Submitted:
30 June 2026
Posted:
01 July 2026
You are already at the latest version
Abstract
Keywords:
1. Introduction
2. Materials and Methods
2.1. Site Description
2.2. Sampling Design
2.3. Microclimatic Measurements
2.4. Preparation of Lichen thalli
2.5. Isolation of Fungal Strains
2.6. Data Analysis
3. Results
3.1. Composition of Endolichenic and Rock-Surface Communities
3.2. Species Richness and Density of Fungal Isolates in Lichen thalli
3.3. Effect of Geographic Region, Season, and Lichen Species on the Characteristics of Endolichenic Communities
4. Discussion
5. Conclusions
Supplementary Materials
Author Contributions
Funding
Acknowledgments
Conflicts of Interest
References
- Li, W-C.; Zhou, J.; Guo, S-Y.; Guo, L-D. Endophytic fungi associated with lichens in Baihua mountain of Beijing, China. Fungal Divers. 2007, 25, 69-80.
- Peršoh, D.; Rambold, G. Lichen-associated fungi of the Letharietum vulpinae. Mycol. Progr. 2012, 11, 753-760. [CrossRef]
- Zhang, T/; Wei, X-L.; Zhang, Y-Q.; Liu, H-Y.; Yu, L-Y. Diversity and distribution of lichen-associated fungi in the Ny-Ålesund Region (Svalbard, High Arctic) as revealed by 454 pyrosequencing. Sci Rep. 2015. Available from: www.nature.com/scientificreports. [CrossRef]
- Suryanarayanan, T.S.; Thirunavukkarasu, N. Endolichenic fungi: the lesser known fungal associates of lichens. Mycology 2017, 8, 189-196. [CrossRef]
- Suryanarayanan, T.S.; Govindarajulu, M.B.; Rajamani, T.; Tripathi, M.; Joshi, Y. Endolichenic fungi in lichens of Champawat district, Uttarakhand, northern India. Mycol. Progr. 2017, 16, 205-211. [CrossRef]
- Chakarwarti, J.; Nayaka, S.; Srivastava, S. Diversity of endolichenic fungi – A review. Asian J. Mycol. 2020, 3(1), 488–509. [CrossRef]
- Oh, S-Y.; Yang, J.H.; Woo, J-J.; Oh, S-O.; Hur, J-S. Diversity and distribution patterns of endolichenic fungi in Jeju Island, South Korea. Sustainability 2020, 12, 3769. [CrossRef]
- Muggia, L.; Quan, Yu.; Gueidan, C.; Al-Hatmi, A.M.S.; Grube, M.; de Hoog, S. Sequence data from isolated lichen-associated melanized fungi enhance delimitation of two new lineages within Chaetothyriomycetidae. Mycol. Progr. 2021, 20, 911–927. [CrossRef]
- Santiago, K.A.A.; Dela Cruz, T.E.E.; Ting, A.S.Y. Diversity and bioactivity of endolichenic fungi in Usnea lichens of the Philippines. Czech Mycol. 2021, 73. 1–19. [CrossRef]
- Oh, S-Y.; Yang, J.H. New Trichoderma species and six unrecorded species of endolichenic fungi isolated from lichens. Mycobiology 2025, 53, 435–449. [CrossRef]
- Gao, H.; Zou, J.; Li, J.; Zhao, H. Endolichenic fungi: a potential treasure trove for discovery of special structures and bioactive compounds. Stud. Nat. Prod. Chem. 2016, 48, 347-397. [CrossRef]
- Kellog, J.; Raja, HA. Endolichenic fungi: a new source of rich bioactive secondary metabolites on the horizon. Phytochem. Rev. 2017, 16, 271–293. [CrossRef]
- Singh, B.N.; Upreti, DK.; Gupta, VK.; Dai, X-F.; Jiang, Y. Endolichenic fungi: a hidden reservoir of next generation biopharmaceuticals. Trends Biotechnol. 2017, 35, 808-813. [CrossRef]
- Tripathi, M.; Joshi, Y. Endolichenic Fungi: Present and Future Trends. Springer Nature Pte Ltd.: Singapore, 2019. [CrossRef]
- Wethalawe, A.N.; Alwi, Y.V.; Udukala, D.N.; Paranagama, P.A. 2021. Antimicrobial compounds isolated from endolichenic fungi: a review. Molecules 2021, 26, 3901. [CrossRef]
- Chang, R.; Yan, Z.; Jiang, J.; Wang, Y.; Si, H.; Bose, T.; Miao, C. Four novel endolichenic fungi from Usnea spp. (Lecanorales, Parmeliaceae) in Yunnan and Guizho, China: Taxonomic description and preliminary assessment of bioactive potentials. MycoKeys 2025, 118, 55-80. [CrossRef]
- Grishkan, I.; Temina. M. Interior of saxicolous lichens on different types of rocks as a habitat for microfungal communities in Upper Galilee, Israel. Acta Mycol. 2019, 54, 1123. [CrossRef]
- Grishkan, I.; Temina. M. Composition and diversity of endolichenic microfungal communities from saxicolous lichens at Nahal Boker, the central Negev Desert, Israel. Fungal Ecol. 2023, 61. [CrossRef]
- Grishkan, I.; Temina. M. Diversity of microfungal communities inside saxicolous lichens from Nahal Oren, Mount Carmel, Israel. Czech Mycol. 2023, 75, 15–34. [CrossRef]
- Atlas of Israel. 3rd ed.; Surveys of Israel: Tel Aviv, Israel, 1985.
