Asian Journal of Mycology 3(1): 490–511 (2020) www.asianjournalofmycology.org ISSN 2651-1339 Article Doi 10.5943/ajom/3/1/18 The diversity of endolichenic fungi – A review Chakarwarti J1,2, Nayaka S1,2* and Srivastava S2,3 1 Lichenology Laboratory, CSIR-National Botanical Research Institute, Rana Pratap Marg, Lucknow-226001, India Academy of Scientific and Innovative Research (AcSIR), CSIR-HRDC Campus, Kamla Nehru Nagar, Ghaziabad 201002, India 3 Plant Ecology and Environmental Technologies Division, CSIR-National Botanical Research Institute, Rana Pratap Marg, Lucknow-226001, India 2 Chakarwarti J, Nayaka S, Srivastava S 2020 – Diversity of endolichenic fungi – A review. Asian Journal of Mycology 3(1), 490–511, Doi 10.5943/ajom/3/1/18 Abstract Endolichenic fungi reside inside the lichen thalli without showing any visible symptoms of their presence. The fossil record of endolichenic fungi dates back to 415 million years, however, they have been successfully isolated only since the 1990s. Several reviews are available dealing with the bioprospecting potential of unique metabolites produced by endolichenic fungi; however, reviews addressing their taxonomic diversity are lacking. In this review, the names and the current taxonomic status of the endolichenic fungi reported so far are compiled and updated. The study indicated that approximately 500 endolichenic fungi are isolated of which only 135 identified up to species level. A large number of endolichenic fungi are either partially identified up to genus level or treated as unidentified. The reasons for such treatment are discussed. The study also revealed that only 114 well-known species of lichens belonging to 22 families are utilized for isolation of endolichenic fungi, while in some cases hosts were not identified. These lichens belong mostly to macro lichens with Parmeliaceae as the dominant members. The occurrence of the endolichenic fungi on different groups of plants indicates that they are not host-specific. Keywords – Endosymbionts – lichenicolous fungi – miniature ecosystem – symbiosis Introduction Lichens are considered as a miniature ecosystem (Honegger 1991) due to their close interaction with a variety of organisms such as fungi, algae, cyanobacteria, eubacteria, and microinvertebrates. While a fungus (Ascomycetes or Basidiomycetes) and a photoautotroph (Chlorophyta or Cyanobacteria) make a primary association and form lichen thallus, others make secondary, obligate or opportunistic associations. Recently, yeasts thought to be obligate, third partners in lichen formation (Spribille et al. 2016), however, this claim is not yet accepted (Mark et al. 2020). Due to growing ambiguity over the presence of yeast and other microbes, the lichens are redefined as ‘a self-sustaining ecosystem formed by the interaction of an exhabitant fungus and an extracellular arrangement of one or more photosynthetic partners and an indeterminate number of other microscopic organisms’ (Hawksworth & Grube 2020). Among the secondary associates, the lichenicolous and endolichenic fungi (hereafter referred to as ELF) are interesting groups due to their taxonomic and evolutionary status (Oh et al. 2020), and for bioprospecting (Maduranga et al. 2018). The ELF, which are endosymbionts of lichens (Miadlikowska et al. 2004) are predominantly filamentous, reside inside the lichen thallus without showing any visible symptoms of their Submitted 8 October 2020, Accepted 12 November 2020, Published 3 December 2020 Corresponding Author: Sanjeeva Nayaka – e-mail – nayaka.sanjeeva@gmail.com 1 presence and apparently, do not harm the host. The ELF closely associate themselves with the photobiont of the lichen for their nutrition (Arnold et al. 2009). They are similar to endophytic fungi of vascular plants. The endophytic fungi play various functional roles in their plant hosts like altering their disease resistance or susceptibility, augmenting their growth, and ability to survive in extreme environments (Yu et al. 2018). However, the role of endolichenic fungi in increasing the fitness of lichens is not known (Suryanarayan et al. 2017). Diversity of ELF The ELF are cosmopolitan in distribution and are known from every lichen species tested from the Arctic to tropics. Around 13,500 species of ELF are estimated to occur in the world (Kellogg & Raja 2016). The ELF exhibits a vast diversity as indicated by Petrini et al. (1990), who isolated 506 fungal taxa from 17 lichen species; of these, 306 were isolated only once. An intensive study of two lichens Peltigera praetextata and Xanthoparmelia taractica from a coniferous forest (Aosta Valley, Italy) yielded different fungi belonging to 117 taxa (Girlanda et al. 1997). Suryanarayanan et al. (2005) isolated 242 endolichenic fungi from 500 thallus segments of five corticolous lichens. Li et al. (2007) isolated a total of 32 taxa from 488 thallus segments, with 31 ascomycetes and one yeast. Zhang et al. (2016) recovered 247 fungal isolates from 2712 thallus segments of 17 lichen species. Govinda Rajulu et al. (2020) isolated a total of 389 species from 10 lichen species of Western Ghats, Southern India. These reports exemplify the tremendous diversity of ELF. Rashmi et al. (2019) provided a lengthy list of endophytic fungi reported from the world, however, it included very few ELFs (eg. Nodulisporium hyalosporum). The compilation of all the ELF studies carried out so far indicated that about 500 ELFs have been isolated till date, among them only 135 were identified up to species level while most of them are either identified up to the genus or class level or treated as unidentified (Table 1). As the ELFs do not produce sufficient morphological characters such as colouration, conidia or spores in culture, it is difficult to identify under routine microscopic studies (Tripathi and Joshi 2019). The DNA based molecular techniques are only partially helpful due to the lack of sequences in gene bank or library related to ELFs. Usullay, the aim of most of the researchers is to isolate ELFs for bioactive molecule and hence not much attention is paid towards their taxonomy. Recent studies have indicated that ELFs are not different from endophytes and the DNA based techniques are able to unravel their identity (Govinda Rajulu et al. 2020). Table 1. List of lichens so far explored for ELFs (name in parenthesis are synonyms as originally annotated by authors) 1 2 3 4 Fungi Acremonium sp. Host lichen Cladonia sp.; Stereocaulon sp. Parmotrema thomsonii (= Parmelia