The Southeast Asian Journal of Tropical Biology Vol. 32 No. 3, 2025: 385 - 393 DOI: 10.11598/btb.2025.32.3.2547 ISSN: 0215-6334 | e-ISSN: 1907-770X 385 NEW RECORD OF THE BIOLUMINESCENT FUNGUS Mycena chlorophos (Berk. & M.A.Curtis) Sacc. in INDONESIA Fathansah Nugeraha1, Silva Ristiana Haryadi1, Wahyu Aji Mahardhika1, Oktan Dwi Nurhayat2, Ivan Permana Putra1* 1Department of Biology, Faculty of Mathematics and Natural Sciences, IPB University, Bogor 16680, Indonesia 2Research Center for Applied Microbiology, National Research and Innovation Agency (BRIN), Bogor 16914, Indonesia ARTICLE HIGLIGHTS • Glowing mushrooms were found at night in the forest of IPB University and showed a bright green light in the dark. • Examination of their morphological features and phylogeny material confirmed they are the species Mycena chlorophos. • This is the first confirmed record of Mycena chlorophos in Indonesia, extending its known distribution worldwide. Article Information Received : 30 May 2025 Revised : 17 September 2025 Accepted : 22 September 2025 *Corresponding author, e-mail: ivanpermanaputra@apps.ipb.ac.id Research Paper ABSTRACT The majority of fungi exhibiting bioluminescence are classified within the Mycenaceae with Mycena chlorophos among the most widely recognized species. Despite its broad distribution in the Asia- Pacific region, no confirmed record of M. chlorophos has previously been documented in Indonesia. During a late-night mushroom foraging in IPB University Campus Forest, glowing basidiomata were encountered and subjected to detailed taxonomic investigation. The current study aimed to clarify the taxonomical identity of our specimens based on morphological and molecular analyses. Fresh fruiting bodies were examined for macroscopic and microscopic features, and the internal transcribed spacer (ITS) region of rDNA was amplified using ITS1/2 primers and analyzed phylogenetically which confirmed our specimens belong to Mycena chlorophos. By morphology, our specimen possess convex pileus with shallow depression at center, viscid, shiny, adnexed lamellae with series of lamellae, ellipsoid and smooth basidiospores. The pileus and hymenophore glow vividly with a bright greenish hue in the darkness. The BLAST analysis indicated a 98% similarity between our specimen and M. chlorophos from China, as the top matches. Furthermore, the phylogenetic tree placed our specimens within the M. chlorophos clade with a 100% bootstrap support value. Our specimen was in subclade with materials from Malaysia. This study represents the first confirmed record of M. chlorophos in Indonesia, extending its known geographic range and contributing to the growing documentation of bioluminescent fungi in this biodiverse yet underexplored region. Keywords: bioluminescent, fungi, Indonesia, Mycenaceae, new record INTRODUCTION Bioluminescence is a product of a chemical reaction in an organism that can produce light (Bechara & Stevani 2018). Bioluminescent organisms are found across diverse biological groups, encompassing marine life such as squids, jellyfish, dinoflagellates, as well as land- dwelling species like fireflies, specific worms, and fungi (Deane et al. 2015; Bessho-Uehara et al. 2020; Ramesh & Meyer-Rochow 2021). Among terrestrial bioluminescent organisms, fungi are particularly noteworthy due to their ecological importance in forest environments and their promising potential in fields such as biotechnology, environmental assessment, and eco- friendly illumination (Ke & Tsai 2022; Cortés-Pérez et al. 2023). The variation in the luminescence of mushrooms is found in the fruiting body and mycelium of the mushroom, depending on the species (Cortés-Pérez et al. 2023; Oba & Hosaka 2023). Copyright (c) 2025@author(s). https://doi.org/10.11598/btb.2025.32.3.2547 https://creativecommons.org/licenses/by-nc-nd/4.0/ BIOTROPIA Vol. 32 No. 3, 2025 386 Bioluminescent fungi are widely distributed across the world and are found in tropical and temperate forests (Dauner et al. 2021). Desjardin et al. (2008) identified three distinct evolutionary lineages of bioluminescent fungi: the Omphalotus lineage (Omphalotaceae), the Armillaria lineage (Physalacriaceae), and the Mycenoid lineage, predominantly consisting of Mycenaceae species. According to Cortés-Pérez et al. (2023), Mycenaceae contain at least 65 bioluminescent species, which are classified under the genera  Favolaschia  (Pat.) Pat., Filoboletus Henn., Mycena (Pers.) Roussel, Panellus P. Karst., Resinomycena Redhead & Singer, and Roridomyces Rexer (Karunarathna et al. 2020; Nimalrathna et al. 2022). In Mycenaceae, Mycena is the most significant genus, with around 40 species (Chang et al. 2020; Oliveira et al. 2021). Mycena chlorophos is reported from Australia, Brazil, Borneo, India, Japan, Malaysia, Papua New Guinea, and Sri Lanka (Cortés-Pérez et al. 2019; Arya et al. 2021; Oba & Hosaka 2023; Koli et al. 2024; Lu et al. 2024). Mycena chlorophos is originally described from the Bonin Island, Japan (Desjardin et al. 2010). However, the published descriptions of M. chlorophos that document taxonomically informative features are based on material collected from Sri Lanka, Malaysia, Borneo and Brazil. Despite its high biodiversity, knowledge of bioluminescent mushrooms in Indonesia remains limited. The only Indonesian authors that reported M. chlorophos are Armadhan et al. (2023), but the report lacked taxonomic evidence. Another Mycena species (M. illumans) was discovered decades ago on Java Island, Indonesia (Hennings 1903). To date, there is no subsequent report regarding the taxonomy and distribution of M. chlorophos in Indonesia. During our regular night foray of bioluminescent fungi at IPB University Campus Forest, some glowing basidiomata were encountered, collected, and studied. This study provides an updated account of M. chlorophos in Indonesia. MATERIALS AND METHODS Specimen Collection Fresh fruiting bodies of the bioluminescent fungus growing on dead bamboo were obtained in 2024 at the Lowland Bamboo Forest, IPB University, Bogor, West Java, Indonesia (Fig. 1), located at 6°32›51.0» S and 106°43’09.4” E (Latitude -6.547497°; Longitude 106.719280°) during a late-night mushroom hunt. The collection and monitoring were done three times (September– December 2024) at the same location. The fruiting bodies were photographed in situ (with and without light/dark condition) using Canon EOS 1500D. Photographs of bioluminescence were taken in complete darkness in the forest and laboratory. Ecological data, including coordinates, substrate type, and nearby vegetation, were recorded. Figure 1 Sampling location of Mycena chlorophos at bamboo forest of IPB University Source: Photo edited from Google Earth. New Record of Mycena chlorophos - Nugeraha et al. 387 Morphological Analysis The morphological characteristics of fresh basidiomata were examined both in situ and in the Mycology Laboratory, Department of Biology, Faculty of Mathematics and Natural Sciences, IPB University, Indonesia. The macroscopic characters assessment was carried out following Putra (2021). Micromorphological features of the basidium, cystidia, hyphae, cheilocystidia, and spores, including their shape, size, and coloration were investigated using OlympusBX-63 light microscope. Specimen identification was carried out using relevant taxonomic references (Arya et al. 2021; Oba & Hosaka 2023). amplification was carried out according to Putra et al. (2024). The sequencing was performed using 1st Base Malaysia. The sequence data were processed using ChromasPro software for assembly, while the finalized alignments were submitted to GenBank (https://www.ncbi.nlm.nih.gov/) to obtain accession numbers. Homology comparisons were conducted through the Basic Local Alignment Search Tool (BLAST) in NCBI, aligning sequences against existing data for taxonomic verification. The BLASTN results were examined to identify the highest sequence similarity (http://blast.ncbi.nlm. nih.gov/Blast.cgi). Sequences from the selected BLAST results in this study (bold), 16 fungal sequences from Chew et al. (2014), GenBank records, and additional relevant sequences were included to construct the phylogenetic tree (Table 1). Mycena noctilucens was designated as the outgroup following Chew et al. (2014). The phylogenetic tree was generated using the maximum likelihood (ML) method in MEGA X software (Kumar et al. 2018). Default parameters in MEGA X were applied for the ML analysis, and bootstrap values of 60% or greater were reported. Molecular Analysis DNA isolation from the fresh specimens was conducted using the Qiagen Dneasy Plant Mini Kit following the manufacturer’s protocol. DNA amplification was performed using a Thermo Scientific Arktik Thermal Cycler (ThermoFisher Scientific). The amplification process employed ITS 5 (5’-GGA AGT AAA AGT CGT AAC AAG G-3’) and ITS 4 (5’-TCC TCC GCT TAT TGA TAT GC-3’) primers (White et al. 1990). PCR Table 1 Species, collection code, and genbank accession numbers used in this study Species Voucher/Isolate ITS accession number Reference Country