- Temina, M.; Kondratyuk, S.Y.; Zelenko, S.D.; Wasser, S.P.; Nevo, E. Lichen-forming, lichenicolous and allied fungi of Israel. Wasser, S.P.; Nevo, E., Eds. A.R.G. Ganter Verlag: Ruggell, Liechtenstein, 2005.
- Bills, G.F.; Christensen, M.; Powell, M.; Thorn, G. Saprobic soil microfungi. In Biodiversity of Microfungi. Inventory and Monitoring Methods. Mueller, G.M; Bills, G.F; Foster, M.S, Eds., Elsevier Academic Press: California, USA, 2004, pp. 271-302.
- Grishkan, I.; Kidron, G.J. Vertical divergence of cultured microfungal communities at the Hallamish dune field, Western Negev Desert, Israel. Geomicrobiol. J 2017, 34, 706–721. [CrossRef]
- Grishkan. I. Ecological stress: Melanization as a response in fungi to radiation. In Extremophiles Handbook. Horikoshi, K.; Antranikian, G.; Bull, A.; Robb, F.; Stetter, K, Eds., Springer DE, 2011, pp. 1136-1148. [CrossRef]
- Treseder, K.K.; Lennon, J.T. Fungal traits that drive ecosystem dynamics on land. Microbiol. Mol. Biol. Rev. 2015, 79, 243-262. [CrossRef]
- Lawrence, D.P.; Rotondo, F.; Gannibal, P.B. Biodiversity and taxonomy of the pleomorphic genus Alternaria. Mycol. Progr. 2016, 15, 3. [CrossRef]
- Ellis, M.B.; Ellis, J.P. 1997. Microfungi on Land Plants. An Identification Handbook. Richmond Publishing Co. Ltd., The England.
- Mapperson, R.R.; Kotiw, M.; Davis, T.A.; Dearnaley, J.D.W. The diversity and antimicrobial activity of Preussia sp. endophytes isolated from Australian dry rainforests. Curr. Microbiol. 2014, 68: 30–37. [CrossRef]
- Schulz, B.; Wanke, U.; Draeger, S.; Aust, H-J, Endophytes from herbaceous plants and shrubs: effectiveness of surface sterilization methods. Mycol. Res. 1993, 97, 1447–1450.
- Domsch, K.H.; Gams, W.; Anderson, T.H. Compendium of Soil Fungi, 2nd revised ed.; Academic Press: NY, USA, 2007.
- Volz, P.A.; Ellanskaya, I.A.; Grishkan, I.; Wasser, S.P.; Nevo E. Soil Microfungi of Israel. Subramanian, C.V.; Wasser, S.P., Eds., A.R.A. Ganter Verlag K.-G.: Ruggell, Liechtenstein, 2001.
- Su, H.; Li, Q-R.; Kang, J-Ch.; Wen, T-Ch.; Hyde, K. Rosellinia convexa sp. nov. (Xylariales, Pezizomycotina) from China. Mycoscience 2016, 57, 164-170.
- Ju, Y-M.; Rogers, J.D. New and interesting Biscogniauxia taxa, with a key to the world species. Mycol. Res. 2001, 105, 1123–1133. [CrossRef]
- Sutton, B.C. The Coelomycetes. Commonwealth Mycological Institute: Kew, Surrey, England. 1980.
- Gams, W. Cephalosporium-Artige. Schimmelpilze (Hyphomycetes). G. Fischer: Stuttgart, Germany, 1971.