thomsonii) Dermatocarpon miniatum; Melanelia sorediata; Parmotrema thomsonii Parmotrema sp. Lobaria kurokawae; Parmotrema thomsonii; P. tinctorum; Punctelia rudecta Lobaria kurokawae; Parmotrema tinctorum; Punctelia rudecta Alternaria sp. (= Ulocladium Peltigera praetextata; Xanthoparmelia taractica sp.) Parmotrema hababianum Hypotrachyna sp.; Unknown lichen A. alternariae (Cooke) Xanthoparmelia taractica (= Parmelia taractica); Woudenb. & Crous (= Peltigera praetextata Ulocladium alternariae (Cooke) Simmons) A. alternata (Fr.) Keissl. Peltigera praetextata; Xanthoparmelia taractica Bulbothrix meizospora; Heterodermia flabellata; H. hypochraea; Parmotrema crinitum; P. Reference Petrini et al. (1990) Suryanarayanan et al. (2005) Li et al. (2007) Kannangara et al. (2008) Tripathi et al. (2014) Suryanarayanan et al. (2017) Girlanda et al. (1997) Suryanarayanan et al. (2017) Wang et al. (2012) Girlanda et al. (1997) Girlanda et al. (1997) Suryanarayanan et al. (2005) 2 Table 1 Continued. Fungi 5 6 7 8 9 10 11 12 A. tenuissima (Kunze) Wiltshire Aphanocladium sp. Apiosordaria sp. Apiotrichum (= Hyalodendron sp.) Arthrinium arundinis (corda) Dyko & B. Sutton (= Apiospora montagnei Sacc.) Arthrinium phaeospermum (Corda) M. B. Ellis Ascochyta sp. Aspergillus sp. (= Eurotium sp.) Host lichen praesorediosum; P. nilgherrense; P. reticulatum; P. thomsonii; Physcia dilatate Heterodermia flabellata; Parmotrema crinitum; P. thomsonii; Physcia dilatate Usnea aciculifera Bulbothrix meizospora; Heterodermia flabellata; H. hypochraea; Flavoparmelia caperata; Leptogium burnetiae; Parmotrema critinum; P. graynum; P. nilgherrense; P. praesorediosum; P. reticulatum; P. thomsonii; Physcia dilatate Cladonia fruticulosa; Parmotrema reticulatum; P. tinctorum #Not specified – list of several lichens given Cladonia sp.; Stereocaulon sp. 15 16 17 18 A. aculeatus Iizuka A. coremiiformis Bartoli & Maggi A. flavus Link A. fumigatus Fresen. A. hiratsukae Udagawa, Tsub. & Y. Horie A. niger Tiegh. Li et al. (2007) He et al. (2012) Tripathi et al. (2014) Vinayaka et al. (2016) Zhang et al. (2016) Petrini et al. (1990) Cladonia sp.; Stereocaulon sp. #Not specified – list of several lichens given Cladonia sp.; Stereocaulon sp. Petrini et al. (1990) Zhang et al. (2016) Petrini et al. (1990) Cladonia sp. Wang et al. (2017) Cladonia sp.; Stereocaulon sp. Peltigera praetextata; Xanthoparmelia taractica Cladonia sp.; Stereocaulon sp. Pseudocyphellaria sp. Peltigera elisabethae var. mauritzii Cetrelia sp. Petrini et al. (1990) Girlanda et al. (1997) Petrini et al. (1990) Kannangara et al. (2008) Zhao et al. (2014) Chen et al. (2014), Chen et al. (2019) Chen et al. (2014) Padhi and Tayung (2015, 2016) Suryanarayanan et al. (2017) Maduranga et al. (2018) Tripathi et al. (2014) Cladina grisea Parmelia sp. 13 14 Reference Phaeophyscia hispidula Porina tetracerae Flavoparmelia caperata Bulbothrix meizospora; Heterodermia flabellata; H. hypochraea; Parmotrema crinitum; P. nilgherrense; P. praesorediosum; P. reticulatum; P. thomsonii; Physcia dilatate Heterodermia flabellata; Parmotrema crinitum; P. thomsonii; Physcia dilatate Bulbothrix meizospora; Flavoparmelia caperata; Heterodermia flabellata; H. hypochraea; Leptogium burnetiae; Parmotrema crinitum; P. graynum; P. nilgherrense; P. praesorediosum; P. reticulatum; P. thomsonii; Physcia dilatate Unidentified Opegrapha medusulina; Pyxine cocoes; Roccella montagnei; Unidentified Heterodermia flabellata; Parmotrema thomsonii; Physcia dilatata; Roccella montagnei Heterodermia flabellata; Parmotrema thomsonii; Physcia dilatate Parmotrema cristiferum; Ramalina arabum; Teloschistes flavicans; Usnea galbinifera Parmotrema ravum Suryanarayanan et al. (2005) Li et al. (2007) Tripathi et al. (2014) Maduranga et al. (2018) Maduranga et al. (2018) Suryanarayanan et al. (2005) Li et al. (2007), Tripathi et al. (2014) Vinayaka et al. (2016) Padhi et al. (2019) 3 Table 1 Continued. 35 Fungi Host lichen A. versicolor (Vuill.) Tirab. Lobaria retigera Ricasolia quercizans (= Lobaria quercizans) Unknown A. quadricinctus E. Yuill Usnea longissima Aureobasidium pullulans (de Canoparmelia texana; Heterodermia diademata; Bary & Lӧwenthal) G. H. podocarpa; Leptogium askotense; Lobaria Arnaud kurokawae; Parmotrema hababianum; P. tinctorum; Phaeophyscia hispidula; Punctelia rudecta; Ramalina conduplicans; Usnea sp. Bacidina sp.; Teloschistes flavicans Bahusakala sp. Cladonia sp.; Stereocaulon sp. Berkeleyomyces basicola Polyblastidium dendriticum (= Heterodermia (Berk. & Broome) W.J. Nel, dendritica); Parmotrema reticulatum; Ramalina Z.W. de Beer, T.A. Duong & arabum; Usnea galbinifera M.J. Wingf (= Thielavia basicola (Berk. & Broome) Zopf) Biatriospora sp. Pseudocyphellaria sp. Biscogniauxia petrensis Z.F. Parmotrema sp. Zhang, F. Liu & L. Cai Bispora sp. Canoparmelia texana Botrytis sp. Roccella montagnei Parmotrema tinctorum Dirinaria consimilis B. cinerea Pers. Cladonia sp.; Stereocaulon sp. Broomella sp. Parmotrema sp. Byssochlamys spectabilis Pyrenula ochraceoflava (Udagawa & Shoji Suzuki) Houbraken & Samson Cadophora bubakii (Laxa) Dermatocarpon miniatum; Melanelia sorediata; Damm & S. Bien (= Ramalina sinensis Phailophora bubakii (Laxa) Schol-Schwarz) Calonectria gracilis Corus, Peltigera praetextata; Xanthoparmelia taractica M.J. Wingf. & Alfenaas (= Cylindrocarpon gracile Bugnic.) Cerrena sp. Pyrenocarp sp.; Pyrenula indet Cerrena unicolor (Bull.) Roccella montagnei Murrill Chaetomella sp. Parmotrema nilgherrense 36 Chaetomium sp. 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 37 C. elatum Kunze Roccella montagnei Punctelia borreri Canoparmelia texana; Heterodermia diademata; H. podocarpa; Leptogium askotense; Lobaria kurokawae; Parmotrema tinctorum; Phaeophyscia hispidula; Punctelia rudecta Heterodermia incana; Parmotrema cristiferum; P. reticulatum; P. tinctorum; Ramalina arabum; Ramalina pacifica; Teloschistes flavicans Bacidina sp.; Dirinaria consimilis; Evernia prunastri; Parmotrema sp.; Usnea sp. Melanelia sorediata Hypotrachyna cirrhata (= Everniastrum cirrhatum) Parmotrema reticulatum; Ramalina arabum; Usnea galbinifera Reference Duo et al. (2014) Li et al. (2015a) Li et al. (2015a) Prateeksha et al. (2020) Suryanarayanan et al. (2017) Govinda Rajulu et al. (2020) Petrini et al. (1990) Vinayaka et al. (2016) Zhou et al. (2016) Oh et al. (2020) Suryanarayanan et al. (2017) Suryanarayanan et al. (2005) Suryanarayanan et al. (2017) Govinda Rajulu et al. (2020) Petrini et al. (1990) Kannangara et al. (2008) Maduranga et al. (2018) Li et al. (2007) Girlanda et al. (1997) Maduranga et al. (2018) Maduranga et al. (2018) Suryanarayanan et al. (2005), Tripathi et al. (2014) Suryanarayanan et al. (2005) Li et al. (2007) Suryanarayanan et al. (2017) Vinayaka et al. (2016) Govinda Rajulu et al. (2020) Li et al. (2007) Chen et al. (2013) Vinayaka et al. (2016) 4 Table 1 Continued. 