species epithet honours of Dr. William C.
Elsik.

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6. Staurosporae

6.1. Genus: EOGLOBELLA W.H. Bradley, U.S. Geological Survey Professional Paper 168: 1–58
(1931); Index Fungorum Registration Identifier: 637509; Type: E. longipes W.H. Bradley 1931.
Original Diagnosis: Thallus of definite shape, radially symmetrical, consisting of a single
spherical cell to which are attached four equally spaced, greatly elongated, cylindrical appendages.
These are apparently single cells that are enlarged and flattened where they join the globular cell
and tapered to a rather blunt conical point at the distal end. They arise from the globular cell a little
below the equator and diverge slightly downward. The globular cell is about 38 μm in diameter; the
appendages are about 6 μm in diameter and range in length from 115 to about 145 μm. The
characters of the genus will also serve to define the species (Bradley 1931).
Classification: Fungi Imperfecti, Staurosporae.
Number of species known: One.
Notes: According to Kalgutkar & Jansonius (2000), Eoglobella has four appendices that grow
from one corner or side whereas Tribolites W.H. Bradley 1964 has four (or three) well developed
filaments, each coming from opposite corners (and one of which, according to Bradley (1931),
tends to be truncated, and possibly served as an attachment). Frasnacritetrus is similar to
Eoglobella, but is larger, and lacks a distinct hilate scar.
6.1.1. Species: E. longipes W.H. Bradley 1931 (Fig. 14O); Index Fungorum Registration Identifier:
637510; Location: Garfield County, Colorado, U.S.A.; Age: Middle Eocene.

6.2. Genus: FRASNACRITETRUS Taug., Cahiers de Micropaléontologie 10: 3 (1968); Index

Fungorum Registration Identifier: 519771; Type: F. josettae Taug. 1968.
Original Diagnosis: Organic-walled microorganisms, generally of subcylindrical shape
tending to a rounded or slightly bell-shaped parallelepiped, in transversal section nearly circular at
one pole, becoming rectangular with rounded corners at the opposite pole which carries four hollow
horns (or “processes”) that extend the ribs of the body (Taugourdeau 1968).
Emended Diagnosis: Microfossils having two to four processes. Body subrectangular,
unicellular or divided into chambers by septa, smooth or variously sculptured. Processes mostly
smooth but may also be sculptured, unicellular or septate. Main body of the microfossils generally
rectangular-subrectangular but variously shaped; either unicellular or divided into longitudinal
chambers by vertical septa or multichambered, being divided by both vertical and transverse septa;
septa may be complete or incomplete, sometimes septa faintly developed; body either smooth or
ornamented with grana, verrucae or coni, etc., sculpturing elements may be closely or sparsely or
evenly distributed. Two to four processes arising from one end of the body (although in
Frasnacritetrus sp. 4, three processes are attached at the end of the body while the fourth one
comes out from the middle of the body); generally broader at the base and tapering towards the
apices; cylindrical or ribbon-like; either aseptate-unicellular or septate, septa one to many in each
process; apex of processes pointed or blunt. Frasnacritetrus is not comparable to any of the known
fossil palynogenera (Saxena & Sarkar 1986).
Classification: Fungi Imperfecti, Staurosporae.
Number of species known: Seven.
Notes: Taugourdeau (1968) stated that this single specimen does not resemble any microfossil
already described. He also did not rule out possibility of contamination. He speculated possibility
of this specimen being either a broken Diacrodian or half an organism of some Conjugales
(Desmidiales) or linear colony such as certain Desmochitina or hydrozoans. But he rejected all the
above possibilities and could not reach to any conclusion. Saxena & Sarkar (1986) emended the
generic diagnosis, to allow inclusion of a number of fungal conidia that show a general similarity to
the morphology of Tetraploa. This fungus generally grows on Poaceae, and the fossil species occur
in association with grass pollen. Saxena & Sarkar (1986) adduced this to support their suggestion
that Frasnacritetrus should not be considered an acritarch but a fossil genus with affinity to
Tetraploa. The generic diagnosis of Frasnacritetrus as originally proposed by Taugourdeau, only

827
accommodates microfossils with four processes, whereas Saxena & Sarkar (1986) also recovered
specimens with three processes. Specimens with only two processes have also been recorded by
Sharma (1976). Except for the difference in number of processes, there is no major morphological
difference between the specimens which could justify the erection of a new genus. Saccardo (1880,
1886) classified similar forms under the Staurosporae, which include spores having a forked or star-
shaped appearance (Subramanian 1971). Kendrick & Carmichael (1973) published a list of
staurosporous genera and their illustrations. A comparison of the present microfossils with these
genera reveals a close resemblance with Tetraploa, a genus belonging to the dematiaceous
hyphomycetes, and in all probability they belong to it; hence their placement under Acritarcha
incertae sedis by Taugourdeau (1968) does not seem justified.
6.2.1. Species: F. conatus R.K. Saxena & S. Sarkar 1986 (Fig. 14P); Index Fungorum Registration
Identifier: 519773; Location: Lower Siwalik, Nalagarh-Ramshahr Road section, Solan
District, Himachal Pradesh, India; Age: Middle-Late Miocene.
6.2.2. Species: F. indicus R.K. Saxena & S. Khare 1991 (Fig. 14Q); Index Fungorum Registration
Identifier: 483898; Location: Jayamkondacholapuram well–12 (JC–12), 45 km south of
Neyveli, Tiruchirappalli District, Tamil Nadu, India; Age: Tertiary.
6.2.3. Species: F. jamtahensis A. Gupta 2002 (Fig. 14R); Index Fungorum Registration Identifier:
540509; Location: Jamtah Road Section, Sirmaur District, Himachal Pradesh, India; Age:
Late Palaeocene-Early Oligocene (Subathu and Dagshai formations).
6.2.4. Species: F. josettae Taug. 1968 (Fig. 14S); Index Fungorum Registration Identifier: 519772;
Location: France; Age: Late Devonian.
6.2.5. Species: F. masolensis R.K. Saxena & S.K.M. Tripathi 2011 (Fig. 14T); Index Fungorum
Registration Identifier: 519941; Location: Masol-Kiratpur Section, Ambala District, Haryana,
India; Age: Pliocene (Tatrot and Pinjor formations).
6.2.6. Species: F. siwalikus R.K. Saxena et al. 1987 (Fig. 14U); Index Fungorum Registration
Identifier: 519774; Location: Upper Siwalik exposed between Masol and Kiratpur in Ambala
District, Haryana, India; Age: Miocene-Pliocene (Tatrot and Pinjor formations).
6.2.7. Species: F. taugourdeaui R.K. Saxena & S. Sarkar 1986 (Fig. 14V); Index Fungorum
Registration Identifier: 519775; Location: Near Banethi, Sirmaur District, Himachal Pradesh,
India; Age: Early Miocene (Kasauli Formation).

6.3. Genus: MOSSOPISPORITES Kalgutkar & Janson., AASP Contributions Series (Dallas) 39:
183 (2000); Index Fungorum Registration Identifier: 28621; Type: M. multicellulus (P. Ke & Z.Y.
Shi) Kalgutkar & Janson. 2000.
Original Diagnosis: Multicellate medium-sized triporate fungal spores with a (concave)
triangular outline. The pores occur at the distal ends of three radial extensions of the outline, where
a series of squat transverse cells are stacked to form short, broad arms. The interior of the spore is
filled with similar cells, that are interlocking to form a kind of mosaic. One of the arms may be
more strongly developed than the other two (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Staurosporae.
Number of species known: One.
Notes: The overall shape of Trihyphites Kalgutkar & Janson. 2000 is triradiate, rather than
triangular; this genus also has a single (large) central cell. Tribolites is inaperturate, and has a very
large central cell. The genus name honours Dr. G. D. Mossop.
6.3.1. Species: M. multicellulus (P. Ke & Z.Y. Shi) Kalgutkar & Janson. 2000 (Fig. 14W); Index
Fungorum Registration Identifier: 483444; Basionym: Triporicellaesporites multicellulus P.
Ke & Z.Y. Shi 1978; Location: Gangzhou, Hebei Province, Coastal region of Bohai, China;
Age: Eocene-Oligocene.

6.4. Genus: PESAVIS Elsik & Janson., Can. J. Bot. 52(5): 955 (1974); Index Fungorum
Registration Identifier: 21231; Type: P. tagluensis Elsik & Janson. 1974.

828
 Original Diagnosis: Multicellular fungal fruiting body consisting of a stalked central cell and
two lateral arms consisting of some 5–10 cells each. Lateral arms may be straight or curved, widely
spread or closely appressed. Two planes of symmetry present. Cells of the primary structure may or
may not have secondary septate hyphae-like filaments (Elsik & Jansonius 1974).
Classification: Fungi Imperfecti, Staurosporae.
Number of species known: Four (but we accept only two species as legitimate because two
species have been transferred to other genera).
6.4.1. Species: P. elongatus (P. Ke & Z.Y. Shi) Z.C. Song in Z.C. Song et al. 1999; Index
Fungorum Registration Identifier: 483850; Current name: Triporicellaesporites elongatus P.
Ke & Z.Y. Shi 1978 fide Kalgutkar & Jansonius (2000).
6.4.2. Species: P. parvus Kalgutkar & Sweet 1988; Index Fungorum Registration Identifier:
637511; Location: Big Fish River area on the Yukon Coastal Plain, the Bonnet Plume Basin
in Central Yukon, the central Alberta Foothills and southwestern Saskatchewan, Canada;
Age: Maastrichtian.
6.4.3. Species: P. simplex Elsik & Janson. 1974; Index Fungorum Registration Identifier: 319408;
Current name: Triporicellaesporites simplex (Elsik & Janson.) Kalgutkar & Janson. 2000 fide
Kalgutkar & Jansonius (2000).
6.4.4. Species: P. tagluensis Elsik & Janson. 1974 (Fig. 14X); Index Fungorum Registration
Identifier: 319409; Location: Washington state, British Columbia, Alaska, U.S.A., and the
Mackenzie delta, Northwest Territorries, Canada; Age: Palaeocene-Eocene.

6.5. Genus: SPEGAZZINITES Félix, Zeitschr. Deutsche Geol. Gesell. 46: 279 (1894); Index
Fungorum Registration Identifier: 21303; Type: S. cruciformis Felix 1894 (lectotype was
designated by Jansonius & Hills 1976).
Original Diagnosis (Combined description): The remains are the conidia of a hyphomycete.
Their shape and size varies more than is usually the case with such structures. However, the
morphology is the same in all of them as far as the somewhat thick polished sections disclose: they
consist of 4 partial cells. In the smaller conidia, these partial cells are of a slightly elongated shape,
and their narrow sides are oriented towards a point in such a way that together they form a regular,
equal-armed cross (figs. 8a-c). In the larger specimens, the individual cells are more roundish, the
arms of the cross therefore shortened, so that the entire structure approaches the shape of a tightly
tied, cube-shaped parcel, a shape found, for instance, in the body of the genus Sarcina. The size of
the smaller conidia is 0.012–0.015 mm [12–15 µm], that of the larger ones 0.021–0.024 mm [21–24
µm]; in between occur numerous transitional forms. Some of the conidia are spinose, and not only
the larger ones as was claimed by Hoffmann, but rather small conidia as well. The spines are of
various length, in the larger conidia they are generally shorter than in the smaller specimens. The
dimensions specified above refer to asetose specimens. In addition to conidia, the respective
polished sections also contain numerous mycelial remains. A connection between them and the
conidia could not be definitely established, but there is one case where a conidium appears to sit at
the end of a hyphal branch. The mycelium is sparsely ramified, septa were not observed. The
thicker filaments are 0.003–0.006 mm (3–6 µm) in diameter (Felix 1894, translated English version
in Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Staurosporae.
Number of species known: Three.
6.5.1. Species: S. cruciformis Félix 1894 (Fig. 14Y); Index Fungorum Registration Identifier:
197021; Location: Germany; Age: Tertiary; Notes: Félix (1894) stated that the fossil conidia
resemble very closely to those of a hyphomycete described by Saccardo as Spegazzinia
ornata which belongs to the family Tubercularieae. Spegazzinia also displays partly parcel-
shaped and partly cruciform conidia which consist of 4 cells and are partly covered with
spines. Jansonius & Hills (1976) designated the lectotype.

829
6.5.2. Species: S. indicus Ramanujam & Srisailam 1980 (Fig. 14Z); Index Fungorum Registration
Identifier: 109246; Location: Kannur Beach area, Palayangadi and Cheruvattur (southern side
of Karingottu River), Kerala, India; Age: Miocene.
6.5.3. Species: S. tetradus (Rouse) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483549; Basionym: Inapertisporites tetradus Rouse 1962; Location: Terminal
Dock, the city of Vancouver, British Columbia, Canada; Age: Late Cretaceous-Middle
Eocene (Burrard Formation).

6.6. Genus: TRIBOLITES W.H. Bradley, Amer. J. Sci. Arts, Ser. 2 262: 413 (1964); Index
Fungorum Registration Identifier: 21338; Type: T. tetrastonyx W.H. Bradley 1964.
Synonym: Trihyphaecites Peppers 1970 fide Kalgutkar & Jansonius (2000), Index Fungorum
Registration Identifier: 21343.
Original Diagnosis: Conidia consisting of four tapered arms that end in an abruptly tapered
sharp point and that radiate from a large central, polyhedral cell; apices of the arms corresponding
to the apices of a more or less regular tetrahedron; arms subdivided by thick septa into three to five
nearly equidimensional cells; one arm tip generally flattened, which presumably represents the
point of attachment to the conidiophore; overall dimensions 60–90 µm (Bradley 1964).
Emended Diagnosis: Inaperturate, medium to large-sized conidia consisting of a large,
inflated polyhedral (generally triangular or tetrahedral) central cell, the corners of which are
extended into tapered arms consisting of some two to six nearly equidimensional cells; the radial
arms closed terminally, rounded to pointed, but one arm generally with a flat (hilar?) end
(Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Staurosporae.
Number of species known: Two.
Notes: According to Bradley (1964), the conidia resemble in structure and general form the
conidia of extant genera Tetrachaetum and Lemonniera.
6.6.1. Species: T. tetrastonyx W.H. Bradley 1964 (Fig. 14AA); Index Fungorum Registration
Identifier: 340260; Location: Wyoming, Colorado, U.S.A.; Age: Eocene; Notes: According to
Bradley (1964), the conidia resemble in structure and general form the conidia of extant
Tetrachaetum elegans and Lemonniera aquatica, but the arms are much shorter, stockier, and
the cell walls much thicker.
6.6.2. Species: T. triangulatus (Peppers) Kalgutkar & Janson. 2000; Index Fungorum Registration
Identifier: 483562; Basionym: Trihyphaecites triangulatus Peppers 1970; Location: North
eastern part of the Illinois Basin, Illinois. U.S.A.; Age: Carboniferous-Pennsylvanian
(Carbondale and Spoon formations).

6.7. Genus: TRIHYPHAECITES Peppers, Bull. Ill. St. geol. Surv. 93: 135 (1970); Index Fungorum
Registration Identifier: 21343; Type: T. triangulatus Peppers 1970; Current name: TRIBOLITES
W.H. Bradley 1964 fide Kalgutkar & Jansonius (2000).
Original Diagnosis: The fossils are composed of radially symmetrical, triangular to roundly
triangular bodies that give rise to septate hyphae from each of the corners. Septa are also present at
the corners where the hyphae are joined to the body. The hyphae may consist of one segment with a
well rounded terminal end or may be of several segments in which the terminal end is either well
rounded or broken. The fossils generally possess minor folds and are slightly torn. They are
laevigate and about 1 μm thick. The triangular central body is generally thicker than the hyphae.
Known size range from one corner to the opposite side of the triangular body is 32.2 to 48.8 μm
(Peppers 1970).
Classification: Fungi Imperfecti, Staurosporae.
Number of species known: Two (both the species have been transferred to other genera).
Notes: According to Peppers (1970), Trihyphaecites is characterized by its triangular body
with septate hyphae at each of the corners.

830
6.7.1. Species: T. fractus Z.C. Song & Liu Cao in Z.C. Song et al. 1989; Index Fungorum
Registration Identifier: 485274; Current name: Trihyphites fractus (Z.C. Song & Liu Cao in
Z.C. Song et al.) Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).
6.7.2. Species: T. triangulatus Peppers 1970; Index Fungorum Registration Identifier: 325019;
Current name: Tribolites triangulatus (Peppers) Kalgutkar & Janson. 2000 fide Kalgutkar &
Jansonius (2000).