Mycena chlorophos CT15101401 MH400938 Wei et al. (2024) China Mycena chlorophos CT151014 MH400939 Wei et al. (2024) China Mycena chlorophos ACL051 KJ206965 Chew et al. (2014) Malaysia Mycena chlorophos ACL055 KJ206967 Chew et al. (2014) Malaysia Mycena chlorophos MMRD-20 KF010856 Chew et al. (2014) India Mycena chlorophos 305759 AB512312 Chew et al. (2014) Japan Mycena chlorophos IPB02 PQ821413 This study Indonesia Mycena illuminans ACL161 KJ206975 Chew et al. (2014) Malaysia Mycena illuminans ACL175 KJ206976 Chew et al. (2014) Malaysia Mycena illuminans ACL212 KJ206980 Chew et al. (2014) Malaysia Mycena deeptha DM334g JX481737 Chew et al. (2014) India Mycena amicta 189f JF908394 Chew et al. (2014) Italy Mycena amicta AFTOL-ID 1908 DQ490645 Chew et al. (2014) USA Mycena nocticaelum ACL258 KC507796 Chew et al. (2014) Taiwan Mycena nocticaelum ACL272 KJ206987 Chew et al. (2014) Taiwan Mycena kentingensis YSH-2014 KC507796 Chew et al. (2014) Taiwan Mycena stylobates 455 JF908439 Chew et al. (2014) Italy Mycena adscendens 35m JF908420 Chew et al. (2014) Italy Mycena noctilucens ACL054 KJ206966 Chew et al. (2014) Malaysia http://blast.ncbi.nlm.nih.gov/Blast.cgi http://blast.ncbi.nlm.nih.gov/Blast.cgi BIOTROPIA Vol. 32 No. 3, 2025 388 RESULTS AND DISCUSSION Taxonomy Photographs of Mycena chlorophos were presented in Figures 2 – 4. Current Name Mycena chlorophos Berk. and M. A. Curtis Sacc., Syll. Fung. (Abellini) 5:301 (1887) Synonym Agaricus chlorophos Berk. and M.A Curtis 1860. Basidiomata encountered in dead bamboo as saprobes in a solitary to scattered manner. Pileus 5 – 12.5 mm in diameter, viscid with a thick gluten, parabolic to convex in maturity, white to cream at straight margin and center is light brown, shallow depression at center, striate, sticky; Lamellae free, entire, with series of lamellulae, concolorous with pileus margin. Stipe 6.8 – 14.1 mm x 0.7 – 1.6 mm central, cylindric, hollow, smooth, glabrous, basal tomentum with a white surface. Basidia 19.24 – 22.40 x 4.43 – 6.25 µm, clavate, hyaline, thin- walled, with 2 – 4 sterigmata, Basidiospores 6.5 – 7.6 x 5.1 – 6.1 µm, smooth, ellipsoid, thin-walled. Cheilocystidia 41.46 – 44.52 x 9.53 – 12.59 µm, fusoid-ventricose, hyaline, thin walled. Hyphae on pileal trama septate, hyaline. Pileipellis ixotrichoderm, hyaline to brownish color, thin walled. Luminescence emitting strongly in pileus and lamellae, weakly in stipe, greenish light in dark. Location of specimens Dramaga, Bogor, West Java, Indonesia, 6°32’51.0” S; 106°43’09.4” E, 145 m asl, on decayed bamboo tree, 2024, collected by Fathansah Nugeraha. Figure 2 Field photograph of Mycena chlorophos Notes: A = upper side of pileus; B = underside of pileus; C = developmental stages of basidiomata; D, E, F = Green light from the fruiting bodies. New Record of Mycena chlorophos - Nugeraha et al. 389 Figure 3 Microscopic features of Mycena chlorophos Notes: A = Hypahe on trama; B = Pileipellis marginal cell (arrows); C = Oleiferous hyphae of pileipellis (arrow). BIOTROPIA Vol. 32 No. 3, 2025 390 Figure 4 Microscopic feature of Mycena chlorophos Notes: A = Cystidia (arrows); B = Basidia (arrow); C = Basidiospores. Molecular Analysis BLAST analysis identified a high degree of similarity (98%) between our specimens and M. chlorophos specimens from China, ranking them among the top matches. In line with the BLAST results, the phylogenetic tree (Fig. 5) constructed from ITS sequences placed specimen IPB02 in the clade of M. chlorophos with a 100% Bootstrap value. The phylogenetic tree displayed that our specimen was closely related to materials from Malaysia collections.  The Mycenaceae family comprises numerous species capable of bioluminescence fungi (Cortés- Pérez et al. 2019; Heinzelmann et al. 2024) globally, including M. chlorophos. This study represents the first comprehensive documentation of M. chlorophos in Indonesia. Our report is not only providing the basic information for taxonomy of bioluminescent fungi in Indonesia, but also the reference for future studies of this species. The distribution of M. chlorophos has recently been reported in several countries including China, Japan, Pacific islands, Sri Lanka (Lu et al. 2024), India (Arya et al. 2021; Koli et al. 2024), Malaysia (Chew et al. 2014), New Record of Mycena chlorophos - Nugeraha et al. 391 Figure 5 The phylogenetic tree depicting Mycena chlorophos Voucher PQ821413 and closely related species Notes: Bootstrap values > 60% are displayed at the corresponding nodes. The scale bar represents an evolutionary distance of 0.050. Southern Asia, Australia, and Brazil (Desjardin et al. 2010). However, the taxonomic understanding of bioluminescent mushrooms in Indonesia is limited. To date, Armadhan et al. (2023) are the only Indonesian researchers who have reported the presence of M. chlorophos from karst forest in Central Java, but their taxonomic evidence was inadequate to confirm the identification with confidence.  