Abstract
During our macrofungal survey, we discovered a Candolleomyces species that did not conform to previously described taxa within the genus. Morpho-anatomical characteristics and phylogenetic analyses based on nrITS, LSU, and tef-1α confirmed the novelty and distinct phylogenetic placement of Candolleomyces lahorensis. Candolleomyces lahorensis is characterized by an umbonate, grayish-brown pileus with uplifted margins, grayish-brown lamellae, light-gray stipe, variable shapes of cheilocystidia such as ellipsoid, globose, and ovoid-pedunculate. Currently, eighty-one species of the genus Candolleomyces are known worldwide; however, with the inclusion of this species, the total number increases to eighty-two. Morpho-anatomical descriptions, phylogenetic analysis results, and comparison with phylogenetically allied species are provided.
References
- Ahmad, S., Iqbal, S.& Khalid, A.N. (1997) Fungi of Pakistan. Sultan Ahmad Mycological Society of Pakistan, Department of Botany, University of the Punjab, Quaid-e-Azam campus, Lahore, pp. 1–248.
- Ahmad, I., Haqnawaz, M., Niazi, A.R.& Khalid, A.N. (2024) A new species of Candolleomyces (Psathyrellaceae) from Pakistan. Phytotaxa 658 (3): 280–288. https://doi.org/10.11646/phytotaxa.658.3.6
- Asif, M., Izhar, A., Niazi, A.R.& Khalid, A.N. (2022) Candolleomyces asiaticus sp. nov. (Psathyrellaceae, Agaricales), a novel species from Punjab, Pakistan. European Journal of Taxonomy 826: 176–187. https://doi.org/10.5852/ejt.2022.826.1845
- Bruns, T.D. (1995) Thoughts on the processes that maintain local species diversity of ectomycorrhizal fungi. Plant Soil 170: 63–73. https://doi.org/10.1007/BF02183055
- Bhunjun, C.S., Niskanen, T., Suwannarach, N., Wannathes, N., Chen, Y.J., McKenzie, E.H., Maharachchikumbura, S.S., Buyck, B., Zhao, C.L., Fan, Y.G. & Zhang, J.Y. (2022) The numbers of fungi: are the most speciose genera truly diverse?. Fungal Diversity 114 (1): 387–462. https://doi.org/10.1007/s13225-022-00501-4
- Cao, B., Phurbu, D., Ralaiveloarisoa, A., Liimatainen, K., Niskanen, T., Ramírez-Cruz, V., Bradshaw, A.J., Dentinger, B.T.M., Ramírez-Guillén, F., Cortés-Pérez, A., Guzmán-Dávalos, L., Villalobos-Arámbula, A.R., Castro-Jauregui, O., Santos da Silva, P., Day, R., Davoodian, N., Lebel, T., Castellano, M., May, T.W., Toome, M., Vasey, J., Hofer, K., Thangavel, R., Braithwaite, M., Braithwaite, L., Dutta, A.K., Chattopadhyay, P., Roy, N., Tanti, B., Biswas, P.R., Arumugam, E., Kezo, K., Kaliyaperumal, M., Murugadoss, R., Ji, J.-X., Kakishima, M., Cooper, J., Nuytinck, J., Luo, Z.-L., Wang, W.-P., Zhang, X., Wen, T.-C., Jia, A.-L., Fan, X.-L., Jiang, N., Jiao, N., Zhang, Y., Manawasinghe, I.S., Li, H., Xiong, Y.-R., Zhou, T., Tan, Q.-L., Hoshino, T., Lagaet, P., De Lange, R., Verbeken, A., Tondeleir, L., Manz, C., Hampe, F., Zhao, C.-L., Yang, Y., Zhang, G.-Q., Dai, D.-Q., Tomšovský, M., Denchev, T.T., Denchev, C.M., Léveillé-Bourret, É., Kemler, M., Chen, Y.-P., Maharachchikumbura, S.S.N., Feng, Z.-X., Wang, L., Wang, Q.-M., Wan, S.-P., Yu, F.-Q., Liu, J.-K., Du, H.-Z., Wu, N., Hu, H.-L., Su, J.-Y., Wang, Z.-H., Hu, Y.-P., Yu, H., Wang, J., Yang, Y.-H., Dong, W., Shu, Y.-X., Zhao, H.-J., Wang, C.-Q., Xia, W.-X., Liao, C.-F., Doilom, M., Li, G.-J., Suwannarach, N., Senwanna, C., Gafforov, Y., Flakus, A., Suchan, T., Plata, O., Rodriguez-Flakus, P., Tamang, J., Acharya, K., Yang, H.-D., Jayawardena, R.S., Tiendrebeogo, A., Decock, C., Somda, I., Legrève, A., Yang, Y., Xiao, Y.-P., Zhou, X.-Z., Li, J.-X., Thongklang, N., Han, X.-X., Yang, W.-Q., Wang, S.-H., He, M.-Q., Hyde, K.D. & Zhao, R.L. (2025) Fungal diversity notes 1919–2016: taxonomic and phylogenetic contributions to fungal taxa. Fungal Diversity 132 (1): 425–651. https://doi.org/10.1007/s13225-025-00556-z
- Ediriweera, A.N., Voto, P., Karunarathna, S.C., Kumla, J., Thiyagaraja, V., Wijesooriya, M.K. & Jianchu, X. (2023) A new species and a new record in the Agaricales from Sri Lanka. Mycological Observations 6: 94.
