Wild edible mushrooms are widely recognized as valuable natural food resources for their nutritional value and culinary importance. They are rich sources of proteins, dietary fiber, vitamins, minerals and numerous bioactive compounds that contribute to human health. In addition to their nutritional benefits, mushrooms have gained increasing attention as environmentally sustainable food resources because they can be cultivated using agricultural wastes and contribute to circular agricultural production systems (Jaiswal et al., 2024). In many rural and forest-dependent communities, wild mushrooms serve as seasonal food resources that supplement household diets and provide opportunities for additional income. Their collection from natural ecosystems also contributes to food security and dietary diversity, particularly in upland and indigenous communities where access to conventional food sources may be limited.
       
In addition to their nutritional importance, wild edible mushrooms play significant ecological roles in forest ecosystems. Many species function as ectomycorrhizal fungi, forming mutualistic relationships with trees and enhancing nutrient and water uptake, while saprotrophic species decompose organic matter and recycle nutrients in forest soils (Kumar et al., 2021). These ecological processes support forest productivity and create favorable conditions for the continued availability of wild edible mushrooms as natural food resources.
       
Despite their ecological and nutritional importance, the diversity and distribution of wild edible mushrooms remain insufficiently documented in many tropical regions. In the Philippines, a country recognized as a global biodiversity hotspot, mycological studies have historically focused on lowland forests and selected protected areas. Consequently, upland and mountainous ecosystems remain underrepresented in national fungal inventories, resulting in limited information on the occurrence and identification of edible macrofungal species in these environments (Hawksworth and Lücking, 2017). Accurate taxonomic identification is particularly important for edible mushrooms to ensure food safety, prevent misidentification of toxic species and support the sustainable utilization of wild mushroom resources (Dulay and Damaso, 2023).
       
Recent surveys of macrofungi in the Philippines have reported considerable species richness in selected forest ecosystems; however, these studies also highlight persistent gaps in geographic coverage and taxonomic resolution. Many investigations rely primarily on morphological identification, which can be challenging due to phenotypic plasticity and overlapping macroscopic characteristics among tropical fungi. The integration of morphological characterization with molecular tools, particularly sequencing of the internal transcribed spacer (ITS) region of ribosomal DNA, has therefore become an important approach for improving taxonomic reliability and strengthening fungal identification studies (Arguelles et al., 2022; Bustillos et al., 2024).
       
Abra Province in Northern Luzon represents a potentially important but understudied area for documenting wild edible mushrooms as natural food resources. The province is characterized by rugged mountainous terrain, steep elevation gradients and heterogeneous vegetation, including pine forests, secondary forests, grasslands and riparian ecosystems. Such environmental diversity creates favorable microhabitats that may support a variety of macrofungal species. Upland forest environments often harbor both ectomycorrhizal fungi associated with woody host plants and saprotrophic species growing on decaying organic substrates.
       
Despite these favorable conditions, systematic documentation of edible macrofungi in Abra Province remains extremely limited and no published species-level records of wild edible mushrooms have been reported for the area. The absence of localized inventories limits understanding of the diversity of edible mushrooms that may contribute to local food resources and sustainable forest-based livelihoods.
       
In this context, the present study documents the occurrence and taxonomic identification of wild edible mushrooms in selected upland municipalities of Abra Province, Northern Philippines. Using integrated macromorphological characterization supported by microscopic observations and ITS sequence similarity analysis, the study identifies edible macrofungal species occurring in these upland ecosystems. The findings provide the first species-level records of wild edible mushrooms for Abra Province and highlight their potential importance as natural food resources for upland communities.
 

Study area and research period
 
The study was conducted in selected upland municipalities of Abra Province, Northern Luzon, Philippines, namely Licuan-Baay, Malibcong, Tubo, Luba and Daguioman. These municipalities are characterized by rugged, mountainous terrain, steep elevational gradients and heterogeneous vegetation, including pine forests, secondary forests, grasslands and riparian ecosystems. Elevations in the surveyed areas range from approximately 800 to 1,600 meters above sea level, creating diverse environmental conditions and microhabitats favorable for the growth and distribution of wild edible macrofungi.
       
The operational base station and specimen processing center for the study were located at Abra State Institute of Sciences and Technology, which served as the primary hub for field coordination, temporary storage of collected specimens, preliminary sorting and documentation.
       
Field surveys were conducted from April to September 2025 to capture seasonal variation in the fruiting patterns of edible macrofungi. Sampling activities were strategically scheduled during periods of increased rainfall, elevated humidity and moderate temperatures, as these environmental factors are widely recognized as stimulating mushroom emergence, sporocarp development and fruiting abundance in tropical upland ecosystems. This survey period was selected to maximize the probability of detecting species that fruit intermittently or only under favorable climatic conditions.
 