38 Fungi C. globosum Kunze 39 Chalara sp. 40 C. constricta NagRaj & W.B. Kendr. C. fusidioides (Corda) Rabenh. Chrysosporium sp. C. pseudomerdarium van Oorschot Cladophialophora chaetospira (Grove) Crous & Arzanlou (= Heteroconium chaetospira (Grove) M. B. Ellis) Cladosporium sp. 41 42 43 44 45 46 C. cladosporioides (Fr.) de Vries 47 48 49 C. herbarum (Pers.) Link Colletotrichum sp. C. dematium (Pers.) Grove 50 Coniochaeta sp. (= Lecythophora sp.) 51 52 Host lichen Cladonia coniocraea; Melanelia sorediata Hypotrachyna nepalensis (= Everniastrum nepalense) Cladonia fruticulosa; Parmotrema cristiferum; P. reticulatum; P. tinctorum; Polyblastidium dendriticum; Ramalina pacifica; Usnea stigmatoides #Not specified – list of several lichens given Cladonia sp.; Stereocaulon sp. Parmotrema reticulatum; Ramalina arabum; Usnea stigmatoides Cladonia sp.; Stereocaulon sp. Reference Li et al. (2007) Zheng et al. (2014) Cladonia sp.; Stereocaulon sp. Petrini et al. (1990) Parmotrema sp.; Usnea sp. Cladonia sp.; Stereocaulon sp. Kannangara et al. (2008) Petrini et al. (1990) Cladonia sp.; Stereocaulon sp. Peltigera praetextata; Xanthoparmelia taractica Petrini et al. (1990) Girlanda et al. (1997) Cladonia sp.; Stereocaulon sp. Roccella montagnei; Parmotrema nilgherrense; P. praesorediosum; Physcia dilatata Physcia dilatata Pseudocyphellaria sp.; Parmotrema sp.; Usnea sp. Parmotrema hababianum; P. nilgherrense; P. praesorediosum; Lobaria kurokawae; Physcia dilatata Lobaria kurokawae Arthonia sp.; Hypotrachyna crenata; Parmotrema tinctorum Cladonia sp.; Stereocaulon sp. Peltigera praetextata; Xanthoparmelia taractica Heterodermia incana; Ramalina arabum; Teloschistes flavicans; Usnea stigmatoides; U. galbinifera Cladonia sp.; Stereocaulon sp. Cladonia sp.; Stereocaulon sp. Parmotrema tinctorum; Polyblastidium dendriticum; Ramalina arabum; R. pacifica; Usnea stigmatoides Dermatocarpon miniatum; Parmelia sp.; Punctelia borreri; Zeroviella mandschurica (= Xanthoria mandschurica) Zeroviella mandschurica #Not specified – list of several lichens given Cladonia evansii Cladonia sp.; Stereocaulon sp. Peltigera praetextata; Xanthoparmelia taractica Petrini et al. (1990) Suryanarayanan et al. (2005) Cordyceps farinose (Holmsk.) Kepler, B. Shrestha & Spatafora (= Paecilomyces farinosus (Holm ex S. F. Gray) Brown et Smith) Corynespora sp. Usnea cavernosa Vinayaka et al. (2016) Zhang et al. (2016) Petrini et al. (1990) Vinayaka et al. (2016) Petrini et al. (1990) Li et al. (2007) Kannangara et al. (2008) Tripathi et al. (2014) Suryanarayanan et al. (2017) Govinda Rajulu et al. (2020) Petrini et al. (1990) Girlanda et al. (1997) Vinayaka et al. (2016) Petrini et al. (1990) Petrini et al. (1990) Vinayaka et al. (2016) Li et al. (2007) Wang et al. (2010) Zhang et al. (2016) Wijeratne et al. (2016) Petrini et al. (1990) Girlanda et al. (1997) Paranagama et al. (2007) 5 Table 1 Continued. 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 Fungi Cosmospora butyri (J.F.H. Beyma) Gräfenhan, Seifert & Schroers (=Acremonium butyri (Beyma) W. Gams) Cryptocline sp. C. fasciculata (Tode) Petr. Curvularia sp. C. australiensis (Bugnic. ex M.B. Ellis) Manamgoda, L. Cai & K.D. Hyde (= Bipolaris australiensis (Bugnic. ex M.B. Ellis) Tsuda & Ueyama) C. lunata (Wakker) Boedijn Host lichen Peltigera praetextata; Xanthoparmelia taractica Reference Girlanda et al. (1997) Cladonia sp.; Stereocaulon sp. Cladonia sp.; Stereocaulon sp. Pseudocyphellaria sp.; Usnea sp. Heterodermia flabellata; Physcia dilatata Petrini et al. (1990) Petrini et al. (1990) Kannagara et al. (2008) Suryanarayanan et al. (2005), Li et al. (2007), Tripathi et al. (2014) Polyblastidium dendriticum; Ramalina pacifica; Usnea stigmatoides Peltigera praetextata; Xanthoparmelia taractica Vinayaka et al. (2016) C. spicifera (Bainier) Boedijn (= Drechslera anamorph Cochliobolus spicifer Nelson) C. trifolii (Kauffman) Usnea sp. Boedijn Cylindromonium lichenicola Parmotrema reticulatum (W. Gams) Crous (=Acremonium lichenicola W. Gams) Cylindrosporium sp. Bulbothrix meizospora Cystobasidium laryngis (Reiersӧl) Yurkov Cytospora sp. Daldinia sp. Daldinia eschscholtzii (Ehrenb.) Rehm #Not specified – list of several lichens given Parmelia sp. Dirinaria picta; Pyxine cocoes Dirinaria picta; Opegrapha medusulina; O. arabica; Porina tetracerae; Roccella montagnei Dirinaria consimilis Leptogium sp. Diaporthe sp. (= Phomopsis Cladonia sp.; Porina tetracerae; Stereocaulon sp. sp.) D. arengae R.R. Gomes, Glienke & Crous Endomelanconiopsis sp. E. endophytica E.I. Rojas & Samuels Epicoccum nigrum Link (= Epicoccum purpurascens Ehrenb. ex Schlecht) Exophiala sp. E. lecanii-corni (Benedek & G. Specht) Haase & de Hoog, (= E. jeanselmei var. lecanii-corvi (Benedeck & Specht) De Hoog) E. mansonii (Castell.) De Hoog Fusarium sp. Arthonia antillarum; Opegrapha medusulina; Unidentified Porina tetracerae; Pyrenula sp. Porina tetracerae; Pyrenula sp.; Unidentified Girlanda et al. (1997) Samanthi et al. (2015a) Suryanarayanan et al. (2005), Tripathi et al. (2014) Suryanarayanan et al. (2005), Tripathi et al. (2014) Zhang et al. (2016) Padhi and Tayung (2015) Maduranga et al. (2018) Maduranga et al. (2018) Govinda Rajulu et al. (2020) Oh et al. (2020) Petrini et al. (1990), Padhi and Tayung (2015), Maduranga et al. (2018) Maduranga et al. (2018) Maduranga et al. (2018) Maduranga et al. (2018) Cladonia sp.; Stereocaulon sp. Petrini et al. (1990) Cladonia sp.; Stereocaulon sp. Cladonia sp.; Stereocaulon sp. Petrini et al. (1990) Petrini et al. (1990) Cladonia sp.; Stereocaulon sp. Petrini et al. (1990) Cladonia sp.; Stereocaulon sp. Peltigera praetextata; Xanthoparmelia taractica Usnea sp. Petrini et al. (1990) Girlanda et al. (1997) Kannagara et al. (2008) 6 Table 1 Continued. Fungi 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 Host lichen Parmelia sp. Arthonia sp.; Bacidina sp. F. lateritium Nees Cladonia sp.; Stereocaulon sp. Heterodermia incana; Polyblastidium dendriticum; Ramalina arabum; Teloschistes flavicans; Usnea stigmatoides F. lolii (Wm.G.Sm.) Sacc. (= Cladonia sp.; Stereocaulon sp. Fusarium heterosporum Cladonia fruticulosa; Parmotrema cristiferum; Nees