Checklist of the agaricoid fungi from Paraguay A. FLECHA RIVAS1* & N. NIVEIRO2 1 Laboratorio de Micología, Facultad de Ciencias Exactas y Naturales 2 Instituto de Botánica del Nordeste-IBONE, Universidad Nacional de Asunción-Paraguay Universidad Nacional del Nordeste-UNNE, Argentina. CORRESPONDENCE TO *: flecharivas@gmail.com ABSTRACT—The aim of this work is to provide a complete literature-based checklist of the agaricoid fungi from Paraguay. For this, an extensive review of the literature and online databases was made. We list 125 species belonging to the agaricoid fungi distributed among 20 families. Agaricaceae, Tricholomataceae, Psathyrellaceae, and Marasmiaceae are the most species-abundant families. Regarding species distribution, the Capital department has the highest number of reported species, and other biogeographical regions as Chaco, Cerrado and Pantanal are practically unexplored. INTRODUCTION The agaricoid fungi are characterized by having ephemeral basidiomata with a putrescent consistency and lamellated hymenophore (Webster & Weber 2007, Niveiro et al. 2014, Piepenbring 2015). Usually, they have the typical umbrella shape, constituted in a pileus supported by a stipe. On the underside of the pileus, the hymenophore is usually formed by radially arranged lamellae (Webster & Weber 2007). The agaricoid fungi belong to the Agaricomycetes clade, and are distributed in 7 orders: Agaricales, Boletales, Cantharellales, Gloeophyllales, Hymenochaetales, Polyporales and Russulales (Hibbett et al. 2007). Together, these include more than 21000 described species (Hibbett et al. 2014), widely distributed in all ecosystems (Singer 1986). The bibliographical references on the mycobiota of Paraguay are limited (Popoff & Wright 1998). The first records of agaricoid fungi in Paraguay date from 1883, when Carlos Spegazzini, botanist-mycologist, made a two-month expedition through the country. During that journey he visited the departments of Central, Cordillera, Caaguazú, Paraguarí, Concepción and Itapúa. The results of this expedition were published by Spegazzini (1922), where he says: "I gathered a collection of both flowering plants and cryptogams, especially fungi, which served as the basis for writing this booklet. In it the reader will not find great novelties, although I think it will not be totally devoid of interest, and I consider it a new grain of sand contributed to the increase of our knowledge of Mycology in general and that of Paraguay in particular" (Spegazzini 1922, p.3). In spite of this statement by Spegazzini, claiming the importance for the mycological studies in the country, for decades there were no mycological investigations in Paraguay. In this hiatus, important contribution was made especially by Rolf Singer, who studied extensively the mushrooms in the South America and revised a lot of material collected by Spegazzini, including some collections and types from Paraguay (Singer 1950, 1970, 1976). Only in the recent 20 years a few studies on the diversity of agaricoid mushrooms from Paraguay had been published. Zanotti-Cavazzoni (1996) made a list of edible fungi known from the Central Department of Paraguay and reported 39 genera of agaricoid fungi. Gullón (2011) cited only 4 species from the Refugio Biológico Tati Yupi. Campi et al. (2013) listed 14 species as occurring