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A first review of Gasteroid fungi (Agaricomycetes, Basidiomycota) in Paraguay

Una primera revisión de los hongos gasteroides (Agaricomycetes, Basidiomycota) en Paraguay

ABSTRACT

Based on the available literature, a list of 40 gasteroid species recorded in Paraguay since the earliest known collection is presented. A total of forty species are reported, distributed among 16 genera and 4 families. This work represents the first compilation of data available on this group of fungi for Paraguay. These numbers reveal the scarce amount of species formally cited for the Paraguayan Funga.

Keywords:
bird-nest fungi; earth-stars; neotropical fungal diversity; puffballs; stinkhorns

RESUMEN

Con base en la literatura disponible, se presenta una lista de 40 especies de gasteroides registradas en Paraguay desde los primeros registros conocidos. Se reporta un total de cuarenta especies, distribuidas en 16 géneros y 4 familias. Este trabajo representa la primera compilación de datos disponibles de este grupo de hongos para Paraguay. Estos números nos revela la escasa cantidad de especies citadas formalmente para la Funga del país.

Palabras clave:
diversidad de Funga neotropical; estrellas de tierra; falo hediondo; hongos nido de pájaro; hongos polvera

Introduction

Gasteroid fungi present a wide range of basidiomata structure and these distinctive characteristics lead to the designation of many genera as monotypic or with few species represented.

The existing bibliographic information on the group of gasteroid Funga in the last century was limited to a few citations made by Spegazzinii (1884Soto, M. & Wright, J.E. 2000. Taxonomía del género Geastrum (Basidiomycetes, Lycoperdales) en la provincia de Buenos Aires, Argentina. Boletín de la Sociedad Argentina de Botánica 34: 185-Spegazzini, C. 1884. Fungi Guaranitici. Pugillus 1. Anales de la Sociedad Científica Argentina 17(3): 119-134., 1888Spegazzini, C. 1888. Fungi Guaranitici. Pugillus II. Anales de la Sociedad Científica Argentina 26(1): 5-74., 1891Spegazzini, C. 1891. Fungi Guaranitici nonnulli novi vel critici. Revista Argentina de Historia Natural 1(3): 168-177.) who recorded 12 species from the country:Geastrum saccatum Fr. [as Geaster saccatus], Tulostoma cyclophorum Lloyd [as Tylostoma berteroanum], Lycoperdon lilacinum (Mont. & Berk.) Speg., Lycoperdon pseudo-lilacinum Speg., Lycoperdon pseudogemmatum Speg., Simblum sphaerocephalum Schltdl., Geastrum saccatum [Geaster spegazzinianus], Lanopila guaranitica Speg., Bovista dubiosa Speg. and Lycoperdon scleroderma Speg.

Most of the new citations were made by contemporary mycologists since the last publication of Spegazzini, 122 years ago. The next recording of gasteroid fungi in the country occurred in 2013 (Campi et al. 2013Campi, M.G., De Madrignac, B., Flecha, A.M. & Ortellado, A. 2013b. Hongos de la Reserva Natural Laguna Blanca. Salpa SA, Paraguay.), and more recent studies have been carried out by Campi and collaborators (Campi et al. 2015Campi, M.G., Maubet, Y. & Britos, L. 2015a. Mycorrhizal fungi associated with plantations of Pinus taeda L. from the National University of Asunción, Paraguay. Mycosphere, 6(4), 486-492., Campi & Maubet 2015aCampi, M.G. & Maubet, Y. 2015a. Cyathus poeppigii (Agaricales, Basidiomycetes): nuevo registro para Paraguay. Steviana 7: 74-78., 2015bCampi, M.G., Maubet, Y. & Weiler, A. 2015b. Podaxis pistillaris (Basidiomycota, Agaricomycetes) nuevo registro para el Chaco paraguayo. Boletín de la Sociedad Micológica de Madrid 39: 77-81., Campi et al. 2017Campi, M.G., Trierveiler-Pereira, L. & Maubet, Y. 2017. New records of Phallales from Paraguay. Mycotaxon, 132(2): 361-372., Maubet et al. 2018Maubet, Y., Campi, M.G., Trierveiler-Pereira, L. & Moreno, G. 2018. Contribución a la Micobiota Gasteroide de Paraguay. Nuevas citas. Bol. Soc. Micol. Madrid 42: 119-128.)

In the last decade the record of gasteroid fungi has increased but despite the efforts of mycologists in recent years, knowledge about the gasteroid mycobiota in Paraguay remains scattered and incomplete since the collections have been made sporadically and in very few areas.

Paraguay comprises two distinct geographical regions, the eastern and western, which are separated by the Paraguay River. The western region is also known as the Chaco. Which include 3 ecoregions: Humid Chaco, Dry Chaco, and Pantanal. The ecoregions of eastern Paraguay includes the Humid Chaco, Atlantic Forest, and Cerrado (Dinerstein et al. 1995Dinerstein, E., Olson, D.M., Graham, D.J, Webster, A.L., Primm, S.A, Bookbinder, M.P. & Ledec, G. 1995. Una Evaluación del Estado de Conservación de las Eco-regiones Terrestres de América Latina y el Caribe. Banco Mundial, Washington DC.).

This study is a revision of the available data and an update of the records of species of gasteroid fungi from Paraguay.

Materials and methods

This checklist has been compiled based on literature records of gasteroid fungi recorded from Paraguay. The information retrieved from the literature includes distribution records per department (figure 1), notes about habitat and substrate, and detailed descriptions and remarks of the basidiomata. The current taxonomic position is indicated, as have been described in previous publications and a brief introduction of the most outstanding characteristics of each species is provided. The classification and nomenclature followed the database Mycobank (http://www.mycobank.org). Genera and species are listed alphabetically (table 1) and the authorities for the binomials, as well as the author names and spellings, are in accordance to Robert et al. (2005Robert, V., Stegehuis, G. & Stalpers, J. 2005. The MycoBank engine and related databases. Available at Available at http://www.mycobank.org (access 14-XII-2019).
http://www.mycobank.org...
) database.

Figure 1.
Map of Paraguay with the Departments where gasteroid mushrooms have been collected are. AP: Alto Paraguay, AR-Alto Paraná, AM: Amambay, BO: Boquerón, CG: Caaguazú, CZ: Caazapá, CA: Canindeyú, CE: Central, CO: Concepción, CR: Cordillera, GU: Guairá, IT: Itapúa, MI: Misiones, ÑE: Neembucú, PR: Paraguarí, PH: Presidente Hayes, SP: San Pedro.

