Fungi are amongst the most diverse groups of organisms on earth. There have been numerous estimates regarding the total number of fungi worldwide. Bisby and Ainsworth (1943) recorded the total number to be about 100,000 and later, Hawksworth (1991) hypothesised the total number of fungal species to be 1.5 million. Later, Blackwell (2011) estimated the total number of fungi to be around 3.5 – 5.1 million. More recently, Hawksworth and Lucking (2017) predicted the total number to be in the range of 2.8 to 3.8 million. To date, 149,974 species have been recognised (Index Fungorum 2021). The current rate of fungal species discovery per year averages at 2,000 as compared to 1,000 to 2,000 a decade ago (Cheek et al. 2020).

Macrofungi form fruiting bodies that can be detected with the naked eye in the field and handled by hand. They mostly consist of basidiomycetes, but also include some ascomycetes. They play many essential roles in ecosystems as mutualists, pathogens, decomposers or saprotrophs (Volk 2013). Some are edible, medicinal or toxic to humans. About 20,000 macrofungal species have been recognised worldwide (Hawksworth 2001), but many belong to cryptic species complexes and many more await discovery, particularly from poorly explored regions of the world.

A major hindrance of traditional systematics in fungal discovery and identification is the presence of limited taxonomic features (Wu et al. 2019). The traditional identification techniques utilised morphological features, ecological characters, physiology and biochemistry of tissues (Wang et al. 2016). The boom in molecular methods in the 1980s and a remarkable paper by White et al. (1990) describing rRNA primers in fungi spurred the beginning of molecular data utilisation in fungal classification and species identification. Phylogenetic studies have shown that morphologically similar taxa might belong to different lineages (e.g. Hibbett et al. 1997; Moncalvo et al. 2002). DNA sequences can also be helpful for detecting and distinguishing amongst cryptic taxa sharing similar morphological traits (e.g. Moncalvo and Buchanan 2008; Schoch et al. 2012; Wu et al. 2019).

Before the partition of British India, mycoflora of the region (presently India and Pakistan) was listed by Butler and Bisby (1931) and Mundkur (1938). These checklists recorded only 198 species of this region expanding on 30,000 square miles (77700 km²). Later, Ahmad et al. (1997) recorded about 4,500 fungal taxa in a list that included all groups of fungi, i.e. macrofungi as well as microfungi and lichens. In the last two decades, many new records and description of new species have been added, based on morphological characters alone or in combination with molecular data (e.g. Sarwar et al. 2011; Saba et al. 2019a; Bashir et al. 2020a; Khalid in press), but none of these was comprehensively addressing macrofungal diversity in Pakistan and the ecoregions of their occurrence.

From a biodiversity conservation perspective, ecologists have been concerned about the factors that affect the delimitation of ecological units and how it affects our knowledge of ecological processes (Weins et al. 1985; Gosz 1991). Numerous efforts have been made to categorise geographical zones with analogous features. In a remarkable paper, Olson et al. (2001) defined ecoregions as broad areas of land or water that consist of geographically distinct assemblages of taxa, natural communities and environmental conditions. They presented an ecoregion map for its utilisation at global as well as regional scales. They based their map on biogeographic information and this was built with the collaboration of more than 1,000 experts in biogeography, taxonomy, conservation biology and ecology from all over the world. Ecoregions were classified by taking into account biogeographic features like endemism, species richness and special evolutionary perspectives. The unique feature of this global biodiversity map is that it focuses on species allocation and communities more precisely than the earlier models, based on biophysical characters, for instance, rainfall and temperature (Holdridge 1967; Walter and Box 1976; Schultz 1995; Bailey 1998) or vegetation structure (UNESCO 1969; de Laubenfels 1975; Schmidthüsen 1976). In Olson et al. (2001), the terrestrial world is divided into 14 biomes, eight biogeographic realms and 867 ecoregions; out of which, nine biomes, two realms and 15 ecoregions are found in Pakistan. This country covers a wide altitudinal range from sea level (Arabian Sea) to the second highest peak of the world, K2. The variety of ecoregions from Himalayan alpine grasslands and subtropical pine forests to deserts and xeric shrublands promotes a great deal of fungal diversity that still remains largely unaccounted for.

In this study we compiled a compendium of macrofungi reported from Pakistan to date from searches in the historical literature as well as in the GenBank database. We have included fungi with prominent fruiting bodies visible to the naked eye in this taxonomic list. We have excluded taxa in Ascomycota which are immersed or half immersed structures, galls or non-prominent fruiting structures on animal dung. We also categorised the reported macromycetes into ecoregions, based on available data.

Table 1 provides a comprehensive record of the macrofungal biota of Pakistan known to date, to the best of our knowledge. It lists 1,293 species belonging to 411 genera, 115 families and 24 orders. Out of which, 1,117 species, 338 genera, 83 families and 16 orders belong to Basidiomycota and 176 species, 73 genera, 32 families and eight orders are from Ascomycota. The source reference in Table 1 indicates that most entries are from the extensive checklist by Ahmad et al. (1997; 874 entries). The highest order recorded is Agaricales (27%) with 31 families, 146 genera and 602 species, followed by Polyporales (11%), Russulales (9%) and Pezizales (8%). The orders of least occurrences are Atheliales, Leotiales and Trechisporales representing one taxon in a single genus and family. The proportion of respective families, genera as well as species are shown in Fig. 1.

Table 2 indicates the ecoregions from which each genus was recorded. Fig. 2 shows that the highest macrofungal diversity is found in the western Himalayan broadleaf forests (36%) followed by north-western thorn scrub forests (25%). In addition, Himalayan subtropical pine forests have rich macrofungal diversity with 17% taxa representation, followed by western Himalayan subalpine conifer forests with 13% distribution. The Karakorem West Tibetan Plateau alpine steppe and Baluchistan xeric woodlands show 5% or lesser distribution. On the other hand, the Indus River Delta, Arabian Sea mangroves, Thar Desert, Sulaiman range alpine meadows, East Afghan montane conifer forests, Registan north Pakistan sandy desert, south Iran Nubo-Sindian desert and semi-desert, Rann of Kutch seasonal marsh, as well as the north-western Himalayan alpine scrub and meadows have 0–2% macromycetes record.

Biomes and macrofungi occurrence details therein are presented in the supplementary document labelled “Macrofungi_list_by_biomes_and_ecoregions_of_Pakistan, whereas supplementary table entitled “Detailed_compendium_of_macrofungi_of_Pakistan” contains thorough information about synonymy, locality and taxa level ecoregion allocation.