Biological characteristics of S. rugosoannulata

Stropharia rugosoannulata Farl. ex Murrill (1922), commonly known as the wine-cap mushroom or giant Stropharia, is a large basidiomycete fungus belonging to the genus Stropharia, family Strophariaceae, and order Agaricales (Huang 1995). In Chinese pronunciation, it is referred to as “Daqiugaigu,” where “Da” means giant and “qiugaigu” means the genus Stropharia. Wild populations of S. rugosoannulata are mainly distributed in temperate regions of Europe, North America, and Asia. Fruiting bodies typically appear in spring and autumn and are commonly found in grassy areas, forest edges, woodchip piles, or compost heaps containing cow or horse manure (Huang et al. 2023b). The fruiting bodies occur singly or in clusters. When immature, the fruit bodies are white; as they mature, the caps become wine-red to reddish-brown. Initially hemispherical, the caps flatten with age and can reach up to 25 cm in diameter, often covered with fibrous white scales. The flesh is thick and white. The stipe is white, up to 15 cm in length and 4 cm in diameter, thickened at the base and featuring a prominent annulus near the apex. The annulus is wrinkled and easily detached (Huang 1995). The gills are initially white, turning purplish-gray upon maturity, and the spore print is dark purplish-brown. The mycelium is white in color (Szudyga 1978; Stamets and Chilton 1983). During development, the stipe morphology changes: in the young stage, the stipe is short and thick with a bulbous base tapering toward the top, while in the mature stage, the stipe elongates and maintains a more uniform diameter throughout (Huang et al. 2023c).

Growing conditions of S. rugosoannulata

S. rugosoannulata is a litter-decomposing fungus that primarily relies on carbon and nitrogen sources for nutrition (Pozdnyakova et al. 2018). Its mycelium efficiently decomposes substrates rich in cellulose, hemicellulose, and lignin, converting them into carbon sources for growth (Guo et al. 2023). Although it exhibits minimal selectivity for nitrogen source types, it is highly sensitive to nitrogen concentration, as low levels of nitrogen promote mycelial branching and proliferation (Zhang et al. 2023a, b, c).

Temperature and moisture are critical environmental factors influencing spore germination, mycelial growth, primordium formation, and fruiting body morphology (Table 1). As a moderate-temperature species, S. rugosoannulata spores require cold stratification (5 °C for 30 days) prior to germination, which occurs optimally at temperatures above 24 °C, with the ideal germination temperature being 28 °C (Bao et al. 2016). Mycelial growth occurs within a wide temperature range (5–35 °C), with an optimum at 26 °C. Growth slows significantly below 10 °C and ceases below 5 °C, though the mycelium remains viable. Temperatures exceeding 35 °C inhibit or kill the mycelium (Chen et al. 2021). Primordium formation is triggered by a drop in temperature to 10–20 °C (Stamets and Chilton 1983). At lower temperatures (around 16 °C), the fruiting bodies are stout and compact, grow slowly, and are less likely to open. In contrast, at higher temperatures (around 24 °C), the fruiting bodies grow rapidly, are slender and loose, and are prone to early cap opening (Huang et al. 2023b).

Table 1 Growing conditions of S. rugosoannulata

Optimal mycelial growth requires a substrate moisture content of 65%–80%, with 75% being ideal (Yan et al. 2001). For fruiting body growth, the preferred ambient relative humidity is 85%–95%, which ensures a moist mushroom surface and prevents cap cracking (Yan et al. 2001). When relative humidity drops below 60%, stipe cracking becomes more likely (Huang et al. 2023c).

The value of S. rugosoannulataEdible value

Unlike other members of the genus Stropharia, S. rugosoannulata does not produce the hallucinogenic neurotoxin psilocybin and is therefore considered a completely safe edible mushroom (Yang et al. 2022). Its fruiting bodies are rich in proteins, amino acids, mineral elements, carbohydrates (including dietary fiber), B vitamins, and unsaturated fatty acids, with an amino acid composition that meets the standards of high-quality protein (Huang et al. 2023a). Compared to commonly cultivated edible mushrooms such as shiitake (Lentinula edodes), oyster mushroom (Pleurotus ostreatus), button mushroom (Agaricus bisporus), and black fungus (Auricularia auricula), S. rugosoannulata contains higher levels of proteins, amino acids, and minerals. Notably, it has the highest potassium content among edible mushrooms, reaching 3.58 g·100 g−1, making it a typical example of a high-protein, low-fat, and mineral-rich mushroom (Liu et al. 2024a, b). In addition to its nutritional value, S. rugosoannulata is rich in taste-enhancing compounds, including umami peptides, free amino acids, soluble sugars, organic acids, and taste-active nucleotides (Hu et al. 2020a; Li et al. 2022a; Lu et al. 2022). It also possesses a variety of characteristic aroma compounds, predominantly alcohols, aldehydes, and ketones (Jiang et al. 2023). With its large size, thick flesh, and crisp texture, S. rugosoannulata can be consumed raw as sashimi or cooked in various ways such as in soups, hot pots, pan-fried, or stir-fried dishes. It is thus recognized as a high-quality edible mushroom that combines health benefits, nutritional richness, and palatable flavor.