Table 1
List of species cited in Paraguay in alphabetical order, the collection locations and the reference.

Results and Discussion

The 40 gasteroid fungi species recorded in Paraguay are distributed among 16 genera and four families. The most highly represented family is Agaricaceae with 14 species (39%), followed by Phallaceae and Geastraceae (with 10 species each respectively, 28%). Geastrum is the genus with the highest number of species with nine taxa (25%). The departments with the most records were the Central department, corresponding to the humid Chaco ecoregion, with 19 records (47.5%) and the Paraguarí department, corresponding to the transition ecoregion between humid Chaco and Atlantic Forest, with 15 records (37.5%), two species of gasteroid fungi were cited for the Atlantic Forest ecoregion (5%), eight species for the Pantanal ecoregion (20%) and two species for the dry Chaco (5%).

It is not possible to speak of the most represented species in terms of distribution since the country lacks extensive samplings to represent all the ecoregions appropriately. Dring (1973Dring, D.M. 1973. Gasteromycetes. In: Ainsworth GC, Sparrow FK, Sussman AS (Eds) The fungi. An advanced treatise. Academic Press, New York and London, 451-478.) and Miller & Miller (1988Miller, H.R. & Miller, O.K. 1988. Gasteromycetes: Morphological and Developmental Features, with Keys to the Orders, Families, and Genera. Eureka Mad River Press, California-USA.) estimated a hundred genera of gasteromycetes, therefore the diversity of this group is still not well represented in Paraguay, since only 15 genera are recorded in the literature.

Checklist for the gasteroid species in Paraguay

Agaricaceae Chevall.

Battarrea phalloides (Dicks.) Pers.

Basionym: Lycoperdon phalloides Dicks.

Locality: BO (Campi et al. 2016bCampi, M.G., Maubet, Y. & Weiler, A. 2016b. Nueva cita de hongo xerófito para el Chaco paraguayo: Batarrea phalloides (Basidiomycota, Agaricomycetes). Bol. Soc. Micol. Madrid 41: 47-52.).

Identification. Species easily recognizable for presenting a high basidiome about 10-30 cm including the spore sac, rapid development, rust in color, stipe expending up to 25 cm in length and bearing a spore sac; exoperidium remaining as a volva and apically as scales on the endoperidium; endoperidium and spore sac hemispherical to conical, 2.8 cm to 7.2 cm in diam., later splitting circular along the middle and deteriorating. Volva membranous up to 15 cm in diameter, presenting similar color and surface as the stipe, partially adhered to the stipe base (Rea 1942Rea, P.M. 1942. Fungi of southern California. I. Mycologia 34: 563-574., Garrido-Benavent 2014Garrido-Benavent, I. 2014. The Battarrea phalloides-stevenii complex: multiple sources of evidence as a strategy to unveil cryptic species within poorly characterized taxa. Bull.Soc. Micol. Valenciana 19: 17-35., Campi et al. 2016bCampi, M.G., Maubet, Y. & Weiler, A. 2016b. Nueva cita de hongo xerófito para el Chaco paraguayo: Batarrea phalloides (Basidiomycota, Agaricomycetes). Bol. Soc. Micol. Madrid 41: 47-52.).

Calvatia cyathiformis (Bosc) Morgan

Basionym: Lycoperdon cyathiforme Bosc

Localities: CE (Campi et al. 2021Campi, M., Maubet, Y., & Trierveiler-Pereira, L. 2021. Rediscovery of Clathrus argentinus and new contributions to the gasteroid mycobiota of Paraguay. Current Research in Environmental & Applied Mycology 11(1): 90-111.)

Identification. The main characteristic of this species is a globose, compact basidiome when young, which develop a basal portion like an inverted pear or a loaf of bread at maturity. The surface can be commonly brown and broken up into mosaic-like scales. The gleba is purplish and pulverulent at maturity (Zeller & Smith 1964Zeller, S.M., Smith, A.H. 1964. The genus Calvatia in North America. Lloydia 27(3): 148-186., Morales & Kimbrough 1978Morales, M.I., Kimbrough, J.W. 1978. The Lycoperdaceae of North Central Florida I. The genera Calvatia and Disciseda. Revista de Biologia Tropical 26(1): 227-236.).

Calvatia rugosa (Berk. & M.A. Curtis) D.A. Reid

Basionym: Lycoperdon rugosum Berk. & M.A. Curtis

Locality: SP, AP (Campi & Maubet 2016Campi, M.G. & Maubet, Y. 2016. Nuevos registros de Calvatia rugosa (Berk. & M.A. Curtis) D.A. Reid y Vascellum pampeanum (Speg.) Homrich (Agaricaceae-Basidiomycota) en Paraguay. Steviana, 8(1): 43-49., Campi et al. 2021Campi, M., Maubet, Y., & Trierveiler-Pereira, L. 2021. Rediscovery of Clathrus argentinus and new contributions to the gasteroid mycobiota of Paraguay. Current Research in Environmental & Applied Mycology 11(1): 90-111.)

Identification. The main characteristic is the presence of the orange pigment that stains the basidioma when it is cut (Cortez et al. 2008Cortez, V.G., Baseia, I.G. & Silveira, R.M. 2008. Gasteromicetos (Basidiomycota) no Parque Estadual de Itapuã, Viamão, Rio Grande do Sul, Brasil. Biociências (Porto Alegre) 6(3): 291-299.). Another distinctive feature is its long rhizomorphic extension of 20-200 mm in diameter and 10-100 mm high, which folds at the base and is attached by one or more thin rhizomorphs (Wright & Albertó 2006Wright, J.E. & Albertó, E. 2006. Guía de los hongos de la Región Pampeana II. Hongos sin laminillas. Editorial L.O.L.A, Buenos Aires-Argentina.).

Calvatia lilacina (Mont. & Berk.) Henn

Basionym: Bovista lilacina Berk. & Mont.

Locality: PR (Spegazzini 1884Soto, M. & Wright, J.E. 2000. Taxonomía del género Geastrum (Basidiomycetes, Lycoperdales) en la provincia de Buenos Aires, Argentina. Boletín de la Sociedad Argentina de Botánica 34: 185-Spegazzini, C. 1884. Fungi Guaranitici. Pugillus 1. Anales de la Sociedad Científica Argentina 17(3): 119-134., 1888Spegazzini, C. 1888. Fungi Guaranitici. Pugillus II. Anales de la Sociedad Científica Argentina 26(1): 5-74.).