ex Fr.) Ramalina pacifica; Usnea galbinifera F. sublunatum var. Cladonia sp.; Stereocaulon sp. sublunatum Nierenb. Geopyxis majalis (Fr.) Sacc. Pseudevernia intensa Geotrichum sp. Cladonia sp.; Stereocaulon sp. Canoparmelia texana; Heterodermia diademata; Lobaria kurokawae; Parmotrema tinctorum Gilmaniella humicola G.L. Bulbothrix meizospora; Parmotrema nilgherrense Barron Bulbothrix meizospora; Leptogium burnetiae; Parmotrema nilgherrense; P. graynum; Punctelia subrudecta Heterobasidion annosum Fr. Cladonia sp.; Stereocaulon sp. Bref. (= Spiniger meineckellus (Olson) Stalpers) Heteroconium sp. Cladonia sp.; Stereocaulon sp. Humicola sp. Heterodermia diademata; H. podocarpa; Lobaria kurokawae; Parmotrema hababianum; P. tinctorum; Punctelia rudecta Hypoxylon sp. (= Cladonia fruticulosa; Parmotrema reticulatum Nodulisporium sp.) Dermatocarpon miniatum; Melanelia sorediata; Punctelia borreri Hypotrachyna sp. (= Everniastrum sp.) Arthonia sp.; Bacidina sp.; Evernia prunastri; Hypotrachyna crenata; Parmotrema tinctorum; Teloschistes flavicans H. anthochroum Berk. & Dirinaria picta; Opegrapha medusulina; Porina Broome tetracerae H. fuscum (Pers.) Fr. Usnea sp. H. fragiforme (Pers.) J. Lobaria discolor; L. japonica Kickx f. H. investiens (Schwein.) Dirinaria consimilis; Evernia prunastri; M.A. Curtis Hypotrachyna crenata; Parmotrema sp.; P. tinctorum; Ramalina pacifica Lasiodiplodia crassispora Roccella montagnei T.I Burgess & P.A. Barber L. theobromae (Pat.) Griffon Roccella montagnei & Maubl. Arthonia antillarum; Opegrapha arabica; Pyrenula ochraceoflava; P. parvinuclea; Roccella montagnei Bacidina sp.; Hypotrachyna crenata L. pseudotheobromae A.J.L. Roccella montagnei; Unidentified Phillips, A. Alves & Crous Monilinia sp. (= Monilia sp.) Heterodermia diademata; Lobaria kurokawae Mucor hiemalis Wehmer f. hiemalis M. racemosus Fresen. Peltigera praetextata; Xanthoparmelia taractica Myrothecium inundatum Tode Ramalina sp. Parmotrema reticulatum Reference Padhi and Tayung (2015) Govinda Rajulu et al. (2020) Petrini et al. (1990) Vinayaka et al. (2016) Petrini et al. (1990) Vinayaka et al. (2016) Petrini et al. (1990) Wijeratne et al. (2012) Petrini et al. (1990) Tripathi et al. (2014), Suryanarayanan et al. (2017) Suryanarayanan et al. (2005) Tripathi et al. (2014) Petrini et al. (1990) Petrini et al. (1990) Tripathi et al. (2014), Suryanarayanan et al. (2017) Vinayaka et al. (2016) Li et al. (2007) Zhao et al. (2015) Govinda Rajulu et al. (2020) Maduranga et al. (2018) Basnet et al. (2019a) Oh et al. (2020) Govinda Rajulu et al. (2020) Maduranga et al. (2018) Suryanarayanan et al. (2005) Maduranga et al. (2018) Govinda Rajulu et al. (2020) Maduranga et al. (2018) Tripathi et al. (2014), Suryanarayanan et al. (2017) Girlanda et al. (1997) Suryanarayanan et al. (2005), Tripathi et al. (2014) Basnet et al. (2019b) 7 Table 1 Continued. 97 98 99 Fungi Myxotrichum sp. Nemania bipapillata (Berk. & M.A. Curtis) N. serpens (Pers.) Gray (= Geniculosporium serpens Chesters & Greenh.) 100 Neocosmospora solani (Mart.) L. Lombard & Crous (= Fusarium solani (Mart.) Sacc.) 101 102 103 104 105 106 107 108 Host lichen Cetraria islandica Bacidina sp.; Dirinaria consimilis; Hypotrachyna crenata; Parmotrema sp.; P. tinctorum; Teloschistes flavicans; Usnea sp. Dermatocarpon miniatum Xanthoparmelia taractica; Peltigera praetextata Bulbothrix meizospora; Heterodermia flabellata; H. hypochraea; Parmotrema crinitum; P. graynum; P. praesorediosum; P. nilgherrense; P. reticulatum; P. thomsonii; Physcia dilatata Heterodermia flabellate: Parmotrema crinitum; P. graynum; P. thomsonii; Physcia dilatata Bulbothrix meizospora; Flavoparmelia caperata; Heterodermia flabellata; H. hypochraea; Leptogium burnetiae; Parmotrema crinitum; P. graynum; P. praesorediosum; P. nilgherrense; P. reticulatum; P. thomsonii; Physcia dilatata; Punctelia subruducta; Usnea sp. Parmotrema cristiferum; P. reticulatum; Ramalina pacifica; Usnea stigmatoides Neonectria ditissima (Tul. & Peltigera praetextata; Xanthoparmelia taractica C. Tul.) Samuels & Rossman (= Cylindrocarpon heteronema (Berk. & Br.) Wollenw.) Neurospora sp. Roccella montagnei N. crassa Shear & B.O. Pyrenula parvinuclea; Pyxine cocoes Dodge N. terricola Goch. & Backus Hypotrachyna cirrhata Nigrospora sp. Usnea sp. Roccella montagnei N. aurantiaca Mei Wang & Lobaria discolor; L. japonica L. Cai N. chinensis Mei Wang & L. Lobaria discolor; L. japonica Cai N. oryzae (Berk. & Broome) Parmotrema reticulatum Petch Leptogium askotense; Parmotrema reticulatum Parmotrema tinctorum 109 N. oryzae (Berk. & Broome) Parmotrema thomsonii Petch (= N. sphaerica (Sacc.) E.W. Mason) Parmelinella wallichiana Opegrapha medusulina; Unidentified 110 Nodulisporium hyalosporum Punctelia borreri S.C. Agarwal & J.K. Misra* 111 N. sylviforme Deighton* Melanelia sorediata 112 Ochrocladosporium elatum Cladonia sp.; Stereocaulon sp. (Harz) Crous & U. Braun (= C. elatum (Harz.) Nannfeldt) 113 Oidiodendron cf. Cladonia sp.; Stereocaulon sp. echinulatum Barron 114 O. flavum Svilv. Cladonia sp.; Stereocaulon sp. 115 Ophiosphaerella korrae (J. Unknown Walker & A.M. Sm. Bis) Physcia sp. Shoemaker & C.E. Babc. Reference Yuan et al. (2013) Govinda Rajulu et al. (2020) Li et al. (2007) Girlanda et al. (1997) Suryanarayanan et al. (2005) Li et al. (2007) Tripathi et al. (2014) Vinayaka et al. (2016) Girlanda et al. (1997) Maduranga et al. (2018) Maduranga et al. (2018) Zhang et al. (2009) Kannangara et al. (2008) Maduranga et al. (2018) Oh et al. (2020) Oh et al. (2020) Suryanarayanan et al. (2005) Tripathi et al. (2014), Suryanarayanan et al. (2017) Govinda Rajulu et al. (2020) Suryanarayanan et al. (2005), Li et al. (2007), Tripathi et al. (2014) He et al. (2012) Maduranga et al. (2018) Li et al. (2007) Li et al. (2007) Petrini et al. (1990) Petrini et al. (1990) Petrini et al. (1990) Li et al. (2018) Li et al. (2019) 8 Table 1 Continued. Fungi 116 Paecilomyces sp. 117 Papulospora sp. Host lichen Roccella montagnei Dirinaria consimilis; Evernia prunastri; Hypotrachyna crenata; Parmotrema sp.; P. tinctorum; Ramalina pacifica; Teloschistes flavicans Heterodermia hypochraea; Parmotrema reticulatum Peltigera praetextata, Xanthoparmelia taractica 118 Paraboeremia putaminum (Speg.) Qian Chen & L. Cai (= P. putaminum Speg.) 119 Paraphaeosphaeria michotii Cladonia sp.; Stereocaulon sp. (Westend) Eriksson 120 Penicillium sp. (= Cladonia sp.; Stereocaulon sp. Torulomyces sp.) Bulbothrix meizospora; Parmotrema reticulatum; Roccella montagnei Leptogium