- Gardes, M. & Bruns, T.D. (1993) ITS primers with enhanced specificity for basidiomycetes‐application to the identification of mycorrhizae and rusts. Molecular Ecology 2 (2): 113–118. https://doi.org/10.1111/j.1365-294X.1993.tb00005.x
- Hall, T.A. (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. In: Nucleic acids symposium series 41: 95–98.
- Haqnawaz, M., Niazi, A.R.& Khalid, A.N. (2023a) A study on the genus Candolleomyces (Agaricales: Psathyrellaceae) from Punjab, Pakistan. BMC Microbiology 23 (1): 1–8. https://doi.org/10.1186/s12866-023-02938-2
- Haqnawaz, M., Niazi, A.R.& Khalid, A.N. (2023b) Two new species of Xanthagaricus from Punjab, Pakistan. Phytotaxa 583 (2): 163–173. https://doi.org/10.11646/phytotaxa.583.2.4
- Haqnawaz, M., Usman, M., Fatima, N., Niazi, A.R.& Khalid, A.N. (2024a) Three new species of the genus Candolleomyces (Psathyrellaceae) from Pakistan. Plant Systematics and Evolution 310 (6): 42. https://doi.org/10.1007/s00606-024-01925-y
- Haqnawaz, M., Fatima, N., Niazi, A.R.& Khalid, A.N. (2024b) Two new species of Moniliophthora from Punjab, Pakistan. Phytotaxa 635 (2): 137–148. https://doi.org/10.11646/phytotaxa.635.2.3
- Haqnawaz, M., Khalid, M., Li, Q., Lu, P., Bibi, A., Naseer, A., Afshan, N.S., Niazi, A.R.& Khalid, A.N. (2025) Morphology and multi-locus phylogeny reveal four new Candolleomyces (Psathyrellaceae) species from Pakistan. MycoKeys 121: 143. https://doi.org/10.3897/mycokeys.121.157282
- Han, X.X., Phurbu, D., Ma, G.F., Li, Y.Z., Mei, Y.J., Liu, D.M., Lin, F.C., Zhao, R.L., Thongklang, N. & Cao, B. (2024) A taxonomic study of Candolleomyces specimens from China revealed seven new species. Journal of Fungi 10 (7): 499. https://doi.org/10.3390/jof10070499
- Hu, Y., Karunarathna, S.C., Li, H., Galappaththi, M.C., Zhao, C.L. & Kakumyan, P. (2022) The impact of drying temperature on basidiospore size. Diversity 14 (4): 239. https://doi.org/10.3390/d14040239
- Hongsanan, S., Khuna, S., Manawasinghe, I.S., Tibpromma, S., Chethana, K.W.T., Xie, N., Bagacay, J.F.E., Calabon, M.S., Chen, C., Doilom, M., Du, H.Y., Gafforov, Y., Huang, S.K., Li, J.X., Luangharn, T., Luo, Z.L., Opiña, L.A.D., Pem, D., Sadaba, R.B., Singh, R., Tan, Q., Tang, S.M., Wang, W.P., Wen, T.C., Xia, G., Zhao, Q., Bhunjun, C.S., Cao, B., Chen, Y.P., de Silva, N.I., Dai, D.Q., Dong, W., Du, T.Y., Ferreira-Sá, A.S., Gao, Y., Gui, H., Han, L.S., Han, M.Y., Han, X.X., Jayawardena, R.S., Khyaju, S., Kumar, S., Lei, L., Leonardo-Silva, L., Li, H., Li, Y.X., Liao, C.F., Liu, J.W., Liu, X.F., Lu, L., Lu, W.H., Luo, M., Maharachchikumbura, S.S.N., Meng, Q.F., Mi, L.X., Norphanphoun, C., Peng, X.C., Su, H.L., Tennakoon, D.S., Thiyagaraja, V., Tun, Z.L., Wijayawardene, N.N., Xavier-Santos, S., Xiong, Y.R., Xu, R.F., Yadav, S., Yang, T., Yang, Y.H., Yarasheva, M., Zeng, X.Y., Zhang, H., Zhang, G.Q., Zhang, X., Zhao, H.J., Zhao, R.L. & Zheng, D.G. (2025) Mycosphere Notes 521–571: A special edition of fungal biodiversity to celebrate Kevin D. Hyde's 70th birthday and his exceptional contributions to Mycology. Mycosphere 16 (2): 2002–2180. https://doi.org/10.5943/mycosphere/16/2/1