Identification. Bates et al. (2009Bates, S.T., Roberson, R.W. & Desjardin, D.E. 2009. Arizona gasteroid fungi I: Lycoperdaceae (Agaricales, Basidiomycota). Fungal Diversity 37: 153-207.) reported Calvatia lilacina as a synonym of Calvatia fragilis in Arizona, USA. On the other hand both species are treated as independent by Trierveiler-Pereira & Baseia (2009Trierveiler-Pereira, L. & Baseia, I.G. 2009. A checklist of the Brazilian gasteroid fungi (Basidiomycota). Mycotaxon 108: 441-444.) in Brazil. Cortez et al. (2012Cortez, V.G., Baseia, I.G. & da Silveira, R.M.B. 2012. Gasteroid mycobiota of Rio Grande do Sul, Brazil: Calvatia, Gastropila and Langermannia (Lycoperdaceae). Kew Bulletin, 67(3): 471-482.) consider this species a synonym of C. cyathiformis but other authors differ and consider C. cyathiformis as a different species (Verma et al. 2018Verma, R.K., Mishra, S.N., Pandro, V. & Thakur, A.K. 2018. Diversity and distribution of Calvatia species in India: a new record from central India. International Journal of Current Microbiology and Applied Science 7(9): 2540-2551.).

Comments: C. lilacina was recorded for Spegazzinii (1884Soto, M. & Wright, J.E. 2000. Taxonomía del género Geastrum (Basidiomycetes, Lycoperdales) en la provincia de Buenos Aires, Argentina. Boletín de la Sociedad Argentina de Botánica 34: 185-Spegazzini, C. 1884. Fungi Guaranitici. Pugillus 1. Anales de la Sociedad Científica Argentina 17(3): 119-134.) as Lycoperdon lilacinum.

Cyathus earlei Lloyd

Locality: PR (Campi et al. 2021Campi, M., Maubet, Y., & Trierveiler-Pereira, L. 2021. Rediscovery of Clathrus argentinus and new contributions to the gasteroid mycobiota of Paraguay. Current Research in Environmental & Applied Mycology 11(1): 90-111.).

Identification. This species is characterized by an exoperidium without plication, covered by a light color tomentum, organized in small tufts, endoperidium smooth or inconspicuously plicate, with a light, shiny color, contrasting with the exterior layer and peridioles with a double-layer cortex (Cruz et al. 2014Cruz, R.H., Assis, S.F., Silva, M.A. & Baseia, I.G. 2014. Revision of the genus Cyathus (Basidiomycota) from the herbaria of northeast Brazil. Mycosphere, 5(4): 531-540.).

Cyathus limbatus Tul. & C. Tul

Basionym: Cyathodes limbatum (Tul. & C. Tul.)

Localities: AP, CE, PR (Maubet et al. 2017Maubet, Y., Campi, M.G., Armoa, J. & Cristaldo, E. 2017. Nuevas citas de Cyathus Haller (Agaricaceae, Basidiomycetes) para Paraguay y ampliación de la distribución del género. Steviana, 9(1): 31-39., Campi et al. 2021Campi, M., Maubet, Y., & Trierveiler-Pereira, L. 2021. Rediscovery of Clathrus argentinus and new contributions to the gasteroid mycobiota of Paraguay. Current Research in Environmental & Applied Mycology 11(1): 90-111.).

Identification. This species is characterized by having plicate peridium on the internal and external surface, large peridioles (7-10 mm × 6-7 mm) with bilayered cortex, dark brown to black peridioles and basidiospores of 10-12 × 16-22 μm. Another distinctive feature of the species is that at the base of the pseudostipe a woolly golden brown subicule is formed (Brodie & Dennis 1954Brodie, H. & Dennis, R. 1954. The Nidulariaceae of the West Indies. Transactions of the British Mycological Society 37(2): 151-160., Trierveiler-Pereira & Baseia 2010Trierveiler-Pereira, L. & Baseia, I.G. 2010. Contribution to the knowledge of gasteroid fungi (Agaricomycetes, Basidiomycota) from the state of Paraíba, Brazil. Revista Brasileira de Biociências, 9(2):167-173.).

Cyathus poeppigii Tul. & C. Tul

Basionym: Cyathodes poeppigii (Tul. & C. Tul.)

Localities: CE (Campi & Maubet 2015aCampi, M.G. & Maubet, Y. 2015a. Cyathus poeppigii (Agaricales, Basidiomycetes): nuevo registro para Paraguay. Steviana 7: 74-78., Maubet et al. 2017Maubet, Y., Campi, M.G., Armoa, J. & Cristaldo, E. 2017. Nuevas citas de Cyathus Haller (Agaricaceae, Basidiomycetes) para Paraguay y ampliación de la distribución del género. Steviana, 9(1): 31-39.).

Identification. The most remarkable diagnostic characteristics of the species are the large size of the basidiospores (20-28 × 30-42 µm) and the deep internal and external striations of the peridium (Brodie and Dennis 1954Brodie, H. & Dennis, R. 1954. The Nidulariaceae of the West Indies. Transactions of the British Mycological Society 37(2): 151-160.).

Cyathus setosus H.J. Brodie

Localities: PR (Maubet et al. 2017Maubet, Y., Campi, M.G., Armoa, J. & Cristaldo, E. 2017. Nuevas citas de Cyathus Haller (Agaricaceae, Basidiomycetes) para Paraguay y ampliación de la distribución del género. Steviana, 9(1): 31-39.).

Identification. C. setosus is characterized by a conical peridium, of intense dark brown coloration, smooth external surface and a slightly plicated internal surface, with dark conspicuous setae 0.5-1 mm long, large peridioles without a tunic, double cortex, and a small subiculum (Brodie 1967Brodie, H.J. 1967. Cyathus setosus, a new member of the Nidulariaceae from Jamaica. Canadian Journal of Botany 45: 1-3., Trierveiler-Pereira & Baseia 2013Trierveiler-Pereira, L. & Baseia, I.G 2013. Cyathus species (Basidiomycota: Fungi) from the Atlantic Forest of Pernambuco, Brazil: taxonomy and ecological notes. Revista Mexicana de Biodiversidad, 84(1): 1-6.).

Cyathus stercoreus (Schwein.) De Toni

Basionym: Nidularia stercorea Schwein.