in the Natural Reserve Laguna Blanca. Some articles reported species not previously known for Paraguay (De Madrignac et al. 2013; Flecha et al. 2013; Campi et al. 2016, 2017a, 2017b). Some genera such as Leucocoprinus (Campi et al. 2015) and Pleurotus (Flecha et al. 2014) have received more detailed study in the country. With the increasing number of investigations being conducted on the diversity of agaricoid fungi in Paraguay, the aim of this work is to provide a checklist of the agaricoid fungi from Paraguay that is up to date, establishing a baseline for future studies. Summary: Mycotaxon 133: 729—Regional mycobiota new to www.mycotaxon.com Expert reviewers: Timothy J. Baroni, Vagner Gularte Cortez, Gonzalo M. Romano Uploaded — February 2019 2 … Flecha Rivas & Niveiro METHODS An extensive review of the literature and online databases was made. All agaricoid fungi described and cited for Paraguay from 1883 until 2017 are cataloged in this work. The online databases MycoBank (www.mycobank.org) and Index Fungorum (www.indexfungorum.org) were used for taxonomic classification, renaming, and updating names of the species if necessary. For authors abbreviations, the Authors of Fungal Names (http://www. indexfungorum.org/names/AuthorsOfFungalNames.asp) was used. Species distribution is cited according to the department (State) where it was collected followed by the specific locality. A map with the political divisions of Paraguay and the abbreviations for each state can be seen in Figure 1. For the biogeographical regions Olson et al. (2001) and Morrone (2001, 2014) were followed. Figure 1: Map of Paraguay. Departments where agaricoid mushrooms have been collected: AM: Amambay, AP: Alto Paraná, CA: Capital, CG: Caaguazú, CE: Central, CO: Concepción, CR: Cordillera, PA: Paraguarí, SP: San Pedro. RESULTS Based on the records found in 33 bibliographic sources, we counted 125 species (with 148 records) belonging to the agaricoid fungi distributed in 20 families has been reported for Paraguay. The list of catalogued species is shown in Table 1 Agaricaceae is the most abundant family with 22 species, followed by Tricholomataceae with 14 species, Psathyrellaceae (13 species), Marasmiaceae (12 species), and Pleurotaceae (11 species), the remaining families have less than 10 species each. Regarding species distribution by location, the Capital Agaricoid fungi from Paraguay … 3 department has the highest number of cited species, with a total of 72 reported species (51 %), followed by Paraguarí, with 15 species (11%), Alto Paraná and Central, with 11 species (10%) (Table 2). Considering the biogeographic regions, we observed that all collections are related to the Parana Forest and Humid Chaco, and there are no collections of the other biogeographic regions present in Paraguay as Dry Chaco, Cerrado (Chacoan domain) and Pantanal (Amazonia region). Table 1: Checklist of agaricoid fungi from Paraguay SPECIES DISTRIBUTION REFERENCE AGARICACEAE Agaricus campestris L. PA Spegazzini 1886 Agaricus pampeanus Speg. CA Spegazzini 1919 Agaricus succineus Speg. CG Spegazzini 1886 Coprinus comatus (O.F. Müll.) Pers. AP Campi et al. 2016 Coprinus fimetarius Fr. CA Spegazzini 1922 Coprinus rufopruinatus Romagn. AP Gullón 2011 Lepiota camporum (Speg.) Speg. CA Spegazzini 1919 (as Annularia camporum Speg.) Lepiota cinerascens