6.8. Genus: TRIHYPHITES Kalgutkar & Janson., AASP Contributions Series (Dallas) 39: 305
(2000); Index Fungorum Registration Identifier: 28625; Type: T. fractus (Z.C. Song & Liu Cao)
Kalgutkar & Janson. 2000
Synonym: Trihyphaecites fractus Z.C. Song & Liu Cao in Z.C. Song et al. 1989 fide
Kalgutkar & Jansonius (2000), Index Fungorum Registration Identifier: 485274.
Original Diagnosis: Fungal spores triradiate; from a small triangular central cell, three arms
radiate out, each consisting of up to a dozen cells, and each terminating with a wide pore-like
structure (Kalgutkar & Jansonius 2000).
Classification: Fungi Imperfecti, Staurosporae.
Number of species known: One.
Notes: The generic name is a contraction of Trihyphaecites, of which the type species, T.
triangulatus, was transferred to Tribolites.
6.8.1. Species: T. fractus (Z.C. Song & Liu Cao) Kalgutkar & Janson. 2000 (Fig. 14AB); Index
Fungorum Registration Identifier: 483569; Basionym: Trihyphaecites fractus Z.C. Song &
Liu Cao in Z.C. Song et al. 1989; Location: Shenxian county of Shandong Province, China;
Age: Late Eocene-Middle Oligocene (Shahejie Formation).

6.9. Genus: TRIPORICELLAESPORITES P. Ke & Z.Y. Shi, Early Tertiary spores and pollen
grains from the coastal region of Bohai: 51 (1978); Index Fungorum Registration Identifier: 21344;
Type: T. triangulus P. Ke & Z.Y. Shi 1978.
Original Diagnosis: Spores triangular-lenticular in shape, outline triangular in polar view.
Triporate, pores situated at corners of triangle, prominent, might be vestibulate. Multicellular, cells
in triangulate [triaxial?] arrangement. Spore wall of medium thickness, surface psilate or provided
with granulate to indistinct finely reticulate sculpturing (Ke & Shi 1978).
Emended Diagnosis: Pluricellate fungal spores with triangular to inverted V-shaped outline,
with a central (stalked) hilum; the two lateral wings or appendages may be closed terminally, but
commonly are preserved with the distalmost cells lacking; spore wall smooth (Kalgutkar &
Jansonius 2000).
Classification: Fungi Imperfecti, Staurosporae.
Number of species known: Four (but we accept only three species as legitimate because one
species has been transferred to Mossopisporites Kalgutkar & Janson. 2000).
Notes: This genus can be distinguished from all other genera of fossil fungal spores by the
fact that its members are both multicellular and triporate and exhibit a triangular to chevron-shaped
outline.
6.9.1. Species: T. elongatus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115699; Synonym: Pesavis elongatus (P. Ke & Z.Y. Shi) Z.C. Song in Z,C, Song et al. 1999
fide Kalgutkar & Jansonius (2000); Location: Cangxian, Hebei Province, Coastal region of
Bohai, China; Age: Eocene-Oligocene.
6.9.2. Species: T. multicellulus P. Ke & Z.Y. Shi 1978; Index Fungorum Registration Identifier:
115651; Current name: Mossopisporites multicellulus (P. Ke & Z.Y. Shi) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).
6.9.3. Species: T. simplex (Elsik & Janson.) Kalgutkar & Janson. 2000; Index Fungorum
Registration Identifier: 483570; Basionym: Pesavis simplex Elsik & Janson. 1974; Location:
Mackenzie Delta, Northwest Territories, Canada; Age: Palaeogene.

831
6.9.4. Species: T. triangulus P. Ke & Z.Y. Shi 1978 (Fig. 14AC); Index Fungorum Registration
Identifier: 115650; Location: Panshan, Liaoning Province, Coastal region of Bohai, China;
Age: Eocene-Oligocene.

7. Fossil fungal spore species assigned to modern genera

7.1. Genus: ACREMONIUM Link, Mag. Gesell. naturf. Freunde, Berlin 3(1-20: 15 (1809); Index
Fungorum Registration Identifier: 7028; Type: A. alternarium Link 1809.
Classification: Bionectriaceae, Hypocreales, Hypocreomycetidae, Sordariomycetes,
Pezizomycotina, Ascomycota.
Number of fossil species known: One.
7.1.1. Species: A. succineum Casp. 1907; Index Fungorum Registration Identifier: 107444;
Location: Baltic area, Poland; Age: Tertiary (Oligocene?).

7.2. Genus: ALTERNARIA Nees, Syst. Pilze (Würzburg): 72 (1816); Index Fungorum Registration
Identifier: 7106; Type: A. tenuis Nees 1816.
Classification: Pleosporaceae, Pleosporales, Pleosporomycetidae, Dothideomycetes,
Pezizomycotina, Ascomycota.
Number of fossil species known: One (the single fossil species has been transferred to
Pluricellaesporites Hammen 1954).
7.2.1. Species: A. malayensis Trivedi & C.L. Verma 1970; Index Fungorum Registration Identifier:
308511; Current name: Pluricellaesporites malayensis (Trivedi & Verma) Kalgutkar &
Janson. 2000 fide Kalgutkar & Jansonius (2000).

7.3. Genus: BOTRYODIPLODIA Sacc., Syll. fung. (Abellini) 3: 377 (1884); Index Fungorum
Registration Identifier: 7420; Type: As per Index Fungorum, Saccardo accepted the type of
Diplodia b Botryodiplodia as Diplodia juglandis (Fr.) Fr. 1849.
Classification: Diporthales, Diaporthomycetidae, Sordariomycetes, Pezizomycotina,
Ascomycota.
Number of fossil species known: One (the single fossil species has been transferred to
Diplodites).
7.3.1. Species: B. mohgaoensis Barlinge & Paradkar 1982; Index Fungorum Registration Identifier:
108725; Current Name: Diplodites mohgaoensis (Barlinge & Paradkar) Kalgutkar et al. 1993
fide Kalgutkar et al. 1993.

7.4. Genus: BRACHYSPORIUM Sacc., Syll. fung. (Abellini) 4: 423 (1886); Index Fungorum
Registration Identifier: 7444; Type: B. obovatum (Berk.) Sacc. 1886.
Classification: Trichosphaeriaceae, Trichosphaeriales, Diaporthomycetidae,
Sordariomycetes, Pezizomycotina, Ascomycota.
Number of fossil species known: One (the single fossil species has been transferred to
Pluricellaesporites Hammen 1954).
7.4.1. Species: B. minutum Trivedi & C.L. Verma 1970; Index Fungorum Registration Identifier:
7444 (nom. inval.); Current name: Pluricellaesporites minutus (Trivedi & Verma) ex
Kalgutkar & Janson. 2000 fide Kalgutkar & Jansonius (2000).

7.5. Genus: CHAETOSPHAERIA Tul. & C. Tul., Select. fung. corpol. (Paris) 2: 252 (1863); Index
Fungorum Registration Identifier: 970; Type: C. innumera Berk. & Broome ex Tul. & C. Tul.
1863.
Classification: Chaetosphaeriaceae, Chaetosphaeriales, Sordariomycetidae,
Sordariomycetes, Pezizomycotina, Ascomycota.
Number of fossil species known: One.

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7.5.1. Species: C. elsikii M.J. Pound et al. 2018; Index Fungorum Registration Identifier: 821981;
Location: Bees Nest Pit, Brassington, Derbyshire, U.K.; Age: Miocene.

7.6. Genus: CLASTEROSPORIUM Schwein., Trans. Am. phil. Soc., New Series 4(2): 300 (1834);
Index Fungorum Registration Identifier: 7685; Type: C. caricinum Schwein. 1834.
Classification: Magnaporthaceae, Magnaporthales, Diaporthomycetidae, Sordariomycetes,
Pezizomycotina, Ascomycota.
Number of fossil species known: One.
7.6.1. Species: C. eocenicum Fritel & R. Vig. 1909; Index Fungorum Registration Identifier:
198755; Location: Brétigny et Muirancourt (Oise), France; as calcareous pseudomorphs in
marls underlying lignites; Age: Sparnacian (Early Eocene).

7.7. Genus: DESMIDIOSPORA Thaxt., Bot. Gaz. 16: 203 (1891); Index Fungorum Registration
Identifier: 7964; Type: D. myrmecophila Thaxt. 1891.
Classification: Ascomycota genera incertae sedis.
Number of fossil species known: Two.
Notes: Desmidiospora Thaxt. 1891 comprises three species. The type species D.
myrmecophila Thaxt. 1891 represents living fungi, whereas the other two, viz. D. willoughbyi
(W.H. Bradley) D.L.E. Glass et al. 1986 and D. marginiconvoluta Kalgutkar 1997, represent fossil
fungi.
7.7.1 Species: D. marginiconvoluta Kalgutkar 1997; Index Fungorum Registration Identifier:
437899; Location: Kanguk Peninsula, Axel Heiberg Island, Northwest Territories, Canada;
Age: Late Palaeocene-Early Eocene (Iceberg Bay Formation, Eureka Sound Group); Notes:
The species epithet indicates convoluted spore margin.
7.7.2. Species: D. willoughbyi (W.H. Bradley) D.L.E. Glass et al. 1986 (Fig. 14AD); Index
Fungorum Registration Identifier: 357741; Basionym: Entophlyctis willoughbyi Bradley
1967; Location: Wyoming, Colorado (Bradley 1967), East and south-central Texas, U.S.A.
(D.L.E. Glass et al. 1986); Age: Eocene (Bradley 1967), Late Eocene (Manning Formation,
Jackson group) (Glass et al. 1986).

7.8. Genus: DIPLODIA Fr., in Montagne, Annls Sci. Nat. Bot., ser. 2 1: 302 (1934); Index
Fungorum Registration Identifier: 8047; Type: D. mutila (Fr.) Mont. 1834.
Classification: Botryosphaeoriaceae, Botryosphaeriales, Dothideomycetes, Pezizomycotina,
Ascomycota.
Number of fossil species known: Two (both the fossil species have been transferred to
Diplodites).
7.8.1. Species: D. rodei Mahab. 1969; Index Fungorum Registration Identifier: 313187; Current
Name: Diplodites rodei (Mahab.) Kalgutkar et al. 1993 fide Kalgutkar et al. 1993.
7.8.2. Species: D. sahnii Singhai 1974; Index Fungorum Registration Identifier: 313191; Current
Name: Diplodites sahnii (Singhai) Kalgutkar et al. 1993 fide Kalgutkar et al. 1993.

7.9. Genus: DIPOROTHECA C.C. Gordon & C.G. Shaw, Mycologia 52(2): 331 (1961); Index
Fungorum Registration Identifier: 1635; Type: D. rhizophila C.C. Gordon & C.G. Shaw 1961.
Classification: Diporothecaceae, Pezizomycotina, Ascomycota.
Number of fossil species known: Two.
7.9.1. Species: D. doniana O’Keefe 2017; Index Fungorum Registration Identifier: 821917;
Location: Tumbes Province, Peru; Age: Miocene (Heath Formation); Notes: The specific
epithet is a feminization of ‘Don’ in honour of Donald W. Engelhardt, Earth Sciences
Resources Institute, University of South Carolina, Columbia, South Carolina, U.S.A.
7.9.2. Species: D. gorda O’Keefe 2017; Index Fungorum Registration Identifier: 821918; Location:
Tumbes Province, Peru; Age: Miocene (Heath Formation); Notes: The specific epithet is

833
 Spanish for ‘fat woman’, given the very boxy, fat shape of the grain relative to other spores of
Diporotheca.

7.10. Genus: ENTOPHLYCTIS A. Fisch. in Winter, Rabenh. Krypt.-Fl., Edn. 2 (Leipzig) 1(4): 114
(1892); Index Fungorum Registration Identifier: 20222; Type: E. cienkowskiana (Zopf) A. Fisch.
1892 (Basionym: Rhizidium cienkowskianum Zopf 1885)
Classification: Chytriomycetaceae, Chytridiales, Chytridiomycetidae, Chytridiomycetes,
Chytridiomycota.
Number of fossil species known: One (the single fossil species has been transferred to
Desmidiospora Thaxt.).
7.10.1. Species: E. willoughbyi W.H. Bradley 1967; Index Fungorum Registration Identifier:
330609; Current name: Desmidiospora willoughbyi (W.H. Bradley) D.L.E. Glass et al. 1986
fide D.L.E. Glass et al. 1986.

7.11. Genus: EPICOCCUM Link, Mag. Gesell. naturf. Freunde, Berlin 7: 32 (1816); Index
Fungorum Registration Identifier: 8188; Type: E. nigrum Link 1816.
Classification: Didymellaceae, Pleosporales, Pleosporomycetidae, Dothideomycetes,
Pezizomycotina, Ascomycota.
Number of fossil species known: One.
7.11.1. Species: E. deccanense R. Srivast. et al. 2009 (Fig. 14AE); Index Fungorum Registration
Identifier: 561524; Location: Jhargad, near Jhadgaon village, Yavatmal District, Maharashtra,
India; Age: Late Maastrichtian-Danian (Deccan Intertrappean Beds).

7.12. Genus: GONATOBOTRYS Corda, Pracht-Fl. Eur. Schimmelbiled.: 9 (1839); Index Fungorum
Registration Identifier: 8374; Type: G. simplex Corda 1839; Current name: MELANOSPORA
Corda 1837 fide Wijayawardene et al. (2020a).
Classification: Ceratostomataceae, Coronophorales, Hypocreomycetidae, Sordariomycetes,
Pezizomycotina, Ascomycota.
Number of fossil species known: One (the single fossil species is here transferred to
Melanospora Corda 1837).
7.12.1. Species: G. primigenius Casp. 1907; Index Fungorum Registration Identifier: 8374; Current
name: Melanospora primigenia (Casp.) R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde &
P.M. Kirk comb. nov. This new combination is described under the section “New species and
new combinations”.

7.13. Genus: MELANOSPORA Corda, Icon fung. (Prague) (1837); Index Fungorum Registration
Identifier: 3085; Type: M. zamiae Corda 1837.
Synonym: Gonatobotrys Corda 1839 fide Wijayawardene et al. 2020a, Index Fungorum
Registration Identifier: 8374.
Classification: Ceratostomataceae, Melanosporales, Hypocreomycetidae, Sordariomycetes,
Pezizomycotina, Ascomycota.
Number of fossil species known: One.
7.13.1. Species: M. primigenia (Casp.) R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk
comb. nov. (Fig. 14AF). This new combination is described under the section “New species
and new combinations”.

7.14. Genus: POTAMOMYCES K.D. Hyde, Nova Hedwigia 61(1-2): 132 (1995); Index Fungorum
Registration Identifier: 27626; Type: P. armatisporus K.D. Hyde 1995.
Synonym: Mediaverrunites Nandi & A. Sinha 2007 fide Nuñez Otaño et al. 2017, Index
Fungorum Registration Identifier: 548478.
Classification: Ascomycota genera incertae sedis.
Number of fossil species known: Seven.

834
7.14.1. Species: P. batii (Sancay) ex Nuñez Otaño et al. 2017; Index Fungorum Registration
Identifier: 814793; Basionym. Mediaverrunites batii Sancay 2014; Location: Turkey; Age:
Late Miocene.
7.14.2. Species: P. elsikii (Nandi & A. Sinha) Nuñez Otaño et al. 2017; Index Fungorum
Registration Identifier: 814695; Basionym. Mediaverrunites elsikii Nandi & A. Sinha 2007;
Location: Rengtekawn-Sherlui Road Traverse, Mizoram, India; Age: Neogene.
7.14.3. Species: P. fournieri (Elsik & Jarzen) Nuñez Otaño et al. 2017; Index Fungorum
Registration Identifier: 814696; Basionym: Mediaverrunites fournierii Elsik & Jarzen 2009;
Location: Colombia, South America; Age: Early Miocene.
7.14.4. Species: P. invaginatus (Elsik & Jarzen) Nuñez Otaño et al. 2017; Index Fungorum
Registration Identifier: 814698; Basionym: Mediaverrunites invaginatus Elsik & Jarzen
2009; Location: Colombia, South America; Age: Early Miocene.
7.14.5. Species: P. magnus (Elsik & Jarzen) Nuñez Otaño et al. 2017; Index Fungorum
Registration Identifier: 814697; Basionym: Mediaverrunites magnus Elsik & Jarzen 2009;
Location: Colombia, South America; Age: Early Miocene.
7.14.6. Species: P. mulleri (Nandi & A. Sinha) Nuñez Otaño et al. 2017; Index Fungorum
Registration Identifier: 814692; Basionym: Mediaverrunites mulleri Nandi & A. Sinha 2007;
Location: Rengtekawn-Sherlui Road Traverse, Mizoram, India; Age: Neogene.
7.14.7. Species: P. pontidiensis (Sancay) ex Nuñez Otaño et al. 2017; Index Fungorum Registration
Identifier: 814794; Basionym: Mediaverrunites pontidiensis Sancay 2014; Location: Turkey;
Age: Late Miocene.