askotense; Lobaria kurokawae; Ramalina conduplicans; Usnea sp. Pseudocyphellaria sp. Bulbothrix meizospora; Canoparmelia texana; Heterodermia podocarpa; Leptogium askotense; Lobaria kurokawae; Parmotrema hababianum; P. reticulatum; Phaeophyscia hispidula; Punctelia rudecta Parmelia sp. Arthonia sp.; Bacidina sp.; Hypotrachyna crenata; Parmotrema tinctorum 121 P. aurantiogriseum Dierckx Xanthoparmelia taractica; Peltigera praetextata (= P. viridicatum Westling) 122 P. canescens Sopp (= P. Xanthoparmelia taractica; Peltigera praetextata raciborskii Zaleski) 123 P. chrysogenum Thom Parmotrema reticulatum; Polyblastidium dendriticum; Usnea stigmatoides 124 P. citrinum Thom Parmotrema sp. 125 P. griseofulvum Dierckx Xanthoparmelia taractica; Peltigera praetextata 126 P. melinii Thorn Xanthoparmelia taractica; Peltigera praetextata 127 P. purpurogenum Stoll Xanthoparmelia taractica; Peltigera praetextata 128 P. glaucoalbidum (Desm.) Cladonia sp.; Stereocaulon sp. Houbraken & Samson (= Thysanophora penicillioides (Roum.) Kendrick) 129 Periconia sp. Parmotrema sp. Heterodermia podocarpa; Leptogium askotense; Lobaria kurokawae; Parmotrema hababianum P. tinctorum; Phaeophyscia hispidula; Punctelia rudecta Parmelia sp. 130 Pestalotiopsis sp. Heterodermia flabellata; Parmotrema thomsonii Multiclavula sp. Cetraria islandica Bacidina sp. Parmotrema reticulatum; P. tinctorum; Teloschistes flavicans Parmotrema reticulatum 131 P. hypodermia (Niessl) Steyaert 132 P. maculans (Corda) Nag Raj 133 Pezicula (= Cryptosporiopsis Cladonia sp.; Stereocaulon sp. sp.) Reference Suryanarayanan et al. (2005) Govinda Rajulu et al. (2020) Suryanarayanan et al. (2005), Tripathi et al. (2014) Girlanda et al. (1997) Petrini et al. (1990) Petrini et al. (1990) Suryanarayanan et al. (2005) Suryanarayanan et al. (2017) Kannangara et al. (2008) Tripathi et al. (2014) Padhi and Tayung (2015) Govinda Rajulu et al. (2020) Girlanda et al. (1997) Girlanda et al. (1997) Vinayaka et al. (2016) Samanthi et al. (2015b) Girlanda et al. (1997) Girlanda et al. (1997) Girlanda et al. (1997) Petrini et al. (1990) Kannangara et al. (2008) Tripathi et al. (2014), Suryanarayanan et al. (2017) Wu et al. (2015) Suryanarayanan et al. (2005), Li et al. (2007), Tripathi et al. (2014) Ding et al. (2009) Yuan et al. (2017) Govinda Rajulu et al. (2020) Vinayaka et al. (2016) Suryanarayanan et al. (2005), Tripathi et al. (2014) Petrini et al. (1990) 9 Table 1 Continued. Fungi 134 Phaeoramularia sp. 135 Phaeosphaeria sp. 136 Phialocephala fortinii C.J.K. Wang & H.E. Wilcox 137 Phialophora sp. 138 139 140 141 142 143 144 145 146 147 148 149 150 151 152 153 154 Host lichen Cladonia sp.; Stereocaulon sp. Heterodermia obscurata Unknown Roccella montagnei Dermatocarpon miniatum Cetrelia braunsiana Cladonia ochrochlora Parmotrema tinctorum; Teloschistes flavicans P. asteris (Dowson) Burge & Cladonia sp.; Stereocaulon sp. I. Isaac (= P. fastigiata (Lagerb. et al) Conant) P. verrucosa Medlar Heterodermia incana; Ramalina pacifica; Usnea galbinifera Phoma sp. Cladonia sp.; Stereocaulon sp. Peltigera praetextata; Xanthoparmelia taractica Punctelia borreri Pseudocyphellaria sp. Heterodermia incana; Parmotrema tinctorum; Teloschistes flavicans Unknown Arthonia sp. P. leveillei Boerema & G.J. Cladonia fruticulosa; Parmotrema reticulatum; Bollen Polyblastidium dendriticum; Ramalina arabum Plagiostoma (= Diplodina Cladonia sp.; Stereocaulon sp. sp.) Preussia sp. Opegrapha medusulina Preussia africana Arenal, Ramalina calicaris Platas & Pelàez P. tenerifae (Arx & Guarro) Arthonia parantillarum Kruys Pseudeurotium sp. #Not specified – list of several lichens given Pseudogymnoascus (Link) Cladonia sp.; Stereocaulon sp. Minnis & D.L. Lindner (= Xanthoparmelia taractica; Peltigera praetextata Geomyces pannorum (Link) Sigler et Carmichael Purpureocillium lilacinum Xanthoparmelia taractica; Peltigera praetextata (Thom) Luangsa-ard, Houbraken, Hywel-Jones & Samson (= Paecilomyces lilacinus (Thorn) Samson) P. variotii Bainier Cladonia sp.; Stereocaulon sp. Ramichloridium cf. Cladonia sp.; Stereocaulon sp. apiculatum (Miller et al.) De Hoog Ramularia sp. Cladonia sp.; Stereocaulon sp. Rigidoporus vinctus (Berk.) Roccella montagnei Ryvarden Rhizoctonia sp. Punctelia subruducta R. solani J.G. Kühn Heterodermia incana; Parmotrema cristiferum; Usnea stigmatoides 155 Rhizopus sp. 156 Rhizosphaera kalkhoffii Bubak 157 Schizophyllum commune Fr. 158 Scopulariopsis sp. Reference Petrini et al. (1990) Li et al. (2012) Xie et al. (2016) Suryanarayanan et al. (2005) Li et al. (2007) He et al. (2012) Ye et al. (2013) Govinda Rajulu et al. (2020) Petrini et al. (1990) Vinayaka et al. (2016) Petrini et al. (1990) Girlanda et al. (1997) Li et al. (2007) Kannangara et al. (2008) Vinayaka et al. (2016) Kim et al. (2018a) Govinda Rajulu et al. (2020) Vinayaka et al. (2016) Petrini et al. (1990) Maduranga et al. (2018) Zhang et al. (2012) Maduranga et al. (2018) Zhang et al. (2016) Petrini et al. (1990) Girlanda et al. (1997) Girlanda et al. (1997) Petrini et al. (1990) Petrini et al. (1990) Petrini et al. (1990) Maduranga et al. (2018) Tripathi et al. (2014) Vinayaka et al. (2016) Roccella montagnei Cladonia sp.; Stereocaulon sp. Suryanarayanan et al. (2005) Petrini et al. (1990) Opegrapha medusulina; Unidentified Cladonia sp.; Stereocaulon sp. Maduranga et al. (2018) Petrini et al. (1990) 10 Table 1 Continued. Fungi 159 Scutula sp. (= Karsteniomyces sp.) 160 Sordaria sp. 161 S. fimicola (Roberge ex Desm.) Ces. & De Not. 162 Spegazzinia tessarthra (Berk. & M.A. Curtis) Sacc. Host lichen Cladonia coniocraea; Dermatocarpon miniatum; Melanelia sorediata; Parmelia sp.; Punctelia borreri; Zeroviella mandschurica Xanthoparmelia taractica; Peltigera praetextata Reference Li et al. (2007) Heterodermia diademata; H. podocarpa; Leptogium askotense; Lobaria kurokawae; Parmotrema tinctorum; Phaeophyscia hispidula; Punctelia rudecta; Usnea sp. Unidentified Tripathi et al. (2014), Suryanarayanan et al. (2017) Parmotrema reticulatum Tripathi et al. (2014), Suryanarayanan et al. (2005) Oh et al. (2020) Myelochroa sp.; Parmotrema sp.; Phaeophyscia sp.; Physcia sp.; Stereocaulon sp. Heterodermia flabellata Girlanda et al. (1997) Maduranga et al. (2018) Suryanarayanan et al. (2005), Li et al. (2007), Tripathi et al. (2014) Tripathi et al. (2014), Suryanarayanan et al. (2017) Petrini et al. (1990) Li et al. (2007) Suryanarayanan et al. (2005) Govinda Rajulu et al. (2020) 163 S. lobulata Thrower Punctelia rudecta 164 Spiniger sp. 165 