- Izhar, A., Asif, M., Khan, Z. & Khalid, A.N. (2023) Introducing two new members of the genus Candolleomyces (Agaricales, Psathyrellaceae) from Punjab, Pakistan. Plant Systematics and Evolution 309 (5): 40. https://doi.org/10.1007/s00606-023-01876-w
- Jabbar, M., Nasar-u-Minallah, M. & Yusoff, M.M. (2024) Predicting the impact of land use changes on thermal environment in Lahore, Pakistan. Implications for urban planning. Geography, Environment, Sustainability 17 (1): 95–109. https://doi.org/10.24057/2071-9388-2023-2862
- Khan, M., Azeem, M., Ijaz, H., Arshad, W., Tufail, A. & Jabeen, S. (2025) Molecular phylogeny and morphological characterization revealed Candolleomyces mianwaliensis Psathyrellaceae, Agaricales, a new species from Punjab, Pakistan. Nordic Journal of Botany e04858. https://doi.org/10.1002/njb.04858
- Kiran, M., Javeed, P., Haqnawaz, M., Fatima, N. & Khalid, A.N. (2025) Two novel species of Candolleomyces (Psathyrellaceae) from Pakistan. Phytotaxa 691 (3): 231–246. https://doi.org/10.11646/phytotaxa.691.3.1
- Li, Q., Haqnawaz, M., Niazi, A.R.& Khalid, A.N. (2025) Phylogenetic studies on the genus Candolleomyces (Psathyrellaceae, Basidiomycota) occurring in the bed of the Indus River, Punjab, Pakistan, reveal three new species. MycoKeys 112: 165. https://doi.org/10.3897/mycokeys.112.141766
- Largent, D. (1986) How to Identify Mushrooms to Genus Vol. I. Macroscopic Features. Mad River Press, Eureka, CA, USA, 166 pp.
- Nie, C., Wang, S.N., Tkalčec, Z., Yan, J.Q., Hu, Y., Ge, Y., Na, Q., Zeng, H., Ding, H., Huo, G., Pošta, A., Pradeep, C.K. & Mešić, A. (2022) Coprinus leucostictus rediscovered after a century, epitypified, and its generic position in Hausknechtia resolved by multigene phylogenetic analysis of Psathyrellaceae. Diversity 14 (9): 699. https://doi.org/10.3390/d14090699
- Nayana, P.K. & Pradeep, C.K. (2023) New species and new record of Candolleomyces (Psathyrellaceae) from India. Botany 101 (11): 472–484. https://doi.org/10.1139/cjb-2023-0066
- Nayana, P.K. & Pradeep, C.K. (2025) New species and new geographic record of Candolleomyces (Psathyrellaceae, Agaricales) from India. Phytotaxa 691 (1): 34–48. https://doi.org/10.11646/phytotaxa.691.1.3
- Nylander, J.A.A. (2004) Mr Modeltest v2. Program distributed by the author. Evolutionary Biology Centre, Uppsala University, Uppsala.