7.15 Genus: RAMULARIA Unger, Exanth. Pflanzen (Wein): 119 (1833); Index Fungorum
Registration Identifier: 9691; Type: R. pusilla Unger 1833.
Classification: Mycosphaerellaceae, Capnodiales, Dothideomycetidae, Dothideomycetes,
Pezizomycotina, Ascomycota.
Number of fossil species known: One
7.15.1. Species: R. oblongispora Casp. 1907; Index Fungorum Registration Identifier: 107582;
Location: Baltic area, Poland; Age: Tertiary (Oligocene?).

7.16. Genus: RHEXOAMPULLIFERA P.M. Kirk, Trans. Br. mycol. Soc. 78(2): 299 (1982); Index
Fungorum Registration Identifier: 9717; Type: R. fagi (M.B. Ellis) P.M. Kirk & C.M. Kirk 1982.
Classification: Ascomycota genera incertae sedis.
Number of fossil species known: Two.
7.16.1. Species: R. stogieana M.J. Pound et al. 2018; Index Fungorum Registration Identifier:
821979; Location: Bees Nest Pit, Brassington, Derbyshire, U.K.; Age: Miocene.
7.16.2. Species: R. sufflata M.J. Pound et al. 2018; Index Fungorum Registration Identifier:
821980; Location: Bees Nest Pit, Brassington, Derbyshire, U.K.; Age: Miocene.

7.17. Genus: SCLERODERMA Pers., Syn. meth. fung. (Göttingen) 1: xiv, 150 (1801); Index
Fungorum Registration Identifier: 19309; Type: S. verrucosum (Bull.) ex Pers. 1801.
Classification: Sclerodermataceae, Boletales, Agaricomycetidae, Agaricomycetes,
Agaricomycotina, Basidiomycota.
Number of fossil species known: One.
7.17.1. Species: S. echinosporites Rouse 1962; Index Fungorum Registration Identifier: 110078;
Location: South shore of Burrard Inlet, British Columbia, Canada; Age: (Middle?) Eocene
(but possibly as old as Late Cetaceous?).

7.18. Genus: SOROSPORIUM F. Rudolphi, Linnaea 4: 116 (1829); Index Fungorum Registration
Identifier: 16318; Type: S. saponariae F. Rudolphi 1829; Current name: THECAPHORA Fingerh
1836 fide Wijayawardene et al. (2020a).

835
 Classification: Glomosporiaceae, Urocystidales, Ustilaginomycetes, Ustilaginomycotina,
Basidiomycota.
Number of fossil species known: One (the single fossil species is here transferred to
Melanospora Corda 1837).
Notes: Sorosporium F. Rudolphi was rejected against the conserved name Thecaphora
Fingerh and its type species S. saponariae F. Rudolphi, was transferred to Thecaphora Fingerh by
Vánky 1998.
7.18.1. Species: S. mohgaoense Chitaley & Yawale 1978; Index Fungorum Registration Identifier:
111035; Current Name: Thecaphora mohgaoensis (Chitaley & Yawale) R.K. Saxena,
Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk comb. nov. This new combination is described
under the section “New species and new combinations”.

7.19. Genus: SPHAEROPHORUS Pers., Ann. Bot. (Usteri) 7: 23 (1794); Index Fungorum
Registration Identifier: 5112; Type: S. coralloides Pers. 1794.
Classification: Sphaerophoraceae, Lecanorales, Lecanoromycetidae, Lecanoromycetes,
Pezizomycotina, Ascomycota.
Number of fossil species known: One.
7.19.1. Species: S. moniliformis Menge 1858; Index Fungorum Registration Identifier: 628321;
Location: Baltic area; Age: Early Tertiary (Oligocene?).
7.20. Genus: SPORIDESMIUM Link, Mag. Gesell. naturf. Freunde. Berlin 3(1-2): 41 (1809);
Index Fungorum Registration Identifier: 10024; Type: S. atrum Link 1809.
Classification: Sporidesmiaceae, Sporidesmiales, Diaporthomycetidae, Sordariomycetes,
Pezizomycotina, Ascomycota.
Number of fossil species known: One.
7.20.1. Species: S. henryense Dilcher 1965; Index Fungorum Registration Identifier: 339523;
Location: Western Tennessee, U.S.A.; Age: Early Eocene.

7.21. Genus: STILBUM Tode, Fung. mecklenb. sel. (Luneburg) 1: 10 (1790); Index Fungorum
Registration Identifier: 18601; Type: S. vulgare Tode 1790.
Classification: Chionosphaeraceae, Agaricostilbales, Agaricostilbomycetes,
Pucciniomycotina, Basidiomycota.
Number of fossil species known: One.
7.21.1. Species: S. succini Casp. 1907; Index Fungorum Registration Identifier: 107596; Location:
Baltic area, Poland; Age: Tertiary (Oligocene); Notes: The species epithet is derived from
Latin succinum = amber.

7.22. Genus: TETRACOCCOSPORIUM Szabó, Hedwigia 44: 77 (1905); Index Fungorum

Registration Identifier: 10192; Type: T. paxianum Szabó 1905
Classification: Ascomycota genera incertae sedis.
Number of fossil species known: One.
7.22.1. Species: T. eocenum Biradar & Mahab. 1974 (Fig. 14AG); Index Fungorum Registration
Identifier: 519781; Location: Mohgaon Kalan, Chhindwara District, Madhya Pradesh, India;
Age: Maastrichtian (Deccan Intertrappean Series).

7.23. Genus: THECAPHORA Fingerh, Linnaea 10: 230 (1836); Index Fungorum Registration
Identifier: 16347; Type: T. hyalina Fingerh 1836.
Synonym: Sorosporium F. Rudolphi 1829 fide Index Fungorum, Index Fungorum
Registration Identifier: 16318.
Classification: Glomosporiaceae, Urocystidales, Ustilaginomycetes, Ustilaginomycotina,
Basidiomycota.
Number of fossil species known: One.

836
7.23.1. Species: T. mohgaoensis (Chitaley & Yawale) R.K. Saxena, Wijayaw., D.Q. Dai, K.D.
Hyde & P.M. Kirk comb. nov. (Fig. 14AH). This new combination is described under the
section “New species and new combinations”. Notes: Kalgutkar & Jansonius 2000 transferred
Sorosporium mohgaoense to fossil fungal spore genus Papulosporonites [as
Papulosporonites mohgaoensis (Chitaley & Yawale) Kalgutkar & Janson. 2000] without
providing any convincing reason for the same. We, therefore, do not accept this transfer.

7.24. Genus: TORULA Pers., Ann. Bot. (Usteri) 15: 25 (1795); Index Fungorum Registration
Identifier: 10248; Type: T. monilis Pers. 1795.
Classification: Torulaceae, Pleosporales, Pleosporomycetidae, Dothideomycetes,
Pezizomycotina, Ascomycota.
Number of fossil species known: Three.
7.24.1. Species: T. globulifera Casp. 1907; Index Fungorum Registration Identifier: 107604;
Location: Baltic area, Poland; Age: Tertiary (Oligocene?).
7.24.2. Species: T. heteromorpha Casp. 1907; Index Fungorum Registration Identifier: 105832;
Location: Baltic area, Poland; Age: Tertiary (Oligocene?).
7.24.3. Species: T. mengeana Casp. & R. Klebs in Casp. 1907; Index Fungorum Registration
Identifier: 634247; Location: Baltic area, Poland; Age: Tertiary (Oligocene?).

7.25. Genus: USTILAGO (Pers.) Roussel, Fl. Calvados, Edn 2: 47 (1806), Index Fungorum
Registration Identifier: 16391; Type: U. hordei (Pers.) Lagerh. 1889.
Classification: Ustilaginaceae, Ustilaginales, Ustilaginomycetidae, Ustilaginomycetes,
Ustilaginomycotina, Basidiomycota.
Number of fossil species known: One (the single species has been transferred to
Inapertisporites Hammen 1954).
7.25.1. Species: U. deccanii Chitaley & Yawale 1978; Index Fungorum Registration Identifier:
111080; Current Name: Inapertisporites deccanii (Chitaley & Yawale) Kalgutkar & Janson.
2000 fide Kalgutkar & Jansonius (2000).

7.26. Genus: ZOPFIELLA G. Winter, Rabenh. Krypt. -Fl., Edn 2 (Leipzig) 1.2: 56 (1884); Index
Fungorum Registration Identifier: 5876; Type: Z. tabulata Zopf ex G. Winter 1884.
Classification: Lasiosphaeriaceae, Sordariales, Sordariomycetidae, Sordariomycetes,
Pezizomycotina, Ascomycota.
Number of fossil species known: One.
7.26.1. Species: Z. neogenica O’Keefe 2017; Index Fungorum Registration Identifier: 821916;
Location: Tumbes Province, Peru; Age: Miocene (Heath Formation); Notes: The specific
epithet refers to the age of the specimen.

New species and new combinations

Hypoxylonites disciformis (Sheffy & Dilcher) R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde &
P.M. Kirk comb. nov. Fig. 6G
Index Fungorum Registration Identifier: 554345.
Basionym – Inapertisporites disciformis Sheffy & Dilcher, Palaeontographica Abt. B 133:
39, pl. 13, fig. 8, pl. 15, fig. 8, 1971; Index Fungorum Registration Identifier: 111544.
Synonym – Spirotremesporonites disciformis (Sheffy & Dilcher) Elsik 1990a.
Holotype – Sheffy & Dilcher 1971: 39, pl. 13, fig. 8, pl. 15, fig. 8; PPS (1) 17.6 × 101.2.
Diagnosis – Disk-shaped spore, size 6.8 × 16.4 µm, a narrow, straight slit present from one
end to the other, spore wall 1 µm thick, psilate, medium pigment
Location – Puryear clay pit, 800 m south of Puryear, Henry County, Tennessee, U.S.A.
Age – Middle Eocene (Claiborne Formation).

837
 Notes – Kalgutkar & Jansonius (2000) stated that the line drawing of Inapertisporites
disciformis given by Sheffy & Dilcher (1971) appears to show similarity to Hypoxylonites, based
on the pattern of the furrow, which is straight and not oblique, wavy or spiral. However, they
tentatively followed synonymy in Elsik (1990a) and described it as Spirotremesporites disciformis.
We interpret the longitudinal slit, from one end to the other, as straight and transfer it to
Hypoxylonites.

Hypoxylonites lanceolatus (Debi Mukh.) R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M.
Kirk comb. nov. Fig. 6N
Index Fungorum Registration Identifier: 554568.
Basionym – Xylariasporites lanceolatus Debi Mukh., International Journal of Geology, Earth
and Environmental Sciences 2(2): 9, fig. 3.13, 2012; Index Fungorum Registration Identifier:
588483.
Holotype – Mukherjee 2012: 9, fig. 3.13; size 60 × 40 µm, slide no. LU-D/6, co-ordinates
(K7/4), Museum, Department of Geology, Lucknow University, Lucknow, India.
Diagnosis – Aeciospore hyaline, brown, one celled, dark brown in colour, germ pore slit-like,
size range 60–70 × 30–40 μm (usually 60 × 40 μm); lanceolate, biconvex, tapering at the poles,
aperture fine, 2 μm wide.
Location – Neyveli Lignite Mine-I, Cuddalore District, Tamil Nadu, India.
Age – Miocene (Neyveli Lignite).
Notes – Mukherjee (2012) stated that this species belongs to family Xylariaceae of
Ascomycota. The present spore having one cell with fine germ pore, shows affinity with Xylaria
spores (as mentioned by Ainsworth et al. 1973). Xylariasporites lanceolatus Debi Mukh. (Type of
Xylariasporites) conforms to Hypoxylonites Elsik 1990a in all the essential characters, hence
transferred to Hypoxylonites. The specific epithet refers to its lens-like shape, lanceolatus = lens-
like.

Melanospora primigenia (Casp.) R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk
comb. nov. Fig. 14AF
Index Fungorum Registration Identifier: IF556982.
Basionym – Gonatobotrys primigenius Casp. (as “primigenia”), in Klebs, Abh.. preuss. geol.
Landesanst. 4(1): 11, taf. 1, fig. 6. 1907, Index Fungorum Registration Identifier: 626875.
Synonym – Gonatobotrytites primigenius (Casp.) Pia 1927; Index Fungorum Registration
Identifier: 115070.
Holotype – Gonatobotrys primigenius Casp. 1907: 11, taf. 1, fig. 6, collected from Tertiary
sediments of Austria.
Location – Location: Baltic area, Poland.
Age – Tertiary (Oligocene?).

Thecaphora mohgaoensis (Chitaley & Yawale) R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde &
P.M. Kirk comb. nov. Fig. 14AH
Index Fungorum Registration Identifier: 556983.
Basionym – Sorosporium mohgaoense Chitaley & Yawale, Botanique 7(4): 190, pl. 1, fig. 1.
1978; Index Fungorum Registration Identifier: 111035.
Synonym: Papulosporonites mohgaoensis (Chitaley & Yawale) Kalgutkar & Janson. 2000:
217, pl. 18, fig. 6; Index Fungorum Registration Identifier: 483503.
Holotype – Chitaley & Yawale 1978: 190; pl. 1, fig. 1; I Fu/NRY deposited with SRII,
Nagpur (designated by Kalgutkar & Jansonius 2000).
Diagnosis – The spore balls are deeply buried in the host tissue and look reddish brown to
pale yellow. They are egg-shaped consisting of 5 to 25 spores. The spores are more or less
permanently united and the balls are not covered by a sterile sheath or any pseudomembrane. No
sterile cells are present inside the spore balls. Size of spore balls varies from 17–21 × 35–46 μm.

838
Individual spores are globose in shape but because of compression they look polyhedral. Germ
pores are observed in many of them. However, the mycelium is not seen. Spores are 5.3 to 10.6 μm
with an average of 8 μm in diameter, globose to ovoid, polyhedral in balls, without any contents,
epispore smooth, 0.3–0.7 μm.
Location – Mohgaonkalan, Chhindwara District, Madhya Pradesh, India.
Age – Late Cretaceous-Maastrichtian.

Dicellaesporites vermae R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk sp. nov.
Fig. 9P
Index Fungorum Registration Identifier: 555416.
Etymology – The species is named in honour of late Professor C.L. Verma of the Department
of Botany, Lucknow University, Lucknow, India.
Holotype – Dicellaesporites sp. A in H.P. Singh, R.K. Saxena & M.R. Rao 1986,
Palaeobotanist 35(1): 98, pl. 2, fig. 18, BSIP slide no. 8123, coordinates 62.4 × 106.5, Museum,
Birbal Sahni Institute of Palaeosciences, Lucknow, India.
Diagnosis – Fungal spores elliptical in shape, size 79–87 × 31–35 µm, dicellate, inaperturate,
uniseptate, septa 1 µm thick, spore wall psilate, hyaline, sometimes with minor folds.
Location – Sonapur-Badarpur Road section, Jaintia Hills, Meghalaya and Cachar, Assam,
India.
Age – Early Miocene (Dona Member, Bhuban Formation, Surma group).
Dyadosporites singhii R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk sp. nov.
Fig. 9AH
Index Fungorum Registration Identifier: 555425.
Etymology – The species is named in honour of late Dr. H.P. Singh of the Birbal Sahni
Institute of Palaeosciences, Lucknow, India.
Holotype – Diporicellaesporites sp. in R.Y. Singh, Dogra & Vimal 1985, Journal of
Palynology 21: 53, pl. 3, fig. 54, slide no. LUGM 44/10/20, Museum, Department of Geology,
Lucknow University, Lucknow, India.
Diagnosis – Fungal spores brown coloured, fusiform with tapering ends, size 78–90 × 23–28
µm, diporate, pores terminal, distinct, large, 6–10 µm in diameter, dicellate, uniseptate, septum 4
µm thick, spore wall 1.5 µm thick, psilate.
Location – Assam and Meghalaya, India.
Age – Oligocene (Barail Group).