Sporormiella sp. 166 S. intermedia (Auersw.) Ahmed & Cain Cladonia sp.; Stereocaulon sp. Zeroviella mandschurica Roccella montagnei Arthonia sp.; Dirinaria consimilis; Hypotrachyna crenata; Parmotrema sp.; P. tinctorum; Ramalina pacifica; Teloschistes flavicans 167 S. minima (Auersw.) S.I. Ahmed & Cain Zeroviella mandschurica Canoparmelia texana; Heterodermia diademata; H. podocarpa; Lobaria kurokawae; Parmotrema hababianum; P. tinctorum; Phaeophyscia hispidula; Punctelia rudecta; Ramalina conduplicans; Usnea sp. Li et al. (2007) Tripathi et al. (2014), Suryanarayanan et al. (2017) 168 S. muskokensis (Cain) S.I. Ahmed & Cain 169 Sporothrix sp. Zeroviella mandschurica Li et al. (2007) Dermatocarpon miniatum; Parmelia sp.; Ramalina Li et al. (2007) sinensis 170 Sydowia polyspora (Bref. & Cladonia sp.; Stereocaulon sp. Petrini et al. (1990) Tavel) E. Müll. (= Hormonema dematioides Lagerberg et Melin) 171 Taeniolella sp. Teloschistes flavicans Govinda Rajulu et al. (2020) 172 Talaromyces sp. Leptogium askotense; Lobaria kurokawae Tripathi et al. (2014) Canoparmelia texana; Heterodermia podocarpa; Suryanarayanan et al. (2017) Leptogium askotense; Lobaria kurokawae; Parmotrema hababianum; Phaeophyscia hispidula; Punctelia rudecta Porina tetracerae Maduranga et al. (2018) Xanthoparmelia angustiphylla Yuan et al. (2018) Parmotrema tinctorum; Ramalina pacifica Govinda Rajulu et al. (2020) 173 T. funiculosus (Thom.) Diorygma hieroglyphicum Samson 174 T. tratensis Manoch, Lecanora sp. Dethoup & N. Yilmaz 175 T. verruculosus (Peryronel) Peltigera praetextata; Xanthoparmelia taractica Samson, N. Yilmaz, Frisvad & Seifert (= Penicillium verruculosum Peyronel) Padhi et al. (2019) Jayakumar et al. (2016) Girlanda et al. (1997) 11 Table 1 Continued. Fungi Host lichen 176 Teichospora striata (Kohlm. Umbilicaria sp. & Volkm. -Kohlm.) Jaklitsch & Voglmayr (= Floricola striata Kohlm. & VolkmKohlm.) 177 Thielavia sp. Melanelia sorediata; Parmelia sp.; Punctelia borreri Heterodermia incana; Polyblastidium dendriticum 178 Tolypocladium sp. (= Peltigera praetextata; Xanthoparmelia taractica Chaunopycnis sp.) Parmelia sp. Cladonia sp.; Stereocaulon sp. 179 T. cylindrosporum W. Gams Lethariella zahlbruckneri 180 T. geodes W. Gams Peltigera praetextata; Xanthoparmelia taractica 181 T. inflatum W. Gams (= T. Peltigera praetextata; Xanthoparmelia taractica niveum (Rostr.) Bissett) 182 Torula sp. Cladonia sp.; Stereocaulon sp. 183 Tremella sp. #Not specified – list of several lichens given 184 Trichobotrys sp. Dermatocarpon miniatum; Melanelia sorediata 185 Trichoderma sp. Roccella montagnei Cladonia coniocraea; Melanelia sorediata; Punctelia borreri Phaeophyscia hispidula; Punctelia rudecta Pamelia sp. 186 T. hamatum (Bon.) Bain. Cladonia sp.; Stereocaulon sp. 187 T. harzianum Rifai Cladonia sp.; Stereocaulon sp.; Xanthoparmelia taractica Peltigera praetextata Heterodermia flabellata; Parmotrema crinitum; P. thomsonii; Physcia dilatata Heterodermia flabellata; Parmotrema crinitum; P. thomsonii; Physcia dilatata Heterodermia flabellata; Parmotrema crinitum; P. thomsonii; Physcia dilatata; Usnea sp. 188 T. koningii Oud. Cladonia sp.; Stereocaulon sp. 189 T. polysporum (Link ex Peltigera praetextata; Xanthoparmelia taractica Pers.) Rifai 190 T. viride Pers. ex S. F. Gray Cladonia sp.; Stereocaulon sp. Peltigera praetextata; Xanthoparmelia taractica Cladonia fruticulosa; Parmotrema tinctorum; Ramalina arabum; Usnea stigmatoides 191 Trichophyton roseum E. Parmotrema graynum Bodin 192 Umbelopsis vinacea (Dixon- Cladonia sp.; Stereocaulon sp. Stew.) Arx (= Mortierella vinacea Dixon-Stewart) 193 Verticillium sp. Cladonia sp.; Stereocaulon sp. 194 Xylaria sp. Leptogium saturninum Canoparmelia texana; Heterodermia diademata; H. podocarpa; Leptogium askotense; Lobaria kurokawae; Parmotrema hababianum; P. tinctorum; Phaeophyscia hispidula; Punctelia rudecta; Ramalina conduplicans; Usnea sp. Opegrapha medusulina 195 X. apiculata Cooke Arthonia sp. 196 X. arbuscula Sacc. Lobaria discolor; L. japonica 197 X. castorea Berk. Roccella montagnei 198 X. feejeensis (Berk.) Fr. Opegrapha medusulina 199 X. grammica (Mont.) Mont. Menegazzia sp. Reference Li et al. (2016) Li et al. (2007) Vinayaka et al. (2016) Girlanda et al. (1997) Hu et al. (2017) Petrini et al. (1990) Li et al. (2015b) Girlanda et al. (1997) Girlanda et al. (1997) Petrini et al. (1990) Zhang et al. (2016) Li et al. (2007) Suryanarayanan et al. (2005) Li et al. (2007) Tripathi et al. (2014) Padhi and Tayung (2015) Petrini et al. (1990) Petrini et al. (1990) Girlanda et al. (1997) Suryanarayanan et al. (2005) Li et al. (2007) Tripathi et al. (2014) Petrini et al. (1990) Girlanda et al. (1997) Petrini et al. (1990) Girlanda et al. (1997) Vinayaka et al. (2016) Tripathi et al. (2014) Petrini et al. (1990) Petrini et al. (1990) Wu et al. (2011) Tripathi et al. (2014), Suryanarayanan et al. (2017) Maduranga et al. (2018) Govinda Rajulu et al. (2020) Oh et al. (2020) Maduranga et al. (2018) Maduranga et al. (2018) Kim et al. (2018b) 12 Table 1 Continued. Fungi 200 X. hypoxylon (L.) Grev. 201 X. primorskensis Y.M. Ju, H.M. Hsieh, Lar. N. Vassiljeva & Akulov 202 X. psidii J.D. Rogers & Hemmes 203 X. vaporaria Berk. Host lichen Parmotrema reticulatum Reference Suryanarayanan et al. (2005), Tripathi et al. (2014) Hypotrachyna crenata; Parmotrema sp.; Ramalina Govinda Rajulu et al. (2020) pacifica; Usnea sp. Amandinea sp. Maduranga et al. (2018) Parmotrema cristiferum; Ramalina arabum; R. Vinayaka et al. (2016) pacifica 204 Zygosporium sp. Parmotrema tinctorum Govinda Rajulu et al. (2020) Note: #List of 17 species utilized by Zhang et al. (2016) for isolation of ELF. Acarospora molybdina (Ach.) Trevis., Allantoparmelia alpicola (Th. Fr.) Essl., Cetraria delisei (Bory ex Schaer.) Nyl. (= Cetrariella delisei (Bory ex Schaer.) Kärnefelt & A. Thell), Cladonia arbuscula (Wallr.) Flot., C. borealis S. Stenroos, C. pocillum (Ach.) O.J. Rich., Nephromopsis nivalis (L.) Divakar, A. Crespo & Lumbsch (= Flavocetraria nivalis (L.) Kärnefelt & A. Thell), Ochrolechia frigida (Sw.) Lynge, Peltigera canina (L.) Willd., Placynthium asperellum (Ach.) Trevis., Pseudephebe pubescens (L.) M. Choisy, Stereocaulon alpinum Laurer, S. botryosum Ach., S. vesuvianum Pers., Umbilicaria aprina Nyl., U. arctica (Ach.) Nyl., U. torrefacta (Lightf.) Schrad. *status of the species is unconfirmed under the genus. The ELFs identified till date belongs to three phyla, 32 orders, 70 families and 112 genera (Table 2, according to Wijayawardene et