- Muggia, L.; Grube, M. Fungal diversity in lichens: from extremotolerance to interactions with algae. Life 2018, 8, 15–28. [CrossRef]
- Kim, H.; You, Y-H.; Yoon, H.; Seo, Y.; Kim, Y-E.; Choo, Y-S.; Lee, I-J.; Shin, J-H.; Kim, J-G. Culturable fungal endophytes isolated from the roots of coastal plants inhabiting Korean east coast. Mycobiology 2014, 42, 100-108. [CrossRef]
- Chen, L.; Guo, W.; Zheng, Y.; Zhou, J.; Liu, T.; Chen, W.; Liang, D.; Zhao, M.; Zhu, Y.; Wu, Q.; Zhang, J. Occurrence and characterization of fungi and mycotoxins in contaminated medicinal herbs. Toxins 2020, 12, 30. [CrossRef]
- Poulsen-Silva, E.; Carolina, Otero, M.; Diaz-Cornejo, S.; Atala, C.; Fuentes, J.A.; Gordillo-Fuenzalida, F. 2025. Secondary metabolites of lichens: the untapped biomedical and pharmaceutical potential of antimicrobial molecules. Fungal Biol. Rev. 2025, 51. [CrossRef]
- Lange, O.L.; Kilian, E.; Zingler, H. Water vapor uptake and photosynthesis of lichens: performance differences in species with green and blue-green algae as phycobionts. Oecologia 1986, 71, 104–110. [CrossRef]





| Location | Coordinates | Altitude, m Above Sea Level | Annual Precipitation | Dominant Vegetation |
|---|---|---|---|---|
| Upper Galilee, Mount Meron (UG1) |
N 33°00.902’ E 35°23.606’ |
890 | 900 mm | Trees: Quercus calliprinos, Q. boissieri, Arbutus andrachne, Eriolobus trilobatus; shrubs: Pistacia terebinthus; grasses: Juniperus oxycedrus |
| Upper Galilee, Wadi Amud (UG2) |
N 32°58.805’ E 35°26.668’ | 635 | 800 mm | Trees: Q. calliprinos. Calicotome villosa, bushes: Rhamnus alaternus, Sarcopoterium spinosum; grasses: Cyclamen persicum, Anemone coronaria |
| Mount Carmel (MC) |
N 32°44.026’ E 35°02.311’ |
500 | 600 mm | Trees: Q. calliprinos; shrubs: Pistacia palaestina, Cistus salviifolius; Calicotome villosa |
| Negev Desert, Sede Boker (ND1) |
N 30°52.478’ E 34°47.013’ |
470 | 95 mm | Semishrubs: Artemisia herba-alba, Zygophyllum dumosum, Hammada scoparia. |
| Negev Desert, Borot Lots (ND2) |
N 30°30.263’ E 34°36.511’ |
1000 | 95 mm | Semishrubs: Artemisia sieberi; grasses: Helianthemum vesicarium |
| Locality | Rock Temperature, oC | Air Humidity, % | ||||
|---|---|---|---|---|---|---|
| Spring | Winter | Summer | Spring | Winter | Summer | |
| Upper Galilee, Mount Meron |
15.4±0.3 | 10.4±0.4 | 36.9±0.1 | 69.2±0.1 | 62.3±0.5 | 57.8±0.2 |
| Upper Galilee, Wadi Amud |
21.6±0.1 | 19.5±0.3 | 39.8±0.2 | 67.2±0.6 | 79.4±1.4 | 40.7±0.7 |
| Mount Carmel | 21.6±0.2 | 17.1±0.3 | 44.3±0.2 | 56.0±0.1 | 65.9±0.9 | 54.2±0.1 |
| Negev Desert, Sede Boker |
34.2±0.3 | 18.2±0.3 | 50.5±0.4 | 57.3±0.1 | 56.4±0.2 | 56.1±0.2 |
| Negev Desert, Borot Lots |
42.6±0.2 | 24.2±0.3 | - | 57.3±0.1 | 47.3±1.6 | - |
| Species | Spring | Winter | Summer | |||||||||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| UG1 | UG2 | MC | ND1 | ND2 | UG1 | UG2 | MC | ND1 | ND2 | UG1 | UG2 | MC | ND1 | |
| Zygomycota | ||||||||||||||
|
Actinomucor elegans |
- | - | - | - | - | 0.4 | - | - | 0.15 | - | - | - | - | - |
| Mortierella humilis | - | - | - | - | - | - | 0.9 | - | - | - | 1.7 | - | - | - |
| Mucor hiemalis | - | - | - | 1.1 | - | - | 0.3 | - | - | - | - | - | - | - |