Fusiformisporites sahii R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk sp. nov.
Fig. 10L
Index Fungorum Registration Identifier: 555491.
Etymology – The species is named in honour of late Dr. S.C.D. Sah of the Wadia Institute of
Himalayan Geology, Dehradun, India.
Holotype – Fusiformisporites sp. in R.K. Saxena & S. Khare 1992, Geophytology 21: 38, pl.
1, fig. 8, BSIP slide no. 10384, coordinates 40.2 × 107.9, Museum, Birbal Sahni Institute of
Palaeosciences, Lucknow, India.
Diagnosis – Spores fusiform with pointed ends, size 103–110 × 35–38 µm, inaperturate,
dicellate, septum 3–4 µm thick, each cell having longitudinal ribs, spore wall 1 µm thick, psilate.
Location – Jayamkondacholapuram Well 12, Tiruchirappalli District, Tamil Nadu, India.
Age – Late Palaeocene-Middle Eocene.

Diporicellaesporites tiruchirappalliensis R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M.
Kirk sp. nov. Fig. 11H
Index Fungorum Registration Identifier: 555593.
Etymology – The species is named after Tiruchirappalli District of Tamil Nadu, India where
the type locality is situated.

839
Figure 14 – A Dictyosporae. B–N Helicosporae. O–AC Staurosporae. AD–AH Fossil fungal spore
species assigned to modern genera. A Staphlosporonites tetracellatus A. Gupta 2002, Bar = 10 μm.
B Colligerites kutchensis (R.K. Kar & R.K. Saxena) K.P. Jain & R.K. Kar 1979, Bar = 8 μm. C
Colligerites trochus B. Samant in R.K. Saxena 2009, Bar = 30 μm. D Elsikisporonites tubulatus P.
Kumar 1990, Bar = 12 μm. E Helicominites salvinites Barlinge & Paradkar 1982, Bar = 10 μm. F
Helicoonites goosii Kalgutkar & Sigler 1995, Bar = 10 μm. G Helicosporiates pirozynskii
Kalgutkar & Sigler 1995, Bar = 8 μm. H Involutisporonites chowdhryi (K.P. Jain & R.K. Kar)
Kalgutkar & Janson. 2000, Bar = 10 μm. I Involutisporonites foraminus R.T. Clarke 1965, Bar = 10
μm. J Palaeocirrenalia elegans Ramanujam & Srisailam 1980, Bar = 8 μm. K Palaeocirrenalia
oligoseptata Ramanujam & Srisailam 1980, Bar = 20 μm. L Paleoslimacomyces canadensis

840
Kalgutkar & Sigler 1995, Bar = 5 μm. M Paleoslimacomyces wilcoxii (Elsik.) Kalgutkar & Janson.
2000, Bar = 8 μm. N Retihelicosporonites elsikii Ramanujam & K.P. Rao 1979, Bar = 20 μm. O
Eoglobella longipes W.H. Bradley 1931, Bar = 8 μm. P Frasnacritetrus conatus R.K. Saxena & S.
Sarkar 1986, Bar = 10 μm. Q Frasnacritetrus indicus R.K. Saxena & S. Khare 1991, Bar = 10 μm.
R Frasnacritetrus jamtahensis A. Gupta 2002, Bar = 15 μm. S Frasnacritetrus josettae Taug. 1968,
Bar = 5 μm. T Frasnacritetrus masolensis R.K. Saxena & S.K.M. Tripathi 2011, Bar = 20 μm. U
Frasnacritetrus siwalikus R.K. Saxena et al. 1987, Bar = 10 μm. V Frasnacritetrus taugourdeaui
R.K. Saxena & S. Sarkar 1986, Bar = 10 μm. W Mossopisporites multicellulus (P. Ke & Z.Y. Shi)
Kalgutkar & Janson. 2000, Bar = 10 μm. X Pesavis tagluensis Elsik & Janson. 1974, Bar = 10 μm.
Y Spegazzinites cruciformis Félix 1894, Bar = 10 μm. Z Spegazzinites indicus Ramanujam &
Srisailam 1980, Bar = 10 μm. AA Tribolites tetrastonyx W.H. Bradley 1964, Bar = 15 μm. AB
Trihyphites fractus (Z.C. Song & Liu Cao) Kalgutkar & Janson. 2000, Bar = 10 μm. AC
Triporicellaesporites triangulus Ke & Shi 1978, Bar = 15 μm. AD Desmidiospora willoughbyi
(W.H. Bradley) D.L.E. Glass, et al. 1986, Bar = 8 μm. AE Epicoccum deccanense R. Srivast. et al.
2009, Bar = 10 μm. AF Melanospora primigenia (Casp.) R.K. Saxena, Wijayaw., D.Q. Dai, K.D.
Hyde & P.M. Kirk comb. nov., Bar = 5 μm. AG Tetracoccosporium eocenum Biradar & Mahab.
1974, Bar = 20 μm. AH Thecaphora mohgaoensis (Chitaley & Yawale) R.K. Saxena, Wijayaw.,
D.Q. Dai, K.D. Hyde & P.M. Kirk comb. nov., Bar = 10 μm.

Holotype – Diporicellaesporites sp. in R.K. Saxena & S. Khare 1992, Geophytology 21: 39,
pl. 1, fig. 6, BSIP slide no. 10381, coordinates 69.2 × 93.2, Museum, Birbal Sahni Institute of
Palaeosciences, Lucknow, India.
Diagnosis – Spores elongated, size 58–68 × 20–22 µm, tetracellate, terminal cells smaller
than the middle ones, septum between two bigger cells 4 µm thick, other septa less than 1 µm thick,
diporate, pores terminal, distinct, ca. 4 µm in diameter, spore wall 1.5 µm thick, psilate.
Location – Jayamkondacholapuram Well 12, Tiruchirappalli District, Tamil Nadu, India.
Age – Late Palaeocene-Middle Eocene.

Multicellites tamilensis R.K. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde & P.M. Kirk sp. nov.
Fig. 11AJ
Index Fungorum Registration Identifier: 555609.
Etymology – The species is named after Tamil Nadu State of India where the type locality is
situated.
Holotype – Multicellaesporites sp. 1 in R.K. Saxena & S. Khare 1992, Geophytology 21: 38,
pl. 1, fig. 9, BSIP slide no. 10381, coordinates 30.1 × 100.0, Museum, Birbal Sahni Institute of
Palaeosciences, Lucknow, India.
Diagnosis – Spores elliptical with rounded ends; size 82–90 × 27–29 µm, octacellate, septa
distinct, 1.5–2 µm thick, no longitudinal slit or furrow present, spore wall 1.5 µm thick, psilate.
Location – Jayamkondacholapuram Well 12, Tiruchirappalli District, Tamil Nadu.
Age – Late Palaeocene-Middle Eocene.

Discussion
It is evident from the fungal spore records that they exhibit a broad range of morphological
variations. Since their assignment to modern fungal taxa is seldom possible, these are placed into
artificial supra-generic taxa based on morphological characters, e.g. number and nature of cells and
characters associated with apertures, septa and spore wall. These may be unicellate, dicellate,
tricellate, tetracellate, multicellate, muriform, filiform, spirally coiled and star-like. Similarly, these
may be inaperturate, monoaperturate, diaperturate, triaperturate and multiaperturate. These artificial
supra-generic taxa are: Amerosporae, Didymosporae, Phragmosporae, Dictyosporae, Helicosporae
and Staurosporae. Under each of these, genera are arranged in alphabetical order. Similarly, species
are also arranged under each genus in alphabetical order. Five new species, viz. Dicellaesporites
vermae, Dyadosporites singhii, Fusiformisporites sahii, Diporicellaesporites tiruchirappalliensis

841
and Multicellites tamilensis, and four new combinations, viz. Hypoxylonites disciformis (Sheffy &
Dilcher), Hypoxylonites lanceolatus (Debi Mukh.), Melanospora primigenia (Casp.) and
Thecaphora mohgaoensis (Chitaley & Yawale) are also proposed. The dominant genera, both in
number and variety, having more than twenty species are: Dicellaesporites (66 species),
Diporicellaesporites (43 species), Diporisporites (35 species), Dyadosporites (43 species),
Hypoxylonites (55 species), Inapertisporites (71 species), Monoporisporites (58 species),
Multicellaesporites (91 species), Multicellites (46 species), Pluricellaesporites (83 species) and
Staphlosporonites (23 species). The diagnostic characters of the most important and commonly
occurring fossil fungal spore genera are summarized in Table 2.

Table 2 Summary of diagnostic characters of important, commonly occurring, fossil fungal spores

Supra-generic Fungal spore genera Diagnostic features

taxa
Amerosporae Basidiosporites Elsik Elongate-variable shaped spores with a single pore offset from
Unicellate one apex, spore wall psilate.
Biporipsilonites Kalgutkar Elongate, fusiform to barrel-shaped spores, generally with a
& Janson. plane of symmetry through the equator, spore wall generally
smooth, occasionally with some subdued sculpture, two terminal
pores, forming pore chambers subtended by a basal septum and
enclosed by thin wall material that further thins centrifugally,
septa thin or thick, may have a central perforation and/or small
septal folds.
Cervichlamydospora R. Chlamydospores sub-circular, dark brown-black, originate from
Kar et al. neck of hyphae; solitary, many hyphae adhere together at base,
branch out laterally at tip; hyphae wall laevigate-granulose, grana
up to 1 μm thick, sparsely placed.
Diporisporites Hammen Elongate, diporate spores, pores on opposite ends.
Exesisporites Elsik Lenticular, monoporate, pore small, pore surrounded by
thickening.
Foveodiporites C.P. Varma Diporate, fusiform to elliptic, spore wall relatively thin,
& Rawat externally smooth, internally smooth, or with punctate, granulate,
foveolate or similar sculpture; pores terminal, complex,
consisting of a thin collar and separated from the spore interior
by one or two septa.
Hypoxylonites Elsik Oval to elongate, bilateral, psilate spores, provided with elongate
scar, slit or furrow.
Inapertisporites Hammen Inaperturate, shape and size variable, spore wall psilate to
variously ornamented.
Lacrimasporonites R.T. Spatulate to elliptical spores, spore wall psilate, monoporate,
Clarke pore apical.
Monoporisporites Hammen Monoporate, spherical to sub-spherical spores, spore wall psilate
to finely punctuate.
Palaeoamphisphaerella Elliptical, oblong or rhomboidal spores with rounded ends,
Ramanujam & Srisailam provided with equatorial pore.
Retidiporites C.P. Varma & Diporate spores with reticulate exine.
Rawat
Spirotremesporites Dueñas Psilate, aseptate, elongate-elliptical to oval spores, aperture in the
form of single furrow at an angle to the axis of the spore, straight
or curved to S-shaped or sigmoidal in outline, or spiral around
the spore axis.
Striadiporites C.P. Varma Oval to fusiform spores, wall longitudinally ribbed to broadly
& Rawat reticulate. Two pores, one at each end of the spore on the long
axis.
Didymosporae Dicellaesporites Elsik Two-celled, uniseptate spores with variable shapes, inaperturate,
spore wall psilate.
Didymoporisporonites Dicellate, uniseptate spores, apex of one cell provided with pore,
Sheffy & Dilcher spore wall psilate to punctuate.
Dyadosporites R.T. Clarke Diporate spores, with a single pore at each end, spore wall psilate
to variously sculptured.

842
Table 2 Continued.

Supra-generic Fungal spore genera Diagnostic features

taxa
Fusiformisporites Rouse Fusiform, inaperturate spores, split into two equal halves by
equatorial wall, bearing characteristic elongate striae, ribs,
ridges or costae oriented parallel to longer axis.
Hilidicellites Kalgutkar & Small to medium-sized dicellate spores, with the proximal
Janson. end flattened or truncate, due to the presence of a hilum or
pore-like structure, the two cells generally of comparable size;
spore wall thin, smooth or with subdued sculpture.
Phragmosporae Alleppeysporonites Spores branched, multicellate, septate, individual cell
Ramanujam & K.P. Rao rectangular, basal and terminal cells provided with a
conspicuous appendage.
Axisporonites Kalgutkar & Inaperturate, tricellate spores, elliptical in shape, two polar
Janson. cells smaller, thicker, triangular, with dark pigmentation,
central cell large, hyaline; septa thick.
Brachysporisporites R.T. Obovate, turbinate or pyriform, multi-celled spores, cells
Lange & P.H. Sm. broader than long, gradually diminishing in size towards the
attachment cell which is the smallest, with very dark, thick
bands of septa similarly reducing in size.
Cannanorosporonites Tetracellate, barrel-shaped spores, basal and terminal cells
Ramanujam & K.P. Rao smaller than central cells.
Ceratohirudispora R. Kar Conidiophore small, growth terminated by production of
et al. apical conidium; conidium enlarges laterally in opposite
direction to produce two-three arms, conidia 5–10 celled,
septa with broad base and narrow tip.
Chaetosphaerites Felix Sporidia strongly obtuse spindle-shaped, almost like that of a
cylinder with rounded ends, consist of 4 segments, two
median, dark coloured cells larger than the two others.
Circinoconites R. Kar et al. Conidia acrogenous, strongly spiraled, 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.
Cladosporiumsporinites Conidia cylindrical, oblong, rounded at both the ends, 3–4
Debi Mukh. septate, fuliginous (dark soot colour), septa slightly
constricted.
Diporicellaesporites Elsik Elongate, diporate spores, one pore at each end of the spore,
wall psilate to finely structured.
Diporipollis S.K. Dutta & Diporate; globular to sub-spheroidal; pores placed one over
S.C.D. Sah the other, circular in shape, and encircled by one or more
thickened rims; exine thin, surface sculpture psilate to
scabrate or finely granulate.
Dwayabeejaesporonites Conidiophores with 14-15 cells arranged in acropetal order,
Debi Mukh. thick walled, enclosed in an undulating conidiophores wall;
conidiophores up to 200 μm size, apical cell mostly smaller.
Edmundmasonaesporites Spore multicellular, septate, dark in colour; 5 celled, apical
Debi Mukh. cell enlarged, globular, vacuolated, apical cell larger than the
basal one, spore wall psilate.
Foveoletisporonites Four or more celled, elongate spores, spore wall foveolate,
Ramanujam & K.P. Rao foveolae irregularly aligned.
Kumarisporites Kalgutkar Tricellate, inaperturate spores, central cell may be larger than
& Janson. the tapering terminal cells; septa thicker than spore wall;
spore wall ornamented by longitudinal ribs running full length
of the spore, tapering towards the poles.
Mathurisporites Kalgutkar Hilate spores, generally consisting of a darker central part
& Janson. with 2–4 cells, and proximal and distal parts of a single to few
hyaline cells. No distal pore.
Multicellaesporites Elsik Three or more celled spores, shape variable, a longitudinal slit
or furrow present, inaperturate, spore wall psilate or
ornamented or differentially coloured or thickened.

843
Table 2 Continued.

Supra-generic Fungal spore genera Diagnostic features

taxa
Multicellites Kalgutkar & Multicellate, uniseriate, inaperturate, number of cells three to
Janson. many, terminal cells usually rounded; spore wall usually
smooth, of medium thickness, septa generally perforate, or
with septal folds.
Nigrosporites Debi Mukh. Conidia oval/ lanceolate, somewhat flattened, large,
absolutely opaque, colour black, spore wall hyaline.
Ornasporonites Fusiform, four-celled, diporate spores, basal and apical cells
Ramanujam & K.P. Rao much small, one pore at each end.
Pluricellaesporites Three or more celled spores, long, monoporate, psilate to
Hammen scabrate.
Quilonia K.P. Jain & R.C. Filamentous, multicellular spores, apical and basal portions
Gupta narrow, central wide. Basal stalk prominent with one or two
rectangular thick-walled cells; apical cell mostly incomplete,
curved, central portion broad, elongate with irregularly
shaped furrow-like suture, inside the filament at different
places occur one to four small circular, ostiolate bodies. Exine
thick, margin undulated.
Ramasricellites Kalgutkar Inaperturate, tetracellate spores, ellipsoidal, with central cells
& Janson. broader, thicker walled and more pigmented than the terminal
cells; terminal cells thin-walled to hyaline, with rounded
ends;septa (or septal bases) thick and dark, evenly spaced.
Reduviasporonites L.R. Conidia-like spores occurring in uniseriate chains,
Wilson subspherical, slightly flattened at the contacts with adjacent
spores, all approximately same diameter, walls 1–2 μm thick,
uniform, smooth or slightly rough, yellow or brown,
translucent.
Scolecosporites R.T. Lange Long to very long, linear filamentous phragmospores, hilate,
& P.H. Sm. with or without distal pore, length many times width of spore.
Spores scalariform, commonly broken and lacking proximal
and/or distal portions; wall and septa commonly thin; septa
often with septal folds. Not or barely indented at septa.
Varmasporites Kalgutkar Fusiform, four-celled spores, inaperturate, with a pronounced
& Janson. constriction at the thick median septum, and with a distinct
ribbed or striate sculpture parallel to the long axis.
Dictyosporae Dictyosporites Felix Inaperturate, multicellate, muriform spores, cells rounded to
rounded polygonal. Overall shape rounded, oval/ovoid to
elongate.
Kutchiathyrites R.K. Kar Hilate conidia, fan shaped, formed by numerous linear
filaments radiating out from the hilum, conidia may be
flattened, filaments may be joined to their neighbours, or
partially free, and may branch towards the periphery.
Lirasporis R. Potonié & Fungal bodies oval-elliptical with equal or unequal, broad,
S.C.D. Sah generally notched ends. Mycelia, long, septate, ± parallel to
one another, extending from one end to other; wall generally
laevigate, sometimes granulose.
Papulosporonites Fungal remains of globular to elongate shape, consisting of
Schmied. & A.J. Schwab numerous more or less polygonal cells that are firmly fused
into mulberry-shaped aggregates. Cells without any regular
order, or concentrically arranged; one to three of the
innermost cells commonly much larger. Occasionally
individual aggregates fused together.
Polyadosporites Hammen Spores subspherical, loosely aggregated in clusters, with
individual cells not connected to others by shared walls;
clusters (colonies?) more or less regularly spherical to
subspherical.
Polycellaesporonites Anil Elongate, multicellate, inaperturate, psilate, one end rounded,
Chandra et al. other end giving rise to a tube-like projection, cells arranged
in clusters.

