Mycosphere Doi 10.5943/mycosphere/4/1/5 Ectomycorrhizal fungi from southern Brazil – a literature-based review, their origin and potential hosts

Sulzbacher MA1*, Grebenc, T2, Jacques RJS3 and Antoniolli ZI3

1Universidade Federal de Pernambuco, Departamento de Micologia/CCB, Av. Prof. Nelson Chaves, s/n, CEP: 50670- 901, Recife, PE, Brazil 2Slovenian Forestry Institute Vecna pot 2, SI-1000 Ljubljana, Slovenia 3Universidade Federal de Santa Maria, Departamento de Solos, CCR Campus Universitário, 971050-900, Santa Maria, RS, Brazil

Sulzbacher MA, Grebenc T, Jacques RJS, Antoniolli ZI 2013 – Ectomycorrhizal fungi from southern Brazil – a literature-based review, their origin and potential hosts. Mycosphere 4(1), 61– 95, Doi 10.5943 /mycosphere/4/1/5

A first list of ectomycorrhizal and putative ectomycorrhizal fungi from southern Brazil (the states of Rio Grande do Sul, Santa Catarina and Paraná), their potential hosts and origin is presented. The list is based on literature and authors observations. Ectomycorrhizal status and putative origin of listed was assessed based on worldwide published data and, for some genera, deduced from taxonomic position of otherwise locally distributed species. A total of 144 species (including 18 doubtfull species) in 49 genera were recorded for this region, all accompanied with a brief distribution, habitat and substrate data. At least 30 collections were published only to the level and require further taxonomic review.

Key words – distribution – habitat – – neotropics – regional list

Article Information Received 28 November 2012 Accepted 20 December 2012 Published online 10 February 2013 *Corresponding author: MA Sulzbacher – e-mail – [email protected]

Introduction work of Singer & Araújo (1979), Singer et al. Ectomycorrhizal fungi (ECM) and their (1983) and Singer & Aguiar (1986) in the partner trees are well studied around the globe. Amazon region. Recent publications with Nevertheless, Europe, North America and surveys of putative ECM in exotic and native several tropical regions have been considerably plantations throughout the country have more explored while ECM dominated habitats contributed to the knowledge of this group in in South America, Southeast Asia, Africa, and different areas, producing new records and remain relative undersampled newly described species (Baseia & Milanez (Tedersoo et al. 2010). In Brazil little is known 2000, 2002, Gurgel et al. 2008, Menolli et al. about the ECM communities. Among the 2009a, 2009b, Wartchow & Maia 2007, -based studies of ECM conducted in Wartchow et al. 2009, Wartchow & Cavalcanti Brazil, focusing both on native as well as 2010, Wartchow et al. 2012a, Wartchow exotic woody plants, we refer mainly to the 2012a, 2012b). All of these works are

61 Mycosphere Doi 10.5943/mycosphere/4/1/5 taxonomic and they do not confirm the semi-desciduous forest, as well as pionner association between plant and fungi. vegetation (de Meijer 2008, Veloso et al. Henkel et al. (2012) presented a 1991). The Campos Sulinos (or Pampa) biome comprehensive study of the diversity of is covered by open grassy formations, used as ectomycorrhizal fungi sporocarps in the Guiana natural pastures (Fiaschi & Pirani 2009). It is Shield. Neves & Capelari (2007) published a estimated that there are around 3,000 grassland Brazilian checklist of , reporting 20 species which belong mainly to the botanical genera and 70 species belonging to Boletales families Poaceae, Asteraceae, Cyperacea, sensu Kirk et al. (2001), excluding the Fabaceae, Apiacea, Oxalidaceae, Verbanaceae, . In a recent checklist and Iridaceae. The main forest formations in Trierveiler-Pereira & Baseia (2009) reported Campus Sulinos are found at the Northern limit 232 taxa of Brazilian . In of the biome, in the transition area to Atlantic southern Brazil, the first contributions were Rain Forest. In other regions, plant formations made by Johannes Rick during the first half of with trees are found mainly as gallery forests the twentieth century (reviewed in Fidalgo (islands of trees within the grassland) and 1962, Mauhs 2000). In the 1950s, Rolf Singer shrub forests (Overbeck et al. 2006, 2007). A reviewed some of the species collected by particular habitat for ECM fungi is forest tree Rick, named Fungi Rickiani (Singer 1953a). plantations. Currently more than 5.98 Mha are Several studies have added to the knowledge of planted with , eucalypts and acacia ECM fungi in Brazil. Putzke (1994) has monocultures in Brazil (SBS 2008). provided a checklist for the Brazilian The central, western and northern , listing 1,011 taxa. Many genera regions of Brazil have a tropical climate with published by Putzke remain unresolved and high annual temperatures averaging near 25° C need additional taxonomic confirmation. with rainfall characterized by a wet season Most of these contributions were from October to March and a dry season from checked by means of morphological features of April to September (Rocha et al. 2009, the sporocarps. Nowadays, however, Vourlitis et al. 2002). In the south of Brazil the systematics and of ECM fungi have subtropical climate is defined by four seasons been under profound changes, mainly due to and uniform annual rainfall. In this region, the use of molecular tools (Binder & Hibbett maximum average temperatures reach 30° C, 2002, Binder & Hibbett 2006, Hibbett et al. and average minimum temperatures in the 1997, Hibbett 2006, Hosaka et al. 2006, winter are around 5° C (annual averages near Matheny et al. 2006, Miller et al. 2006, 18° C). As a consequence of its subtropical Moncalvo et al. 2002, 2006). Despite the fact, climate, this region has less intemperized soil data on tropical and subtropical fungi were than tropical Brazil and higher organic carbon rarely included in DNA-based studies, content (Dieckow et al. 2009). compromising the knowledge about the In view of the differences in climate, identification and phylogenetic placement of soil and vegetation in the Southern part of those fungi (Rinaldi et al. 2008). Brazil, this survey aimed to provide Brazil is the fifth largest country in the information on the diversity of ECM fungi in world, with more than 8.5 million km2, with Southern Brazil (Paraná, Santa Catarina and two recognized hotspots for the conservation of Rio Grande do Sul), including a summary of biodiversity, the Atlantic Forest and the information on putative hosts, nativeness, and a Cerrado (Myers et al. 2000). The southern part literature-based review of the area of origin. of Brazil is characterized by two domains, This study will, potentially, facilitate future Atlantic Forest and Campos Sulinos or studies on the systematics and biogeography of southern grasslands (Fiaschi & Pirani 2009). ECM fungi in Brazil. The Atlantic forest is characterizated by high species diversity and endemism. It includes Methods different forest types: dense ombrophilous The checklist and meta-analyses were forest, mixed ombrophilous forest (including based on a literature survey of scientific papers Araucaria forest), seasonal and citing species of putative ectomycorrhizal fungi

62 Mycosphere Doi 10.5943/mycosphere/4/1/5 from Southern Brazil. The following chrysoleuca Pegler information was retrieved: distribution of each Distribution – PR – Antonina. recorded species (per state), potential ECM Habitat and substrate – dense partners, the nativeness or potential origin, ombrophilous forest; terricolous. general habitat requirements and known Literature – de Meijer (2006: 11). substrates. The ectomycorrhizal status for each Comments – this species is apparantly species is based either on the available distributed in (sub) tropical America. It was description of ectomycorrhizae or on the cited from the US Virgin Islands (Miller et al. taxonomic position of the species (Rinaldi et al. 2000). 2008, Tedersoo et al. 2010). The list and correlations also include several records by Amanita grallipes Bas & de Meijer Rick (1961a, b) which may represent nomina Distribution – PR – Curitiba. dubia, synonyms or species of other genera. Habitat and substrate – mixed Genera and species are listed ombrophilous forest, seasonal semi-deciduous alphabetically according to the MycoBank alluvial forest; terricolous. database Literature – Bas & de Meijer (1993: (http://www.mycobank.org/MycoTaxo.aspx), 345), de Meijer (2001: 112, 2006: 11, 2008: 44, and the Index Fungorum 140). (http://www.indexfungorum.org/names/names. Comments – de Meijer (2008) refers asp). The authority of the species names are that A. grallipes belongs to sect. Lepidella given as in the original publications. The area subsect. Vittadiniae Bas. It is not known if A. of „Southern Brazil‟ included in the study is grallipes is ectomycorrhizal, but many species represented by the States of Rio Grande do Sul in the subsect. Vittadiniae are most certainly (RS), Santa Catarina (SC) and Paraná (PR). non-ectomycorrhizal. Wolfe et al. (2012) have shown that some species of Amanita (A. thiersii Results and Discussion Bas) present saprotrophic nutrition. The following ECM fungi have been recorded from Southern Brazil. Amanita muscaria (L.: Fr.) Lam. sensu lato Distribution – PR – Colombo; SC – Alnicola spadicea (D.A. Reid) Bon Correia Pinto, Joinville, Três Barras; RS – Distribution – PR – Rio Negro. Nova Petrópolis. Habitat and substrate – planted Salix; Habitat and substrate – plantations of terricolous. Pinus taeda L.; terricolous. Literature – de Meijer (2001: 113, Literature – de Meijer (2001: 113, 2006: 11). 2006: 11, 2008: 142), Giachini et al. (2000: Comments – The genus Alnicola 1168), Guerrero & Homrich (1999: 39), Kühner, includes 60 specific taxa, all Karstedt & Stürmer (2008: 1039), Sobestiansky mycorrhizal and mainly associated with (2005: 442), Stijve & de Meijer (1993: 322). Betulaceae and Salicaceae, most species Comments – this species has a global originally described from Europe and later distribution, at least for the Southern revised by Moreau (2005). The number of hemisphere, due to human activities. As endemic species is undoubtedly large. suggested by Wartchow (pers. comm.), A. Although, with our growing knowledge of the muscaria growing under P. taeda corresponds tropical fungi of other continents, the to „ssp. flavivolvata‟. „neotropical‟ and endemic element is bound to become gradually less conspicuous in the lists Amanita multisquamosa Peck of native South American species. Examples of Distribution – SC – Correia Pinto. truly native species of the La Plata region seem Habitat and substrate – plantations of to be Alnicola devia, tigrensis, Pinus taeda L.; terricolous. Paxillus argentinus, Friesula platensis, but Literature – Giachini et al. (2000: 1168, their precise area in South America cannot yet as A. pantherina var. multisquamosa (Peck) be determined (Singer 1953b, Moreau 2005). Jenkins).