al. 2020). Majority of them belong to Ascomycota, followed by Basidiomycota, and very few belong to Mucoromycota. Within the Ascomycota, the classes Dothideomycetes and Sordariomycetes represent the largest number of ELFs reported so far. Alternaria, Aspergillus, Chaetomium, Cladosporium, Curvularia, Fusarium, Penicillium, Trichoderma, and Xylaria are some of the most common genera. Alternaria alternata, Aspergillus flavus, A. niger, Neocosmospora solani, and Trichoderma harzianum are the most frequently occurring ELF species. Table 2 Classification of endolichenic fungi (sensu Wijayawardene et al. 2020). Phylum Ascomycota Class Dothideomycetes Order Family Botryosphaeriales Botryosphaeriaceae Capnodiales Cladosporiaceae Dissoconiaceae Mycosphaerellaceae Dothideales Dothideaceae Saccotheciaceae Dothideales genus incertae sedis Kirschsteiniotheliales genus incertae sedis Pleosporales Biatriosporaceae Coniothyriaceae Corynesporascaceae Didymellaceae Dothideomycetes genera incertae sedis Eurotiomycetes Chaetothyriales Eurotiales Genus Endomelanconiopsis; Lasiodiplodia Cladosporium Ramichloridium Phaeoramularia; Ramularia Sydowia Aureobasidium Rhizosphaera Taeniolella Biatriospora Ochrocladosporium Corynespora Ascochyta; Epicoccum; Paraboeremia; Phoma Didymosphaeriaceae Paraphaeosphaeria; Spegazzinia Periconiaceae Periconia Pleosporaceae Alternaria; Curvularia Phaeosphaeriaceae Ophiosphaerella; Phaeosphaeria Sporormiaceae Preussia; Sporormiella Teichosporaceae Teichospora Torulaceae Torula Bahusakala; Heteroconium Herpotrichiellaceae Cladophialophora; Exophiala; Phialophora Aspergillaceae Aspergillus; Penicillium Thermoascaceae Paecilomyces 13 Table 2 Continued. Phylum Class Order Family Trichocomaceae Genus Talaromyces; Thysanophora; Byssochlamys Onygenales Arthrodermataceae Trichophyton Onygenaceae Chrysosporium Lecanoromycetes Lecanorales Ramalinaceae Scutula Leotiomycetes Chaetomellales Chaetomellaceae Chaetomella Helotiales Amorphothecaceae Myxotrichum Dermateaceae Pezicula Mollisiaceae Phialocephala Myxotrichaceae Oidiodendron Ploettnerulaceae Cadophora; Cylindrosporium Sclerotiniaceae Botrytis; Monilinia Helotiales genus incertae sedis Cryptocline Thelebolales Pseudeurotiaceae Pseudeurotium; Pseudogymnoascus Leotiomycetes family and genus incertae Hamatocanthoscypha Chalara sedis ceae Pezizomycetes Pezizales Tarzettaceae Geopyxis Saccharomycetales Dipodascaceae Saccharomycetes Geotrichum Sordariomycetes Amphisphaeriales Apiosporaceae Arthrinium; Nigrospora Sporocadaceae Broomella; Pestalotiopsis Coniochaetales Coniochaetaceae Coniochaeta Diaporthales Cytosporaceae Cytospora Diaporthaceae Diaporthe Gnomoniaceae Plagiostoma Glomerellales Glomerellaceae Colletotrichum Plectosphaerellaceae Verticillium Hypocreales Bionectriaceae Acremonium; Cylindromonium Cordycipitaceae Cordyceps Hypocreaceae Trichoderma Nectriaceae Aphanocladium; Calonectria; Cosmospora; Fusarium; Neocosmospora; Neonectria Ophiocordycipitaceae Purpureocillium; Tolypocladium Stachybotryaceae Myrothecium Microascales Ceratocystidaceae Berkeleyomyces Microascaceae Scopulariopsis Ophiostomatales Ophiostomataceae Sporothrix Sordariales Chaetomiaceae Chaetomium; Humicola; Thielavia Lasiosphaeriaceae Apiosordaria Sordariaceae Neurospora; Sordaria Xylariales Graphostromataceae Biscogniauxia Hypoxylaceae Daldinia; Hypoxylon Xylariaceae Nemania, Xylaria Zygosporiaceae Zygosporium Sordariomycetes genus incertae sedis (?) Papulospora Ascomycota genera incertae sedis Bispora; Gilmaniella; Trichobotrys Basidiomycota Agaricomycetes Cantharellales Ceratobasidiaceae Rhizoctonia Agaricales Schizophyllaceae Schizophyllum Bartheletiomycetes Russulales Bondarzewiaceae Heterobasidion; Spiniger Cystobasidiomycetes Cystobasidiales Cystobasidiaceae Cystobasidium Tremellomycetes Tremellales Tremellaceae Tremella Trichosporonales Trichosporonaceae Apiotrichum Polyporales Polyporaceae Cerrena Meripilaceae Rigidoporus Mucoromycota Mucoromycetes Mucorales Mucoraceae Mucor Rhizopodaceae Rhizopus Umbelopsidomycetes Umbelopsidales Umbelopsidaceae Umbelopsis 14 ELF host diversity and ecology The ELFs are not host-specific and behave as generalists (Chagnon et al. 2016). Studies have shown that certain endolichenic fungal species are widespread and occur in different lichen species (Tripathi et al. 2014). The ELF communities, their incidence, diversity, and composition reflect the interplay of climatic patterns, geographic separation, host type, and host lineage (U’Ren et al. 2012). Their composition depends on the environment of the lichens and the chemical ambience of the thalli (Arnold and Herre 2003, Suryanarayanan and Thennarasan, 2004). In addition, the altitude and the geographical location also affect the ELF composition as disclosed by 18S rDNA sequencing study (Wang et al. 2016). The compilation of ELF studies carried out to date shows that a total of 114 species of lichens belonging to 22 families were explored for ELFs (see name list). In few cases, the host species are partially identified (Petrini et al. 1990, Kannangara et al. 2008, Chen et al. 2014, 2019, Samanthi et al. 2015a, Wang et al. 2017, Govinda Rajulu et al. 2020) or unidentified (Wang et al. 2012, Kim et al. 2014, 2018a,b, Li et al. 2015a, Xie et al. 2016, Li et al. 2018). The present study shows that only a fraction of the 20000 species of lichens (Lücking et al. 2017) have been explored for their ELF. A majority of the host lichens studied for their ELF belong to the macro lichens (foliose and fruticose), probably because they are easy to handle and identify. Among them, members of families Parmeliaceae, Physciaceae, Peltigeraceae and Cladoniaceae have been studied more intensely. Also, species of Parmotrema were most preferred for isolation of ELF (Name list. Lichen hosts of endolichenic fungi). It is clear from the species diversity of ELF that most of the taxa (Alternaria, Aspergillus, Chaetomium, Cladosporium, Curvularia, Fusarium, Penicillium, Trichoderma, and Xylaria) are generalists that occur as saprotrophs (Arnold 2009). Their invasion into lichen thallus and association with the photobiont of the lichen suggests that they form an opportunistic interaction. It is speculated that the high level of fungal diversity may be due to highly porous and heterogeneous nature of the lichen thalli (Stone et al. 2012). Petrini et al. (1990) categorized ELFs into three ecological groups, of which the first one comprised of the species known as soil and litter inhabitants. Several of these belonged to genera