Speg. CA Spegazzini 1919 Lepiota fuscoroceola Speg. CA Spegazzini 1919 Lepiota lycoperdinea Speg. CA Spegazzini 1919 Lepiota micromyces Speg. CA Spegazzini 1922 Lepiota pluvialis Speg. CA Spegazzini 1919 Leucoagaricus erythrellus (Speg.) Singer CA Spegazzini 1919 (as Lepiota erythrella Speg.) Leucoagaricus rickianus (Speg.) Singer CA Spegazzini 1919 (as Lepiota rickiana Speg.) Leucoagaricus rubrosquamosus (Rick) Singer AP Gullón 2011 Leucoagaricus lilaceus Singer AP, CE, SP Flecha et al. 2013 Leucocoprinus birnbaumii (Corda) Singer CE Campi et al. 2015 Leucocoprinus cepistipes (Sowerby) Pat. CE Campi et al. 2015 Leucocoprinus cretaceus (Bull.) Locq. AP Campi et al. 2015 Leucocoprinus straminellus (Bagl.) Narducci & Caroti AP Campi et al. 2015 Macrolepiota bonaerensis (Speg.) Singer CA, CE, PA Spegazzini 1886 (as Agaricus bonaerensis Speg.), 1919 (as Lepiota bonaerensis Speg.) Macrolepiota excoriata (Schaeff.) Wasser CA Spegazzini 1919 [as Lepiota excoriata (Schaeff.) P. Kumm.] AMANITACEAE 3 4 … Flecha Rivas & Niveiro Limacella laeviceps (Speg.) Raithelh. CA, CE Spegazzini 1919 (as Lepiota laeviceps Speg.), Raithelhuber1980 Conocybe antipus (Lasch) Fayod CA Spegazzini 1922 [as Galera antipus (Lasch) Quél.] Conocybe rickeniana P.D. Orton CA Spegazzini 1922 [as Galera teneroides (Peck) Sacc.] Conocybe tenera (Schaeff.) Fayod CA Spegazzini 1919, 1922 [as Galera tenera (Schaeff.) P. Kumm.]. CA Spegazzini 1919 (as Clitocybe himantiigena Speg.], Silva-Filho et al. 2018. CR Campi et al. 2017a. Galera nemoricola (Speg.) Speg. PA Spegazzini 1886 (as Agaricus nemoricola Speg.). Hebeloma fastibile (Pers.) P. Kumm. CA Spegazzini 1919. Naucoria cerodes (Fr.) P. Kumm. CA Spegazzini 1922. Naucoria melinoides (Bull.) P. Kumm. CA Spegazzini 1919 Crepidotus mollis (Schaeff.) Staude CA Spegazzini 1919 Flammulaster muricatus (Fr.) Watling CA Spegazzini 1922 (as Pholiota muricata Fr.) Lactocollybia carneipes (Speg.) Singer CA Spegazzini, 1919 (as Omphalia carneipes Speg.), Singer 1949 (as Macrocystidia carneipes Singer (1986). Marasmius balansae Speg. PA Spegazzini 1891 Marasmius caespitosus Velen. CA Spegazzini 1919 Marasmius cohortalis var. hymeniicephalus (Speg.) Singer CE, PA Spegazzini 1886, 1919 [as Agaricus hymeniicephalus Speg.; Heliomyces hymeniicephalus (Speg.) Speg.], Singer 1976 [as Marasmius hymeniicephalus (Speg.) Singer] Marasmius crinis-equi F. Muell. ex Kalchbr. CA, CG Spegazzini 1886 (as Marasmius trichorhizus Speg.), Spegazzini 1922 (as Marasmius equicrinis Rüll.) Marasmius ferrugineus Berk. SP De Madrignac et al. 2013 BOLBITIACEAE ENTOLOMATACEAE Clitocella himantiigena (Speg.) Silva-Filho & Cortez HYDNANGIACEAE Laccaria fraterna (Sacc.) Pegler HYMENOGASTRACEAE INOCYBACEAE MARASMIACEAE Agaricoid fungi from Paraguay … Marasmius glabellus Peck CA Spegazzini 1919 Marasmius haematocephalus (Mont.) Fr. AP, CA, SP Spegazzini 1919, Gullón 2011, De Madrignac et al. 2013 Marasmius pallipes Speg. PA Spegazzini 1886 Marasmius petalinus Berk. & M.A. Curtis CA Spegazzini 1919 Marasmius siccus (Schwein.) Fr. SP Campi et al. 2013 Marasmius spegazzinii Sacc. & Sydow PA Spegazzini 1891 (as M. balansae Speg.) Trogia icterina (Singer) Corner SP Campi et al. 2013 [as Gerronema icterinum (Singer) Singer] Mycena corticola (Pers.) Gray PA Spegazzini 1886 (as Agaricus corticola Speg.) Mycena juncicola (Fr.) Gillet CA