63 Mycosphere Doi 10.5943/mycosphere/4/1/5 Comments – Amanita multisquamosa Austroboletus festivus (Singer) Wolfe occurs in mixed coniferous and deciduous Distribution – PR – Paranaguá. forest. In North America it occurs in the Habitat and substrate – „restinga‟; eastern parts and it may have been found in the terricolous. Pacific Northwest as well (Jenkins 1986). This Literature – de Meijer (2001: 112), species appear to have mesophilous Singer et al. (1983: 137), Watling & de Meijer distribution in both Americas (continents). (1997: 232). Comments – The ectomycorrhizal Amanita petalinivolva Wartchow status is based on it fruiting close to Ocotea Distribution – RS – Viamão. pulchella (Nees & Mart.) Mez, but a Habitat – subtropical rain forest mycorrhizal connection has not been „restinga‟; terricolous. determined (Watling & de Meijer 1997). Other Literature – Wartchow et al. (2012b). potential plant hosts are Guettardia angelica Comments – The type for this taxon Martius, Eschweilera ovata (Cambess.) Mart. was collected in Brazil. ex Miers., and in particular, Hymenaea microphylla Barb. Rodr. (Singer et al. 1983). Amanita rubescens Pers. The genus Austroboletus is treated by Tedersoo Distribution – RS – Gramado. et al. (2010) as potentially ectomycorrhizal Habitat and substrate – Pinus based in its phylogenetic placement. plantations; terricolous. Literature – Sobestiansky (2005: 443). exiguus (Singer & Digilio) Comments – It is common throughout Watling much of Europe and eastern North America, Distribution – PR – Piraquara; RS – growing on poor soils as well as in deciduous Sapiranga. or coniferous woodlands. It has also been Habitat and substrate – dense recorded from South Africa, where it is thought ombrophilous forest, mixed ombrophilous to have been accidentally introduced with trees forest; terricolous, occurring on dead as well as imported from Europe (Reid & Eicker 1991). living wood. Literature – de Meijer (2001: 112, Amanita spissa (Fr.) Bertill. 2006: 12, 2008: 44, 130), Putzke et al. (1994: Distribution – RS – locality unknown. 82, as exiguus Singer & Digilio), Habitat and substrate – unknown Watling & de Meijer (1997: 243). habitat; terricolous Comments – This species is putative Literature – Rick (1961a: 301). ectomycorrhizal. It grows on soil, often in Comments – Amanita spissa (and disturbed sites and it appears widely distributed allies) is commonly found in Europe and in tropical America (Halling & Mueller 1998, eastern North America growing with both 2005). Also treated as Gyrodon exiguus (Singer broad leaved as well as coniferous trees et al. (1983) in Watling & de Meijer (1997). (Phillips 2006). Several species from the genus can alter root morphology (Gruhn et al. 1992) or are Amanita strobiliformis (Paulet ex Vittad.) mycorrhizal (Gyrodon). However, to the best Bertill. of our knowledge, no ectomycorrhizal Distribution – RS – locality unknown. association has been described for B. exiguus. Habitat – unknown. The non obligatory ectomycorrhizal status of Literature Rick (1930, 1937, 1961a) – B. exiguous is discussed in de Meijer (2008). all cited in Menolli et al. (2009b). Comments – This species is Boletinellus rompelii (Pat. & Rick) Watling predominantly distributed in Europe. It is more Distribution – PR – Curitiba, Piraquara, common in the Mediterranean region. São José dos Pinhais; RS – Liberato Salzano,

64 Mycosphere Doi 10.5943/mycosphere/4/1/5 Nova Petrópolis, Parecí, São Leopoldo, Literature – Singer (1953a: 102 as Viamão. Braunia alba Rick), Singer (1962: 66). Habitat and substrate – mixed Comments: Brauniella is a South American ombrophilous forest, seasonal semi-deciduous genus and its mycorrhizal status remains alluvial forest; terricolous. unknown. Literature – de Meijer (2001: 112, 2006: 12, 2008: 44, 128), Putzke, et al. [1994: zanchianum (Rick) Baseia & 81, as Gyrodon rompelii (Pat. & Rick) Sing.], Calonge Singer (1953a: 101, as Phylloporus rompelii Distribution – RS – Cachoeira do Sul. Pat. & Rick, 117 and 128, as Gyrodon rompelii Habitat and substrate – unknown (Pat. & Rick) Sing., Singer & Digilio (1957: habitat; roots. 256, as Gyrodon rompelii (Pat. et Rick) Sing.), Literature – Baseia et al. (2006: 114, Singer [1964: 118, as Gyrodon rompelii (Pat. 2007: 278), Cortez (2009: 03), Rick (1961b: & Rick) Sing.], Sobestiansky (2005: 443), 456, as Mitremyces zanchianus Rick). Watling & de Meijer (1997: 243). Comments – This species is considered Comments – This species is putative ectomycorrhizal since its close relative facultatively ectomycorrhizal with Allophylus. C. cinnabarinum was proven to be mycorrhizal It was recorded near Allophylus edulis (A.St.- with (Wilson et al. 2007). However, no Hil. et al.) Hieron. ex Niederl., Trichilia direct evidence exists for the ectomycorrhizal elegans A. Juss. (Watling & de Meijer 1997, de status of C. zanchianum. Meijer 2008), and near an Inga and Allophylus edulis site (Putzke et al. 1994). Singer (1953) Cantharellus guyanensis Mont. referred to this species as ocurring near Acacia Distribution – PR – Campina Grande do woods, while its mycorrhizal condition was Sul, Colombo, São José dos Pinhais. discussed by Singer et al. (1983). Boletinellus Habitat and substrate – dense rompelii ocurrs in South Texas, Mexico and ombrophilous forest, mixed ombrophilous South America (Bessette et al. 2000). No forest; unknown substrate. records have been published for Asia or Literature – de Meijer (2006: 13, 2008: Europe. 372). Comments – Cantharellus guyanensis edulis Bull. sensu Singer et al. (1983) form ECM (a typical Distribution – RS – Gramado. mantle and Hartig-net was observed) with the Habitat and substrate – Pinus roots of Glycoxylon inophyllum (Mart. ex Miq.) plantations; terricolous. Ducke and Aldina heterophylla ex Literature – Sobestiansky (2005: 443). Benth., and possibly also with Psychotria and Comments: Boletus edulis is widely distributed some Sapindaceae (Singer et al. 1983). This in the Northern Hemisphere across Europe, species was recorded in Guiana Shield region Asia, and North America. It does not occur (Henkel et al. 2012). There are no records naturally in the Southern Hemisphere, although available on the distribution of this species for it has been introduced to Southern Africa, other continentes. Australia and . In North America this species was described as Boletus edulis Cantharellus xanthoscyphus R. H. Petersen var. grandedulis with distribution linked to Distribution – PR – Antonina, Quatro Pinus, Quercus and Abies spp. (Arora 2008). Barras. This species forms ectomycorrizae with a range Habitat and substrate – dense of and hardwood species (Agerer 1987– ombrophilous forest; terricolous, occurring on 2006). dead wood. Literature – de Meijer (2001: 112, Brauniella alba (Rick) Rick ex Singer 2006: 13). Distribution – RS – Cacequí, São Leopoldo. piperatus (Bull.) Bataille Habitat and substrate – on sandy earth. Distribution – PR – Curitiba; SC –