Medicinal value

S. rugosoannulata is rich in a variety of bioactive compounds with significant pharmacological potential, including polysaccharides, polyphenols, sterols, peptides, lectins, flavonoids, and ergothioneine (Huang et al. 2023a). Studies have shown that these functional components exhibit a wide range of pharmacological effects, such as immunomodulatory, antitumor, antioxidant, antiviral, hypoglycemic, hypolipidemic, and anti-obesity activities. In particular, the polysaccharides derived from S. rugosoannulata are abundant and have been demonstrated to exert preventive or adjuvant therapeutic effects against several chronic metabolic disorders and cancers, including hyperglycemia, coronary heart disease, hepatic steatosis, obesity, and various malignancies (He et al. 2012; Liu et al. 2019a, b; Wang et al. 2021; Li et al. 2022b). Thus, S. rugosoannulata is regarded as a promising natural medicinal resource with substantial potential for development.

Environmental remediation value

The mycelia of S. rugosoannulata have demonstrated considerable potential in soil and water bioremediation. Studies have shown that it can effectively degrade wood preservatives such as polychlorophenols, polychlorinated dibenzo-p-dioxins, and polychlorinated dibenzofurans from sawmill soils and timber (Valentín et al. 2013). During the degradation of tree litter, the mycelium employs laccase-mediated oxidation to break down organic pollutants including nonylphenol, bisphenol A, and anthracene (Pozdnyakova et al. 2018). Moreover, the fungus is capable of degrading 2,4,6-trinitrotoluene (TNT) in soil (Weiß et al. 2004) and accumulating toxic heavy metals such as mercury (Hg) and cadmium (Cd) from soil or substrate matrices (Gabriel et al. 2016; Dong et al. 2022). In aquatic systems, it has been reported to efficiently decolorize industrial effluents containing azo and anthraquinonic dyes (Jarosz-Wilkołazka et al. 2002), degrade pharmaceutical contaminants like carbamazepine in domestic wastewater (removal rate exceeding 70%), and partially remove cyclophosphamide (Castellet-Rovira et al. 2017). It also exhibits antibacterial activity against common waterborne pathogens such as Escherichia coli, Pseudomonas aeruginosa, and antibiotic-resistant bacteria (Sen et al. 2023).

Ecological conservation value

The cultivation of S. rugosoannulata can utilize nearly all types of commonly available agricultural biomass waste (e.g., crop straw, tree litter, livestock and poultry manure) as growth substrates (Liu et al. 2021). This practice reduces air pollution and CO₂ emissions caused by the open burning of agroforestry residues (Hu et al. 2025). It avoids deforestation and pesticide use, helping to conserve forests and reduce soil contamination. The cultivation process is simple and labor-intensive but physically undemanding, making it particularly suitable for the participation of physically challenged individuals in rural areas (Huang et al. 2023b). Moreover, the spent substrate can be directly applied to the field, where it significantly improves soil structure, enhances soil fertility, and increases microbial diversity, reducing the need for chemical fertilizers (Liu et al. 2021). Therefore, S. rugosoannulata cultivation represents an ecologically sound agricultural practice and a model of circular economy well-suited for broad adoption in developing countries (Liu et al. 2021; Hu et al. 2025). It also contributes to achieving two major goals of the United Nations: poverty reduction and climate change mitigation (FAO 2022).