| M. plumbeus | 0.6 | - | - | - | - | - | 0.8 | - | - | - | - | - | - | - |
| M. racemosus | - | - | - | - | - | 1.7 | 0.8 | 1.7 | - | - | 1.7 | - | - | - |
| Rhizopus arrhizus | - | 1.1 | - | - | - | - | 1.7 | - | - | - | - | 0.8 | - | - |
| Ascomycota | ||||||||||||||
|
Acaulium acremonium |
- | 1.1 | - | - | - | - | - | - | - | - | - | - | - | - |
|
Alternaria alternata |
0.7 | 13.9 | 4.2 | 9.3 | 12.3 | - | 4.2 | 3.3 | 17.2 | 18 | - | 3.5 | 1.3 | 5.7 |
| A. atra | 1 | - | 7.2 | 11.5 | 10.3 | 2.2 | 5 | 3.1 | 21.2 | 26 | - | - | - | 13.3 |
| A. botrytis | - | - | - | - | - | - | 1.2 | - | - | - | - | - | - | - |
| A. chartarum | - | - | - | - | - | - | - | - | - | 0.8 | - | - | - | - |
| A. chlamydospora | - | 0.6 | - | - | 0.7 | - | - | - | - | 1.2 | - | - | - | - |
| A. chlamydo-sporigena | - | - | - | - | 0.4 | - | - | - | - | 0.8 | - | - | - | - |
| A. destruens1 | - | - | - | - | - | - | - | - | - | - | - | 3.5 | 9.6 | 1.1 |
| A. phragmospora | 0.6 | 1.2 | 1.7 | 6.8 | 4 | 4.5 | 3 | - | 9.6 | 0.25 | - | - | - | - |
| A. raphani | - | 0.5 | - | - | - | 0.5 | - | 0.5 | 0.35 | 0.35 | - | - | - | - |
|
Anthostomella limitata |
- | 25 | - | - | - | - | - | - | - | - | - | - | - | - |
|
Aphanocladium album |
- | - | - | 4.2 | - | 4.9 | - | - | - | - | - | - | - | - |
|
Apiospora phaeosperma |
- | - | - | - | - | - | 2 | - | - | - | - | - | - | - |
|
Ascochyta medicaginis |
- | - | - | 0.5 | - | 2.3 | - | - | - | - | - | - | - | - |
| Aspergillus flavus | - | - | - | 0.5 | - | - | 0.4 | - | - | - | - | - | - | - |
| A. fisheri1 | 1.2 | - | - | 0.6 | 0.7 | - | - | - | - | - | - | 0.8 | - | - |
| A. niger | 0.3 | 2.2 | - | - | - | 1.3 | 0.8 | - | 0.15 | - | 2 | - | 2.6 | - |
| A. ustus | - | 0.6 | - | - | - | - | 0.4 | - | 0.35 | - | - | - | - | - |
|
Biscognialixia meditterranea1 |
3.2 | 0.4 | - | - | 0.9 | - | - | - | - | - | - | - | - | - |
| Bisifusarium dimerum | - | - | 0.6 | - | - | - | - | - | - | - | - | - | 5 | - |
| Boeremia exigua | - | 0.6 | - | - | - | - | 0.6 | - | 3.6 | 1.6 | - | - | - | - |
| Camarosporium aequivocum | - | 2.8 | - | - | 0.8 | - | - | - | - | - | - | - | - | - |
|
Canariomyces notabilis |
- | - | - | - | - | - | - | - | - | - | 7 | - | - | - |
| Cephalotrichum stemonitis | - | - | - | - | - | 2.9 | - | - | - | - | - | - | - | - |
|
Chaetomium bostrichoides |
- | - | - | - | - | - | - | - | - | - | - | 2.2 | - | - |
| Ch. cochliodes | - | 1.7 | - | - | - | - | 0.35 | - | 1.3 | - | - | - | - | - |
| Ch. elatum | - | - | - | - | - | 7.8 | 3.4 | - | - | - | - | - | 1.9 | - |
| Ch. fusisporum | - | 3.3 | - | - | - | 0.35 | 0.7 | - | - | - | - | - | - | - |
| Ch. globosum | 0.8 | - | 3.1 | 3.3 | - | - | 0.7 | - | - | - | - | - | - | 1.1 |
| Ch. madrasense | - | - | 0.8 | - | - | - | - | - | - | - | - | - | - | - |
| Ch. strumarium | - | - | - | - | - | 1.5 | - | 3.3 | - | - | - | - | - | - |
| Ch. succinum | - | - | - | - | 0.5 | 0.4 | - | - | - | - | - | - | - | 0.8 |
| Chaetomium subaffine | 0.6 | - | - | - | - | - | - | - | 2 | - | - | - | - | - |
| Chaetomium sp. | - | - | - | - | - | - | - | - | - | - | - | - | 1.3 | 1.3 |
|
Chrysosporium pannorum |
- | - | - | - | - | - | - | - | - | 3.5 | - | - | - | - |