65 Mycosphere Doi 10.5943/mycosphere/4/1/5 Joinville, Três Barras. Literature – de Meijer (2006: 14). Habitat and substrate – plantations of P. Comments: puiggarii (Speg.) Corner taeda; terricolous. is also known from the West Indies, South Literature – de Meijer (2001: 113, America, and Australia (Henkel et al. 2006: 13), Giachini et al. (2000: 1168); 2011). Karstedt & Stürmer (2008: 1039, as Chalciporus cf. piperatus (Bull.: Fr.) Bat.); Coltricia barbata Ryvarden & de Meijer Watling & de Meijer (1997: 233). Distribution – PR – Antonina, Comments – This species is potentially Morretes, Paranaguá. ectomycorrhizal (Rinaldi et al 2008, Watling & Habitat and substrate – dense de Meijer 1997). However, Tedersoo et al. ombrophilous forest; unknown substrate. (2010), suggest that there is no evidence that C. Literature – de Meijer (2006: 15). piperatus is ectomycorrhizal. Chalciporus Comments: Coltricia barbata is only known piperatus, also frequently recorded as Boletus from the type locality in the State of Paraná piperatus Bull. ex Fr., is distributed in America (Baltazar et al. 2010). and Europe (Phillips 2006). It occurs in coniferous, and forests, while in Coltricia cinnamomea (Jacq.) Murrill New Zealand it was recorded with Distribution – PR – Curitiba, (Fuhrer & Robinson 1992). Guaraqueçaba, São Mateus do Sul. Habitat and substrate – mixed Chondrogaster angustisporus Giachini, ombrophilous forest, seasonal semi-deciduous Castellano, Trappe & V.L. Oliveira alluvial forest, „restinga‟ seasonal semi- Distribution – SC – Correia Pinto, Três deciduous alluvial forest; terricolous, occurring Barras. also in living wood. Habitat and substrate – plantations of Literature – de Meijer (2001: 113, dunnii Maid.; terricolous with 2006: 15, 2008: 372), Rajchenberg & de Meijer hypogeous sporocarps. (1990: 177, as Coltricia duportii). Literature – Giachini et al. (2000: Comments – Coltricia cinnamomea is a 1168). common and cosmopoliton polypore with Comments – Chondrogaster records in North America (Phillips 2006), angustisporus is a hypogeous ectomycorrhizal South America (Baltazar et al. 2010), Europe described from fruiting bodies collected and Asia (Jülich 1984, Ryvarden 1976). This under Eucalyptus spp. in Brazil, Uruguay and species is recorded as ectomycorrhizal Australia (Giachini et al. 2000). In Europe this (Tedersoo et al. 2010, Thoen 1993). species was recorded in Spain (Lago & Castro 2004) next to Eucalyptus globulus Labill. Coltricia duportii (Pat.) Ryvarden Distribution – PR – Morretes. Chondrogaster pachysporus Maire Habitat and substrate – dense Distribution – RS – Santa Maria. ombrophilous forest; unknown substrate. Habitat and substrate – plantations of Literature – de Meijer (2006: 15). Eucalyptus saligna Sm.; terricolous with Comments – This species is putative hypogeous sporocarps. ectomycorrhizal (Tedersoo et al. 2008). Literature – Sulzbacher et al. (2010: 378). Coltricia perennis (L.) Murrill Comments – This species was recorded Distribution – PR – Lapa. in Spain (Lago & Castro 2004) next to Habitat and substrate – Pinus Eucalyptus sp. plantations; terricolous. Literature – de Meijer (2001: 113, Clavulina puiggarii (Speg.) Corner 2006: 15). Distribution – PR – Antonina. Comments – In Brazil, this species is Habitat and substrate – dense only known from a Pinus plantation in the ombrophilous forest; „restinga‟; terricolous. State of Paraná (Baltazar et al. 2010).

66 Mycosphere Doi 10.5943/mycosphere/4/1/5 Coltriciella oblectabilis (Lloyd) Kotl., Pouzar North Africa, Spain, UK and the USA (Keane & Ryvarden et al. 2000, Moreno-Arroyo et al. 2005). Distribution – PR – Paranaguá. Habitat and substrate – dense Descomyces giachinii Trappe, V.L. Oliveira, ombrophilous forest, „restinga‟; terricolous. Castellano & Claridge Literature – de Meijer (2001: 113, Distribution – SC – Correia Pinto. 2006: 15), Rajchenberg & de Meijer (1990: Habitat and substrate – plantations of 178). Eucalyptus dunnii; terricolous. Comments – Known from Brazil (de Literature – Giachini et al. (2000: Meijer 2001, 2006, Rajchenberg & de Meijer 1168). 1990) and the South-Eastern United States Comments – The species is recorded (Kotlába et al. 1984). Recent records were also from Australia, Brazil, the USA and New published for China (Dai et al. 2001). Zealand (Giachini et al. 2000).

Cortinarius castaneofulvus Cleland Entoloma bloxamii (Berk. & Broome) Sacc. Distribution – SC – Três Barras. Distribution – PR – São José dos Habitat and substrate – plantations of Pinhais, São Mateus do Sul. Eucalyptus dunnii; terricolous. Habitat and substrate – mixed Literature – Giachini et al. (2000: ombrophilous alluvial forest, mixed 1168). ombrophilous montane forest; terricolous. Comments – This species was first Literature – de Meijer (2006: 19, 2008: described from an eucalypt site in Australia 271, 372). (Cleland 1928). The only other collections Comments – In Europe, Asia and North known are from Brazil (Giachini et al. 2000). America this species inhabits unimproved grasslands such as old meadows and hayfields Descomyces albellus (Massee & Rodway) and is known for its indication value of Bougher & Castellano extensive grasslands (Newton et al. 2003). It is Distribution – SC – Correia Pinto; RS – rare throughout Europe (ECCF 2001). Records Santa Maria. were published also from other areas (e.g. Habitat and substrate – plantations of Japan – Kasuya et al. 2010). In general this Eucalyptus dunnii and Eucalyptus spp.; species is regarded as saprobic, although there terricolous. are several representatives of this genus that Literature – Cortez et al. (2008a: 514), can form mycorrhizae (Kasuya et al. 2010) or Giachini et al. (2000: 1168). transitional structures as describes by Agerer & Comments – This species was Waller (1993). Species of Entoloma sect. originally delimited from Hymenogaster by Entoloma are suspected to be ectomycorrhizal Bougher and Castellano in Australia. (Noordeloos 2002, in de Meijer 2008). Additional locations are also known for New Zealand (Bougher & Castellano 1993). The Gloeocantharellus corneri (Singer) Corner species was formerly restricted to Australasia. Distribution – PR – Campina Grande do Nowadays, however, it spreads to areas Sul. covered by Eucalyptus forest (Kirk et al. 2008). Habitat and substrate – dense ombrophilous forest, mixed ombrophilous Descomyces albus (Klotzsch) Bougher & forest; terricolous. Castellano Literature – de Meijer (2001: 112, Distribution – SC – Correia Pinto. 2006: 23, 2008: 372), Watling & de Meijer Habitat and substrate – plantations of (1997: 245). Eucalyptus dunnii; terricolous. Comments – The type for Literature – Giachini et al. (2000: Gloeocantharellus corneri was collected in 1168). Brazil. The genus has been recorded in North Comments – Descomyces albus is and South America (Corner 1969). recorded in Australia, Germany, New Zeland,

67 Mycosphere Doi 10.5943/mycosphere/4/1/5 castaneus (Bull) Quél. Distribution – RS – Santa Maria. Distribution – RS – Pelotas. Habitat and substrate – Eucalyptus Habitat and substrate – planted plantations; terricolous. Quercus; terricolous. Literature – Cortez et al. (2011a: 221). Literature – Putzke et al. (1994: 79). Comments – Hysterangium affine is Comments: The species is commonly found in recorded from Australia (Beaton et al. 1985), oak forests in continental Europe and in eastern North (Zeller & Dodge 1929) and South North America while rare in western North America. It is widespread in Eucalyptus America (Arora 1986, Phillips 2006). plantations (Cortez et al. 2011a).