Cultivation history of S. rugosoannulata

Before human cultivation began, wild S. rugosoannulata fruiting bodies were frequently foraged due to their large size, attractive wine-red cap, mild flavor, and edibility. In 1922, American mycologist William A. Murrill first scientifically described this mushroom species (Murrill 1922). The earliest record of artificial cultivation appeared in 1969, when German farmers successfully domesticated the mushroom using straw. Subsequently, countries such as Poland, Czechoslovakia, Hungary, Netherland, and the former Soviet Union introduced and cultivated the species (Sharma et al. 2007). By 1989, production of S. rugosoannulata in Europe had reached approximately 1300 tons (Domondon et al. 2000), though no further production data were reported thereafter. In 1983, mycologist Paul Stamets described the cultivation characteristics of the mushroom in his book The Mushroom Cultivator and introduced European strains of cultivated S. rugosoannulata into the USA, marking the transition of the species from wild to cultivated status there (Stamets and Chilton 1983). In the 1980 s, a strain was introduced from Poland to Shanghai, China, with successful trial cultivation but no subsequent promotion. In 1992, Fujian province reintroduced the species and, after successful trials, began to promote its cultivation (Huang 1995) (Fig. 1).

Fig. 1

Timeline of S. rugosoannulata cultivation

With the increasing recognition of S. rugosoannulata’s value in environmental protection and bioremediation, the cultivation purpose in some countries has gradually shifted from the production of edible fruiting bodies to the utilization of its mycelium for the efficient absorption and degradation of pollutants. Globally, growers in North America or New Zealand primarily cultivate S. rugosoannulata as a landscape mushroom in the city garden or in the backyard, using hardwood chips or straw beds as substrates (Stamets and Chilton 1983). In Europe, commercial cultivation has ceased, and the species is now primarily used for the remediation of contaminated soil and water bodies (Valentín et al. 2013). In these regions, the scale of cultivation is relatively small, and there is a lack of statistical data. At present, China is the only country to have achieved large-scale commercial cultivation of S. rugosoannulata, and since 2013, its production data have been included in the national annual statistical reports on edible fungi (China Edible Fungi Association 2025). China has become the world’s leading producer of S. rugosoannulata.

S. rugosoannulata industry in China

The cultivation characteristics of S. rugosoannulata align well with China’s abundant natural resources, an aging rural labor force with low incomes, and predominantly temperate to subtropical geographic zones. Since 2013, driven by the Chinese government’s targeted poverty alleviation and rural revitalization strategies, the cultivation of S. rugosoannulata has been rapidly promoted as a flagship project. Its cultivation areas have expanded across all provinces in the country, and its production volume has increased sharply. According to statistics, S. rugosoannulata is now cultivated in all 31 provincial-level regions in mainland China (Huang et al. 2023b), and the total fresh mushroom yield reached 494,745 tons in 2023 (China Edible Fungi Association 2025), more than 17 times the yield in 2017 (Fig. 2A). The total cultivation area exceeded 6000 hectares, with Hunan, Sichuan, and Jiangxi being the major producing provinces (Fig. 3). However, China is the world’s largest producer of edible fungi, with an annual output exceeding 40 million tons (Li and Xu 2022). Compared with major species such as L. edodes, A. auricula, P. ostreatus, and A. bisporus, each with an annual production of over 1.5 million tons, S. rugosoannulata is still considered a minor edible fungus species in China (Fig. 2B).

Fig. 2

A Annual yield of fresh S. rugosoannulata in China from 2017 to 2023. B Comparison of the yield of fresh S. rugosoannulata and bulk edible fungi in China in 2023

Fig. 3

Discovery sites of wild S. rugosoannulata strains in China and fresh S. rugosoannulata yield of each province in 2023 (note: green dots mark discovery sites; yield data comes from China Edible Fungi Association and is measured in kilotons(kt); the shade of red represents the yield level; gray areas indicate provinces without data)

With continuous efforts from Chinese edible mushroom partitioners, advancements in food processing technologies, and growing public awareness of nutrition and health, significant progress has been made in the processing and product development of S. rugosoannulata. A growing variety of S. rugosoannulata products is now available in the market. A relatively complete industry chain has taken shape, covering all stages from strain selection, cultivation to post-harvest processing and product sales.

Strain selection and breeding of S. rugosoannulata

Germplasm resources serve as the foundation and origin of the S. rugosoannulata industry. With the increasing scope and depth of wild fungal resource surveys in China, wild S. rugosoannulata was first discovered in Tibet in 1993. Since then, new wild strains have been continuously identified in regions such as Yunnan, Jilin, Taiwan, Sichuan, Gansu, Shaanxi, Hunan, Ningxia, and Shandong (Huang 1995; Huang et al. 2024; Fu et al. 2025) (Fig. 3). Concurrently, as the cultivation scale of S. rugosoannulata has expanded, color-mutant strains with stable genetic traits have been successively identified in various regions (Fu et al. 2025).