844
Table 2 Continued.

Supra-generic Fungal spore genera Diagnostic features

taxa
Staphlosporonites Sheffy Shape variable, four or more irregular cells arranged in
& Dilcher clusters along more than one axis, inaperturate, psilate to
punctate.
Helicosporae Colligerites K.P. Jain & Multicellular, coiled spores, cells generally smaller, rounded
R.K. Kar in central region and bigger, rectangular in outer region.
Spore wall mostly psilate, sometimes granulose. Pore may be
present or absent in each cell
Elsikisporonites P. Kumar Tubular and coiled spores, monoporate, pore at outer end,
non-septate, spore wall smooth and hyaline.
Helicominites Barlinge & Mycelium septate, branched, hyphae faint in colour;
Paradkar pycnidium and acervulus absent; conidia coiled in loose
spirals and narrow at both ends.
Involutisporonites R.T. Coiled, transversely septate spores, monoporate, psilate to
Clarke variously ornamented.
Palaeocirrenalia Inaperturate, helicoid spores, 1 to 1.25 times loosely coiled,
Ramanujam & Srisailam multicellular, 2- to 6- septate, septa transverse, prominent, as
thick and dark bands, cells of unequal size, terminal cell
dome-shaped and broader, basal cell usually cuneate, pale-
coloured, surface psilate.
Retihelicosporonites Uniseriate, multicellular, inaperturate spores, basal cell
Ramanujam & K.P. Rao cuneate, other cells rectangular, apical part of spore helical.
Spore wall reticulate.
Staurosporae Frasnacritetrus Taug. Main body rectangular, spherical or oval, psilate to variously
ornamented, body provided with four unicellular processes.
Spegazzinites Felix Hyphomycetous conidia consist of 4 partial cells, individual
cells more roundish, some conidia spinose, spines of various
lengths.

Relationship of fossil fungal spore genera with extant fungal taxa

The major applications of any kind of fossil study are in biostratigraphic zonation, correlation
and dating of the sedimentary rock successions and in determination of palaeoclimate,
palaeoecology and environment of deposition. For palaeoclimatic and palaeoecological
interpretations, “Uniformitarian Principle” (Uniformitarianism) is applied. This principle is based
on the assumption that the same natural laws and processes that operate in our present-day
scientific observations have always operated in the universe in the past and apply everywhere in the
universe. This principle is largely accepted by earth scientists the world over and is simplified as
“The Present is the key to the Past”.
The primary prerequisite for applying this principle is to relate fossils with their modern
counterparts. Since habitats, climate and ecology of present day organisms is known, it can easily
be suggested to be the same for their ancient fossil relatives. This prompts fossil workers, including
palaeomycologists, to trace affinity of fossil taxa to their present day representatives. An attempt
has been made here to trace relationship of fossil fungal spore taxa with extant fungi. This also
helps in interpreting evolutionary trend of a taxon or group of taxa.
Palaeomycologists and fungal taxonomists have been using molecular clock methods and
fossil records for ‘dating evolutionary divergences in the fungal tree of life’ (Lücking et al. 2009).
However, some studies suggested that the early fungi could have existed as lichens (Eriksson
2005). Taylor et al. (2015) accepted fossil fungal genera in six different phyla (viz.
Chytridiomycota, Blastocladiomycota, Glomeromycota, Ascomycota, Basidiomycota, Zygomycota),
according to the modern classification in the tree of life. Even though fossil fungal genera have
been reported in six different phyla, fungal spores are mainly restricted to Ascomycota and
Basidiomycota. Moreover, in many cases, the spores have been compared with hyphomycetous or

845
coelomycetous taxa, the two main artificial groups in traditional mycology (i.e. only morphological
characters have been considered in taxonomy).
Among the linked modern genera, hyphomycetous taxa which have separate conidiophores or
rarely synnemata are more important than other morphological groups. For examples, Alternaria
Nees, Dictyosporium Corda, Macrosporium Bon., Septosporium Corda and Stemphylium Wallr.
have been repeatedly compared with fossil fungal genera. Moreover, Helicoma Corda, Helicomina
L.S. Olive, Slimacomyces Minter and Trochophora R.T. Moore have also been compared with
fossil genera which have ‘helicoid’ conidia. Recent DNA based phylogenetic studies showed that
Tubeufia Penz. & Sacc., a sexually typified genus, has helicoma-like asexual morphs thus we can
assume some of these helicoma-like fossil asexual genera might have links with other sexual
morphs.
Fossil fungal spore taxa and their counterparts according to modern taxonomy are
summarized in Tables 3, 4. Table 3 presents fungal spore species assigned to extant fungal genera.
Altogether, 24 genera have been recorded, 19 belonging to Ascomycota, four to Basidiomycota and
one to Chytridiomycota. Table 4 presents fossil fungal spore genera and species and their probable
relationship with extant taxa. Here also, Ascomycota constitute the bulk of assemblage, with more
dominant classes being Dothideomycetes and Sordariomycetes.

Palaeoenvironmental Implications
Fungi, being heterotrophic in nature, are found in close association with specific plants and
animals, and when found in fossil state are indicative of similar kind of situations during the
geological past. Fossil fungi, therefore, may provide useful information about the
palaeoenvironment, past habitats and their hosts. In this regard, fossil epiphyllous fungi can be
more reliable and advantageous. Fossil spororocarps of microthyriaceous taxa are generally
considered to be reliable palaeoenvironmental indicators. Their occurrences are, generally,
correlated with moist, humid climates and tropical to subtropical temperatures. The role of fossil
fungi in interpreting past environment is now receiving increased attention (Kar & Saxena 1976,
Ramanujam & Rao 1978, van Geel et al. 1981, Pirozynski et al. 1988, Stubblefield & Taylor 1988,
Saxena & Tripathi 2011). Wolf (1966a, b, 1967a, b, 1968) interpreted the fluctuating abundance of
fungal spores in lakes as reflecting environmental perturbations and vegetation changes caused by
changes in climate. Wolf (1967a), in particular, restated the importance of fungal spore analyses
complement in pollen analyses in interpreting floristic changes and the sequence of vegetational
modifications in ancient times. The fossil peltate fungi are generally identified to the extant
Microthyriaceae which are ectoparasites on leaves of higher plants of tropical to subtropical zones
growing particularly in areas with high humidity. Edwards (1922) reported the occurrence of this
group on conifer needles. Microthyriaceous fungi grow best in rain forests, rain forest margins and
along creek banks (Ramanujam 1982). Hence their presence is generally indicative of a wet tropical
climate with heavy precipitation.
The palaeohabitat interpretations based on fossil epiphyllous microthyriaceous fungi and their
germlings is well established through the studies on their modern equivalents growing on leaf litter
from various Australian regions. These studies have shown the occurrence of microthyriaceous
germlings in greater number on the plants growing in moist tropical habitats. Such studies have
great potential in interpreting the palaeoclimate and should be undertaken for other geographical
areas. However, the ecological interpretations based on epiphyllous fungi should be made with
caution because some of these are reported to occur in wider latitudinal ranges (Dilcher 1965,
Selkirk 1975). It is therefore, advisable to take into consideration the complete palynological
assemblage for palaeoenvironmental interpretations. In most of the cases, coordinated studies on
megafossils in association with palynological assemblages may provide more accurate information
about the palaeoenvironmental conditions. Dilcher (1965) published an account of epiphyllous
fungi from Microthyriales, Erysiphales and Meliolales, thriving on leaves of different plants, from
the middle Eocene of Tennessee, U.S.A. Such studies bear great potential for determining the
regional palaeoclimate by comparing the fossils with extant taxa of known habitats. Environmental

846
interpretations based on the presence of Microthyriaceae may, however, sometimes be hampered
due to the incorrect identification of the material. Their presence in dispersed fossil assemblages
should, therefore, be ascertained before deciphering the past climate. The red alga Caloglossa
leprieurii, generally found on grasses of brackish water marshes may be confused with
Trichopeltinites due to morphological resemblance. Similarly, marine green alga Ulvella lens also
resembles the fructifications of Microthyriaceae.
Studies, particularly focusing on host-fungus relationship, are also of great significance in
attempting the palaeoenvironmental interpretations. Chitaley (1978) and Chitaley and Yawale
(1978) provided valuable palaeoecological information based on the presence of fossil fungal
spores in petrified plant materials from the Deccan Intertrappean beds of Central India.
Similar kinds of interpretations were published by Kar et al. (2003, 2004a, b, 2005, 2006) and
Sharma et al. (2005). These studies emphasize the importance of some fungal spores in evaluation
of palaeoenvironment. Kar et al. (2003) reported a sporocarp assignable to Polyporaceae
(Basidiomycota) from the Lameta Formation (Late Cretaceous) exposed in Madhya Pradesh, India.
This fossil, called Lithopolyporales zeerabadensis, resembles the modern Fomes which are
saprobes on dead wood of various trees. Kar et al. (2004a) described a fossil fungus showing
affinity to Colletotrichum Corda (Melanconiaceae), from an Intertrappean bed exposed at
Mohgaon-Kalan Village, Chhindwara District, Madhya Pradesh, India. The modern species of this
genus causes red rot in the economically important plants. The fossil of this fungus shows setae on
the margins of the acervuli and was found to be preserved on a leaf cuticle. It was called
Protocolletotrichum deccanensis. Kar et al. (2004b) described fossil parasitic fungi and epiphyllous
fruiting bodies from the coprolite of dinosaurs. The coprolite yielding these fossils was collected
from the Lameta Formation (Maastrichtian) of Central India. Occurrence of these fungi indicates
that the plant leaves infected by the recovered fungi were part of dinosaurs’ diet. Kar et al. (2006)
reported two types of fossil Ingoldian aquatic fungi from the Miocene sediments of Mizoram, India.
The first type, comparable to the extant Tetrachaetum, is needle-shaped and belongs to the
scolicospores whereas the other type, comparable to the extant Ceratosporella, possesses globular
to triangular body belongs to staurospores.
On the basis of fossil fungi, Kar & Saxena (1976) interpreted a warm and humid, tropical
climate during the Palaeocene (Matanomadh Formation) of Kutch, western India with the support
from spores and pollen of vascular plants. Ramanujam & Srisailam (1980) recorded a prevalence of
Palaeocirrenalia, the helicoid spore, in Neogene sediments of Kerala, India and interpreted
brackish to marine conditions by comparing them to the similar modern dematiaceous
hyphomycete, Cirrenalia, which is commonly found in such an environment. The presence of other
spores in the same strata, affiliated to Grallomyces, Sporidesmium, Spegazzinia, Amphisphaerella
and Isthmospora, also supports this interpretation of a tropical climate. This conclusion was
corroborated by pteridophytic spores and angiospermous pollen from the same strata and a tropical
climate was concluded (Ramanujam & Rao 1978, Ramanujam & Srisailam 1980). A warm and
humid environment has been interpreted by Kalgutkar & McIntyre (1991) in the Canadian Arctic
due to the presence of helicosporous fungal types. Studies on certain fungal assemblages,
sporomorphs and sporocarps, in coordination with micro- and megafossils of other groups, are used
to infer the palaeoenvironment (Dilcher 1973). Pirozynski (1976a, b) and Ramanujam (1982)
stressed the importance of coordinating the study of fossil fungi with their modern counterparts, in
order to realize the full potential of fossil fungal spores as indicators of ancient environment.
Ramanujam (1982) further urged that only those types, clearly related to modern taxa of which the
environmental requirements are known, are relevant in such studies. These assessments are based
on the assumption that the palaeoclimatic sensitivity of fossil taxa was similar to that of the
comparable modern counterparts. In this regard special stress was laid to explore the possibility of
relating fossil fungal spores with those of modern fungi so as to realize their full potential in
determining the ancient environment. However, only those types that could be related to the
modern forms with certainty should be taken into account for this specific purpose. Jarzen & Elsik

847
(1986) showed how the habitat and host preference of fungal spores recovered from recent deposits along the Luangwa River in Zambia, conceivably
can be used to deduce the environment of Neogene sediments with similar fossil fungal spores.

Table 3 Fossil fungal spore species assigned to extant fungal genera

Phylum Class Order Family Extant genera Fossil species

Ascomycota Caval.- Dothideomycetes Botryosphaeriales Botryosphaeriaceae Theiss. Diplodia Fr. D. rodei Mahab. [Current name: Diplodites
Sm. O.E. Erikss. & Winka C.L. Schoch & P. Syd. rodei (Mahab.) Kalgutkar et al.]; D. sahnii
Singhai [Current name: Diplodites sahnii
(Singhai) Kalgutkar et al.]
Capnodiales Mycosphaerellaceae Lindau Ramularia Sacc. R. oblongispora Casp.
Woron.
Pleosporales Luttr. Didymellaceae Gruyter Epicoccum Link. E. deccanense R. Srivast. et al.
ex M.E, Barr
Pleosporaceae Nitschke Alternaria Nees ex Fr. A. malayensis Trivedi & C.L. Verma [Current
name: Pluricellaesporites malayensis (Trivedi
& C.L. Verma) Kalgutkar & Janson.]
Torulaceae Corda Torula Pers. ex Fr. T. globulifera Casp., T. heteromorpha Casp.,
T. mengeana Casp. & R. Klebs
Sporidesmiales Sporidesmiaceae Fr. Sporidesmium Link ex S. henryense Dilcher
Crous Fr.
Lecanoromycetes Lecanorales Nannf. Sphaerophoraceae Fr. Sphaerophorus Pers. S. moniliformis Menge
O.E. Erikss. & Winka
Sordariomycetes O.E. Chaetosphaeriales Chaetosphaeriaceae Réblová Chaetosphaeria Tul. & C. elsikii M.J. Pound et al.
Erikss. & Winka Huhndorf. et al. et al. C. Tul.
Coronophorales Ceratostomataceae G. Melanospora Corda G. primigenius Casp.
Nannf. Winter
Diaporthales Botryosphaeriaceae Theiss. Botryodiplodia Sacc. B. mohgaoensis Barlinge & Paradkar [Current
Nannf. & P. Syd. name: Diplodites mohgaoensis (Barlinge &
Paradkar) Kalgutkar et al.]
Hypocreales Nannf. Bionectriaceae Samuels & Acremonium Link ex A. succineum Casp.
ex Korf & Lizon Rossman Fries.
Magnaporthales Magnaporthaceae P.F. Clasterosporium C. eocenicum Fritel & R. Vig.
Thongk. et al. Cannon Schwein.
Sordariales Chadef. Lasiosphaeriaceae Nannf. Zopfiella G. Winter Z. neogenica O’Keefe
ex D. Hawksw. &
O.E. Erikss.
Trichosphaeriales Trichosphaeriaceae G. Brachysporium Sacc. B. minutum Trivedi & C.L. Verma [Current
M.E, Barr Winter name: Pluricellaesporites minutus