Hebeloma sacchariolens Quél. Hysterangium australe Speg. Distribution – PR – Almirante Distribution – RS – São Leopoldo. Tamandaré, Rio Negro; RS – Nova Petrópolis. Habitat and substrate – Eucalyptus Habitat and substrate – pasture, Pinus plantations; terricolous. plantations, planted Salix; terricolous. Literature – Rick (1961b: 457). Literature – de Meijer (2001: 113, 2006: 24), Sobestiansky (2005: 447, as Hysterangium inflatum Rodway Hebeloma cf. sacchariolens Quél.). Distribution – RS – Santa Maria. Comments – Hebeloma sacchariolens Habitat and substrate – Eucalyptus is ectomycorrhizal (Marmeisse et al. 1997). In plantations; terricolous. general Hebeloma species are rare in the Literature – Cortez et al. (2011a: 221). tropics but common in temperate Notheren Comments – Hysterangium inflatum is hemisphere (Singer 1986) with Salix and other frequently found in Eucalyptus forest in broadleaf species. Australia (Beaton et al. 1985), New Zealand (Castellano & Beever 1994), France and Hydnodon thelephorus (Lév.) Banker Ecuador (Castellano & Muchovej 1996), Spain Distribution – PR – Antonina, (Lago & Castro 2004, Moreno-Arroyo et al. Campina Grande do Sul, Curitiba, General 2005), United States (Zeller & Dodge 1929), Carneiro, Mandirituba, Morretes, Piraquara, South Europe (Montecchi & Sarasini 2000). São José dos Pinhais, São Mateus do Sul; RS – This species is widespread in Eucalyptus Nova Petrópolis. plantation (Cortez et al. 2011a). Habitat and substrate – dense ombrophilous forest, mixed ombrophilous Hysterangium gardneri E. Fisch. montane forest, gallery forest in area of Distribution – SC – Correia Pinto, Três savanna; terricolous. Barras. Literature – de Meijer (2001: 113, as Habitat and substrate – plantations of Phellodon tenuis Baird, 2006: 24, 2008: 89, Eucalyptus dunnii Maid.; terricolous. 372), Sobestiansky (2005: 447). Literature – Giachini et al. (2000: Comments: This species is putative 1168). ectomycorrhizal. It was noted as Comments – Hysterangium gardneri “ectomycorrhizal?” (Sobestiansky 2005). has a wide distribution, growing especially in Eucalyptus plantations. Nouhra et al. (2008) Hymenogaster vulgaris Tul. & C. Tul. cited this species from Argentina in plantations Distribution – RS – locality unknown. of Eucalyptus camaldulensis and E. cinerea. Habitat and substrate – unknown habitat; terricolous. Inocybe australiensis Cleland & Cheel Literature – Rick (1961b: 456). Distribution – PR – São José dos Comments – In Spain this species Pinhais; SC – Correia Pinto. occurs under Quercus ilex L. (Moreno-Arroyo Habitat and substrate – Eucalyptus et al. 2005). plantations, plantations of Eucalyptus dunnii Maid.; terricolous. Hysterangium affine Massee & Rodway Literature – de Meijer (2001: 113,

68 Mycosphere Doi 10.5943/mycosphere/4/1/5 2006: 26, both as Inocybe cf. australiensis amethystina Cooke Cleland & Cheel), Giachini et al. (2000: 1168). Distribution – SC – Correia Pinto. Comments: Inocybe australiensis occurrs in Habitat and substrate – plantations of Australia (Grgurinovic 1997). Eucalyptus dunnii Maid. and Pinus taeda L.; terricolous. Inocybe curvipes P. Karst. Literature – Giachini et al. (2000: Distribution – PR – Colombo, Curitiba; 1168). RS – Nova Petrópolis, Santa Maria. Comments – Laccaria amethystina is a Habitat and substrate – Pinus common species in most temperate zones of plantations; terricolous. Europe, Asia, Central, South, and eastern North Literature – Cortez & Coelho (2005: America. It forms with a 71), de Meijer (2001: 113, 2006: 26), variety of deciduous and coniferous trees, Sobestiansky (2005: 448), Stijve & de Meijer though it most commonly occurs associated (1993: 321). with Fagales (Laessoe 1998, Mueller 1992). Comments – In Europe, Inocybe curvipes occurs with broadleaf species (birch, poplar, Laccaria bicolor (Maire) P.D. Orton hazel) in rich, humic soils (Stangl 1991). In Distribution – SC – Correia Pinto, Três Africa it is seen close to exotic pine plantations Barras. (Buyck & Eyssartier 1999), and in Australia on Habitat and substrate – plantations of soil under introduced Quercus sp., Pinus Eucalyptus dunnii Maid. and Pinus taeda L.; radiata D. Don., and probably Salix sp. terricolous. (Bougher & Matheny 2011). Literature – Giachini et al. (2000: 1168). Inocybe hyperythra Rick Comments – Laccaria bicolor is found Distribution – RS – São Leopoldo. throughout the temperate zones of the world Habitat and substrate – subtropical (Phillips 2006), and also in Australia (Dunstan forest; terricolous. et al. 1998). Literature – Rick (1961a: 406, as I. hypererythra Rick), Singer (1953a: 94), Singer (Scop.) Cooke var. laccata et al. (1983: 181). Distribution – SC – Correia Pinto, Três Comments – Mycorrhizal. However, Barras; RS – Canela, Ijuí, Pelotas, Porto association and accompanying trees are Alegre, Rio Pardo, Santa Maria, Santa Vitória unknown (Singer et al. 1983). do Palmar, Vera Cruz. Habitat and substrate – Pinus Inocybe violaceolamellata Rick plantations, and plantations of P. taeda L., and Distribution – RS – São Leopoldo. Eucalyptus dunnii Maid.; terricolous. Habitat and substrate – unknown Literature – Giachini et al. (2000: habitat; terricolous, mossy trunk. 1168); Guerrero & Homrich (1999: 41); Putzke Literature – Rick (1961a: 407), Singer (1999: 09; 2003: 286, as Laccaria laccata (1953a: 94). (Scop.:Fr.) Ber. & Br.). Comments – Laccaria laccata is found Labyrinthomyces varius (Rodway) Trappe often in poor soil. It is very common for the Distribution – SC – Correia Pinto. Northern temperate zones. It is ectomycorrhizal Habitat and substrate – plantations of with Pinaceae, and Betulaceae across Eucalyptus dunnii Maid.; terricolous. Europe and North America (Mueller 1991). It Literature – Giachini et al. (2000: is recorded also from Australia, Costa Rica, 1168). South Africa and New Zealand (Arora 1986, Comments – The whole genus is Dunstan et al. 1998). primarilly distributed in Australia in association with Eucalyptus spp. (Kirk et al. Laccaria laccata var. pallidifolia (Peck) Peck 2008). Distribution – SC – Córrego Grande,

69 Mycosphere Doi 10.5943/mycosphere/4/1/5 Rio Vermelho, Três Barras. Habitat and substrate – plantations of Habitat and substrate – Eucalyptus Eucalyptus dunnii Maid. and P. taeda L.; plantations, plantations of Pinus elliottii terricolous. Engelm. and P. taeda L.; terricolous. Literature – Giachini et al. (2000: Literature – Giachini et al. (2000: 1168). 1168). Comments – Laccaria pumila is recorded from Europe, North America (Mueller Laccaria lateritia Malençon 1991, 1992) and from Australasia (Vellinga Distribution – PR – Colombo, Curitiba, 1986). São José dos Pinhais; SC – Três Barras; RS – Barros Cassal, Nova Petrópolis, Porto Alegre, Laccaria tetraspora (Scop.) Fr. Rio Pardo, Salvador do Sul, São Francisco de Distribution – SC – Joinville; RS – São Paula, Santa Cruz do Sul, Sinimbu, Vale do Leopoldo. Sol, Venâncio Aires, Vera Cruz. Habitat and substrate – Pinus plantations; Habitat and substrate – Eucalyptus terricolous. plantations, plantations of E. dunnii; Literature – Karstedt & Stürmer (2008: terricolous. 1039), Putzke (1999: 06, 2003: 280). Literature – de Meijer (2001: 113, 2006: 27, both as Laccaria fraterna (Cooke & Laccaria tortilis (Bolton) Cooke Massee) Pegler), Giachini et al. (2000: 1168), Distribution – SC – Correia Pinto, Três Guerrero & Homrich (1999: 41, as L. fraterna), Barras. Putzke (1999: 08, 2003: 283, both as L. Habitat and substrate – plantations of fraterna), Rick (1961a: 329, as Eucalyptus dunnii Maid. and Pinus taeda L.; laccata Scop.), Singer (1953a: 110, as L. terricolous. tetraspora Sing.), Sobestiansky (2005: 448, as Literature – Giachini et al. (2000: L. fraterna). 1168). Comments – The Laccaria lateritia Comments – Laccaria tortilis is a group is common in Australia forming common species in North America and Europe with a number of plants, but (Phillips 2006), but also recorded from frequently associated with eucalypts Australia and New Zealand (Dunstan et al. (http://www.blueswami.com/laccaria_lateritia. 1998). html). Lactarius argillaceifolius Hesler & A.H. Sm. Laccaria proxima (Boud.) Pat. var. argillaceifolius Distribution – PR – Colombo; SC – Distribution – SC – Joinville, Rio Três Barras; RS – Nova Petrópolis. Vermelho. Habitat and substrate – Pinus plantations, Habitat and substrate – plantations of plantations of P. taeda L.; terricolous. Pinus elliottii Engelm.; terricolous. Literature – de Meijer (2001: 113, Literature – Giachini et al. (2000: 2006: 27), Giachini et al. (2000: 1168), 1168). Sobestiansky (2005: 448). Comments – Lactarius argillaceifolius Comments – Laccaria proxima is ocurrs in North America (USA, Canada) and ectomycorrhizal with in North America. Mexico, mainly associated with oaks (Hesler & It is also common in Europe and was Smith 1979). introduced in New Zealand and Australia (Dunstan et al. 1998). Lactarius braunii Rick Distribution – RS – Porto Alegre, São Laccaria pumila Fayod Leopoldo. Distribution – SC – Correia Pinto, Três Habitat and substrate – unknown Barras. habitat; terricolous. Literature – Raithelhuber (1991: 190), Rick (1961a: 365), Singer (1953a: 84).