Field sampling and specimen collection
 
Wild edible mushroom specimens were collected using a combination of systematic transect walks and opportunistic sampling across representative habitat types, including forest interiors, forest edges, grasslands and riparian zones. Transect walks were used to systematically explore microhabitats within each municipality, while opportunistic sampling collected additional specimens encountered outside designated transects. This combined approach was employed to maximize the detection of edible macrofungal species across heterogeneous landscapes.

Only mature and intact fruiting bodies with clearly observable diagnostic characteristics were collected. Each specimen was photographed in situ prior to collection to document its natural growth habit and surrounding environment. Ecological metadata were recorded for each collection, including habitat type, substrate (soil, decaying wood, or forest litter), growth habit (solitary, gregarious, or clustered) and associated vegetation.
       
Specimens were carefully excavated using a knife or small digging tool to preserve basal structures such as the volva and stipe base, which are important for accurate taxonomic identification and differentiation of edible and potentially toxic species. Collected specimens were placed in labeled paper containers to prevent moisture accumulation and morphological deterioration during transport.
 
Morphological characterization and preliminary identification
 
Macromorphological examination followed standard mycological identification protocols. Diagnostic characters assessed included pileus shape, size, color and surface texture; hymenophore type (gills, pores, or ridges); lamella or pore attachment; stipe morphology; presence or absence of annulus and volva; context coloration; odor; and substrate association. Spore prints were obtained when feasible to determine spore color, an important taxonomic character in macrofungal identification.
       
Microscopic observations were conducted to supplement macromorphological identification of selected specimens. Small portions of the hymenium were mounted on glass slides and examined using a compound microscope. Diagnostic microcharacters such as basidiospore shape, relative size and surface ornamentation were observed when possible to support species identification.
       
Preliminary taxonomic identification was conducted using standard macroscopic keys and diagnostic descriptions provided by Largent (1986), supplemented with relevant literature on tropical and Southeast Asian macrofungi. Species were identified to the lowest possible taxonomic rank based on observable morphological characteristics. Accurate identification was considered essential for determining edible species and ensuring food safety.
 
Molecular verification and taxonomic confirmation
       
To strengthen taxonomic accuracy and validate preliminary morphological identifications, selected representative mushroom specimens were subjected to molecular characterization. Genomic DNA was extracted from preserved tissue samples and the internal transcribed spacer (ITS) region of ribosomal DNA was amplified through polymerase chain reaction (PCR) and subsequently sequenced. The ITS region was targeted because it is internationally recognized as the universal DNA barcode for fungi due to its high discriminatory power across a wide range of fungal taxa (Schoch et al., 2012).
       
Molecular analyses were conducted at the Fungal Laboratory of the Institute of Biology at the University of the Philippines Diliman. Obtained nucleotide sequences were compared against authenticated reference sequences available in publicly accessible databases, particularly NCBI GenBank and UNITE, using the Basic Local Alignment Search Tool (BLAST). Closest taxonomic matches were determined based on sequence homology, alignment coverage and database reliability.
       
Species-level identification was assigned using an integrative taxonomic approach that combined molecular evidence with diagnostic macromorphological and micromorphological characters. In general, sequence similarity values of approximately 97–99% or higher were interpreted as supportive of species-level identity, subject to genus-specific variation and database quality.
       
The resulting DNA sequences were used to confirm the identity of selected wild edible mushroom specimens and were archived at the Fungal Laboratory of the Institute of Biology, University of the Philippines Diliman. Certificates of identification issued by the laboratory served as formal verification of the combined molecular and morphological determinations of the collected specimens (Fig 1).


 
Voucher specimens and documentation
 
Representative voucher specimens of the collected edible macrofungi were preserved and deposited in the institutional fungal reference collection of the Abra State Institute of Sciences and Technology (ASIST). These specimens serve as permanent taxonomic references and provide material for future verification and comparative studies related to edible mushroom resources.
 
Data analysis and classification
 
Species richness was determined as the total number of confirmed wild edible mushroom species recorded across all sampling sites. Taxonomic composition was summarized at the genus, family and order levels to describe the hierarchical structure of the recorded edible macrofungal assemblage.
       
Each species was further classified into ecological functional groups, specifically ectomycorrhizal or saprotrophic fungi, based on published ecological literature and known life history traits. Functional group analysis was conducted to provide insight into the ecological roles of recorded edible mushroom species within upland forest ecosystems.
       
To assess the novelty of the records, documented species were compared with existing national compilations and published literature on Philippine macrofungi. This comparison identified species that represent new provincial-level records for Abra Province and highlights their potential contribution as locally available forest-based food resources.