848
Table 3 Continued.

Phylum Class Order Family Extant genera Fossil species

(Trivedi & C.L. Verma) ex Kalgutkar &
Janson.]
Incertae sedis Incertae sedis Diporothecaceae Mibey & Diporotheca C.C. D. doniana O’Keefe, D. gorda O’Keefe
D. Hawksw. Gordon & C.G. Shaw
Incertae sedis Incertae sedis Incertae sedis Desmidiospora Thaxt. D. marginiconvoluta Kalgutkar, D.
willoughbyi (W.H. Bradley) D.L.E. Glass, et
al.
Incertae sedis Incertae sedis Incertae sedis Potamomyces K.D. Hyde P. batii (Sancay) ex Nuñez Otaño et al.; P.
elsikii (Nandi & A. Sinha) Nuñez Otaño et
al.;
P. fournieri (Elsik & Jarzen) Nuñez Otaño et
al.; P. invaginatus (Elsik & Jarzen) Nuñez
Otaño et al.; P. magnus (Elsik & Jarzen)
Nuñez Otaño et al.; P. mulleri (Nandi & A.
Sinha) Nuñez Otaño et al.; P. pontidiensis
(Sancay) ex Nuñez Otaño et al.
Incertae sedis Incertae sedis Incertae sedis Rhexoampullifera P.M. R. stogieana M.J. Pound et al., R. sufflata
Kirk M.J. Pound et al.
Incertae sedis Incertae sedis Incertae sedis Tetracoccosporium T. eocenum Biradar & Mahab.
Szabó.
Basidiomycota Agaricomycetes Boletales E.-J. Sclerodermataceae Corda Scleroderma Pers. S. echinosporites Rouse
Whittaker ex R.T. Doweld Gilbert
Moore
Agaricostilbomycetes Agaricostilbales Chionosphaeraceae Oberw. & Stilbum Tode ex Fr. S. succini Casp.
R. Bauer et al. Oberw. & R. Bauer Bandoni
Ustilaginomycetes Urocystidales R. Glomosporiaceae Cif. Thecaphora Fingerh. T. mohgaoensis (Chitaley & Yawale) R.K.
Warm. Bauer & Oberw. Saxena, Wijayaw., D.Q. Dai, K.D. Hyde &
P.M. Kirk comb. nov.
[Synonyms: Sorosporium mohgaoense
Chitaley & Yawale; Papulosporonites
mohgaoensis (Chitaley & Yawale) Kalgutkar
& Janson.]
Ustilaginaceae Tul. & C. Tul. Ustilago (Pers.) Roussel. U. deccani Chitaley & Yawale [Current
name: Inapertisporites deccani (Chitaley &
Yawale) Kalgutkar & Janson.]
Chytridiomycota Chytridiomycetes M. Chytridiales Cohn Chytriomycetaceae Letcher Entophlyctis A. Fisch. E. willoughbyi W.H. Bradley [Current name:
Doweld Möbius Desmidiospora willoughbyi (W.H. Bradley)
D.L.E. Glass et al.]

849
Table 4 Fossil fungal spore genera and species and their probable relationship with extant taxa (*when the equivalent extant genus name is not
available, higher rank taxon name is given)

Phylum Class Order Family Modern genus Fossil genus Fossil species
Ascomycota Caval.- Dothideomycetes Capnodiales Cladosporiaceae Cladosporium Cladosporites Félix C. bipartitus Félix;
Sm. O.E. Erikss. & Woron. Chalm. & Archibald. Link C. fasciculatus E.W. Berry; C.
Winka oligocaenicus E.W. Berry
Cladosporiumsporinite C. cylindricus Debi Mukh.
s Debi Mukh.
Mycosphaerellaceae Helicomina L.S. Involutisporonites R.T. I. chowdhryi (K.P. Jain & R.K.
Lindau Olive Clarke Kar) Kalgutkar & Janson.
Piedraiaceae Viégas Trichosporium P. Trichosporites Félix T. conwentzii Félix
ex Cif. et al. Karst.
Microthyriales G. – – *Mariusia D. Pons & M. andegavensis D. Pons &
Arnaud Boureau Boureau
Pleosporales Dictyosporiaceae Dictyosporium Dictyosporites Félix D. dicotylophylli (Paradkar)
Luttr. ex M.E, Boonmee & K.D. Hyde Corda Kalgutkar & Janson.; D. dictyosus
Barr (Sal.-Cheb. & Locq.) Kalgutkar &
Janson.; D. firbasii (Hammen)
Kalgutkar & Janson.; D.
garciabarrigae (Hammen)
Kalgutkar & Janson.; D.
globimuriformis Kalgutkar; D.
hyalinus (R.T. Lange & P.H. Sm.
1971) Kalgutkar & Janson.; D.
loculatus Félix; D. morularis (Sal.-
Cheb. & Locq.) Kalgutkar &
Janson.; D. moruloides (Sal.-Cheb.
& Locq.) Kalgutkar & Janson.; D.
ovalis (Sheffy & Dilcher)
Kalgutkar & Janson. D. ovoideus
Sal.-Cheb. & Locq.; D.
paradkariae Kalgutkar & Janson.;
D. symmetricus (V.S. Ediger)
Kalgutkar & Janson.; D.
tirumalacharii (Ramanujam &
Ramachar) Kalgutkar & Janson.;
D. tristratosus (Sheffy & Dilcher)
Kalgutkar & Janson.
Dictyosporiuminites D. intermedius Debi Mukh.
Debi Mukh.

850
Table 4 Continued.

Phylum Class Order Family Modern genus Fossil genus Fossil species
Pleosporaceae Curvularia Palaeocurvularia D. variabilis Dörfelt & A.R.
Boedijn Dörfelt & A.R. Schmidt
Schmidt
Pleosporaceae Alternaria Nees Polycellaesporonites P. acuminatus (Rouse & Mustard)
Nitschke Anil Chandra et al. Kalgutkar & Janson.;
P. alternariatus (Kalgutkar &
Sigler) Kalgutkar & Janson.; P.
bellus Anil Chandra et al.; P.
clavellatus (Z.C. Song & G.X. Li in
Z.C. Song et al.) Kalgutkar &
Janson.; P. psilatus A. Gupta;
P. saxenae A. Gupta;
P. sirmaurensis A. Gupta.
Tetraplosphaeriaceae Tetraploa Berk. & Frasnacritetrus Taug. F. conatus R.K. Saxena & S.
Kaz. Tanaka & Hiray. Broome Sarkar; F. indicus R.K. Saxena & S.
Khare; F. jamtahensis A. Gupta; F.
josettae Taug.; F. masolensis R.K.
Saxena & S.K.M. Tripathi; F.
siwalikus R.K. Saxena et al.;
F. taugourdeaui R.K. Saxena & S.
Sarkar
Incertae sedis Periconia Tode Palaeopericonia C. G. P. fritzschei C.G. Ibáñez &
Ibáñez & Zamuner Zamuner
Tubeufiales Tubeufiaceae M.E, Slimacomyces Paleoslimacomyces P. canadensis Kalgutkar & Sigler;
Boonmee & K.D. Barr Minter Kalgutkar & Sigler P. minutus (Rouse & Mustard)
Hyde Kalgutkar & Janson.; P. wilcoxii
(Elsik.) Kalgutkar & Janson.
Helicoon Morgan Helicoonites Kalgutkar H. goosii Kalgutkar & Sigler
& Sigler
– *Retihelicosporonites R. elsikii Ramanujam & K.P. Rao
Ramanujam & K.P.
Rao
Eurotiomycetes O.E. Chaetothyriales Herpotrichiellaceae Phialophora Phialophoronites Debi P. magnus Debi Mukh.
Erikss. & Winka M.E, Barr Munk Medlar Mukh.
Lecanoromycetes Incertae sedis Xylographaceae Tuck. Haplographa (Fr.) Haplographites Félix H. cateniger Félix;
O.E. Erikss. & E.W. Mason H. xylophagus Félix
Winka

851
Table 4 Continued.

Phylum Class Order Family Modern genus Fossil genus Fossil species
Leotiomycetes O.E. Helotiales Nannf. Discinellaceae Tetrachaetum Tribolites W.H. T. triangulatus (Peppers) Kalgutkar
Erikss. & Winka ex Korf & Lizoň Ekanayaka & K.D. Ingold/Lemonnier Bradley & Janson.; T. tetrastonyx W.H.
Hyde a De Wild. Bradley
Pezizomycetes Pezizales J. Schrot. Pezizaceae Dumort. – *Pezizasporites T.C. P. taiwanensis T.C. Huang
Huang
Sarcoscyphaceae Le Cookeina Kuntze Fusiformisporites F. acutus P. Kumar; F, Barmerensis
Gal ex Eckblad Rouse R.K. Saxena & S.K.M. Tripathi; F.
elongatus Ramanujam & K.P. Rao;
F. striaoctoformis Mart.-Hern. &
Tom.-Ort.
Saccharomycetes G. Saccharomycetales Dipodascaceae Engl. Geotrichum Link Geotrichites Stubblef. G. glaesarius Stubblef. et al.
Winter Luerss. & E. Gilg et al.
Sordariomycetes Amphisphaeriales Amphisphaeriaceae G. Ceratosporium Ceratohirudispora R. C. miocenica R. Kar et al.
O.E. Erikss. & D. Hawksw. & Winter Schwein. Kar et al.
Winka O.E. Erikss.
Amphisphaerella Palaeoamphisphaerell P. keralensis Ramanujam &
(Sacc.) Kirschst. a Ramanujam & Srisailam; P. pirozynskii
Srisailam Ramanujam & Srisailam; P.
tankensis (G. Norris) Kalgutkar &
Janson.
Hypocreales Incertae sedis Gliomastix Guég. Parapotamomyces P. maydiformis O’Keefe
Lindau O’Keefe
Microascales Halosphaeriaceae E. Cirrenalia Meyers Palaeocirrenalia P. elegans Ramanujam &
Luttr. ex Benny & Müll. & Arx ex & R.T. Moore Ramanujam & Srisailam;
R.K. Benj. Kohlm. Srisailam P. oligoseptata Ramanujam &
Srisailam
Sordariales Chad. Sordariaceae G. Neurospora Shear Diploneurospora K.P. D. tewarii K.P. Jain & R.C. Gupta
ex D. Hawksw. & Winter & B.O. Dodge Jain & R.C. Gupta
O.E. Erikss.
Incertae sedis Edmundmasonia Edmundmasonaesporit E. globulatus Debi Mukh.
Subram. es Debi Mukh.
Trichosphaeriales Trichosphaeriaceae G. Brachysporium Brachysporisporites B. antarcticus Z.C. Song & Liu
M.E, Barr Winter Sacc. R.T. Lange & P.H. Sm. Cao; B. atratus Kalgutkar; B.
bullatus Kalgutkar; B. catinus
(Elsik & Janson.) Kalgutkar &
Janson.; B. cotalis (Elsik & Janson.)
Norris; B. fustitudinus G. Norris;

852
Table 4 Continued.

Phylum Class Order Family Modern genus Fossil genus Fossil species
B. grandus Z.C. Song & Liu Cao;
B. inuvikensis M.G. Parsons & G.
Norris; B. lageniformis Z.C. Song;
B. longovatus Z.C. Song & Liu
Cao; B. magnus B. Samant in R.K.
Saxena; B. maximus (P. Ke & Z.Y.
Shi) Kalgutkar & Janson.; B.
opimus (Elsik & Janson.) G. Norris;
B. ovoidus Z.C. Song & Liu Cao; B.
pyriformis R.T. Lange & P.H. Sm.;
B. tenuis P. Kumar; B. thraceous
(V.S. Ediger) Kalgutkar & Janson.
Xylariales Nannf. Xylariaceae Tul. & Hypoxylon Hypoxylonites Elsik and H. africanus Sal.-Cheb. ex
C. Tul. Adans./Xylaria Hill Xylariasporites Debi Kalgutkar & Janson.; H.
ex Schrank Mukh. armentroutii Elsik; H. asymetricus
(Sal.-Cheb. & Locq.) Elsik; H. ater
(P. Kumar) R.K. Saxena; H.
brazosensis Elsik; H. bhubanensis
Nandi & Subhra Banerjee; H.
chaiffetzii Elsik; H. chuittensis
Elsik; H. claibornensis Elsik; H.
curvatus (Ramanujam & K.P. Rao)
Elsik; H. edigeri Elsik; H.
elongatioides Elsik; H. ellipsoideus
Sal.-Cheb. & Locq. ex Kalgutkar &
Janson; H. eopleistocenicus Elsik;
H. felixii Elsik; H. foldexinus Elsik;
H. fusiformis Elsik; H. foyelensis
Bianchin. et al.; H. gulfensis Elsik;
H. horowitzii Elsik; H. kumarii
Kalgutkar & Janson.; H. lammonsii
Elsik; H. lineatus Elsik; H. magnus
Elsik; H. megaexinus Elsik; H.
minutus Elsik; H. miocenicus Elsik;
H. neogenicus Nandi & Subhra
Banerjee; H. ovaloides Elsik; H.
pirozynskii Elsik; H.
pirozynskioides Elsik;

853
Table 4 Continued.

Phylum Class Order Family Modern genus Fossil genus Fossil species
H. pleistocenicus Elsik; H.
pulvinatus (Sheffy & Dilcher)
Elsik; H. ramanujamii Elsik; H.
subuliformis Elsik; H. sulekii Elsik;
H. truncatus Elsik; H. xylarioides
Sal.-Cheb. & Locq. ex Kalgutkar &
Janson.
Spirotremesporites S. simplex Dueñas, S. duenasii
Dueñas Elsik, S. ecuatorialis Dueñas; S.
tertiarus Nandi et al.; S. clinatus
Elsik; S. neogenicus Elsik
Incertae sedis Apiosporaceae K.D. Spegazzinia Petr. & Spegazzinites Félix S. cruciformis Félix; S. indicus
Hyde et al. Syd. Ramanujam & Srisailam; S.
tetradus. (Rouse) Kalgutkar &
Janson
Incertae sedis Incertae sedis Nigrospora Zimm. Nigrosporites Devi N. neyveliensis Debi Mukh.
Mukh.
Exesisporites Elsik E. neogenicus Elsik; E. psilatus
R.K. Saxena; E. verrucatus P.
Kumar; E. annulatus Kalgutkar
Incertae sedis Incertae sedis Incertae sedis Grallomyces F. Alleppeysporonites A. scabratus Ramanujam & K.P.
Stevens Ramanujam & K.P. Rao
Rao
Incertae sedis Incertae sedis Incertae sedis Xylohypha (Fr.) Xylohyphites Kalgutkar X. verrucosus Kalgutkar & Sigler
E.W. Mason & Sigler
Incertae sedis Incertae sedis Incertae sedis Mycoenterolobium Kutchiathyrites R.K. K. canadensis Kalgutkar & Janson.;
Goos Kar K. mehrotrae R.K. Saxena &
S.K.M. Tripathi; K. perfectus (R.
Kar et al.) R.K. Saxena & S.K.M.
Tripathi; K. eccentricus R.K. Kar;
K. palmatus (P. Ke & Z.Y. Shi)
Kalgutkar & Janson.
Incertae sedis Incertae sedis Incertae sedis Dwayabeeja Dwayabeejaesporonite D. undulatus Debi Mukh.
Subram. s Debi Mukh.
Incertae sedis Incertae sedis Incertae sedis Ampulliferina B. Ampulliferinites A. axelheibergi Kalgutkar & Sigler
Sutton Kalgutkar & Sigler

854
Table 4 Continued.

Phylum Class Order Family Modern genus Fossil genus Fossil species
Basidiomycota Exobasidiomycetes Exobasidiales Graphiolaceae Clem. Graphiola Poit. Graphiolites Fritel G. sabaleos Fritel
Whittaker ex R.T. Begerow et al. Henn. & Shear
Moore
Pucciniomycetes R. Pucciniales Caruel Pucciniaceae Puccinia Pers. Pucciniasporonites P. arcotensis Ramanujam &
Bauer et al. Chevall. Ramanujam & Ramachar
Ramachar
Basidiomycota – – – – *Basidiosporites Elsik B. fournieri Elsik
(unclassified)
*Diporisporites D. ellipsoides (Sal.-Cheb. & Locq.)
Hammen Kalgutkar & Janson.; D.
naviculoides (Sal.-Cheb. & Locq.)
Kalgutkar & Janson.
Blastocladiomycota Blastocladiomycetes Blastocladiales Coelomomycetaceae Coelomomyces Striadiporites C.P. S. asper (P. Ke & Z.Y. Shi)
T.Y. James Doweld H.E. Petersen Couch Keilin Varma & Rawat Kalgutkar & Janson.; S. bistriatus
(P. Ke & Z.Y. Shi) G. Norris; S.
boletelloides Sal.-Cheb. & Locq.; S.
californicus Elsik & Janson.; S.
dolium (P. Ke & Z.Y. Shi)
Kalgutkar & Janson.; S. inflexus (P.
Ke & Z.Y. Shi) G. Norris; S.
irregularis Kalgutkar; S. minor
(Z.C. Song & Z.H. Sun) Z.C. Song
et al.; S. multistriatus (P. Ke & Z.Y.
Shi) G. Norris; S. reticulatus C.P.
Varma & Rawat; S. retistriatus (P.
Ke & Z.Y. Shi) Kalgutkar &
Janson.; S. sanctaebarbarae Elsik &
Janson.; S. spiralis Kalgutkar &
Janson.; S. taiwanensis T. C. Huang
Incertae sedis – – – – Eoglobella W.H. E. longipes W.H. Bradley
Bradley

Kalgutkar & McIntyre (1991) described two helicosporous fungal types (now known as Helicoonites and Helicosporiates Kalgutkar & Sigler,
1995) from the Eocene Eureka Sound Formation in the Canadian Arctic, that are morphologically similar to the extant warm-climate,
subaqueousgeneric groups Helicoon-Helicodendron and Helicosporium, and used their presence to postulate pools of open water in a warm, humid
palaeoenvironment of the region.