Locality: CG, PR, SP (Maubet et al. 2017Maubet, Y., Campi, M.G., Armoa, J. & Cristaldo, E. 2017. Nuevas citas de Cyathus Haller (Agaricaceae, Basidiomycetes) para Paraguay y ampliación de la distribución del género. Steviana, 9(1): 31-39., Campi et al. 2013bCampi, M.G., De Madrignac, B., Flecha, A.M. & Ortellado, A. 2013b. Hongos de la Reserva Natural Laguna Blanca. Salpa SA, Paraguay.).

Identification. The morphological characteristics of the species are the absence of folds, either internal or external, in the peridium, the presence of hairs grouped into golden brown strands covering the entire basidioma in the immature state, these hairs fall with age leaving the peridium with smooth appearance at maturity and taking a dark color ranging from light brown to gray (Maubet et al. 2017Maubet, Y., Campi, M.G., Armoa, J. & Cristaldo, E. 2017. Nuevas citas de Cyathus Haller (Agaricaceae, Basidiomycetes) para Paraguay y ampliación de la distribución del género. Steviana, 9(1): 31-39.).

Cyathus striatus (Huds.) Willd.

Basionym: Peziza striata Huds.

Locality: AR (Gullón 2011Gullón, M. 2011. Hongos Superiores del Refugio biológico Tati Yupi. Hernandarias. Paraguay. Biota 15: 1-68.).

Identification. Easy to recognize by the hirsute exoperidium with a shiny and striate inner surface. Microscopically, it has large ovoid basidiospores (14-17 × 7-9) (Baseia & Milanez 2001Baseia, I.G. & Milanez, A.I. 2001. Cyathus (Gasteromycetes) in areas of the Brazilian cerrado region, São Paulo State. Mycotaxon 80: 493-502.).

Podaxis pistillaris (L.) Fr.

Basionym: Lycoperdon pistillare L.

Localities: BO, CR (Campi et al. 2015bCampi, M.G., Maubet, Y. & Weiler, A. 2015b. Podaxis pistillaris (Basidiomycota, Agaricomycetes) nuevo registro para el Chaco paraguayo. Boletín de la Sociedad Micológica de Madrid 39: 77-81., Campi et al. 2021Campi, M., Maubet, Y., & Trierveiler-Pereira, L. 2021. Rediscovery of Clathrus argentinus and new contributions to the gasteroid mycobiota of Paraguay. Current Research in Environmental & Applied Mycology 11(1): 90-111.).

Identification. This species is characterized by the hard and fibrous pseudostipe that penetrates the gleba. The gleba is covered by a scaly, sub-cylindrical peridium which turns dark and powdery in the mature stage (Morse 1933Morse, E.E. 1933. A study of the genus Podaxis. Mycologia 25(1): 1-31.).

Lycoperdon pseudogemmatum Speg.

Locality: PR (Spegazzinii 1884Soto, M. & Wright, J.E. 2000. Taxonomía del género Geastrum (Basidiomycetes, Lycoperdales) en la provincia de Buenos Aires, Argentina. Boletín de la Sociedad Argentina de Botánica 34: 185-Spegazzini, C. 1884. Fungi Guaranitici. Pugillus 1. Anales de la Sociedad Científica Argentina 17(3): 119-134.).

Identification. Cortez et al. (2013Cortez, V.G., Baseia, I.G. & da Silveira, R.M. 2013. Gasteroid mycobiota of Rio Grande do Sul, Brazil: Lycoperdon and Vascellum. Mycosphere 4(4): 745-758.) consider this species a possible synonym of Lycoperdon perlatum, a highly variable species, which is characterized by the presence of spines in the conical exoperidium with a rounded apex.

Lycoperdon pseudolilacinum Speg.

Locality: PR (Spegazzinii 1884Soto, M. & Wright, J.E. 2000. Taxonomía del género Geastrum (Basidiomycetes, Lycoperdales) en la provincia de Buenos Aires, Argentina. Boletín de la Sociedad Argentina de Botánica 34: 185-Spegazzini, C. 1884. Fungi Guaranitici. Pugillus 1. Anales de la Sociedad Científica Argentina 17(3): 119-134.).

Identification. This species resembles L. lilacinum but differs in the color of the gleba and the very fragile peridium (Spegazzini 1884Soto, M. & Wright, J.E. 2000. Taxonomía del género Geastrum (Basidiomycetes, Lycoperdales) en la provincia de Buenos Aires, Argentina. Boletín de la Sociedad Argentina de Botánica 34: 185-Spegazzini, C. 1884. Fungi Guaranitici. Pugillus 1. Anales de la Sociedad Científica Argentina 17(3): 119-134.).

Tulostoma cyclophorum Lloyd

Basionym: Tulostoma berteroanum f. pampeanum Speg.

Locality: CE, PR (Spegazzini 1884Soto, M. & Wright, J.E. 2000. Taxonomía del género Geastrum (Basidiomycetes, Lycoperdales) en la provincia de Buenos Aires, Argentina. Boletín de la Sociedad Argentina de Botánica 34: 185-Spegazzini, C. 1884. Fungi Guaranitici. Pugillus 1. Anales de la Sociedad Científica Argentina 17(3): 119-134., Campi et al. 2016aCampi, M.G., Maubet, Y., Moreno, G. & Altés, A. 2016a. Tulostoma cyclophorum (Basidiomycota, Agaricomycetes) new record for Paraguay. Bol. Soc. Micol. Madrid 40: 129-132.).

Identification. This species is easily distinguishable by persistent membranous exoperidium in old specimens, endoperidium with a velvety texture and abundant mycosclereids well visible to the stereoscope on its surface, a mamiform and fimbriated mouth, and by the globose basidiospores with ridges which offer a characteristic cross-linked to sub-crosslinked morphology (Campi et al. 2016aCampi, M.G., Maubet, Y., Moreno, G. & Altés, A. 2016a. Tulostoma cyclophorum (Basidiomycota, Agaricomycetes) new record for Paraguay. Bol. Soc. Micol. Madrid 40: 129-132.).

Comments: This species was recorded as Tulostoma berteroanum by Spegazzini (1884Soto, M. & Wright, J.E. 2000. Taxonomía del género Geastrum (Basidiomycetes, Lycoperdales) en la provincia de Buenos Aires, Argentina. Boletín de la Sociedad Argentina de Botánica 34: 185-Spegazzini, C. 1884. Fungi Guaranitici. Pugillus 1. Anales de la Sociedad Científica Argentina 17(3): 119-134.).