70 Mycosphere Doi 10.5943/mycosphere/4/1/5 Lactarius deliciosus (L.) Gray Distribution – SC – Correia Pinto, Três Distribution – PR – Colombo; SC – Barras. Correia Pinto; RS – “highlands” as Habitat and substrate – plantations of Sobestiansky (2005: 448). Pinus taeda L.; terricolous. Habitat and substrate – Pinus Literature – Giachini et al. (2000: plantations and P. taeda L.; terricolous. 1168). Literature – de Meijer (2006: 27), Giachini et al. (2000: 1168), Guerrero & Lactarius russula Rick Homrich (1999: 40), Sobestiansky (2005: 448). Distribution – RS – São Leopoldo. Comments – Lactarius deliciosus is Habitat and substrate – forest; widely distributed in Europe, Asia and North terricolous. America (Hesler & Smith 1979). It has been Literature – Rick (1961a: 364), Singer introduced in Chile, Australia and New (1953a: 83), Singer et al. (1983: 311). Zealand in association with Pinus radiata D. Don. plantations (Phillips 2006, Dunstan et al. Lactarius venezuelanus Dennis 1998). Distribution – PR – Antonina, Campina Grande do Sul, Morretes, Quatro Barras. Lactarius fragilis (Burl.) Hesler & A.H. Sm. Habitat and substrate – dense var. fragilis Distribution: SC – Correia Pinto, ombrophilous forest, mixed ombrophilous Joinville, Três Barras. forest; occurring on dead wood, terricolous. Habitat and substrate – Pinus Literature – Buyck & de Meijer (1999: plantations, plantations of P. elliottii Engelm. 270), de Meijer (2001: 113, 2006: 27, 2008: and P. taeda L.; terricolous. 372). Literature – Giachini et al. (2000: Comments – The species occurs under 1168), Karstedt & Stürmer (2008: 1039, as leguminous trees of campinarana vegetation Lactarius cf. fragilis). (Buyck & de Meijer 1999). The Comments – Lactarius fragilis is found ectomycorrhizal status of this species is not in North America, frequently with Douglas fir clear. It also grows on the bases of living trees (Hesler & Smith 1979). such as Sloanea (Elaeocarpaceae) and Talauma (Magnoliaceae) (Singer et al. 1983). Lactarius paulensis Singer Distribution – PR – locality unknown. Leucogaster braunii Rick Habitat and substrate: dense Distribution – RS – São Leopoldo. ombrophilous forest; terricolous, occurring on Habitat and substrate – Eucalyptus dead wood. plantations; terricolous. Literature – de Meijer (2001: 113). Literature – Rick (1961b: 455). Comments – The genus was recognised Lactarius rufus (Scop.) Fr. as ectomycorrhizal from Australia Distribution – PR – Mandirituba, (http://mycorrhizas.info/ecmf.html). Piraquara; SC – Correia Pinto, Três Barras; RS – Nova Petrópolis. Modicella reniformis (Bres.) Gerd. & Trappe Habitat and substrate – Pinus Distribution – SC – locality unknown. plantations; plantations of P. taeda L.; Habitat and substrate – on or just under terricolous. the leaf cover. Literature – de Meijer (2001: 114, Literature – Thaxter (1922: 321, as 2006: 27), Giachini et al. (2000: 1168, as L. Endogone reniformis Bres.). rufus var. rufus), Sobestiansky (2005: 448). Comments – This species forms Comments – Lactarius rufus is frequent endomycorrhiza (Trappe & Schenck 1982). in Europe and North America, mostly with pines, birch, or spruce (Arora 1986). Neopaxillus echinospermus (Speg.) Singer Distribution – PR – Colombo, General Lactarius rufus var. parvus Hesler & A.H. Sm. Carneiro, São José dos Pinhais; RS – Couto

71 Mycosphere Doi 10.5943/mycosphere/4/1/5 (this locality is not in RS state, but probably in Habitat and substrate – unknown the state of Minas Gerais. Minas Gerais has the habitat; pasture. town of Couto de Magalhães de Minas, as Literature – Rick (1961a: 405). referred by Maria Marchioretto from Instituto Comments – Paxillus alexandri was Anchietano de Pesquisas/UNISINOS (pers. reported from several areas in Europe and com.). North America published as a synonym Habitat and substrate – mixed Clitocybe alexandri (Gillet) Gillet. (Bigelow & ombrophilous montane forest and upper- Smith 1962). montane and dense ombrophilous submontane forests; terricolous. Paxillus involutus (Batsch) Fr. Literature – de Meijer (2001: 112, Distribution – RS – São Salvador, São 2006: 35, 2008: 44, 131, 373), Singer (1950: Leopoldo. 221, as Naucoria echinosperma Speg, 231, Habitat and substrate – subtropical 1953a: 92, as Tubaria crobula Fr., 108, 1964: forest; terricolous. 114), Singer & Digilio (1951: 436), Watling & Literature – Guerrero & Homrich de Meijer (1997: 241). (1999: 40), Rick (1961a: 404), Singer (1964: Comments – Singer et al. (1983) noted 112, as Paxillus aff. involutus). that “this species is terrigenous and there is Comments – Paxillus involutus is reason to believe that it is facultatively widely distributed across the Northern ectomycorrhizal although no convincing Hemisphere (incl. China, India, Japan and ecological or experimental data are available”. North America (Breitenbach & Kränzlin 1991). Based on general observation on the genus It was also recorded in Greenland (Knudson Neopaxillus, de Meijer (2008) stated that it is 2006). It has been introduced to Australia, New not known if it is ectomycorrhizal or not. The Zealand, and South America with European genus is a member of Paxillaceae, associated trees. Paxillus involutus is treated as with Araucaria forest (Issac et al. 1993). It has ectomycorrhizal with a range of coniferous and also been recorded from (Issac et al. deciduous tree species (Taylor et al. 2000). 1993). A similar species (N. dominicanus Angelini & Vizzini) was described recenty Phaeoclavulina cyanocephala (Berk. & M.A. from the Dominican Republic (Vizzini et al. Curtis) Giachini 2012). Distribution – PR – Colombo, Piraquara, Quatro Barras, São José dos Pinhais. Octaviania carnea (Wallr.) Corda. Habitat and substrate – dense Distribution – RS – locality unknown. ombrophilous forest, mixed ombrophilous Habitat and substrate – unknown forest; terricolous. habitat; terricolous. Literature – de Meijer (2001: 112, as Literature – Rick (1961b: 457). grandis (Peck) Corner f. cyanocephala (Berk. & Curtis) R.H. Petersen, Octaviania radicans Rick 2006: 41, 2008: 99, 373, both as Ramaria Distribution – RS – locality unknown. cyanocephala (Berk. & M.A. Curtis) Corner). Habitat and substrate – unknown habitat; dead wood. Phaeoclavulina pancaribbea (R.H. Petersen) Literature – Rick (1961b: 457). Giachini Distribution – PR – Piraquara. Octaviania stillingerii Lloyd Habitat and substrate – dense Distribution – RS – locality unknown. ombrophilous forest, mixed ombrophilous Habitat and substrate – unknown forest; terricolous. habitat; terricolous. Literature – de Meijer (2001: 112, Literature – Rick (1961b: 457). 2006: 41, as Ramaria cf. pancaribbea R.H. Petersen f. caerulea R.H. Petersen). Paxillus alexandri Gillet. Comments – Petersen (1988) described Distribution – RS – locality unknown. Ramaria pancaribbea var. zealandica which