855
 The importance of fossil fungal distribution and their relative abundance in palaeoecological
studies was recognized also by Elsik (1969). He interpreted the relative abundance of Exesiporites
during the Pliocene of northern Gulf of Mexico, to indicate a warmer period, and the fluctuations
before and afterward, therefore, as expressions of a cyclic climate. With reference to the ubiquitous
Hypoxylon, Elsik (op. cit.) wrote that “The genus Hypoxylon, and even some of its species, are
distributed world-wide. Some living species are restricted to the North Temperate zone, or occur
outside that area only as varieties. Some species are restricted to local geographic regions. The
tropics apparently support more varieties. These conclusions regarding Hypoxylon (Miller 1961)
will have validity in the fossil record, once it is demonstrated that species diversity is reflected in
spore morphology.”

Biostratigraphic Implications
It is an established fact that fossils of all kinds, especially microfossils, are indispensable in
biostratigrapic studies, e.g. biostratigraphic zonation and correlation, of sedimentary stratigraphic
successions. These may either be subsurface stratigraphic sequences obtained from the boreholes or
exposed sedimentary rocks obtained from variety of locations, e.g. road and stream cuttings,
escarpments, sea cliffs and mine face sections. The main types of microfossils found in slides
prepared for palynological studies are bryophytic and pteridophytic spores, gymnospermous and
angiospermous pollen, dinoflagellate cysts and other micro-algae and fungal remains. In
biostratigraphic studies, it is better to consider and apply data derived from all kind of microfossils.
The inferences derived from multi-disciplinary cumulative data, i.e. a synergistic approach, are
always more sound and reliable.
Fossil fungi are found in the form of spores, mycelia, sporophores and symbiotic
associations, and are commonly observed in macerated residues prepared for palynological studies.
Fossil records indicate that microthyriaceous fungi occur in the Cenozoic sedimentary rocks all
over the world. Saxena & Tripathi (2011) made an attempt to summarize the stratigraphic
distribution of different fossil sporocarp genera recorded from the Indian Tertiary sequences
(Saxena & Tripathi 2011: 198, fig. 392). This shows that the taxa assigned to Callimothallus
Dilcher 1965 and Cucurbitariaceites R.K. Kar et al. 1972 are long ranging and are recorded from
Palaeocene to Pliocene sediments. Different species of Phragmothyrites W.N. Edwards mark their
presence in Palaeocene to Miocene and Microthyriacites Cookson in Eocene to Miocene. Forms
restricted to Miocene sequences only are: Asterothyrites Cookson, Euthythyrites Cookson,
Parmathyrites K.P. Jain & R.C. Gupta, Plochmopeltinites Cookson, Ratnagiriathyrites R.K.
Saxena & N.K. Misra, Trichopeltinites Cookson and Trichothyrites Rosend.
Although most of the fungal spore genera are long ranging and do not bear much stratigraphic
significance, some are morphologically distinct and have a restricted range in geological time.
Applicability of fungal spores in biostratigraphy has, therefore, increased with the record of such
characteristic spores with limited stratigraphic ranges (Kalgutkar & Jansonius 2000). Because of
their wide geographic distributions, fungal spores commonly occur even in those samples in which
other biostratigraphic markers are not present. The differentiation of dispersed ascospores and
conidia becomes easier with increasing complexity of their morphology. Coiled helicospores and
stellate staurospores are more distinctive than didymospores or amerospores. Some fungal spores,
such as species of Ctenosporites Elsik & Janson., Foveodiporites C.P. Varma & Rawat,
Dictyosporites Félix, Frasnacritetrus Taug., Fusiformisporites Rouse, Palaeoamphisphaerella
Ramanujam & Srisailam, Pesavis Elsik & Janson., Striadiporites C.P. Varma & Rawat, and
helicoid spores like Involutisporonites R.T. Clarke, Helicosporiates Kalgutkar & Sigler,
Helicoonites Kalgutkar & Sigler, Palaeocirrenalia Ramanujam & Srisailam, Paleoslimacomyces
Kalgutkar & Sigler are so discrete that they are readily identified, and generally accepted as index
fossils. However, detailed information on their stratigraphic ranges is a prerequisite for their
application in biostratigraphic studies. (Kalgutkar & Jansonius 2000).
Graham (1962) was amongst the pioneers to suggest the possibility of using fungal spores for
supplementing age determinations in palynological studies. According to Elsik (1970), although a

856
variety of fungal spores have been recorded from Mesozoic strata worldwide, their morphological
complexity and frequency increase in the Cenozoic. He noted that Fusiformisporites Rouse and
similar longitudinally ribbed forms appear to be restricted to the Cenozoic. Elsik (1970) further
observed that fossil fungal spores described as Exesisporites which resemble the extant Hypoxylon
type are more frequently recorded in Neogene sediments. Kalgutkar (1993) showed how some
fungal species with records restricted to the Palaeocene through Eocene in other Palaeogene strata
of Arctic Canada, were useful in arriving at an age of the Bonnet Plume Formation.
According to Kalgutkar & Jansonius (2000), the highly distinct Pesavis tagluensis Elsik &
Janson. has been important in stratigraphic studies (Jansonius 1976, Staplin, 1976, Ioannides &
McIntyre 1980, Norris 1982, Young & McNeil 1984). Elsik & Jansonius (1974) and Lange (1978a,
b) pointed out the importance of forms like P. tagluensis Elsik & Janson. and Ctenosporites
eskerensis Elsik & Janson. in dating sediments in which other palynological fossils are rare.
Jansonius (1976) reported the characteristic association of P. tagluensis Elsik & Janson. with
Striadiporites C.P. Varma & Rawat spores in the Beaufort region, Mackenzie Delta region, Canada.
Their highest occurrences were used to define the top of the Pesavis zone. The lower part of the
zone exhibits great numbers and variety of septate, diporate ascospores, and the upper part an
abundance of one-celled spores. Norris (1986) assigned a middle Eocene age to the Pesavis zone
within the lower part of the Richards Formation, Mackenzie Delta region, Canada. Kalgutkar &
Sweet (1988) documented the first Maastrichtian occurrence of Pesavis in north-western Canada.
They further developed the stratigraphic usefulness of Pesavis by documentating a phylogenetic
lineage starting in the Maastrichtian with P. parvus Kalgutkar & Sweet and extending into the
Eocene with P. tagluensis Elsik & Janson. Transitory stages, with changes in the morphology from
P. parvus through P. tagluensis, were found in the Palaeocene. White (1990) used the presence of
Ctenosporites eskerensis and Pesavis tagluensis, in combination with Tilia spp., to determine the
age of an unnamed (Cretaceous-Tertiary) sedimentary unit in the Union Port Louis well (west coast
of Graham Island, Queen Charlotte Islands, B.C., Canada), as well as of a float from the nearby
Mud Bay Creek, which he estimated to be between early Eocene and early Oligocene in age. There
is indication that the top of the range of P. tagluensis is somewhat younger in the more southern
British Columbia sediments than it is in the Beaufort Sea region.
A similar observation of fungal spore populations in and above the Pesavis zone was
presented by Staplin (1976). Fournier & Elsik (1984) revealed the existence of hundreds of well
preserved species of fungal spores from DSDP Site 493, Leg 66, and showed the stratigraphic value
of some characteristic spores in early Miocene to Recent assemblages.
Kar (1979) recognized an assemblage zone, viz. Aplanosporites robustus Cenozone in the
Maniyara Fort Formation (Oligocene) of Kutch, Gujarat, India on the basis of abundant occurrence
of Aplanosporites robustus [Current name: Palaeomycites robustus (Kar) Kalgutkar & Janson.
2000]. Kar (1990) also recognized this assemblage zone from the borehole sequences of Tripura,
North-eastern India.
Ramanujam (1982) opined that maximum diversity in the morphology of fungal spores was
attained by late Cretaceous and Early Tertiary. While evaluating the stratigraphic potential of
fungal remains in Indian stratigraphic sequences, he further observed that spores with relatively
simpler morphology (aporate, aseptate and essentially smooth) were recorded from the early
Mesozoic and earlier strata. Ornamented spores appeared mostly in late Cretaceous. On the
discoveries of diverse Tertiary horizons of India, Ramanujam (1982), noted that ornately sculptured
walls were encountered consistently in the Neogene sediments, and that Neogene assemblages
appeared to be different from those present in the Palaeogene.
Stratigraphic distribution of selected, commonly occurring, fossil fungal spore taxa (arranged
age-wise, oldest at the bottom and youngest at the top) and their probable relationship with modern
counterparts and location of occurrence is summarized in Table 5.

857
Table 5 Stratigraphic distribution of selected, commonly occurring, fossil fungal spore taxa
(arranged age-wise, oldest at the bottom and youngest at the top) and their probable relationship
with modern counterparts and location of occurrence

Age Fossil fungal spore taxa, their affinity with modern taxa and location of occurrence
Quaternary Hypoxylonites pleistocenicus Elsik, Texas, U.S.A., affinity with Xylariaceae, Xylariales.
Polycellaesporonites bellus Anil Chandra et al., Arabian Sea, affinity with Alternaria
(Pleosporaceae, Pleosporales).
Spirotremesporites simplex Dueñas, S. duenasii Elsik, S. ecuatorialis Dueñas, Colombia, South
America, affinity with Xylariaceae, Xylariales.
Tertiary Spegazzinites cruciformis Félix, Germany, affinity with Spegazzinia (Apiosporaceae,
(undifferentiated) Sordariomycetidae).
Spirotremesporites tertiarus Nandi et al., Mizoram, India, affinity with Xylariaceae, Xylariales.
Neogene Hypoxylonites chaiffetzii Elsik, H. foldexinus Elsik, H. fusiformis Elsik, H. gulfensis Elsik, H.
(undifferentiated) lammonsii Elsik, H. lineatus Elsik, H. pirozynskii Elsik, H. ramanujamii Elsik,
H. truncatus Elsik, The Gulf Coast, U.S.A., H. chuittensis Elsik, Southern Alaska, U.S.A., H.
horowitzii Elsik, Israel, H. bhubanensis Nandi & Subhra Banerjee, H. neogenicus Nandi &
Subhra Banerjee, Mizoram, India, affinity with Xylariaceae, Xylariales.
Spirotremesporites clinatus Elsik, S. neogenicus Elsik, The Gulf Coast, U.S.A., affinity with
Xylariaceae, Xylariales.
Miocene Hirudinaria (Pezizomycotina).
Cladosporiumsporinites cylindricus Debi Mukh., Tamil Nadu, India, affinity with Cladosporium
(Cladosporiaceae, Capnodiales).
Dictyosporiuminites intermedius Debi Mukh., Tamil Nadu, India, affinity with
(Dictyosporiaceae, Pleosporales).
Diploneurospora tewarii K.P. Jain & R.K. Gupta, Western Ghats, South India, affinity with
Neurospora (Sordoriaceae, Sordariales).
Diporotheca doniana O’Keefe, D. gorda O’Keefe; Tumbes Province, Peru; affinity with
Diporthaceae, Pezizomycotina.
Dwayabeejaesporonites undulatus Debi Mukh., Tamil Nadu, India, affinity with genus
Dwayabeeja (Pezizomycotina).
Edmundmasonaesporites globulatus Debi Mukh., Tamil Nadu, India, affinity with genus
Edmundmasonia (Sordariales, Sordariomycetidae).
Exesisporites neogenicus Elsik, Northern Gulf of Mexico, E. psilatus R.K. Saxena, Maharashtra,
India, E. verrucatus Kumar, Kerala, India, probable affinity with the extant Nigrospora Zimm.
(Sordariomycetes).
Fusiformisporites acutus P. Kumar, F. elongatus Ramanujam & K.P. Rao, Kerala, India, affinity
with Cookeina (Sarcoscyphaceae, Pezizales).
Geotrichites glaesarius Stubblef. et al., Dominican Republic; Late Oligocene or Early Miocene,
affinity with Geotrichum (Dipodascaceae, Saccharomycetales) (Stubblefield et al. 1985).
Haplographites cateniger Félix, H. xylophagus Félix, near Baku, Azerbaijan, affinity with
Xylographaceae, Ostropomycetidae.
Hypoxylonites eopleistocenicus Elsik, H. megaexinus Elsik, H. minutus Elsik,
H. miocenicus Elsik, H. sulekii Elsik, The Gulf Coast, U.S.A., H. felixii Elsik, Bristol Bay,
Alaska, H. magnus Elsik, Venezuela, H. ater (P. Kumar) R.K. Saxena, H. curvatus (Ramanujam
& K.P. Rao) Elsik, H. kumarii Kalgutkar & Janson., Kerala, India,
H. subuliformis Elsik, H. sulekii Elsik, Venezuela, H. xylarioides Sal.-Cheb. & Locq. ex
Kalgutkar & Janson., Cameroon, affinity with Xylariaceae, Xylariales.
Kutchiathyrites mehrotrae R.K. Saxena & S.K.M. Tripathi, Meghalaya and Assam, India,
K. perfectus (R. Kar et al.) R.K. Saxena & S.K.M. Tripathi, Mizoram, India, affinity with
Mycoenterolobium Goos (Pezizomycotina).
Nigrosporites neyveliensis Debi Mukh., Tamil Nadu, India, affinity with Nigrospora
(Sordariomycetes, Pezizomycotina).
Palaeoamphisphaerella keralensis Ramanujam & Srisailam, P. pirozynskii Ramanujam &
Srisailam, Kerala, India, affinity with Amphisphaerella (Amphisphaeriaceae, Amphisphaeriales).
Phialophoronites magnus Debi Mukh., Tamil Nadu, India, affinity with genus Phialophora
(Herpotrichiellaceae, Chaetothyriales). Potamomyces batii (Sancay) ex Nuñez Otaño et al., P.
pontidiensis (Sancay) ex Nuñez Otaño et al., Turkey, P. fournieri (Elsik & Jarzen) Nuñez Otaño
et al. P. invaginatus (Elsik & Jarzen) Nuñez Otaño et al. P. magnus (Elsik & Jarzen) Nuñez
Otaño et al., Colombia, South America, P. elsikii (Nandi & A. Sinha) Nuñez Otaño et al., P.
mulleri (Nandi & A. Sinha) Nuñez Otaño et al. Mizoram, India, affinity with Pezizomycotina.
Pucciniasporonites arcotensis Ramanujam & Ramachar, Tamil Nadu, India, affinity with