Tulostoma pygmaeum Lloyd

Locality: PR (Campi et al. 2021Campi, M., Maubet, Y., & Trierveiler-Pereira, L. 2021. Rediscovery of Clathrus argentinus and new contributions to the gasteroid mycobiota of Paraguay. Current Research in Environmental & Applied Mycology 11(1): 90-111.)

Identification. This species is characterized by its tubular mouth, hyphal exoperidium, and basidiospores with a distinct verrucose ornamentation (Moreno et al. 1995Moreno, G., Altés, A., Ochoa, C. & Wright, J.E. 1995. Contribution to the study of the Tulostomataceae in Baja California, Mexico. I. Mycologia 87(1): 96-120.).

Vascellum pampeanum (Speg.) Homrich, in Homrich & Wright

Basionym: Lycoperdon pampeanum Speg.

Locality: CE (Campi & Maubet 2016Campi, M.G. & Maubet, Y. 2016. Nuevos registros de Calvatia rugosa (Berk. & M.A. Curtis) D.A. Reid y Vascellum pampeanum (Speg.) Homrich (Agaricaceae-Basidiomycota) en Paraguay. Steviana, 8(1): 43-49.).

Identification. V. pampeanum is characterized by having a diaphragm that separates the gleba from the subgleba, the presence of mycosclereids on the surface of the exoperidium and the presence of eucapilicium (Homrich & Wright 1988Homrich, M.H. & Wright, J.E. 1988. South American Gasteromycetes. II. The genus Vascellum. Canadian Journal of Botany 66: 1285-1307.).

Geastraceae Corda

Geastrum argentinum (Speg.)

Basionym: Geastrum argentines Speg.

Locality: AP (Campi et al. 2018Campi, M., Maubet, Y. & Trierveiler-Pereira, L. 2018. “Estrellas de tierra” Geastrum (Geastraceae, Basidiomycota): nuevas citas para el Pantanal paraguayo. Steviana 10(2): 3-16.).

Identification. G. argentinum is characterized by emerging from a whitish subicule and by an external surface of the velvety mycelial layer which is detached from fibrous tissue (Zamora et al. 2014Zamora, J.C., Calonge, F.D. & Martín, M.P. 2014. Integrative taxonomy reveals an unexpected diversity in Geastrum section Geastrum (Geastrales, Basidiomycota). Persoonia 34: 130-165.).

Geastrum coronatum Pers.

Basionym: Geastrum limbatum Fr.

Locality: CE (Campi & Maubet 2015bCampi, M.G., Maubet, Y. & Weiler, A. 2015b. Podaxis pistillaris (Basidiomycota, Agaricomycetes) nuevo registro para el Chaco paraguayo. Boletín de la Sociedad Micológica de Madrid 39: 77-81.).

Identification. This species is mainly characterized by arched basidiomata, dark brown and asperulate endoperidium, a myceliar layer covered with abundant debris, a fibrillose peristome, larger pedicel (up to 4 mm high), and basidiospores with prominent truncate warts (Sunhede 1989Sunhede, S. 1989. Geastraceae (Basidiomycotina) morphology, ecology and systematics with emphasis on the north Europearn species. Synopsis Fungorum 1. Oslo: Fungiflora., Soto & Wright 2000Soto, M. & Wright, J.E. 2000. Taxonomía del género Geastrum (Basidiomycetes, Lycoperdales) en la provincia de Buenos Aires, Argentina. Boletín de la Sociedad Argentina de Botánica 34: 185-Spegazzini, C. 1884. Fungi Guaranitici. Pugillus 1. Anales de la Sociedad Científica Argentina 17(3): 119-134., Alves & Cortez 2016Alves, C.R. & Cortez, V.G. 2016. Gasteroid Phallomycetidae (Basidiomycota) from the Parque Estadual São Camilo, Paraná, Brazil. Iheringia Série Botânica 71: 27-42.).

Geastrum hariotii Lloyd

Basionym: Geaster harioti Lloyd

Locality: AP (Campi et al. 2018Campi, M., Maubet, Y. & Trierveiler-Pereira, L. 2018. “Estrellas de tierra” Geastrum (Geastraceae, Basidiomycota): nuevas citas para el Pantanal paraguayo. Steviana 10(2): 3-16.).

Identification. The species is characterized by having non-hygroscopic exoperidium, endoperidium sessile or with a very short stipe, peristoma strongly plicated and well defined. (Trierveiler-Pereira & Silveira 2012Trierveiler-Pereira, L. & Silveira, R.M.B. 2012. On the Geastrum species (Geastraceae, Basidiomycota) described by Rick. Phytotaxa, 61(1): 37-46.).

Geastrum minimum Schwein.

Locality: CE (Campi et al. 2015aCampi, M.G. & Maubet, Y. 2015b. Especies de Geastrum (Geastraceae, Basidiomycota) nuevos registros para Paraguay. Steviana 7: 79-88.).

Identification. G. minimum is normally treated as a single, rather variable species, mostly small-sized and often showing quite large crystals of calcium oxalate on the endoperidial surface (Zamora et al. 2014Zamora, J.C., Calonge, F.D. & Martín, M.P. 2014. Integrative taxonomy reveals an unexpected diversity in Geastrum section Geastrum (Geastrales, Basidiomycota). Persoonia 34: 130-165.)

Geastrum pampeanum var. pallidum Speg.

Basionym: Geastrum pampeanus var. pallidus Speg.

Locality: AP (Campi et al. 2018Campi, M., Maubet, Y. & Trierveiler-Pereira, L. 2018. “Estrellas de tierra” Geastrum (Geastraceae, Basidiomycota): nuevas citas para el Pantanal paraguayo. Steviana 10(2): 3-16.).

Identification. This species is characterized by a small basidiomata (17-30 mm wide), non-hygroscopic exoperidium, sessile endoperidium and basidiospores of 3.5-5 μm diam. (Campi et al. 2018Campi, M., Maubet, Y. & Trierveiler-Pereira, L. 2018. “Estrellas de tierra” Geastrum (Geastraceae, Basidiomycota): nuevas citas para el Pantanal paraguayo. Steviana 10(2): 3-16.).

Geastrum pusillipilosum J.O. Sousa, Alfredo, R.J. Ferreira, M.P Martín & Baseia

Locality: PR (Campi et al. 2021Campi, M., Maubet, Y., & Trierveiler-Pereira, L. 2021. Rediscovery of Clathrus argentinus and new contributions to the gasteroid mycobiota of Paraguay. Current Research in Environmental & Applied Mycology 11(1): 90-111.).