Spegazzini 1922 Mycena paraguaya Speg. CA Spegazzini 1922 Mycena paraguariensis (Speg.) Niveiro, Popoff & Albertó PA Spegazzini 1886 (as Agaricus paraguariensis Speg.), Buyck et al. 2017 Mycena sosarum Singer SP Campi et al. 2013 Xeromphalina tenuipes (Schwein.) A.H. Sm. AM, CG Pegler 1983, Campi et al. 2017b Gymnopus peronatus (Bolton) Gray CA Spegazzini 1919 [as Marasmius urens (Bull.) Fr.] Lentinula guarapiensis (Speg.) Pegler PA Spegazzini 1886 (as Agaricus guarapiensis Speg.) Marasmiellus eburneus (Theiss.) Singer CA Spegazzini 1919 (as Marasmius eburneus Theiss.) Marasmiellus microscopicus (Speg.) Singer CG Spegazzini 1886 (as Agaricus microscopicus Speg.), Singer 1973. Marasmiellus tropicalis (Speg.) Singer CR Spegazzini 1888 (as Pleurotus tropicalis Speg.) Neonothopanus nambi (Speg.) R.H. Petersen & Krisai PA Spegazzini 1886 (as Agaricus nambi Speg.) Cyptotrama asprata (Berk.) Redhead & Ginns SP Campi et al. 2013 Dactylosporina steffenii (Rick) Dörfelt SP Campi et. al. 2013 [as Oudemansiella steffenii (Rick) Singer] Oudemansiella platensis (Speg.) Speg. AP, SP Gullón 2011, Campi et al. 2013 [as Oudemansiella canarii (Jungh.) Höhn.] Oudemansiella orinocensis (Pat.) Speg. CA Spegazzini 1919 5 MYCENACEAE OMPHALOTACEAE PHYSALACRIACEAE 5 6 … Flecha Rivas & Niveiro Xerula pudens (Pers.) Singer CA Spegazzini 1919 (as Collybia longipes P. Kumm.) Hohenbuehelia paraguayensis Speg. CA Spegazzini 1922 (as Pleurotus paraguayensis Speg.) Hohenbuehelia petaloides (Bull.) Schulzer PA Spegazzini 1888 (as Pleurotus petaloides Speg. Hohenbuehelia portegna (Speg.) Singer CR, PA Spegazzini 1886, 1888 [as Agaricus portegnus Speg.; Pleurotus portegnus (Speg.) Speg.] Pleurotus albidus (Berk.) Pegler CA, PA Spegazzini 1886, 1919 [as Agaricus laciniatocrenatus (Speg.) Speg.; Pleurotus laciniatocrenatus (Speg.) Speg.], Flecha et al. 2014 Pleurotus cystidiosus O.K. Miller CE Flecha et al. 2014 Pleurotus djamor var. cyathiformis Corner CA Flecha et al. 2014 Pleurotus djamor var. roseus Corner AP Flecha et al. 2014. Pleurotus elegantissimus Speg. CA Spegazzini 1922 Pleurotus guaraniticus Speg. CA Spegazzini 1922 Pleurotus microspermus Speg. PA Spegazzini 1891 Pleurotus spodoleucus (Fr.) Quél. CA Spegazzini 1922 Pluteus fibulatus Singer SP Campi et al. 2013 Volvariella paraguayensis Speg. CA Spegazzini 1922 (as Volvaria paraguayensis Speg.) CA Spegazzini 1919 [as Mycena epiphloea (Fr.) Sacc.] Coprinellus disseminatus (Pers.) J.E. Lange CA, SP Spegazzini 1919, 1922 [as Psathyrella disseminata (Pers.) Quél.], Campi et al. 2013 (as Coprinus disseminatus (Pers.) Gray] Coprinellus ephemerus (Bull.) Redhead, Vilgalys & Moncalvo CA Spegazzini 1922 [as Coprinus ephemerus (Bull.) Fr.] Coprinopsis extinctoria (Fr.) Redhead, Vilgalys & Moncalvo CA Spegazzini 1922 (as Coprinus extinctorius Fr.) Panaeolina foenisecii (Pers.) Maire CA Spegazzini 1922 [as Psilocybe foenisecii (Pers.) Quél.] Panaeolus papilionaceus (Bull.) Quél. CA Spegazzini 1922 [as Panaeolus campanulatus (L.) Quél.] Parasola hemerobia (Fr.) Redhead, Vilgalys & Hopple CA Spegazzini 1922 (as Coprinus hemerobius Fr.) Psathyrella argentina Speg. CA Spegazzini 1919 Psathyrella conopilea (Fr.) A. CA Spegazzini 1922 (as Psathyra PLEUROTACEAE PLUTEACEAE POROTHELEACEAE Phloeomana hiemalis (Osbeck) Redhead PSATHYRELLACEAE Agaricoid fungi from Paraguay … Pearson & Dennis 7 conopilus (Fr.) P. Kumm.] Psathyrella asperella Quél. & Schulzer CA Spegazzini 1922 Psathyrella candolleana (Fr.) Maire CA Spegazzini 1922 [as Hypholoma appendiculatum (Bull.). Quél.] Psathyrella gyroflexa (Fr.) Konrad & Maubl. CA Spegazzini 1922 [as Psathyra gyroflexa (Fr.) P. Kumm.] Psathyrella microsperma (Speg.) Guzmán CA Spegazzini 1919 (as Psilocybe microsperma Speg.), Guzmán & Vergeer 1978 Psathyrella subcorticalis Speg. CA Spegazzini 1922 SCHIZOPHYLLACEAE Schizophyllum commune Fr CA, CE, GR, IP, SP Spegazzini 1922 [as Schizophyllum alneum (L.) Schört], Campi et al. 2013 STROPHARIACEAE Agrocybe pediades (Fr.) Fayod CA Spegazzini 1922 [as Naucoria semiorbicularis (Bull.) Quél.] Agrocybe retigera (Speg.) Singer CA Spegazzini 1922 (as Naucoria retigera Speg.); Singer 1950 Agrocybe vervacti (Fr.) Singer CA Spegazzini 1919 [as Naucoria vervacti (Fr.) Quél.] Cyclocybe cylindracea (DC.) Vizzini & Angelini CR Spegazzini 1886 (as Agaricus pudicus Bull.) Deconica merdaria (Fr.) Noordel. CA Spegazzini 1922 [as Stropharia merdaria (Fr.) Quél.] Flammula papillosispora Speg. CE Spegazzini 1919 Galerina sideroides (Bull.) Kühner CA Spegazzini 1919 [as Naucoria sideroides (Bull.) Quél.] Gymnopilus earlei Murril. AP Gullón 2011 Gymnopilus picreus (Pers.) P. Karst. CO Spegazzini 1919 (as Flammula picrea (Pers.) Quél.] Gymnopilus sapineus (Fr.) Murrill CA Spegazzini 1922 [as Flammula sapinea (Fr.) P. Kumm.] Clitocybe balansae Speg. PA Spegazzini 1891 Collybia auriantiella (Speg.) Speg. CG Spegazzini 1886 (as Agaricus auriantiellus Speg.), Spegazzini 1888 Collybia brasiliensis (Berk. & Mont.) Dennis CA Spegazzini 1919 (as Marasmius brasiliensis Berk. & Mont.) Collybia micheliana (Fr.) Quél. CA Spegazzini 1922 Collybia muscigena (Schumach.) P. Karst. CE Spegazzini 1919 Collybia nivea (Mont.) Dennis AP Gullón 2011 (as Marasmius niveus Mont.) TRICHOLOMATACEAE 7 8 … Flecha Rivas & Niveiro Melanoleuca balansae (Speg.) Singer PA Spegazzini 1886 (as Agaricus balansae Speg.) Melanoleuca excissa (Fr.) Singer CA Spegazzini 1922 [as Tricholoma exscissum (Fr.) Quél.] Melanoleuca phaeopodia (Bull.) Murrill CA Spegazzini 1922 [as Collybia phaeopodia (Bull.) Sacc.] Omphalia liliputiiana Speg. CA Spegazzini 1919 Omphalia succinea Sacc. CE Spegazzini 1919 Resupinatus alboniger (Pat.) Singer SP Campi et al. 2013 (as Resupinatus argentinus Singer) Resupinatus applicatus (Batsch) Gray CA, CG Spegazzini 1886 [as Agaricus applicatus Batsch; Pleurotus applicatus (Batsch) P. Kumm.] Resupinatus striatulus (Pers.) Murril. CA Spegazzini 1919 (as Pleurotus striatulus (Pers.) P. Kumm.) CA Spegazzini 1919 Lentinus berteroi (Fr.) Fr. CA, SP, PA, CO Spegazzini 1883 (Lentinus nigripes Fr.), Spegazzini 1919 (Lentinus nigripes Fr.; and Lentinus tener Klotzsch), Campi et al. 2013 Lentinus fasciatus Berk. PA Spegazzini 1888 (as Lentinus fuscopurpureus Kalchbr.) Lentinus patulus Lév. PA Spegazzini 1883 (as Agaricus calyx Speg.), Pegler 1983a [as Lentinus calix (Speg.) Pegler], Pegler 1983b Lentinus scleropus Speg. CA, PA Spegazzini 1883 (as Lentinus paraguayensis Speg.), Pegler 1983b Lentinus striguellus Berk. PA Spegazzini 1883 (as Panus guaraniticus Speg.), Pegler 1983b Lentinus tigrinus (Bull.) Fr. SP Campi et al. 2013 Lentinus velutinus Fr. CR Spegazzini 1883 (as Lentinus fallax Speg.), Pegler 1983b Lentinus villosus Klotzsch PA Spegazzini 1883 