858
Table 5 Continued.

Age Fossil fungal spore taxa, their affinity with modern taxa and location of occurrence
Puccinia (Pucciniales, Pucciniaceae).
Retihelicosporonites elsikii Ramanujam & K.P. Rao, Kerala, India, helical spores (conidia) are
found in various hyphomycetes, viz. Helicoma, Helicomina, Helicoon, Helicodendron,
Xenosporella and Hiospira.
Spegazzinites indicus Ramanujam & Srisailam, Kerala, India, affinity with Spegazzinia
(Apiosporaceae, Sordariomycetidae).
Striadiporites californicus Elsik & Janson., California, U.S.A., S. reticulatus C.P. Varma &
Rawat, West Bengal and Assam, India, S. taiwanensis T. C. Huang, Taiwan, affinity with
Coelomomyces (Coelomomycetaceae, Blastocladiales).
Zopfiella neogenica O’Keefe, Tumbes Province, Peru, Lasiosphaeriaceae, Sordariales.
Gonatobotrys simplex Corda, Baltic area, Poland, Ceratostomataceae, Melanosporales.
Hypoxylonites ellipsoideus Sal.-Cheb. & Locq. ex Kalgutkar & Janson., Cameroon, affinity with
Xylariaceae, Xylariales.
Kutchiathyrites eccentricus R.K. Kar, Kutch, India, K. palmatus (P. Ke & Z.Y. Shi) Kalgutkar &
Janson., Coastal region of Bohai, China (affinity with hyphomycetous Mycocenterolobium
platysporum).
Polycellaesporonites clavellatus (Z.C. Song & G.X. Li in Z.C. Song et al.) Kalgutkar & Janson.,
Shandong Province, China,affinity with Alternaria (Pleosporaceae, Pleosporales).
Ramularia pusilla Unger, Baltic area, Poland, Mycosphaerellaceae, Capnodiales.
Sphaerophorus coralloides Pers., Baltic area, Poland, Sphaerophoraceae, Lecanorales.
Stilbum vulgare Tode, Baltic area, Poland, Chionosphaeraceae, Agaricostilbales.
Torula globulifera Casp., T. heteromorpha Casp., T. mengeana Casp. & R. Klebs in Casp.,
Torulaceae, Pleosporales.
Eocene Ampulliferinites axelheibergi Kalgutkar & Sigler, North-western Canada, affinity with
Ampulliferina (Pezizomycotina).
Cladosporites bipartitus Félix, near Baku, Azerbaijan, C. fasciculatus E.W. Berry, Texas,
U.S.A., C. oligocaenicus E.W. Berry, Mississippi, U.S.A., affinity with Trichothecium and
Cladosporium (Hypocreales, Hypocreomycetidae).
Clasterosporium eocenicum Fritel & R. Vig., France, Magnaporthaceae, Magnaporthales.
Desmidiospora marginiconvoluta Kalgutkar, Northwest Territories, Canada, D. willoughbyi
(W.H. Bradley) D.L.E. Glass, et al., Texas, U.S.A., Pezizomycotina.
Dictyosporites loculatus Félix, near Baku, Azerbaijan, Félix, D. ellipsoides (Sal.-Cheb. & Locq.)
Kalgutkar & Janson., D. naviculoides (Sal.-Cheb. & Locq.) Kalgutkar & Janson.), Cameroon,
different species have affinity with Dictyosporium, Stemphylium, Septosporium and Alternaria,
all belonging to dematiaceous Hyphomycetes, and the ascospores of Pleospora.
Exesisporites annulatus Kalgutkar, Yukon Territory, Canada (Glass et al. 1986 cited possible
affinity with the extant fungus Nigrospora Zimm. (Sordariomycetes, Pezizomycotina).
Graphiolites sabaleos Fritel, France, fossil spores of Graphiola (Graphiolaceae, Exobasidiales).
Hypoxylonites africanus Sal.-Cheb. ex Kalgutkar & Janson., H. asymetricus (Sal.-Cheb. & Locq.)
Elsik, Cameroon, H. armentroutii Elsik,. Washington, U.S.A., H. brazosensis Elsik, Texas,
U.S.A., H. claibornensis Elsik, H, elongatioides Elsik, H. pirozynskioides Elsik, H. pulvinatus
(Sheffy & Dilcher) Elsik, Tennessee, U.S.A., H. ramanujamii Elsik, Green River section, U.S.A.,
H. edigeri Elsik, Turkey, H. ovaloides Elsik, Venezuela, affinity with Hypoxylon, Xylaria
(Xylariaceae, Xylariales).
Palaeoamphisphaerella tankensis (G. Norris) Kalgutkar & Janson., North West Territory,
Canada, affinity with Amphisphaerella (Amphisphaeriaceae, Amphisphaeriales).
Pluricellaesporites infacetus (Kalgutkar) Kalgutkar & Janson., Northwest Territories, Canada, P.
malayensis Malaya, affinity with Brachysporium Sacc. (Trichosphaeriaceae, Trichosphaeriales).
Polycellaesporonites acuminatus (Rouse & Mustard) Kalgutkar & Janson. P. alternariatus
(Kalgutkar & Sigler) Kalgutkar & Janson. Northwest Territories, Canada, and U.S.A., P. psilatus
A. Gupta, P. saxenae A. Gupta, P. sirmaurensis A. Gupta, Himachal Pradesh, India, affinity with
Alternaria (Pleosporaceae, Pleosporales).
Spegazzinites tetradus (Rouse) Kalgutkar & Janson., British Columbia, Canada, affinity with
Spegazzinia (Apiosporaceae, Sordariomycetidae).
Sporidesmium Link, Western Tennessee, U.S.A., S. henryense Dilcher, Pleosporomycetidae,
Dothideomycetes.
Tribolites tetrastonyx W.H. Bradley, Colorado, U.S.A., affinity with Tetrachaetum and
Lemonniera (Bradley 1964).

859
Table 5 Continued.

Age Fossil fungal spore taxa, their affinity with modern taxa and location of occurrence
Palaeocene Basidiosporites fournieri Elsik, Texas, U.S.A., B. anceps (G. Norris) Kalgutkar & Janson.,
Mackenzie River delta, Canada, B. belluloides (Z.C. Song) Kalgutkar & Janson., Qinghai
Province, China, Basidiomycota.
Biporisporites rotundus P. Ke & Z.Y. Shi, Coastal region of Bohai, China.
Diporisporites hammenii Elsik, Milam County, Texas, U.S.A.
Desmidiospora marginiconvoluta Kalgutkar, Northwest Territories, Canada, Pezizomycotina.
Fusiformisporites barmerensis R.K. Saxena & S.K.M. Tripathi, Rajasthan, India, affinity with
Cookeina (Sarcoscyphaceae, Pezizales).
Hypoxylonites foyelensis Bianchin. et al., Argentina, affinity with Hypoxylon (Xylariaceae,
Xylariales).
Kutchiathyrites canadensis Kalgutkar & Janson., affinity with Mycoenterolobium platysporum
(Pezizomycotina).
Striadiporites irregularis Kalgutkar, Yukon Territory, Canada, hyphomycetous fungus.
Paleoslimacomyces canadensis Kalgutkar & Sigler, P. minutus (Rouse & Mustard) Kalgutkar &
Janson., P. wilcoxii (Elsik.) Kalgutkar & Janson., Texas, U.S.A., Tubeufiaceae, Tubeufiales.
Cretaceous Anatolinites antarcticus (Z.C. Song & Liu Cao) Kalgutkar & Janson., A. megaporus (Z.C. Song
(mostly Late & Liu Cao) Janson. et al., A. tenuis (Z.C. Song & Liu Cao) Kalgutkar & Janson., King George
Cretaceous) Island, Antarctica.
Brachysporisporites antarcticus Z.C. Song & Liu Cao, B. grandus Z.C. Song & Liu Cao, B.
longovatus Z.C. Song & Liu Cao, B. ovoidus Z.C. Song & Liu Cao, King George Island,
Antarctica.
Ctenosporites sherwoodiae Kalgutkar & Janson., Fremount County, Colorado, U.S.A.
Dicellaesporites antarcticus Z.C. Song & Liu Cao, D. oblongatus Z.C. Song & Liu Cao, King
George Island, Antarctica.
Dictyosporites dicotylophylli (Paradkar) Kalgutkar & Janson., D. paradkariae Kalgutkar &
Janson., Madhya Pradesh, India.
Diporicellaesporites antarcticus Z.C. Song & Liu Cao, King George Island, Antarctica.
Dyadosporites antarcticus Kalgutkar & Janson., D. obscurus Z.C. Song & Liu Cao, King George
Island, Antarctica, D. ellipsus R.T. Clarke, Fremont County, Colorado, U.S.A.
Fractisporonites canalis R.T. Clarke, F. moniliformis R.T. Clarke, Fremont County, Colorado,
U.S.A.
Fusiformisporites striaoctoformis Mart.-Hern. & Tom.-Ort., Coahuila State, Mexico,
Maastrichtian; affinity with Cookeina (Sarcoscyphaceae, Pezizales).
Helicominites salvinites Barlinge & Paradkar, Madhya Pradesh, India.
Heterocystinella bulbosa Cookson & Eisenack, Eucla basin, Western Australia.
Inapertisporites clarkei Kalgutkar & Janson., Fremont County, Colorado, U.S.A., I. deccanii
(Chitaley & Yawale) Kalgutkar & Janson., Madhya Pradesh, India, I. elongatus Rouse, I.
laevigatus Rouse, I. pseudoreticulatus Rouse, I. punctatus Rouse, British Columbia, Canada, I.
solidus Z.C. Song & Liu Cao, King George Island, Antarctica.
Involutisporonites crassus Z.C. Song & Liu Cao, Antarctica, I. foraminus R.T. Clarke, Fremont
County, Colorado, U.S.A.
Lacrimasporonites levis R.T. Clarke, Fremont County, Colorado, U.S.A.
Magnosporites staplinii Rouse, British Columbia, Canada.
Mariusia andegavensis D. Pons & Boureau, Maine-and-Loire, France, Late Cretaceous (Middle
Cenomanian), (Microthyriaceae, Microthyriales).
Monoporisporites scabratus (Z.C. Song & Liu Cao) Kalgutkar & Janson., King George Island,
Antarctica. Pluricellaesporites cooksoniae Kalgutkar & Janson., Eucla basin, Western Australia,
P. ocellatus Z.C. Song & Liu Cao, King George Island, Antarctica, P. psilatus R.T. Clarke, P.
scabiosus R.T. Clarke, Fremont County, Colorado, U.S.A.
Reduviasporonites catenarius (G. Playford) Kalgutkar & Janson., Carnarvon Basin, Western
Australia. Senegalosporites costatus Jardiné & Magloire, Senegal.
Spegazzinites tetradus (Rouse) Kalgutkar & Janson., British Columbia, Canada, affinity with
Spegazzinia (Apiosporaceae, Sordariomycetidae).
Spirotremesporites costatus Jardiné & Magloire, Senegal, S. ellipticus Nandi & Shubhra
Banerjee, Mizoram, India, affinity with Hypoxylon (Xylariaceae, Xylariales).
Tetracoccosporium eocenum Biradar & Mahabalé, Madhya Pradesh, India, Maastrichtian,
Pezizomycotina. Thecaphora mohgaoensis (Chitaley & Yawale) R.K. Saxena, Wijayaw., D.Q.
Dai, K.D. Hyde & P.M. Kirk comb. nov., Madhya Pradesh, India, Glomosporiaceae,
Urocystidales.

860
Table 5 Continued.

Age Fossil fungal spore taxa, their affinity with modern taxa and location of occurrence
Trichosporites conwentzii Félix, Sweden, Late Cretaceous, affinity with Trichosporium
(Piedraiaceae, Capnodiales).
Valsarites senonianus Puri, Nigeria, affinity with Endothia, Didymosphaeria, and Valsaria (Puri
1963).
Xylohyphites verrucosus Kalgutkar & Sigler, Kalgutkar & Sigler, Madhya Pradesh, India,
Maastrichtian, affinity with Xylohypha (Pezizomycotina).
Jurassic Diporicellaesporites serratulus Traverse & Ash, Idaho, U.S.A.
Fractisporonites pittsburgensis Traverse & Ash, Traverse & Ash, Idaho, U.S.A.
Palaeopericonia fritzschei C. G. Ibáñez & Zamuner, Santa Cruz Province, Argentina, Middle
Jurassic, affinity with Periconia, Torula (Pleosporales).
Pluricellaesporites idahoensis Traverse & Ash, Idaho, U.S.A.
Triassic Inapertisporites argentinus (R.K. Jain) Kalgutkar & Janson., I. cacheutensis (R.K. Jain)
Kalgutkar & Janson., western Argentina.
Microsporonites cacheutensis R.K. Jain, western Argentina.
Carboniferous Cadyexinis tenuis Stach, C. vulgaris Stach, Germany.
Felixites playfordii Elsik, Spitsbergen, F. pollenisimilis (Horst) Elsik, Spitsbergen, Poland,
Scotland.
Parapotamomyces maydiformis O’Keefe, Saudi Arabia, Early Carboniferous, cf. Gliomastix
(Jarzen & Elsik 1986).
Portalites confertus Hemer & Nygreen, Saudi Arabia.
Tribolites triangulatus (Peppers) Kalgutkar & Janson. Illinois Basin, U.S.A., Carboniferous-
Pennsylvanian, affinity with Tetrachaetum and Lemonniera (Discinellaceae, Helotiales) (Bradley
1964).
Devonian Frasnacritetrus josettae Taug. France, Late Devonian (Frasnian), affinity with Tetraploa
(Tetraplosphaeriacea, Pleosporales).
Reduviasporonites catenarius (G. Playford) Kalgutkar & Janson., Carnarvon Basin, Western
Australia, affinity with Torulaceae, Pleosporales.

Suggestions for future studies

1. All microfossil (including fossil fungi) studies begin with field work and collection of
samples. Therefore, a sound knowledge of basic principles of field geology and clear concept of
various types of sedimentary rocks and stratigraphic units are essential for reliable results.
However, it has been observed that in a number of cases fossil fungal remains have been recorded
from the grab samples without their accurate stratigraphic locations. This makes them of limited
value in biostratigraphic studies. Exploring the stratigraphic significance of fossil fungal taxa is an
inviting task. In order to achieve this, it is necessary that samples for their studies must be collected
from the measured stratigraphic sections with precise location of their positions in the stratigraphic
column. Data obtained from such sections only can be relied for biostratigraphic inferences.
2. It is strongly felt that the data gathered so far from microfossil studies require more
precision. To achieve this, a synergistic approach must be adopted. Fossil fungal data need be
tagged, as far as possible, with data obtained from other disciplines in order to obtain better and
more dependable results and conclusions.
3. Use of obsolete and invalidly published names of palynotaxa create problems and must be
avoided. It is suggested that only current legitimate names of fossil fungal taxa must be used.
Nomenclatural repositories, viz. Index Fungorum and MycoBank, are indispensable to find out the
correct status of names of fungal taxa.
4. Sincere attempts must be made by expert mycologists to standardize the morphotaxonomy
and character evaluation of fossil fungal taxa and to trace the link of fossil fungal remains with
modern fungi. Such endeavour will help in elucidating the palaeoecology and evolutionary trends
within this group.
5. Data interpretation for stratigraphy and palaeoecology must also take into account various
factors operating for dispersal and deposition of microfossils.

861
 6. Recognition of recycled microfossils is very helpful in locating source area/ formation and
in interpreting palaeogeography. However, non-recognition of such elements may lead to erroneous
conclusions. Fluorescence microscopy is useful in detecting such fossils.
7. Subsurface samples, obtained from boreholes, usually represent uninterrupted sections,
almost completely unaffected by weathering agencies. Such samples are particularly useful where
some part of the sequence is eroded away or covered either by thick vegetation or by modern
sediments. Their study requires to be intensified to evaluate subsurface data vis-à-vis that from
surface sections.
8. A sound knowledge of various rules and regulations as laid down in the "International
Code of Nomenclature for algae, fungi and plants" (Turland et al. 2018) is essential for bringing
stability and discipline in the studies on fossil fungi. In addition, the articles and recommendations
contained in Chapter F are indispensable and deserve special attention.
9. All fossil fungal taxa must be registered with fungal nomenclatural repositories, e.g. Index
Fungorum/ MycoBank/ FungalNames to make its reference accessible world-wide. This will
certainly help in ensuring that they are validly published and in avoiding unnecessary introductions
of later homonyms.
10. Host-pathogen interaction is another aspect, which does not have basic information in the
form of fossil evidence. The interaction of fungi with higher plants with reference to the
palaeobotanical evidences need to be documented in appropriate manner by exploring more fossil
fungi along with chemical and geological aspects.

Acknowledgements
Ramesh K. Saxena is grateful to the authorities of the Birbal Sahni Institute of
Palaeosciences, Lucknow, India for library facilities and to Dr. S.K.M. Tripathi for making
drawings of some fungal spores. Nalin N. Wijayawardene and Dong-Qin Dai would like to thank
the National Natural Science Foundation of China (No. NSFC 31950410558, NSFC 31760013) and
the High-Level Talent Recruitment Plan of Yunnan Provinces (“Young Talents” Program and
“High-End Foreign Experts” Program) for support. This study was supported by the project of State
Key Laboratory of Functions and Applications of Medicinal Plants，Guizhou Medicial University
(No. FAMP201906K). Kevin D Hyde is thankful for the financial support from the Thailand
Research grant “Impact of climate change on fungal diversity and biogeography in the Greater
Mekong Subregion (grant no: RDG6130001)”.

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