|
Cladorrhinum hyalocarpum |
1.1 | - | - | 1.6 | 6 | - | - | - | - | - | - | - | - | - |
| Cladosporium cladosporioides | 32.3 | 18.1 | 23.2 | 13.2 | 6.3 | 28.7 | 33.5 | 22.3 | 21.8 | 9.5 | 9.6 | 25.9 | 2.6 | 14.7 |
| C. sphaerospermum | - | - | - | - | - | - | - | 1.8 | - | - | - | - | - | - |
| Clonostachys rosea | 0.9 | 0.5 | - | - | - | - | - | - | - | - | - | - | - | - |
| Coniochaeta sp.2 | 2.1 | 6.6 | 1.8 | - | - | 4.5 | - | 0.5 | - | - | 12.7 | 0.8 | 9 | - |
| Corniculantispora aranearum1 | - | 7.8 | - | - | - | - | - | 1.4 | - | 3.3 | - | - | - | - |
|
Daldinia concentrica1 |
- | - | - | - | - | 6.8 | 1.2 | 2.6 | - | - | - | 13.1 | 1.3 | - |
| Diplodia seriata1 | - | 3.1 | - | 1.1 | - | - | - | - | - | - | - | - | - | - |
| Epicoccum nigrum | - | - | - | - | - | 0.35 | - | - | 0.35 | - | - | - | - | - |
| Exserohilum rostratum | - | - | 0.9 | - | - | - | - | - | - | 0.2 | - | - | - | - |
| Fusarium equiseti1 | - | - | 4.8 | - | - | 3.3 | 10.5 | 1.5 | 1.1 | 0.55 | 2.0 | 2.4 | - | - |
| F. longifundum1 | - | - | - | - | - | 2.1 | - | - | - | - | 2.1 | 3.6 | 10.3 | 5.7 |
| F. iranicum1 | - | - | 1.4 | - | - | - | - | - | - | - | 1.7 | - | - | - |
| F. oxysporum | 2.1 | 0.6 | - | - | 1 | - | 1.2 | - | - | - | - | - | - | - |
| F. redolens | - | - | - | - | - | 3.5 | 2.5 | - | 0.7 | - | - | - | - | - |
| F. sporotrichoides | - | - | - | - | - | 0.35 | 0.4 | 1.4 | - | - | - | - | - | - |
| F. tricinctum1 | - | 5.6 | 1.4 | - | - | - | 2.4 | 4.2 | 1.3 | 0.2 | - | - | - | 8.5 |
| Hansfordia ovalispora | - | 0.3 | - | 1 | - | - | - | - | - | - | - | - | - | - |
|
Monochaetia kansensis |
- | - | - | 0.9 | - | - | - | - | - | - | 1.9 | - | - | - |
|
Monodictys castaneae |
- | - | - | - | - | 2.2 | - | - | - | - | 1.9 | - | - | - |
| M. fluctuata | 3.3 | - | - | - | - | - | - | - | - | - | - | - | - | - |
| Neocucurbitaria cava | - | - | 2.1 | - | 3.4 | - | - | - | - | 1.2 | - | 5.6 | - | - |
|
Neoscytalidium dimidiatum |
- | 3.9 | - | - | - | - | - | - | - | - | - | - | - | - |
| Nigrospora oryzae | - | 1.7 | - | - | 5.7 | 5.7 | 2.1 | - | 1.4 | 3.2 | 6.3 | - | 1.9 | - |
|
Papulaspora pannosa |
- | - | 0.9 | - | - | - | 0.15 | - | - | - | - | - | - | - |
| Paraboeremia putaminum | - | - | - | 1.1 | - | - | - | - | - | - | - | - | - | - |
|
Parathielavia hyrcaniae |
- | - | - | - | - | - | - | 2.4 | - | 2.3 | 3.2 | 1.1 | 1.3 | - |
|
Penicillium brevicompactum |
- | 0.9 | 0.8 | - | - | - | - | - | - | 2.2 | - | - | - | - |
| P. glabrum | 1.2 | 0.5 | - | - | - | 1.7 | 0.6 | 1.7 | - | - | 6.5 | 1.7 | - | - |
| P. paxillii | - | - | - | - | - | - | - | 4.4 | - | - | - | - | - | - |
| P. restrictum | - | - | - | - | - | - | - | - | - | - | - | - | - | 4.1 |
| P. sacculum | - | - | - | - | - | 2.2 | - | - | - | - | - | - | - | 8.7 |
| P. simplicissimum | - | - | 2.5 | - | - | 0.35 | - | - | 0.2 | 2.2 | - | - | - | - |
| P. waksmanii | - | - | - | - | - | - | - | - | - | - | 5.3 | 4.1 | - | - |
| P. viridicatum | 1.8 | 1.6 | - | - | - | - | - | - | - | - | - | - | - | - |
| Pestalotiopsis guepinii | - | - | 2.4 | - | 0.7 | - | - | 6.7 | - | - | - | - | - | - |
|
Pseudothielavia terricola |