Identification. Recognized by a small basidiomata (up to 17 mm wide), an exoperidium covered with short hairs (up to 1 mm), a fibrillose peristome and basidiospores of 4.4-6 × 4.3-5.9 (-6.8) μm diam (Crous et al. 2016Crous, P.W., Wingfield, M.J., Burgess, T.I., Hardy, G.S.J., Crane, C., Barrett, S., ... Stchigel A.M. 2016. Fungal Planet description sheets: 469-557. Persoonia: Molecular Phylogeny and Evolution of Fungi, 37: 218., Lima & Baseia 2018Lima, A.A. & Baseia, I.G. 2018. Gasteroid fungi (Basidiomycota) from two protected natural areas in Rio Grande do Norte State, Brazil. Journal of Fungal Biology, 8(6): 585-605.).

Geastrum saccatum Fr.

Localities: AP, CE, PR (Spegazzini 1884Soto, M. & Wright, J.E. 2000. Taxonomía del género Geastrum (Basidiomycetes, Lycoperdales) en la provincia de Buenos Aires, Argentina. Boletín de la Sociedad Argentina de Botánica 34: 185-Spegazzini, C. 1884. Fungi Guaranitici. Pugillus 1. Anales de la Sociedad Científica Argentina 17(3): 119-134., Campi et al. 2018Campi, M., Maubet, Y. & Trierveiler-Pereira, L. 2018. “Estrellas de tierra” Geastrum (Geastraceae, Basidiomycota): nuevas citas para el Pantanal paraguayo. Steviana 10(2): 3-16.).

Identification. This species is characterized by a large basidiomata (up to 8 cm in diameter), involute lacinias, non-hygroscopic, a prominent conspicuous collar around the endoperidium from the pseudoparenchymal layer of exoperidium, sessile endoperidium, delimited or unbounded fibrous peristoma (Sunhede 1989Sunhede, S. 1989. Geastraceae (Basidiomycotina) morphology, ecology and systematics with emphasis on the north Europearn species. Synopsis Fungorum 1. Oslo: Fungiflora.).

Geastrum schweinitzii (Berk. & M.A. Curtis) Zeller

Basionym: Coilomyces schweinitzii Berk. & M.A. Curtis

Locality: CE (Campi & Maubet 2015bCampi, M.G., Maubet, Y. & Weiler, A. 2015b. Podaxis pistillaris (Basidiomycota, Agaricomycetes) nuevo registro para el Chaco paraguayo. Boletín de la Sociedad Micológica de Madrid 39: 77-81.).

Identification. This species is recognized by small cespitose basidiomata and by the presence of a white to yellowish, subiculum that spreads throughout the substrate, which is generally decomposing wood (Baseia et al. 2003Baseia, I.G., Cavalcanti, M.A. & Milanez, A.I. 2003. Additions to our knowledge of the genus Geastrum (Phallales: Geastraceae) in Brazil. Mycotaxon 85: 409-416.). Some specimens previously identified as G. schweinitzii for the neotropics were different species based on phylogenetic analysis and thus the species could be endemic of the region were the type material was collected (Accioly et al. 2019Accioly, T., Sousa, J.O., Moreau, P.A., Lécuru, C., Silva, B.D., Roy, M., Gardes, M., Baseia, I.G. & Martín, M.P. 2019. Hidden fungal diversity from the Neotropics: Geastrum hirsutum, G. schweinitzii (Basidiomycota, Geastrales) and their allies. PloS one 14(2): e0211388.)

Geastrum triplex Jungh. in Tijdschr

Locality: SP, CE, AP (Campi et al. 2013bCampi, M.G., De Madrignac, B., Flecha, A.M. & Ortellado, A. 2013b. Hongos de la Reserva Natural Laguna Blanca. Salpa SA, Paraguay., Campi et al. 2018Campi, M., Maubet, Y. & Trierveiler-Pereira, L. 2018. “Estrellas de tierra” Geastrum (Geastraceae, Basidiomycota): nuevas citas para el Pantanal paraguayo. Steviana 10(2): 3-16.).

Identification. This species is characterized by large basidiomata (up to 8 cm in diameter), involute lacinias, non-hygroscopic, prominent conspicuous collar around the endoperidium from the pseudoparenchymal layer of exoperidium, sessile endoperidium, delimited or unbounded fibrous peristoma. (Campi et al. 2018Campi, M., Maubet, Y. & Trierveiler-Pereira, L. 2018. “Estrellas de tierra” Geastrum (Geastraceae, Basidiomycota): nuevas citas para el Pantanal paraguayo. Steviana 10(2): 3-16.). Recent phylogenetic studies suggest that the morphological concept of the species actually represents an assemblage of closely related species which vary in the distribution patterns (Kasuya et al. 2012Kasuya, T., Hosaka, K., Uno, K. & Kakishima, M. 2012. Phylogenetic placement of Geastrum melanocephalum and polyphyly of Geastrum triplex. Mycoscience 53: 411-426.).

Geastrum violaceum Rick

Locality: AR (Campi et al. 2013aCampi, M.G., De Madrignac, B., Flecha, A.M. & Gullón, M. 2013a. Geastrum violaceum Rick (Geastraceae, Basidiomycota) nuevo registro para Paraguay. Reportes Científicos 4(2): 15-18.).

Identification. This species is mainly distinguished by the pink, red to pale violet color of the exoperidium, a character that easily separates it from other species of the genus (Trierveiler-Pereira & Silveira 2012Trierveiler-Pereira, L. & Silveira, R.M.B. 2012. On the Geastrum species (Geastraceae, Basidiomycota) described by Rick. Phytotaxa, 61(1): 37-46.).

Myriostoma calongei Baseia, J.O. Sousa & M.P. Martín

Locality: PR (Campi et al. 2021Campi, M., Maubet, Y., & Trierveiler-Pereira, L. 2021. Rediscovery of Clathrus argentinus and new contributions to the gasteroid mycobiota of Paraguay. Current Research in Environmental & Applied Mycology 11(1): 90-111.).

Identification. This species is characterized by a verrucose endoperidium, with prominent triangular processes and could be restricted to South America (Sousa et al. 2017Sousa, J., Suz, L.M., García, M.A., Alfredo, D.S., Conrado, L.M., Marinho, P., Ainswoth, A.M., Baseia, I.G. & Martín, M.P. 2017. More than one fungus in the pepper pot: Integrative taxonomy unmasks hidden species within Myriostoma coliforme (Geastraceae, Basidiomycota). PloS one 12(6): e0177873. ).