frequently identified as plant pathogens or plant symbionts. The other groups reported as epiphytes or pathogens of higher plants (Dettki 1998, Arnold 2003). Several examples can be cited to show ELFs are not specific to lichens. Alternaria alternata isolated from a large number of lichens found in angiosperm Hedychium spicatum Ham ex Sm. (Sarma et al. 2020) and gymnosperm Pinus tabulaeformis Carr. (Guo et al. 2004). Cladosporium cladosporioides is also isolated from the leaves of medicinally important pteridophyte Huperzia serrata (Thunb. ex Murray) Trev. (Zhang et al. 2011). Name list. Lichen hosts of endolichenic fungi Acarosporaceae Acarospora molybdina (Ach.) Trevis. Arthoniaceae Arthonia sp. Arthonia antillarum (Fée) Nyl. Arthonia parantillarum Aptroot Caliciaceae Dirinaria consimilis (Stirt.) D.D. Awasthi Cladoniaceae Cladonia sp. C. arbuscula (Wallr.) Flot. C. borealis S. Stenroos, C. grisea (Ahti) Trass C. coniocraea (Flӧrke) Spreng. C. evansii Abbayes C. fruticulosa Kremp. 15 Name list continued. C. ochrochlora Flӧrke C. pocillum (Ach.) O.J. Rich. Stereocaulon sp. Stereocaulon alpinum Laurer S. botryosum Ach. S. vesuvianum Pers. Collemataceae Leptogium askotense D.D. Awasthi L. burnetiae Dodge L. saturninum (Dicks.) Nyl. Graphidaceae Diorygma hieroglyphicum (Pers.) Staiger & Kalb Hydnaceae Multiclavula sp. Lecanoraceae Lecanora sp. Lobariaceae Lobaria discolor (Bory) Hue Lobaria japonica (Zahlbr.) Asahina Ochrolechiaceae Ochrolechia frigida (Sw.) Lynge Parmeliaceae Allantoparmelia alpicola (Th. Fr.) Essl. Bulbothrix meizospora (Nyl.) Hale Canoparmelia texana (Tuck.) Elix & Hale Cetraria islandica (L.) Ach. Cetraria delisei (Bory ex Schaer.) Nyl. Cetrelia sp. C. braunsiana (Müll. Arg.) W.L. Culb. & C.F. Culb. Evernia prunastri (L) Ach. Flavoparmelia caperata (L.) Hale Hypotrachyna cirrhata (Fr.) Divakar et al. H. crenata (Kurok.) Hale H. nepalensis (Taylor) Divakar et al. Lethariella zahlbruckneri (Du Rietz) Krog. Melanelia sorediata (Ach.) Goward & Ahti Menegazzia sp. Nephromopsis nivalis (L.) Divakar et al. Parmelia sp. Parmelinella wallichiana (Taylor) Elix & Hale Parmotrema sp. P. austrosinense (Zahlbr.) Hale P. cristiferum (Taylor) Hale P. critinum (Ach.) Choisy P. graynum (Hue) Hale P. hababianum (Gyeln.) Hale P. nilgherrense (Nyl.) Hale P. praesorediosum (Ny) Hale P. ravum (Krog Swinscow) Sérus. P. reticulatum (Taylor) Choisy 16 Name list continued. P. thomsonii (Stirt.) A. Crespo, Divakar & Elix P. tinctorum (Despr. ex Nyl.) Hale Pseudevernia intensa (Nyl.) Hale & W.L. Culb. Pseudephebe pubescens (L.) M. Choisy Punctelia borreri (Turner) Krog P. rudecta (Ach.) Krog. P. subrudecta (Nyl.) Krog Usnea sp. U. aciculifera Vain. U. cavernosa Tuck. U. galbinifera Asahina U. longissima Ach. U. stigmatoides G. Awasthi Xanthoparmelia angustiphylla (Gyelnik) Hale. X. taractica (Kremp.) Hale Peltigeraceae Lobaria kurokawae Yoshim. L. retigera (Bory) Trevis. Peltigera canina (L.) Willd. P. elisabethae var. mauritzii (Gyeln.) J.C. Wei P. praetextata (Flӧrke ex Sommerf.) Zopf Pseudocyphellaria sp. Ricasolia quercizans (Michx.) Stizenb. Pertusariaceae Pertusaria laeviganda Nyl. Physciaceae Amandinea sp. Dirinaria picta (Sw.) Clem. & Shear Heterodermia diademata (Taylor) D.D. Awasthi H. flabellata (Fée) D.D. Awasthi H. hypochraea (Vain.) Swinsc & Krog H. incana (Stirt.) D.D. Awasthi H. obscurata (Nyl.) Trevis. H. podocarpa (Bél) D.D. Awasthi Phaeophyscia hispidula (Ach.) Moberg Physcia sp. P. dilatata Nyl. Polyblastidium dendriticum (Pers.) Kalb Pyxine cocoes (Sw.) Nyl. Placynthiaceae Placynthium asperellum (Ach.) Trevis. Porinaceae Porina tetracerae (Ach.) Müll. Arg. Pyrenulaceae Pyrenula sp. P. ochraceoflava (Nyl.) R.C. Harris P. parvinuclea (Meyen & Flot.) Aptroot Ramalinaceae Bacidina sp. Ramalina sp. 17 Name list continued. R. arabum (Dill. ex Ach.) Meyen & Flot. R. calicaris (L.) Rӧhl. R. conduplicans Vain. R. pacifica Asahina R. sinensis Jatta Roccellaceae Opegrapha arabica (Müll. Arg.) Vain. O. medusulina Nyl. Roccella montagnei Bél. Teloschistaceae Teloschistes flavicans (Sw.) Norman Zeroviella mandschurica (Zahlbr.) S.Y. Kondr. & Hur Umbilicariaceae Umbilicaria sp. U. aprina Nyl. U. arctica (Ach.) Nyl. U. torrefacta (Lightf.) Schrad. Verrucariaceae Dermatocarpon miniatum (L.) W. Mann. Conclusion Lichens establish successful symbiotic associations for over 600 million years (Yuan et al. 2005). However, the origin and evolution of ELFs is not well-understood as lichens, and their components do not preserve well as fossils. Some of the fossil lichen thalli obtained from the Lower Devonian period showed evidence of endolichenic fungal association, which suggests that ELF association existed since 415 million years ago. Such a long association of these fungi with lichens indicate that they have evolved a successful strategy for survival within the lichen thallus (Honegger et al. 2013). Apart from being poorly understood for their diversity and ecology, the ELFs are still unexplored treasures for biomolecules which could lead to the development of novel drugs. More studies are needed with culture-dependent and culture-independent approaches to estimate the true diversity of ELFs. Furthermore, lichens in different environments must be explored to understand their distribution patterns, the influence by abiotic and biotic factors, and host recurrence of ELFs. The estimated diversity of ELFs is undocumented while many sequences deposited in GenBank typically provide limited taxonomic details. Furthermore, the interaction of ELF with the host lichenized fungi as well as with the co-occurring symbionts needs to be studied using modern techniques such as isotopologue profiling. This result will be useful to understand the cross-relationship between the different constituencies of the microbiome of lichens (Götz et al. 2010). The culture-independent techniques and high-throughput sequencing methods are helpful in taxonomic placement, understanding ecological role and evolutionary details of endophytes. Acknowledgments We are thankful to Director, CSIR-NBRI, Lucknow for providing laboratory facilities and partial financial support under project OLP101 and to Council of Scientific and Industrial Research, New Delhi, for financial support under JRF/SRF Fellowship. (CSIR-NBRI manuscript number CSIR-NBRI_MS/2020/08/02). References Arnold AE, Mejía LC, Kyllo D, Rojas EI, Maynard Z, Robbins N, Herre EA. 2003- Fungal endophytes limit pathogen damage in a tropical tree. 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