Species richness and taxonomic composition
 
A total of seven wild edible mushroom species were documented from the upland municipalities of Licuan-Baay, Malibcong, Tubo, Luba and Daguioman in Abra Province. These species belong to seven genera, six families and four taxonomic orders, indicating moderate taxonomic breadth despite relatively low species richness (Table 1). The occurrence of these taxa demonstrates the presence of diverse edible macrofungal groups within the upland forest ecosystems of Abra Province and highlights their potential importance as locally available forest-based food resources.


       
The relatively small number of species recorded in this study may be attributed to the limited sampling duration and the seasonal nature of macrofungal fruiting in tropical forest ecosystems. Many mushrooms produce fruiting bodies only under specific environmental conditions, particularly during periods of high humidity and rainfall. Consequently, short-term surveys often yield modest species counts while still capturing representatives from different taxonomic groups. Similar observations have been reported in preliminary macrofungal surveys conducted in tropical upland forests, where early-stage inventories frequently reveal broad taxonomic representation despite limited species numbers (Nilsson et al., 2019).
       
Comparison with existing Philippine fungal inventories indicates that all seven species documented in this study represent the first formally reported species-level records of wild edible mushrooms for Abra Province. This finding highlights the limited documentation of edible macrofungi in upland regions of Northern Luzon and suggests that additional edible species may occur in these understudied landscapes.
       
At the national level, the seven species recorded in this study represent approximately 1.6% of the 447 wild mushroom species currently reported for the Philippines (Dulay and Damaso, 2023). Although this proportion is relatively small, it is significant considering that no previous species-level records of edible macrofungi have been documented for Abra Province. Beyond documenting biodiversity, inventories of wild edible mushrooms provide an important foundation for identifying species with nutritional, medicinal and economic potential. Such baseline information is valuable for selecting promising species for future domestication, nutritional characterization and sustainable utilization programs (Jaiswal et al., 2024).
 
Functional group composition
 
Functional group analysis revealed a predominance of ectomycorrhizal fungi, which accounted for five of the seven recorded species (71.4%), while saprotrophic fungi comprised two species (28.6%) (Table 2). This functional composition reflects the ecological conditions of forest environments where both symbiotic and decomposer fungi contribute to ecosystem processes that sustain mushroom production.


       
The ectomycorrhizal taxa identified in this study include species belonging to the genera Amanita, Boletellus, Pseudosperma and Scleroderma. Ectomycorrhizal fungi form mutualistic associations with woody plants that enhance nutrient and water uptake, thereby supporting forest productivity and indirectly promoting the growth of edible mushrooms (Hawksworth and Lücking, 2017).
       
In contrast, saprotrophic species such as Macrocybe gigantea and Sparassis latifolia were observed growing on organic-rich soils and decaying wood substrates. Saprotrophic fungi play an essential role in decomposing organic materials and recycling nutrients within forest ecosystems. These processes contribute to the availability of substrates that support the growth of edible mushrooms in forest habitats.
 
Annotated species accounts and ecological implications
 
The diagnostic morphological characteristics used for the identification of the recorded wild edible mushrooms are summarized in Table 3. These include macromorphological characters observed during field and laboratory examination, supported in selected specimens by microscopic observations and ITS sequence similarity analysis. The integration of morphological and molecular approaches improves species identification accuracy, particularly in edible mushrooms, where correct identification is essential for food safety.


       
Several documented taxa provide insight into the ecological conditions that support the occurrence of edible mushrooms in upland forests. The ectomycorrhizal species Boletellus areolatus and Scleroderma citrinum were collected from forest soils associated with woody vegetation, reflecting the ecological characteristics of ectomycorrhizal fungi that form symbiotic relationships with host plants (Nilsson et al., 2019).
       
Amanita javanica, characterized by its diagnostic volva and free lamellae, was observed in shaded forest habitats. Although species of Amanita have been documented in several regions of the Philippines, confirmed species-level records from upland areas of Northern Luzon remain limited. The documentation of this species, therefore, contributes to improving regional knowledge of edible macrofungi.
       
The identification of Pseudosperma yunnanense was supported by ITS sequence similarity analysis, further strengthening its taxonomic placement. Species of Pseudosperma often exhibit subtle morphological features that make field identification challenging, highlighting the importance of combining molecular and morphological approaches for reliable species identification.
       
Among the saprotrophic taxa, Sparassis latifolia was observed growing at the base of decaying wood, while Ramaria thindii occurred on forest litter and soil substrates. Both species are associated with forest environments where organic matter accumulation supports fungal growth. These ecological conditions contribute to the natural availability of edible mushrooms that may serve as seasonal food resources for local communities.
 
Comparison with existing philippine records
 
National compilations of Philippine macrofungi indicate that documented mushroom records are heavily concentrated in lowland forests and protected landscapes, while upland and mountainous areas remain comparatively underexplored. Dulay and Damaso (2023) reported 447 mushroom species across the Philippines but noted that many records lack detailed taxonomic confirmation or molecular support.
       