- | - | 0.5 | 7.7 | 1 | 8.3 | 5 | - | - | 1.2 | 1.7 | - | - | 1.1 |
| Rosellinia convexa1 | - | - | - | - | - | - | - | 3.3 | 1.4 | 0.8 | - | 1.6 | 1.9 | 10.5 |
| Scedosporium desertorum | - | - | - | - | - | - | - | - | - | 7.1 | - | 19.6 | - | - |
| Sordaria fimicola | 4.4 | 1.1 | 14 | 3.2 | - | 12.8 | 6.3 | 14.9 | 0.15 | 0.4 | 28.3 | 7.5 | 40.1 | 1.6 |
|
Sporormiella australis |
32.5 | 12.5 | 3.1 | 28.5 | 49.9 | - | - | - | - | 1 | - | - | 1.9 | 5.4 |
| S. minima | - | - | - | - | - | 2.6 | 4.2 | 8.9 | 6.8 | 9.1 | 7.1 | 6.9 | - | 4 |
|
Stachybotrys c hartarum |
- | - | - | - | - | - | - | 0.3 | 1.2 | 1 | 1.8 | - | - | 0.8 |
| Stenocarpella maydis | - | 1.1 | - | - | - | - | - | - | - | - | - | - | - | - |
| Subramaniula thielavioides | - | - | - | - | - | - | - | 1 | - | - | - | - | - | - |
|
Stemphylium vesicarium |
- | 0.6 | 7.2 | 1.1 | - | 0.5 | 2 | 3.1 | 3.8 | 1.3 | - | - | 1.9 | 1.6 |
| Talaromyces ruber | - | - | 1.2 | - | - | - | - | - | - | - | - | - | - | - |
| T. variabiles | - | - | - | - | - | - | - | - | - | 2.8 | - | - | - | - |
|
Triangularia setosa |
- | - | - | - | - | - | - | - | - | - | - | - | 1.9 | - |
| Trichocladium griseum | - | - | 0.8 | - | - | - | 0.4 | - | - | - | - | - | - | - |
|
Trichoderma koningii |
1.2 | 3.3 | 1 | - | - | - | 1 | 4.2 | - | - | - | 0.8 | 1.4 | - |
| Zasmidium cellare | - | 1.2 | - | - | - | - | - | - | - | - | - | - | - | - |
| Xylaria sp.2 | - | - | - | - | - | - | - | - | 0.15 | 0.8 | - | - | 2.6 | - |
| Overall number of species | 22 | 243 | 23 | 20 | 22 | 34 | 27 | 30 | 30 | 26 | 24 | 17 | 22 | 21 |
| Species | Spring | Winter | Summer | |||||||||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| UG1 | UG2 | MC | ND1 | ND2 | UG1 | UG2 | MC | ND1 | ND2 | UG1 | UG2 | MC | ND1 | |
| Zygomycota | ||||||||||||||
| Cunninghamella echinulata | - | - | - | - | - | - | - | 0.7 | 0.4 | - | - | - | - | - |
| Mortierella humilis | 4.7 | 4.4 | 0.5 | 0.5 | 1.7 | 4.9 | 4.8 | 0.7 | 5.5 | 1.4 | - | - | - | - |
| Mucor hiemalis | - | - | - | 0.3 | - | 0.4 | - | - | - | 0.45 | - | - | - | - |
| M. plumbeus | 0.1 | 0.55 | - | - | - | - | - | 0.7 | - | - | - | - | - | - |
| M. racemosus | - | - | - | - | - | 0.4 | 2.9 | - | - | - | 0.4 | 0.55 | - | 2 |
| Rhizopus arrhizus | 0.4 | 0.25 | 0.6 | 0.4 | 0.6 | 0.2 | 0.2 | 2.2 | 0.4 | - | 0.7 | 0.2 | - | - |
| Ascomycota | ||||||||||||||
|
Alternaria alternata |
7.8 | 9.3 | 19.6 | 28.5 | 31.6 | 2 | 1.7 | 7.6 | 9.7 | 26.5 | 6.4 | 11.2 | 10.5 | 22.3 |
| A. atra | 5 | 3.2 | 11.1 | 9.6 | 19.1 | 7.6 | 7 | 4.4 | 23.6 | 18.2 | 8.3 | 7.6 | 1.9 | 23.7 |
| A. chlamydospora | - | - | - | - | 0.2 | - | 0.4 | - | - | - | - | - | - | 0.15 |
| A. destruens1 | - | - | - | - | - | - | - | - | - | - | 0.4 | - | - | 1 |
| A. phragmospora | 0.5 | 0.13 | 1.5 | 0.3 | - | 3.8 | - | - | - | - | - | - | - | - |
| A. raphani | - | - | 2 | - | - | 0.2 | - | 0.9 | - | - | - | - | - | - |
|
Ascochyta medicaginis |
- | - | - | 0.4 | - | - | - | - | - | - | - | 0.3 | - | - |
| Aspergillus flavus | 0.13 | - | - | - | - | - | - | - | 0.13 | - | 0.3 | 0.3 | - | 0.8 |
| A. fisheri1 | 0.23 | - | - | - | - | - | - | - | - | - | 0.2 | 0.6 | - | 0.15 |