Phallaceae Corda

Blumenavia rhacodes Möller

Locality: CR (Campi et al. 2021Campi, M., Maubet, Y., & Trierveiler-Pereira, L. 2021. Rediscovery of Clathrus argentinus and new contributions to the gasteroid mycobiota of Paraguay. Current Research in Environmental & Applied Mycology 11(1): 90-111.).

Identification. The species is characterized by a pale orange to greyish orange (beige) receptacle consisting of 3-5 columns, and glebifers occuring on lateral expansions (“teeth”) distributed along the column’s margins (Trierveiler-Pereira et al. 2019Trierveiler-Pereira, L., Meijer, A.A.R. & Silveira, R.M.B. 2019. Phallales (Agaricomycetes, Fungi) from Southern Brazil. Studies in Fungi 4(1): 162-184., Melanda et al. 2020Melanda, G.C., Accioly T., Ferreira, R.J., Rodrigues, A.C., Cabral, T.S., Coelho, G., ... & Baseia, I.G. 2020. Diversity trapped in cages: Revision of Blumenavia Möller (Clathraceae, Basidiomycota) reveals three hidden species. PloS one 15(5): e0232467.).

Clathrus argentinus Domínguez

Locality: CE (Campi et al. 2021Campi, M., Maubet, Y., & Trierveiler-Pereira, L. 2021. Rediscovery of Clathrus argentinus and new contributions to the gasteroid mycobiota of Paraguay. Current Research in Environmental & Applied Mycology 11(1): 90-111.).

Identification. This species is characterized by having the receptacle arms not joined at the base and a strongly yellow basal mycelium, and glebiferous zones with digitiform extensions, are at the intersection of the arms that form the network (Domínguez de Toledo 1985).

Clathrus chrysomycelinus Möller

Basionym: Clathrella chrysomycelina (Möller) E. Fisch.

Locality: PR (Maubet et al. 2018Maubet, Y., Campi, M.G., Trierveiler-Pereira, L. & Moreno, G. 2018. Contribución a la Micobiota Gasteroide de Paraguay. Nuevas citas. Bol. Soc. Micol. Madrid 42: 119-128.).

Identification. This species is characterized by obovoid, perforate, white to yellowish receptacle, isodiametric meshes, having the receptacle arms joined at the base and a strong yellow basal mycelium. In addition, the glebe is confined to glebifers at the inner side of the arm’s intersections (Dring et al. 1971Dring, D.M., Meeker, J. & Goos, R. 1971. Clathrus oahuensis, a new species from Hawai. Mycologia 63(4): 893-897., Dring 1980Dring, D.M. 1980. Contributions towards a rational arrangement of the Clathraceae. Kew Bulletin 35(1): 1-96.).

Comments: Maubet et al. (2018Maubet, Y., Campi, M.G., Trierveiler-Pereira, L. & Moreno, G. 2018. Contribución a la Micobiota Gasteroide de Paraguay. Nuevas citas. Bol. Soc. Micol. Madrid 42: 119-128.) mention that the material was collected in Ybycui city, reviewing the herbarium material, the right collection site corresponds to Piribebuy city, Cordillera Department.

Clathrus columnatus Bosc

Basionym: Laternea columnata (Bosc) Nees

Locality: CE (Campi et al. 2017Campi, M.G., Trierveiler-Pereira, L. & Maubet, Y. 2017. New records of Phallales from Paraguay. Mycotaxon, 132(2): 361-372.).

Identification. Species characterized by having 2-5 robust spongy reddish to orange columns free at the base and fused at the apex. The gleba is spread on the internal portion of the columns and not confined to a glebifer (Sandoval-Leiva et al. 2014Sandoval-Leiva, P., Henríquez, J.L. & Trierveiler-Pereira, L. 2014. Additions to the Chilean phalloid mycota. Mycotaxon 128: 45-54., Magnago et al. 2013Magnago, A.C., Trierveiler-Pereira, L. & Neves, M.A. 2013. Phallales (Agaricomycetes, Fungi) from the tropical Atlantic Forest of Brazil. Journal of the Torrey Botanical Society 140 (2): 236-244. ).

Clathrus crispus Turpin

Basionym: Clathrella crispa (Turpin) E. Fisch.

Locality: AP (Maubet et al. 2018Maubet, Y., Campi, M.G., Trierveiler-Pereira, L. & Moreno, G. 2018. Contribución a la Micobiota Gasteroide de Paraguay. Nuevas citas. Bol. Soc. Micol. Madrid 42: 119-128.).

Identification. This species is characterized by having a small receptacle (5 cm or less, some larger up to 15 cm), globose to subglobose of reddish coloration, with somewhat regular meshes (usually round at the apex and more elongated towards the base) and with a corrugated and folded membrane surrounding the meshes where the gleba is located (Dring 1980Dring, D.M. 1980. Contributions towards a rational arrangement of the Clathraceae. Kew Bulletin 35(1): 1-96.).

Itajahya galericulata Möller, Bras.

Localities: CE, PR (Campi et al. 2017Campi, M.G., Trierveiler-Pereira, L. & Maubet, Y. 2017. New records of Phallales from Paraguay. Mycotaxon, 132(2): 361-372., 2021Campi, M., Maubet, Y., & Trierveiler-Pereira, L. 2021. Rediscovery of Clathrus argentinus and new contributions to the gasteroid mycobiota of Paraguay. Current Research in Environmental & Applied Mycology 11(1): 90-111.).

Identification. Characterized by massive ‘eggs’ (3‒8 cm high), a white pseudostipe, a hemispherical, wig-like receptacle with a flat apex that is covered with a calyptra (membranous cap) (Trierveiler-Pereira et al. 2019Trierveiler-Pereira, L., Meijer, A.A.R. & Silveira, R.M.B. 2019. Phallales (Agaricomycetes, Fungi) from Southern Brazil. Studies in Fungi 4(1): 162-184.).

Itajahya rosea (Delile) E. Fisch.

Basionym: Phallus roseus Delile

Localities: CE (Campi et al. 2017Campi, M.G., Trierveiler-Pereira, L. & Maubet, Y. 2017. New records of Phallales from Paraguay. Mycotaxon, 132(2): 361-372.)

Identification. This species, which has a phalloid shape, is characterized by the presence of a calyptra at the apex of the receptacle and the pinkish pseudostipe (Ottoni et al. 2010Ottoni, B.S., Silva, B.D.B., Fazolino, E.P. & Baseia, I.G. 2010. Phallus roseus, first record from the neotropics. Mycotaxon 112: 5-8.).