TUBARIACEAE Tubaria trigonophylla (Lasch) Fayod POLYPORACEAE Species registered by Department: AM: Amambay, AP: Alto Paraná, CA: Capital, CG: Caaguazú, CE: Central, CO: Concepción, CR: Cordillera, GR: Guaira, IP: Itapuá, PA: Paraguarí, SP: San Pedro. Agaricoid fungi from Paraguay … 9 Table 2. Number of species registered by Department* RECORDS BY DEPARTMENT N° species AM AP CA CG CE CO CR GR IP PA SP Agaricaceae 22 — 6 12 1 4 — — — — 2 1 26 Amanitaceae 1 — — 1 — 1 — — — — — — 2 Bolbitiaceae 3 — — 3 — — — — — — — — 3 Entolomataceae 1 — — 1 — — — — — — — — 1 Hydnangiaceae 1 — — — — — — 1 — — — — 1 Hymenogastraceae 4 — — 3 — — — — — — 1 — 4 Inocybaceae 2 — — 2 — — — — — — — — 2 Maramiaceae 13 — 1 6 1 1 — — — — 4 4 17 Mycenaceae 6 1 — 2 1 — — — — — 2 1 7 Omphalotaceae 6 — — 2 1 — — 1 — — 2 — 6 Physalacriaceae 5 — 1 2 — — — — — — — 3 6 Pleurotaceae 11 — 1 6 — 1 — 1 — — 4 — 13 Pluteaceae 2 — — 1 — — — — — — — 1 2 Porotheleaceae 1 — — 1 — — — — — — — — 1 Psathyrellaceae 13 — — 13 — — — — — — — 1 14 Schizophyllaceae 1 — — 1 — 1 — — 1 1 — 1 5 Strophariaceae 10 — 1 6 — 1 1 1 — — — — 10 Tricholomataceae 14 — 1 7 2 2 — — — — 2 1 15 Tubariaceae 1 — — 1 — — — — — — — — 1 Polyporaceae 8 — — 2 — — 1 1 — — 6 2 12 125 1 11 72 6 11 2 5 1 1 23 15 148 FAMILY TOTAL N° records * AM: Amambay, AP: Alto Paraná, CA: Capital, CG: Caaguazú, CE: Central, CO: Concepción, CR: Cordillera, GR: Guaira, IP: Itapuá, PA: Paraguarí, SP: San Pedro 9 10 … Flecha Rivas & Niveiro CONCLUSION This work constitutes a first attempt to provide a general overview of the distribution of agaricoid fungi in Paraguay. Of the 125 species known from Paraguay, 103 were cited by Spegazzini, with 26 of them being described and proposed by him as new species. Although this indicates the incomparable value of Spegazzini's work in Paraguay, it also highlights the scarcity of modern studies on the diversity of agaricoid fungi in Paraguay. We can observe that the highest proportion of the species cited from Paraguay are historical species, many of which were found only once, or that were described in genera that have now been re-delimited, and also several names are doubtful and without a modern taxonomic approach. Another point to consider is the concentration of information in a limited geographical area, such as the Capital department, the most urbanized of all, having large areas and biogeographical zones of Paraguay such as Chaco, Cerrado (Chacoan domain) and Pantanal (Amazonia region) which are practically unexplored in mycological terms. Since the study of agaricoid mushrooms in Paraguay was resumed in 2011, numerous novelties have been found. Some projects are being developed in environments that were never visited by other mycologists, such as Pantanal and Chaco. However, the research team is young, and access to information and literature on South American species is scarce. Therefore, this work serves as a baseline to which future studies on diversity and taxonomy of the agaricoid fungi of Paraguay can continue to add new information in the progress towards, fully understanding the biodiversity of Paraguay. ACKNOWLEDGMENTS The authors wish to thank T.J. Baroni, V.G. Cortez, and G. Romano for the critical revision of the manuscript, and to Dra. A. Michlig for the first revision of the manuscript. 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