| A. nidulans | - | - | - | - | - | - | - | 1.9 | - | - | - | - | - | 0.8 |
| A. niger | 0.25 | 0.9 | 1.3 | 0.25 | 0.9 | 0.1 | 1.2 | 0.7 | 0.45 | 0.5 | 0.4 | 0.8 | 4.8 | 2.5 |
| A. ustus | 0.13 | - | - | - | - | 0.7 | 0.9 | - | - | - | 0.25 | 0.45 | 0.15 | - |
| Beauveria bassiana | - | 0.6 | - | - | - | - | - | - | - | - | - | 0.7 | - | - |
| Bisifusarium dimerum | - | - | - | - | - | - | - | 2.5 | - | - | - | - | - | - |
| Boeremia exigua | 0.3 | 1.9 | - | - | - | 0.23 | 0.85 | 0.7 | 5.2 | 0.13 | 0.13 | 0.7 | 0.65 | - |
| Camarosporium aequivocum | - | 0.35 | - | - | 1.3 | - | - | - | - | - | - | - | - | - |
|
Chaetomium fusisporum |
- | 0.55 | - | - | - | - | 0.15 | - | - | - | - | - | - | - |
| Ch. globosum | - | - | 0.2 | 0.13 | 0.3 | - | - | - | - | - | - | - | - | - |
| Ch. piluliferum | 0.8 | - | - | - | - | - | 0.55 | - | - | - | - | - | - | - |
| Chaetomium sp. | - | - | 0.7 | - | - | - | - | - | 0.35 | - | - | - | - | 1.9 |
| Cladosporium cladosporioides | 64.8 | 49.4 | 41.9 | 38.3 | 24.5 | 71 | 57.9 | 48 | 40.2 | 34.2 | 62.3 | 53.6 | 60.2 | 15.5 |
| Clonostachys rosea | 0.13 | - | 0.3 | - | - | 0.7 | 0.3 | 0.25 | - | - | 1.3 | - | - | - |
| Coleophoma empetri | 0.13 | - | - | - | 0.3 | - | - | - | - | - | - | - | - | - |
| Coniochaeta sp.2 | 1.6 | 1.5 | - | - | - | - | - | - | 0.15 | 1 | 0.7 | - | - | - |
|
Culvularia spicifera |
- | - | 0.3 | 0.13 | - | - | - | - | - | 0.13 | - | - | - | - |
|
Daldinia concentrica1 |
- | - | - | - | - | - | - | - | - | - | - | 0.1 | 0.15 | - |
| Diplodia seriata1 | - | 1.3 | 0.6 | - | - | - | - | - | - | - | - | - | - | 0.15 |
| Exserohilum rostratum | - | - | 0.2 | 0.25 | - | - | - | - | - | - | - | - | - | - |
| Epicoccum nigrum | - | - | - | 0.12 | - | - | 0.12 | - | 0.55 | - | - | - | - | - |
| Epicoccum sp. | - | 0.25 | - | - | - | - | - | - | - | - | - | - | - | 0.15 |
| Fusarium equiseti1 | - | 5 | 6.0 | 4.2 | 2.5 | 2.7 | 6.8 | 4.2 | 2.7 | 5.0 | - | 0.6 | - | 5.5 |
| F. longifundum1 | - | - | 0.3 | 0.7 | - | - | 1.4 | 0.7 | 2.7 | 2.7 | 3.4 | 4.2 | 10.7 | 13.2 |
| F. iranicum1 | - | - | - | - | - | - | - | - | - | - | 1.1 | - | 0.8 | - |
| F. oxysporum | 3.9 | 0.25 | 0.4 | 7.5 | - | 1.1 | 1 | 0.95 | - | 0.4 | 1.2 | 0.15 | 1.6 | - |
| F. tricinctum1 | 2.2 | 3.8 | 3.8 | 1.9 | 4.3 | 0.23 | 3.5 | 5.4 | 1.1 | 2.9 | 2.1 | 2.7 | 3.1 | 2.7 |
|
Lecanicillium psaliotae |
0.4 | 0.6 | 0.25 | 0.25 | - | 0.4 | - | - | - | - | 0.25 | - | - | - |
|
Monodictys fluctuata |
0.4 | - | - | 1.5 | - | - | - | - | - | - | - | - | - | - |
| Neocamarosporium betae | - | - | - | 0.13 | - | - | - | - | - | - | 0.3 | - | - | - |
| Nigrospora oryzae | 0.1 | - | - | - | 0.2 | - | - | - | 0.23 | 1.8 | 5.1 | - | - | - |
|
Penicillium aurantiogriseum |
1.2 | 0.6 | - | 2.2 | 3.4 | - | - | - | - | 3.2 | - | - | - | - |
| P. brevicompactum | - | - | 1.9 | - | - | - | 2.9 | - | - | 1.4 | - | - | - | - |
| P. scabrosum | - | - | - | 1.4 | - | - | - | - | - | - | - | - | - | - |
| P. sclerotiorum | - | - | - | - | - | 0.15 | 0.1 | - | - | - | - | - | - | - |
| P. simplicissimum | - | 2.6 | 0.5 | - | 2.4 | 2 | - | 1.5 | 1.6 | 7.7 | 0.5 | 1.1 | 0.65 | |
| Pestalotiopsis guepinii | - | 1.4 | - | 0.2 | 1.9 | - | - | 2.2 | - | - | - | - | 0.15 | - |