In the current study, the basidiomata of M. chlorophos were found colonizing decayed bamboo. Previous reports have noted that M. chlorophos lives saprobically on decayed bamboo (Arya et al. 2021; Koli et al. 2024.). Armadhan et al. (2023) also obtained M. chlorophos from weathered logs in Indonesia, though they did not specify the tree species. Our specimens were observed growing solitary to scattered on bamboo, which was also covered by bryophytes. This observation is consistent with the report of M. cristinae from Brazil (Oliveira et al. 2021). Desjardin et al. (2010) reported M. chlorophos from topotypical growth on wood instead of bamboo. The specimens in our study emit the greenish light from whole basidiome including the cap, lamellae, and stipe-with the stipe exhibiting a weaker intensity. Several researchers have also reported that bioluminescent fungi can emit light from their mycelium (Weitz et al. 2001; Oba & Hosaka 2023; Perry et al. 2024). This remarkable biological feature is not only scientifically fascinating but also deeply rooted in cultural traditions. For instance, ethnomycological records cited by Havey (1957) in A History of Luminescence indicate that Georg Everard Rumph (1637 – 1706) documented the use of luminescent fungi by indigenous communities on Ambon Island, Maluku, Indonesia. These fungi were used as handheld lanterns, aiding individuals navigate forest paths at night and avoid disorientation. Mycena chlorophos is morphologically similar to M. illuminans based on several morphological BIOTROPIA Vol. 32 No. 3, 2025 392 characters: small basidiome, gelatinous pellicle on pileus, basal disk composed of inflated hyphae of stipe (Chew et al. 2014). However, the latter species usually found in palm trees. In Indonesia, M. illuminans was originally described and found on Java Island (Hennings 1903). Due to the morphological similarity, M. chlorophos and M. illuminans once are acknowledged as synonyms (Pegler 1986; Maas Geesteranus 1992). The morphology of M. chlorophos IPB02 is highly similar to the materials found in Malaysia. Macromorphologically, they appear similar, but the size of the pileus and stipe in the species found in Malaysia (Chew et al. 2014) are larger compared to those found in this study. In addition, other specimens found in India (Arya et al. 2021) showed a larger pileus size, more gelatinous on the pileus surface, and a longer stipe compared to the species found in Malaysia and Indonesia. The basidiospores dimension of specimens from Indonesia and Malaysia are slightly smaller to those reported from India.  The classification of Mycena s. l. is highly intricate, and its subgeneric categorization typically depends on morphological traits. The identification of fungi based on morphology is considered challenging, especially when non- experts are dealing with some fungi (Stephenson 2010; Raja et al. 2017). Therefore, in our study we combined the morphological and molecular analyses. The BLAST analysis revealed a strong genetic resemblance between our specimens and M. chlorophos specimens from China. Based on ITS tree, our specimen exhibited phylogenetic affinity (Bootstrap value of 100%) with the clade consisting of specimen of M. chlorophos from China, India, Japan, and Malaysia. In the subclade of M. chlorophos, specimen IPB02 was displayed as closely related to materials originating from Malaysia (Bootstrap value of 89%). Our ITS phylogenetic tree is in line to those reported (LSU, ITS, and RPB2 tree) by Chew et al. (2013; 2014) and confirming that the phylogenetic tree clearly separates M. chlorophos and M. illuminans into two distinct clades, thus verifies that they are not synonymous taxa. CONCLUSION This study provides the first confirmed record of Mycena chlorophos in Indonesia, verified through morphological characterization and molecular phylogenetic analysis. The findings extend the known geographic range of this bioluminescent species within the Asia-Pacific region and highlight Indonesia’s potential as a reservoir for undocumented fungal diversity. 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