At present, Chinese researchers in the field of edible fungi have developed numerous new S. rugosoannulata cultivars (strains) through various approaches, including domestication of wild strains, systematic selection, mutagenesis breeding, and hybridization. These new strains exhibit advantageous traits such as high yield, strong resistance, wide adaptability, and diverse coloration, and they largely meet the cultivation requirements across different regions of China, which vary greatly in climatic conditions (Table 2). In 2024, Chinese Academy of Agricultural Sciences reported that, based on genetic diversity analysis of 50 S. rugosoannulata strains collected from across China, these strains could be divided into two groups: cultivated and wild. The cultivated strains exhibited high genetic similarity, whereas the wild strains showed significant genetic diversity (Gu et al. 2024).

Table 2 Major cultivars of S. rugosoannulata in China developed through different breeding methodsCultivation models of S. rugosoannulata

S. rugosoannulata is considered one of the easiest edible mushrooms to cultivate globally. Unlike wood-rotting fungi such as L. edodes and A. auricula, S. rugosoannulata can utilize almost all types of agricultural and forestry residues as substrates. Its cultivation does not require bagging, sterilization, or aseptic inoculation equipment. Instead, raw or fermented materials can be directly spread as substrate beds, manually inoculated with spawn, and covered with a simple casing layer (e.g., soil or leaf litter) (Stamets and Chilton 1983). Due to its minimal technical requirements and high edible value, it has been recommended by the Food and Agriculture Organization of the United Nations (FAO) for cultivation in developing countries (Huang et al. 2023b).

Currently, S. rugosoannulata is cultivated in China using three main models: field cultivation, forest cultivation, and facility cultivation (Guo et al. 2025). Field cultivation involves rotation with crops such as wheat, rice, or vegetables, utilizing farmland during the off-season after harvest (Hu et al. 2025). Forest cultivation entails intercropping with trees to make full use of space beneath forest canopies or in orchards (Zhang et al. 2017; Gong et al. 2018). These two models dominate S. rugosoannulata cultivation in China and typically rely on natural climate conditions for seasonal fruiting. They are characterized by low production costs, ease of management, and favorable economic returns. Facility cultivation is primarily adopted in northern China and high-altitude regions, enabling off-season production during summer and winter (Wei et al. 2021).

Based on reported data from various regions, all three cultivation models predominantly involve sowing in autumn. The average yields are ranked as follows: Facility cultivation (7.89 kg·m−2) > forest cultivation (7.24 kg·m−2) > field cultivation (6.47 kg·m−2). Similarly, the average net profits are ranked: facility cultivation (25,293.5 CNY·667 m−2) > forest cultivation (15,316.51 CNY·667 m−2) > field cultivation (9,003.4 CNY·667 m−2). The amount of substrate used is a key factor affecting yield, with all three cultivation models showing a linear increase in yield with increased substrate input (Guo et al. 2025).

Existing products of S. rugosoannulata

At present, the consumer-end products of S. rugosoannulata in China include fresh mushrooms as well as various levels of processed products (Fig. 4), forming a multi-level product structure.

Fig. 4

The classification of current S. rugosoannulata products in China

Fresh mushrooms

Fresh S. rugosoannulata mushrooms are characterized by vibrant coloration, thick and meaty texture, pleasant aroma, and crisp mouthfeel, making them suitable for various cooking methods such as soups, hotpots, stir-frying, and deep-frying. They can also be consumed raw as sashimi. Fresh mushrooms constitute the primary consumer product form of S. rugosoannulata in China and hold a notable position in the fresh edible mushroom market.

Color varieties

Although three color varieties of S. rugosoannulata have been bred in China, only the red and yellow types are currently available in the mushroom market (Wang et al. 2025a, b) (Fig. 5A, B). The red type, representing the wild phenotype, is widely cultivated and marketed across the country. The yellow variety, derived from a spontaneous color mutation during the cultivation of the red type, was first reported in 2021 and has since entered the market on a small scale in provinces such as Yunnan and Sichuan (Huang et al. 2024). The white variety, obtained through radiation-induced mutagenesis of the red strain, was first reported in 2025 and has completed demonstration cultivation but has not yet been commercially promoted (Song et al. 2025) (Fig. 5C).

Fig. 5

S. rugosoannulata mushroom with red cap (A), yellow cap (B), and white cap (C)

Nutrition-enriche