- Örstadius, L., Ryberg, M. & Larsson, E. (2015) Molecular phylogenetics and taxonomy in Psathyrellaceae (Agaricales) with focus on psathyrelloid species: introduction of three new genera and 18 new species. Mycological progress 14: 1–42. https://doi.org/10.1007/s11557-015-1047-x
- Rathnayaka, A.R., Tennakoon, D.S., Jones, G.E., Wanasinghe, D.N., Bhat, D.J., Priyashantha, A.H., Stephenson, S.L. & Tibpromma, S. (2025) Significance of precise documentation of hosts and geospatial data of fungal collections, with an emphasis on plant-associated fungi. New Zealand Journal of Botany 63 (2–3): 462–489. https://doi.org/10.1080/0028825X.2024.2381734
- Ronquist, F., Teslenkop, M., Van Der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–542. https://doi.org/10.1093/sysbio/sys029
- Shaheen, S., Jaffer, M., Khan, F., Hussain, K., Hanif, U., Younis, S., Ilyas, S. & Ishtiaq, S. (2018) Morpho-phylonogical assessment of medicinal flora of district Lahore, Pakistan, based on LM and SEM. Microscopy Research & technique 81: 1397–1405. https://doi.org/10.1002/jemt.23096
- Tian, X., Bao, D., Han, J., Dai, D., Lu, Y., Han, M., Zheng, D., Tarafder, E., Acharya, K., Dovana, F., Boccardo, F., Latha, K.P.D., Manimohan, P., Raj, K.N.A., Nayana, P., Kunjan, P.C., Kaliyaperumal, M., Murugadoss, R., Yuan, Q., Zhao, C., Abhinav, Singh, R., Rajeshkumar, K.C., Sruthi, O.P., Ansil, P.A., Khalid, A.N., Naseer, A., Afshan, N.U.S., Khurshid, R., Ashraf, U., Verma, R.K., Elgorban, A.M., Chattopadhyay, P., Dutta, A.K., Jayasiri, S.C., Gautam, A.K., Suwannarach, N., Kumla, J., Liu, R., Li, Q., Yang, J.-Y., Karunarathna, S.C. & Tibpromma, S. (2026) Morphology and multi-locus phylogeny reveal 23 fungal taxa from terrestrial habitats. Mycology 1–73. https://doi.org/10.1080/21501203.2025.2594942
- Usman, M. & Khalid, A.N. (2020) Termitomyces acriumbonatus sp. nov. (Lyophyllaceae, Agaricales) from Pakistan. Phytotaxa 477: 217–228. https://doi.org/10.11646/phytotaxa.477.2.6
- Vellinga, E.C. (2001) Agaricaceae. In: Noordeloos, M.E., Kuyper, T.W. & Vellinga, E.C. (Eds.) Flora Agaricina Neerlandica 5. A.A. Balkema Publishers, Rotterdam, 220 pp.
- Vilgalys, R. & Hester, M. (1990) Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. Journal of Bacteriology 172: 4238–4246. https://doi.org/10.1128/jb.172.8.4238-4246.1990
- Wächter, D. & Melzer, A. (2020) Proposal for a subdivision of the family Psathyrellaceae based on a taxon-rich phylogenetic analysis with iterative multigene guide tree. Mycological Progress 19 (11): 1151–1265. https://doi.org/10.1007/s11557-020-01606-3
- White, T.J., Bruns, T., Lee, S.J.W.T. & Taylor, J. (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. PCR protocols: a guide to methods and applications 18 (1): 315–322. https://doi.org/10.1016/B978-0-12-372180-8.50042-1
- Yang, K.L., Xiong, X., Luo, Z., Huang, Y., Huang, R., Chen, H., H., Lin, J.Y., Yang, Z.L., Li, G.-M. & Jia, X. (2025) Human Activity Impacts on Macrofungal Diversity: A Case Study of Grazing in Subtropical Forests. Journal of Fungi 11 (10): 749. https://doi.org/10.3390/jof11100749
- Zheng, D.G., Voto, P., Tibpromma, S., Lu, W.H., Han, M.Y., Kumla, J., Suwannarach, N., Dai, D.-Q., Zhang, L.-J., Elgorban, A.M., Chukeatirote, E. & Karunarathna, S.C. (2025) Candolleomyces yunnanensis (Psathyrellaceae, Agaricales), a novel species from Yunnan Province, China. Phytotaxa 711 (3): 281–292. https://doi.org/10.11646/phytotaxa.711.3.3
