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Diversity in Fossil Fungal Spores Article 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 1Birbal Sahni Institute of Palaeosciences, 53 University Road, Lucknow–226007, India 2State Key Laboratory of Functions and Applications of Medicinal Plants, Guizhou Medical University, Guiyang 550014, P.R. China 3Center for Yunnan Plateau Biological Resources Protection and Utilization, College of Biological Resource and Food Engineering, Qujing Normal University, Qujing, Yunnan 655011, P.R. China 4Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand 5Royal 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. 671 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
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