72 Mycosphere Doi 10.5943/mycosphere/4/1/5 was recently renamed as Phaeoclavulina tropical regions, and contains 12 species. The zealandica (R.H. Petersen) Giachini (Giachini species are saprobic, although some may be & Castellano 2011). Phaeoclavulina potentially able to form mycorrhizae with pancaribbea is considered putative exotic trees in certain conditions (Kirk et al. ectomycorrhizal, yet likely to be 2008). ectomycorrhizal as other species from the genus (Agerer & Rambold 2004-2010). tropicus (Rick) Heinem. & Rammeloo Phaeoclavulina subclaviformis (Berk.) Distribution – RS – São Leopoldo, Vera Giachini Cruz; SC – Itapiranga. Distribution – PR – Campina Grande do Habitat and substrate – subtropical Sul, Quatro Barras, Piraquara, São José dos forest; terricolous. Pinhais. Literature – Putzke et al. (1994: 86), Habitat and substrate – dense Singer (1950: 233, 1953a: 101, as Boletus ombrophilous forest, mixed ombrophilous tropicus Rick), Singer & Digilio (1957: 253, as forest, „restinga‟; terricolous. Phaeogyroporus tropicus (Rick) Singer). Literature – de Meijer (2001: 112, Comments – This species is putative 2006: 23, both as Gomphus subclavaeformis ectomycorrhizal, forming mycorrhiza with (Berk.) Corner, de Meijer 2008: 97, 372, as Allophylus spp. (Putzke et al. 1994). There is Gomphus cf. subclaviformis (Berk.) Corner. no evidence that this species forms typical ectomycorrhizae (Singer et al. 1983). Phlebopus beniensis (Singer & Digilio) Heinem. & Rammeloo Phyllobolites miniatus (Rick) Singer Distribution – PR – Paranaguá, Pontal Distribution – RS – locality unknown. do Paraná, Vera Cruz do Oeste. Habitat and substrate – subtropical Habitat and substrate – seasonal semi- forest; terricolous. deciduous submontane/montane forest, Literature – Rick (1961a: 405 as „restinga‟; terricolous. Paxillus miniatus Rick), Singer (1953a: 93, Literature – de Meijer (2001: 112, 1964: 131), Singer et al. (1983: 30). 2006: 36), Watling & de Meijer (1997: 235). Comments – Singer et al. (1983) refer it Comments – Phlebopus beniensis has under Leguminosae trees, as a putative been reported from Argentina, Bolivia, Brazil, ectomycorrhizal species. This is the only Costa Rica, Ecuador and Martinique, in the species in the genus found in tropical South neotropics, and from Liberia in Africa America (Kirk et al. 2008). (Guzmán et al. 2007). In Brazil, specimens are found growing close to Laguncularia racemosa Phylloporia spathulata (Hook.) Ryvarden (L.) C.F. Gaertn. and Hibiscus tiliaceus L. Distribution – PR – Campina Grande do (Watling & Meijer 1997). Its mycorrhizal Sul, Quatro Barras, São Mateus do Sul. association remains doubtful (Singer et al. Habitat and substrate – dense 1983). As Tedersoo et al. (2010), suggested ombrophilous forest, mixed ombrophilous that Phlebopus, as well as Boletinellus and forest, mixed ombrophilous alluvial forest, Phylloboletellus are not ectomycorrhizal but mixed ombrophilous montane forest; facultatively or obligately biotrophic taxa. terricolous and on decayed wood, sometimes on the trunk base of an unidentified living Phlebopus braunii (Bres.) Heinem. Cyatheaceae tree. Distribution – RS – São Leopoldo, Vera Literature – de Meijer (2006: 37, 2008: Cruz. 373). Habitat and substrate – subtropical forest; Comments – This species is putative terricolous. ectomycorrhizal (de Meijer, 2008). Phylloporia Literature – Putzke et al. (1994: 84). spathulata has a pantropical distribution Comments – The genus has a (Wagner & Ryvarden 2002). widespread distribution in subtropical and

73 Mycosphere Doi 10.5943/mycosphere/4/1/5 albus (Cooke & Massee) Priest habitat; occurring on dead wood. Distribution – SC – Correia Pinto. Literature – Rick (1961b: 459). Habitat and substrate – plantations of Eucalyptus dunnii Maid.; terricolous. Ramaria anziana R.H. Petersen Literature – Giachini et al. (2000:1168). Distribution – SC – Correia Pinto, Rio Comments – Pisolithus albus is a Vermelho. eucalyptus symbiont and has been found Habitat and substrate – Eucalyptus predominatly in Europe and North America, plantations; terricolous. even though it originated from Australia Literature – Giachini et al. (2000: (Bougher & Syme 1998, Martin et al. 2002). 1168). Comments – This species is native to Pisolithus arhizus (Scop.) Rauschert Australia and New Zealand (http://www.gwa- Distribution – RS – Capão do Leão, nnon.com/species/Ramaria-anziana). Porto Alegre, Rio Grande, Santa Maria, Viamão. Ramaria geesterani de Meijer Habitat and substrate – Eucalyptus Distribution – PR – São José dos plantations, plantations of Pinus elliottii Pinhais. Engelm., near Tabebuia heptaphylla (Vell.) Habitat and substrate – mixed Toledo; terricolous. ombrophilous montane forest; terricolous. Literature – Guerrero & Homrich (1999: Literature – de Meijer (2008: 44, 101, 48, as Pisolithus tinctorius), Cortez (2011b: 373). 45). Comments – de Meijer (2008: 103) Comments – This species was refered concluded that the ectomycorrhizal status of all by Cortez et al. (2011b) growing in eucalypt ten native species of Ramaria occurring in the and pine (Pinus elliottii Engelm.) plantations as state of Paraná is unknown despite the fact that well near a native Tabebuia heptaphylla (Vell.) the genus is generally considered Toledo. The species is distributed in the ectomycorrhizal in Europe and North America Northern Hemisphere (Europe and North (Agerer & Rambold 2004–2010). America). P. arhizus can be also found in the Southern Hemisphere, growing with introduced Ramaria junquilleovertex R.H. Petersen pine species (Martin et al. 2002). Distribution – SC – Rio Vermelho. Habitat and substrate – Eucalyptus Pisolithus microcarpus (Cooke & Massee) G. plantations; terricolous. Cunn. Literature – Giachini et al. (2000: Distribution – PR – Curitiba, São José 1168). dos Pinhais; SC – Córrego Grande, Rio Comments – Found in New Zealand, as Vermelho, Três Barras. native, on the ground under Leptospermum sp. Habitat and substrate – Eucalyptus and Nothofagus sp. Nationally critically plantations, plantations of E. citriodora Hook., threatened in New Zealand (McKenzie et al. E. dunnii Maid., E. robusta Sm.; terricolous. 2006). Literature – Giachini et al. (2000: 1169), de Meijer (2001: 114, as Pisolithus sp., Ramaria moelleriana var moelleriana (Bres. & 2006: 37, as Pisolithus cf. microcarpus Roum.) Corner (Cooke & Massee) G. Cunn.). Distribution – PR – São José dos Comments – Although now distributed Pinhais. with Eucalyptus plantations worldwide, P. Habitat and substrate – mixed microcarpus is considered to be a native ombrophilous montane forest; decayed Australian taxon (Martin et al. 2002). dicotyledonous branch. Literature – de Meijer (2008: 44, 373). Pisolithus pisiformis (Lloyd) Rick Comment – The species was also Distribution – RS – locality unknown. recorded in Malaysia Habitat and substrate – unknown (http://malaysianfungi.webs.com).