Although several genera identified in this study have previously been reported in other regions of the Philippines, no published species-level records of wild edible mushrooms have been reported for Abra Province. The documentation of species such as Pseudosperma yunnanense and Ramaria thindii, therefore, expands the known geographic distribution of these taxa and improves the taxonomic coverage of edible macrofungi in the country.
       
The occurrence of forest-associated taxa such as Boletellus areolatus and Sparassis latifolia further indicates that upland ecosystems may support additional edible mushroom species that remain undocumented. Continued exploration of mountainous areas in Northern Luzon may therefore reveal a wider diversity of edible macrofungi with potential importance for food resources and rural livelihoods.
 
Food resource and utilization implications
 
The edible mushroom species documented in this study represent natural forest-based food resources that may contribute to dietary diversity in upland communities. Wild edible mushrooms are recognized as valuable sources of protein, dietary fiber, essential amino acids, vitamins, minerals and bioactive compounds that promote human health.
       
Recent nutritional investigations have demonstrated that edible mushroom species differ considerably in their proximate composition, antioxidant activity, mineral content and functional properties. These findings highlight the importance of species-specific nutritional evaluation before recommending indigenous mushrooms for wider food utilization and value-added product development (Anusha et al., 2023). Collectively, these nutritional characteristics reinforce the importance of documenting indigenous edible mushroom resources and their potential contribution to sustainable food systems. Similar observations were reported by Jaiswal et al., (2024), who highlighted the nutritional and environmental significance of edible mushrooms and their role in converting agricultural wastes into nutritious food products.
       
Accurate taxonomic identification of edible mushrooms is essential to prevent accidental consumption of toxic species and to promote the safe use of wild mushrooms as food. The documentation generated by the present study also provides baseline information for future domestication and cultivation studies. Successful future domestication of promising wild edible mushrooms will require the development of suitable protocols for substrate preparation, sanitation and cultivation to ensure consistent production and product quality. Reviews on mushroom cultivation have highlighted that effective substrate disinfection and management are essential components of sustainable mushroom production systems (Gowda and Dronachari, 2019).
       
Although ITS sequence similarity analysis was used to support species identification, the generated sequences were archived at the Fungal Laboratory of the Institute of Biology at the University of the Philippines Diliman. Future studies incorporating sequence deposition and phylogenetic analyses would further strengthen molecular verification of edible macrofungi.
       
In addition to improving taxonomic knowledge, future research may also explore the enterprise potential of these indigenous edible mushrooms. The development of mushroom-based enterprises has been recognized as an effective strategy for livelihood diversification, employment generation and rural income enhancement, particularly when supported by appropriate production technologies and entrepreneurial capacity-building programs (Shirur et al., 2019).
               
Moreover, longer-term, multi-seasonal sampling across a broader range of habitats may reveal additional edible mushroom species and provide a more comprehensive assessment of forest-based food resources in the upland ecosystems of Abra Province.

This study documented seven species of wild edible mushrooms from upland municipalities of Abra Province, Northern Philippines, representing seven genera, six families and four taxonomic orders. These findings constitute the first species-level records of edible macrofungi for the province and highlight the presence of naturally occurring forest-based food resources in upland ecosystems. Accurate taxonomic identification using morphological and molecular approaches supports safe recognition of edible species and helps prevent misidentification of potentially toxic mushrooms. The documented species demonstrate the potential of wild mushrooms to contribute to dietary diversity and sustainable food resources for upland communities. Further studies involving broader sampling and nutritional evaluation are recommended to better explore the food and economic potential of these edible macrofungi.

The authors express their sincere appreciation to the administration of the Abra State Institute of Sciences and Technology for providing logistical support and research facilities necessary for the conduct of this study. Special thanks are extended to the local government units and community members of Licuan-Baay, Malibcong, Tubo, Luba and Daguioman for their cooperation during field surveys. The authors also acknowledge the Fungal Laboratory of the Institute of Biology, University of the Philippines Diliman, for molecular identification and verification of mushroom specimens.
 
Disclaimers
 
The views and conclusions expressed in this article are solely those of the authors and do not necessarily reflect the official policies or positions of Abra State Institute of Sciences and Technology or any affiliated institutions. Any errors or omissions remain the responsibility of the authors.
 
Informed consent
 
The study was conducted with institutional ethical clearance. Gratuitous permits were obtained from the Department of Environment and Natural Resources and prior informed consent and authorization were secured from the National Commission on Indigenous Peoples for research conducted within ancestral domains.

The authors declare that there are no conflicts of interest, financial or otherwise, that could have influenced the conduct or outcomes of this research.