Lysurus sphaerocephalum (Schltdl.) Hern. Caff., Urcelay, Hosaka & L.S. Domínguez

Basionym: Simblum sphaerocephalum Schltdl.

Localities: CE, PR (Spegazzini 1891Spegazzini, C. 1891. Fungi Guaranitici nonnulli novi vel critici. Revista Argentina de Historia Natural 1(3): 168-177., Campi et al. 2021Campi, M., Maubet, Y., & Trierveiler-Pereira, L. 2021. Rediscovery of Clathrus argentinus and new contributions to the gasteroid mycobiota of Paraguay. Current Research in Environmental & Applied Mycology 11(1): 90-111.)

Identification. The diagnostic features of the species are the fertile portion of the receptacle that is an irregular net, reddish, enclosing the stipe, which exceeds the stipe width. Stipe reddish and stylized, length clearly exceeds width, not striate (Hernández-Caffot et al. 2018Hernandez Caffot, M.L., Hosaka, K., Domínguez, L.S. & Urcelay, C. 2018. Molecular and morphological data validate the new combination of Lysurus sphaerocephalum from Argentina, with some additional records on Phallales (Agaricomycetes). Mycologia, 110(2): 419-433.).

Mutinus argentinus Speg.

Localities: CE, CR (Campi et al. 2017Campi, M.G., Trierveiler-Pereira, L. & Maubet, Y. 2017. New records of Phallales from Paraguay. Mycotaxon, 132(2): 361-372., Maubet et al. 2018Maubet, Y., Campi, M.G., Trierveiler-Pereira, L. & Moreno, G. 2018. Contribución a la Micobiota Gasteroide de Paraguay. Nuevas citas. Bol. Soc. Micol. Madrid 42: 119-128.).

Identification. The species is characterized by a pinkish pseudostipe that tapers towards the apex, and a granulose, bright red fertile part with a small pore at the tip when mature (Trierveiler-Pereira et al. 2019Trierveiler-Pereira, L., Meijer, A.A.R. & Silveira, R.M.B. 2019. Phallales (Agaricomycetes, Fungi) from Southern Brazil. Studies in Fungi 4(1): 162-184.).

Comments: Maubet et al. (2018Maubet, Y., Campi, M.G., Trierveiler-Pereira, L. & Moreno, G. 2018. Contribución a la Micobiota Gasteroide de Paraguay. Nuevas citas. Bol. Soc. Micol. Madrid 42: 119-128.) mention that the material was collected in Ybycui city, reviewing the herbarium material the right collection site corresponds to Piribebuy city, Cordillera Department.

Phallus campanulatus Berk.

Locality: CE (Campi et al. 2021Campi, M., Maubet, Y., & Trierveiler-Pereira, L. 2021. Rediscovery of Clathrus argentinus and new contributions to the gasteroid mycobiota of Paraguay. Current Research in Environmental & Applied Mycology 11(1): 90-111.)

Identification. The species is characterized by a whitish pseudostipe with small perforations, an off white, perforate, narrow campanulate, wrinkled to minutely rugulose receptacle with a toothed margin. The pore at the receptacle apex is so prominent that a ring-like structure can be observed around it (Trierveiler-Pereira et al. 2019Trierveiler-Pereira, L., Meijer, A.A.R. & Silveira, R.M.B. 2019. Phallales (Agaricomycetes, Fungi) from Southern Brazil. Studies in Fungi 4(1): 162-184.).

Phallus indusiatus Vent.

Localities:CE (Maubet et al. 2018Maubet, Y., Campi, M.G., Trierveiler-Pereira, L. & Moreno, G. 2018. Contribución a la Micobiota Gasteroide de Paraguay. Nuevas citas. Bol. Soc. Micol. Madrid 42: 119-128.).

Identification. The species is characterized by a whitish pseudostipe, a reticulate, white receptacle and the long, white indusium. Mycelial strands are often abundant and have purplish pigments (Trierveiler-Pereira et al. 2019Trierveiler-Pereira, L., Meijer, A.A.R. & Silveira, R.M.B. 2019. Phallales (Agaricomycetes, Fungi) from Southern Brazil. Studies in Fungi 4(1): 162-184.).

Sclerodermataceae Corda

Pisolithus arhizus (Scop.) Rauschert

Basionym: Lycoperdon arrizon Scop.

Locality: CE (Campi et al. 2015aCampi, M.G. & Maubet, Y. 2015a. Cyathus poeppigii (Agaricales, Basidiomycetes): nuevo registro para Paraguay. Steviana 7: 74-78.)

Identification. This species is recognized by its epigeous, globose basidiomata, echinate basidiospores (Campi et al. 2015aCampi, M.G., Maubet, Y. & Weiler, A. 2015b. Podaxis pistillaris (Basidiomycota, Agaricomycetes) nuevo registro para el Chaco paraguayo. Boletín de la Sociedad Micológica de Madrid 39: 77-81.).

Scleroderma bovista Fr.

Locality: CE (Campi et al. 2015aCampi, M.G. & Maubet, Y. 2015a. Cyathus poeppigii (Agaricales, Basidiomycetes): nuevo registro para Paraguay. Steviana 7: 74-78.)

Identification. The species is characterized by the presence of a smooth yellowish-brown peridium, with some reddish dark brown areas, basidiospores (11‒) 12‒14 (‒16) µm in diameter, with a well-developed crosshair (Nouhra et al. 2012Nouhra, E., Hernandez Caffot, M., Pastor, N. & Crespo, E. 2012. The species of Scleroderma from Argentina, including a new species from the Nothofagus forest. Mycologia 104(2): 488-495. ).

Acknowledgements

This work was carried out within the framework of the project PINV18-699 “Development of molecular tools for the taxonomic identification of fungi from 3 Ecoregions of Paraguay”, funded by the Consejo Nacional de Ciencias y Tecnología (CONACYT). The authors thanks to Professor Andrea Weiler, for the contribution with the map. Anonymous reviewers are generous acknowledged, for discussions and comments that improved the manuscript. Sc. Andrew G. Cervantes is acknowledged, for the revision of the manuscript’s English and for the valuable contribution to the writing.

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Edited by

Associate Editor:

Alice da Cruz Lima Gerlach

Publication Dates

  • Publication in this collection
    02 Sept 2022
  • Date of issue
    2022

History

  • Received
    05 Jan 2021
  • Accepted
    20 Dec 2021
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