74 Mycosphere Doi 10.5943/mycosphere/4/1/5 Ramaria toxica Toledo & R.H. Petersen Comments – nigrescens Distribution – PR – Curitiba; RS – was found in sandy soil under pines in the Porto Alegre, “highlands and interior of Rio Southeastern USA (Bessette et al. 2007). Grande do Sul” as Sobestiansky (2005: 453). Habitat and substrate – Eucalyptus Rhizopogon roseolus (Corda) Th. Fr. plantations; terricolous. Distribution – PR – Colombo, Curitiba; Literature – de Meijer (2001: 113, SC – Correia Pinto, Três Barras; RS – Itaara, 2006: 41), Guerrero & Homrich (1999: 51), Santa Maria. Sobestiansky (2005: 453). Habitat and substrate – Pinus plantations, Comments – Ramaria toxica is referred plantations of P. taeda L.; terricolous. to Argentina and Brazil under Eucalyptus Literature – Cortez (2011b: 45), de (Toledo & Petersen 1989). Meijer (2001: 113, as R. luteorubescens A.H. Sm.; 2006: 41, as Rhizopogon roseolus (Corda) Redeckera fulvum (Berk. & Broome) C. Th. Fr. sensu Martín (1996) or R. Walker & A. Schüßler luetorubescens A.H. Sm.), Giachini et al. Distribution – (?) RS – locality (2000: 1168, as Rhizopogon rubescens (Tul. & unknown. C. Tul.) Tul. & C. Tul. and Rhizopogon Habitat and substrate – occurring on vulgaris (Vittad.) M. Lange). dead bamboo debris, as well as under the leaf Comments – Rhizopogon roseolus is cover. the species most common in Europe. The Literature – Thaxter (1922: 319, as basidiomes are semihypogeous on sandy or Endogone fulva (Berk. & Broome) Pat. calcareous soil in association with Abies spp., Comment: This species forms endomycorrhiza Picea spp., Pinus spp. and also under Quercus (Trappe & Schenck 1982). spp. (Martín et al. 2000). R. roseolus (synonym Rhizopogon rubescens Tul.), an economically Rhizopogon fuscorubens A. H. Sm. important edible mushroom associated with the Distribution – SC – Correia Pinto, Rio Pinaceae (mostly Pinus sp.), has a global Vermelho, Três Barras. distribution. In the Southern Hemisphere it Habitat and substrate – plantations of P. results from the introduction of exotic trees in elliottii Engelm. and P. taeda L.; terricolous. reforestation programs (Visnovsky et al. 2010). Literature – Giachini et al. (2000: 1168). Rhizopogon zelleri A.H. Sm. Comments – Rhizopogon fuscorubens Distribution – SC – Três Barras. has been found in the United States (Smith & Habitat and substrate – plantations of P. Zeller 1966) under especially pines taeda; terricolous. (Pinus rigida Miller and P. contorta Douglas Literature – Giachini et al. (2000: ex Loudon). Molina & Trappe (1994) 1168). mentioned other probable ectomycorrhizal Comments – Rhizopogon zelleri is hosts: Arbutus menziesii Pursh, Arctostaphylos native to North America where the type was uva-ursi (L) Sprengel, Pinus monticola collected by Zeller in 1966 in pure or mixed Douglas ex D. Don, P. ponderosa Douglas ex pine forests (Martín 1996). Lawson & C. Lawson and P. serotina Michaux. The species was also recorded in Russula consobrina (Fr.) Fr. Spain with an unknown host (Martín 1996). Distribution – PR – São José dos Pinhais; RS – Nova Petrópolis. Rhizopogon nigrescens Coker & Couch Habitat and substrate – Pinus Distribution – SC – Rio Vermelho, Três plantations; terricolous. Barras. Literature – de Meijer (2001: 114, Habitat and substrate – plantations of P. 2006: 42), Sobestiansky (2005: 453). elliottii Engelm. and P. taeda L.; terricolous. Comments – It grows in coniferous Literature – Giachini et al. (2000: forests, in association with Picea. Widespread 1168). in Europe. In the USA known from the

75 Mycosphere Doi 10.5943/mycosphere/4/1/5 Northwest under conifers. Also recorded in Comments – The habitat of Russula China (Zhishu et al. 1993). velenovskyi is under broad-leaved trees and pine. It is found in Europe (Phillips 2006) with Russula puiggarii (Speg.) Singer holarctic, subboreal and suboceanic species. Distribution – PR – Campina Grande do The species is mycorrhizal with Fagales Sul, Quatro Barras, Morretes, Piraquara; RS – (Betula, Fagus, Carpinus, Quercus) and only São Leopoldo. rarely with conifers (Krieglsteiner et al. 2000). Habitat and substrate – dense ombrophilous forest, mixed ombrophilous Sarcodon atroviridis (Morgan) Banker forest; terricolous, sandy soil, occurring on Distribution – PR – Colombo, General dead or rotten wood. Carneiro, Quatro Barras, Morretes, Piraquara, Literature – de Meijer (2001: 113, São José dos Pinhais. 2006: 42, 2008: 373), Rick (1961a: 363, as R. Habitat and substrate – dense pectinata (Bull.) Fr.), Singer & Digilio (1951: ombrophilous forest, mixed ombrophilous 442), Singer (1953a: 83, as R. pectinata (Bull.) montane forest; terricolous. Fr.), Singer et al. (1983: 214). Literature – de Meijer & Baird (1992: Comments – Occurring in all types of 639), de Meijer (2001: 113, 2006: 42, 2008: rain forests, mainly in the tropical lowlands, 373, both as Sarcodon bambusinus (R.E.D. probably non-mycorrhizal or only facultatively Baker & W.T. Dale) Maas Geest. mycorrhizal (Singer et al. 1983). Haug et al. Comments – Ectomycorrhizal with (2005) observed R. puiggarii forming a mantle leguminous trees (Singer et al. 1983). Sarcodon and a Hartig‟s net in roots of Neea sp. atroviridis ocurrs in mixed woods. Found in (Nyctaginaceae). eastern and southeastern North America (Phillips 2006). Collections also known from Russula riograndensis Singer Japan and India but probably introduced since Distribution – RS – São Leopoldo. collected in a botanical garden (Leelavathy et Habitat and substrate – forest; al. 1986). terricolous. Literature – Rick (1961a: 363, as R. albidum Pat. & Trab. subfragilis Rick), Singer (1953a: 81, as R. Distribution – SC – Córrego Grande, subfragilis), Singer et al. (1983: 236, as R. Correia Pinto, Rio Vermelho, Três Barras; RS subfragilis). – Capitão, Minas do Leão, Pareci Novo, Rio Grande, Santa Cruz do Sul, Santa Maria, São Russula theissenii Rick Leopoldo, Viamão. Distribution – PR – Morretes; RS – São Habitat and substrate – Eucalyptus Leopoldo. plantations, plantations of Pinus elliottii Habitat and substrate – dense Engelm. and P. taeda L.; terricolous. ombrophilous forest; terricolous. Literature: Cortez et al. (2008b: 293), Cortez Literature – de Meijer (2001: 113, (2011b: 47), Giachini et al. (2000: 1169), 2006: 42, 2008: 373, both as Russula aff. Guzmán (1970: 301). theissenii Rick), Rick (1961a: 363), Singer Comments – Scleroderma albidum occurs (1953a: 81), Singer et al. (1983: 209). in SW Australia Comments: The position and ecological (http://bie.ala.org.au/species/urn:lsid:biodiversi significance of this species is not clear (Singer ty.org.au:apni.taxon:268116) and Argentina et al. 1983: 211). with introduced pine and eucalypts species (Nouhra et al. 2012). In Brazil this species is Russula velenovskyi Melzer & Zvára frequently collected under Eucalyptus spp., and Distribution – PR – locality unknown. according to Cortez et al. (2011b) is apparently Habitat and substrate – planted associated with several other tree species. Castanea sativa Mill.; terricolous. Literature – de Meijer (2001: 114, Scleroderma areolatum Ehrenb. 2006: 42). Distribution – SC – Córrego Grande,