| Rosellinia sp.2 | - | - | - | - | - | - | - | - | - | - | 0.15 | 0.25 | - | - |
| Sordaria fimicola | - | - | 0.4 | - | - | - | - | 0.4 | - | 0.15 | 0.6 | 0.6 | 0.6 | - |
|
Sporormiella australis |
- | - | 0.6 | 1.7 | - | - | 0.15 | - | 0.35 | - | - | - | - | - |
|
Stachybotrys chartarum |
- | 4 | 0.35 | 0.25 | 0.6 | - | 0.25 | - | 0.5 | - | 0.13 | - | 0.5 | 0.15 |
|
Stemphylium vesicarium |
0.4 | 10.9 | - | 1 | 0.1 | - | 0.55 | - | 1.8 | 0.12 | 0.7 | 1.5 | 0.75 | 2.3 |
|
Talaromyces variabiles |
- | - | - | - | 2.8 | 0.5 | - | 0.2 | - | - | - | - | - | - |
| Trichocladium griseum | 0.5 | - | - | - | - | - | 0.9 | 6.9 | - | - | 0.8 | - | - | - |
|
Trichoderma koningii |
- | - | 1.4 | - | - | - | 0.8 | 2 | - | 1.1 | - | 0.15 | - | - |
| T. viride | 0.3 | 3 | 0.5 | - | - | - | - | - | 0.4 | - | - | - | 0.8 | - |
| Venturia carpophila | - | 5 | - | - | - | - | - | - | - | - | - | - | - | - |
| Immature fruit bodies | 0.1 | 0.35 | - | - | - | - | - | - | - | - | - | - | - | - |
| Site | Lichen Species | Spring | Winter | Summer | |||
|---|---|---|---|---|---|---|---|
| SR1 | CFU/spe-cimen | SR | CFU/spe-cimen | SR | CFU/spe-cimen | ||
| Upper Galilee, Mount Meron |
Circinaria calcarea | 2.2(6) | 6.0 | 2.7(8) | 3.5 | 2.8(11) | 4.8 |
| Lathagrium cristatum | 6.0(17) | 15.8 | 4.7(18) | 13.2 | 2.5(7) | 5.6 | |
| Variospora aurantia | 4.8(12) | 13.2 | 5.8(13) | 11.7 | 2.3(6) | 7.3 | |
| Upper Galilee, Wadi Amud |
Circinaria calcarea | 0.7(3) | 1.3 | 4.2(12) | 7.2 | 2.3(11) | 2.3 |
| Lathagrium cristatum | 6.3(20) | 15.6 | 5.5(19) | 14.2 | 4.3(14) | 6.8 | |
| Variospora aurantia | - | - | 6.8(17) | 19.3 | 2.5(8) | 6.3 | |
| Mount Carmel | Circinaria calcarea | 2.0(8) | 2.7 | 2.0(9) | 2.3 | 0.8(6) | 1.5 |
| Lathagrium cristatum | 4.7(13) | 10.8 | 4.2(11) | 6.5 | 2.7(10) | 4.3 | |
| Variospora aurantia | 3.8(15) | 6.1 | 7.2(16) | 21.8 | 2.7(9) | 3.7 | |
| Negev Desert, Sede Boker |
Circinaria calcarea | 1.0(6) | 1.8 | 4.2(11) | 8.3 | 2.0(6) | 2.3 |
| Lathagrium cristatum | 5.7(15) | 10.0 | 7.5(16) | 15.0 | 2.5(11) | 4.3 | |
| Variospora aurantia | 4.0(13) | 5.7 | 6.8(19) | 31.5 | 4.2(13) | 5.8 | |
| Negev Desert, Borot Lots |
Circinaria calcarea | 2.5(9) | 5.2 | 3.3(12) | 5.7 | - | - |
| Lathagrium cristatum | 4.3(11) | 8.2 | 6.3(15) | 13.8 | - | - | |
| Variospora aurantia | 3.0(10) | 10.0 | 6.3(18) | 27.7 | - | - | |
| Source | Species Richness per Specimen | CFU per Specimen | Contribution of Melanin-Containing spp. | Contribution of m-c spp. with Large Multicellular Spores | ||||
|---|---|---|---|---|---|---|---|---|
| DF | F Pr>F | DF | F Pr>F | DF | F Pr>F | DF | F Pr>F | |
| region | 2 | 0.90ns | 2 | 1.96ns | 2 | 1.76 ns | 2 | 6.27@ |
| season | 2 | 7.70@ | 2 | 13.85* | 2 | 0.18 ns | 2 | 0.77 ns |
| lichen species | 2 | 12.11* | 2 | 14.55* | 2 | 0.51 ns | 2 | 1.14 ns |
| region*season | 4 | 0.52 ns | 4 | 4.26@ | 4 | 1.83 ns | 4 | 1.64 ns |
| region*species | 4 | 1.43 ns | 4 | 0.74ns | 4 | 2.76 ns | 4 | 0.09 ns |
| season*species | 4 | 1.65 ns | 4 | 5.83@ | 4 | 2.52 ns | 4 | 1.92 ns |
| region*season* species |
8 |
1.20ns |
8 |
0.86ns |
8 |
0.76 ns |
8 |
0.63 ns |
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