76 Mycosphere Doi 10.5943/mycosphere/4/1/5 Correia Pinto, Rio Vermelho, Três Barras. Literature – Giachini et al. (2000: Habitat and substrate – Eucalyptus 1169). plantations; terricolous. Comments – Scleroderma cepa is a Literature – Giachini et al. (2000: widespread ectomycorrhizal species on 1169). hardwoods (Quercus, Populus), found in North Comments – This species is America (Coker & Couch 1928), South Africa mycorrhizal with hardwoods and conifers in (Bottomley 1948) and Australia (under moist, shady woods. S. areolatum is widely Eucalyptus - distributed in Europe (Fagus, Pinus) and North http://australianfungi.blogspot.com/2010/07/39 America (Castanea) (Phillips 2006). In -scleroderma-cepa.html). Argentina recorded under planted Pinus spp., Populus sp. and Quercus sp. (Nouhra et al. Scleroderma citrinum Pers. 2012). Distribution – PR – Castro; SC – Córrego Grande, Correia Pinto, Joinville, Rio Scleroderma bougheri Trappe, Castellano & Vermelho, Três Barras; RS – Nova Petrópolis, Giachini Santa Maria, São Francisco de Paula, São Distribution – SC – Correia Pinto, Rio Leopoldo. Vermelho, Três Barras. Habitat and substrate – Pinus Habitat and substrate – Eucalyptus plantations, plantations of P. elliottii Engelm. plantations; terricolous. and P. taeda L.; occurs in living wood, Literature – Giachini et al. (2000: terricolous. 1169). Literature – Cortez (2011b: 47), de Comments – Scleroderma bougheri is a Meijer (2001: 114, 2006: 43), Giachini et al. hypogeous ectomycorrhizal fungus described (2000: 1169), Rick (1961b: 458, as from Australia and Brazil (Giachini et al. Scleroderma vulgare Hornem.), Sobestiansky (2000). (2005: 453). Comments – Scleroderma citrinum is Scleroderma bovista Fr. common in Europe, also recorded in South Distribution – SC – Córrego Grande, America, South Africa and New Zealand Correia Pinto, Joinville, Rio Vermelho, Três (Dunstan et al. 1998). The species is Barras; RS – Porto Alegre, Santa Maria. ectomycorrhizal with a range of broadleaf Habitat and substrate – Eucalyptus species and conifers. plantations, plantations of Pinus elliottii Engelm., and P. taeda L.; terricolous. Scleroderma dictyosporum Pat. Literature – Cortez (2011b: 47), Distribution – RS – Santa Maria. Giachini et al. (2000: 1169), Guzmán (1970: Habitat and substrate – near Acacia 344, the author did not mention the name of the caven (Molina) Molina; terricolous. state but the collector is Rick). Literature – Cortez et al. (2011b: 49). Comments – Scleroderma bovista Comments – Cortez et al. (2011b) occurs in Europe and North America usually in reported this species distributed across dry hardwood forests. In Argentina it has been regions of Africa, Asia and America, as well as observed with exotic planted trees (Nouhra et the subtropical zone. The Brazilian specimens al. 2012). The species was also recorded in were found growing near the base of Acacia New Zealand (Dunstan et al. 1998). In Brazil caven (Molina) Molina, a native species from the species was found growing associated with Southern South America. Authors also reported a native tree (Gomidesia spectabilis [DC.] that in Africa S. dictyosporum has been found Berg.) in the northeast (Gurgel et al. 2008). as an ectomycorrhizal partner of other acacia species, such as A. holosericea A.Cunn. ex G. Scleroderma cepa Pers. Don, and A. mangium Willd. (see also Distribution – SC – Correia Pinto. Founoune et al. 2002, Duponnois et al. 2005, Habitat and substrate – plantations of Sanon et al. 2009). Scleroderma dictyosporum Eucalyptus dunnii Maid.; terricolous. is ectomycorrhizal with Afzelia africana Sm.

77 Mycosphere Doi 10.5943/mycosphere/4/1/5 from West Africa (Ba & Thoen 1990) and with RS – Nova Petrópolis, Santa Maria. Uapaca guineensis Müll. Arg., from Southern Habitat and substrate – pasture, Senegal (Thoen & Ba 1989). It has also been plantations of Pinus elliottii Engelm., described associated with Eucalyptus from Eucalyptus plantations; terricolous. Congo (Garbaye et al. 1988). Literature – Cortez et al. (2011b: 50), de Meijer (2001: 114, 2006: 43, as Scleroderma floridanum Guzmán Scleroderma cf. verrucosum), Giachini et al. Distribution – SC – Córrego Grande, (2000: 1169), Rick (1961b: 458), Sobestiansky Correia Pinto, Rio Vermelho, Três Barras. (2005: 454, as Scleroderma cf. verrucosum). Habitat and substrate – Eucalyptus Comments: Scleroderma verrucosum is a plantations, plantations of Pinus elliottii cosmopolitan species (Guzmán 1970). It is Engelm. and P. taeda L.; terricolous. commonly found in Europe in mixed or Literature – Giachini et al. (2000: broadleaf forests. The species was introduced 1169). to New Zealand with Quercus spp. and Pinus radiata. S. verrucosum is ectomycorrhizal with Scleroderma fuscum (Corda) E. Fisch. Afzelia africana Sm., in West Africa (Ba & Distribution – RS – Porto Alegre, Santa Thoen 1990) and cultivated in Burkina Faso Maria; SC – Correia Pinto, Joinville, Rio with the native vegetation - Uapaca somon Vermelho, Três Barras. (http://senegal.ird.fr). In Brazil it was reported Habitat and substrate – plantations of from several States (Cortez 2011b). Pinus elliottii Engelm. and P. taeda L.; terricolous. Sclerogaster luteocarneus (Bres.) Zeller & Literature – Giachini et al. (2000: C.W. Dodge 1171), Cortez (2011b: 49). Distribution – PR – Fênix; RS – Poço Comments – Scleroderma fuscum forms das Antas. ectomycorrhizae with Pinus spp. and is Habitat and substrate – seasonal semi- reported from several regions, including South deciduous submontane/montane forest; on dead America (Cortez et al. 2011b). wood, terricolous. Literature – de Meijer (2001: 113, Scleroderma laeve Lloyd 2006: 43), Dodge & Zeller (1936: 570), Rick Distribution – RS – Porto Alegre. (1961b: 457, as Octaviania luteocarnea Bres.), Habitat and substrate – Eucalyptus Singer (1962: 54, as Sclerogaster cf. plantations; terricolous. luteocarneus). Literature – Cortez et al. (2011b: 49). Comments – Sclerogaster luteocarneus Comments – This species forms is found in tropical America (Dodge & Zeller ectomycorrhizae with Eucalyptus spp. (Cortez 1936). et al. 2011b) in a non-specific manner (Malajczuk et al. 1982). Setchelliogaster tenuipes (Setch.) Pouzar Distribution – SC – Correia Pinto; RS – Scleroderma uruguayense (Guzmán) Guzmán Guaíba, Viamão. Distribution – SC – Córrego Grande, Habitat and substrate – Eucalyptus Correia Pinto, Rio Vermelho. plantations, plantations of E. dunnii Maid.; Habitat and substrate – plantations of terricolous. Pinus elliottii Engelm. and P. taeda L.; Literature – Cortez et al. (2008a: 514), terricolous. Giachini et al. (2000: 1168). Literature – Giachini et al. (2000: Comments – Setchelliogaster tenuipes 1169). ocurs in Argentina (Nouhra et al. 2008), Australia (Grgurinovic 1997) and Europe Scleroderma verrucosum (Bull.) Pers. (Montecchi & Sarasini 2000). This species is Distribution – PR – Colombo, Curitiba, associated with Eucalyptus spp. (Lago & São José dos Pinhais; SC – Córrego Grande; Castro 2004).

78 Mycosphere Doi 10.5943/mycosphere/4/1/5 cothurnatus Singer plantations, especially young forests. The Distribution – SC – Correia Pinto, Rio species forms mycorrhizal with various Pinus Vermelho, Três Barras; RS – Nova Petrópolis; species, including P. sylvestris L., P. nigra PR – Colombo. J.F.Arnold or P. peuce Griseb. in Europe, and Habitat and substrate – Pinus P. resinosa Sol. ex Aiton and P. strobus L. in plantations, plantations of P. elliottii Engelm., North America. It has also been introduced P. patula Schiede ex Schltdl. & Cham. and P. with pines to Australia, New Zealand and taeda L.; terricolous. Argentina (Moser 1980). Singer (1945, 1949) Literature – de Meijer (2001: 113, and Putzke et al. (1994) refer this species as 2006: 44), Giachini et al. (2000: 1168), occurring only in P. nigra J.F. Arnold, P. pinea Sobestiansky (2005: 454), Watling & de Meijer L., P. silvestris L. and P. resinosa Sol. ex (1997: 236). Aiton. Comments – Occurring in the vicinity of Pinus palustris Mill. or P. taeda L., though Suillus subaureus (Peck) Snell occasionally as far as 11 m from the nearest Distribution – RS – Vera Cruz. pine tree (Singer et al. 1983). S. cothurnatus is Habitat and substrate – Pinus native to North America (Singer 1945). plantations; terricolous. Literature – Putzke et al. (1994: 91). Suillus granulatus (L.) Kuntze Comments – Suillus subaureus is Distribution – PR – Colombo; SC – mycorrhizal with Populus spp., Quercus Córrego Grande, Correia Pinto; RS – Santa berberidifolia Liebm and L. in Cruz do Sul, Venâncio Aires. North America (Roody 2003). Habitat and substrate – Pinus plantations, plantations of P. elliottii Engelm., americana Lloyd P. patula Schiede ex Schltdl. & Cham., P. Distribution – SC – Rio Vermelho, Três taeda L.; terricolous. Barras. Literature – de Meijer (2001: 113, Habitat and substrate – Eucalyptus 2006: 44), Giachini et al. (2000: 1168), Putzke plantations, plantations of Pinus elliottii et al. (1994: 90), Watling & de Meijer (1997: Engelm. and P. taeda L.; terricolous. 238). Literature – Giachini et al. (2000: Comments – Suillus granulatus grows 1169). with Pinus spp. on calcareous and acid soils, Comments – Thelephora americana is sometimes in large numbers. It is native to the ectomycorrhizal with Abies alba Mill. based on Northern hemisphere and common in Britain, a sequence similarity search (Grebenc et al. continental Europe, and North America. It has 2009). The species occurs in Canada, USA, also been introduced to Australia (Phillips Mexico, Japan and China (Corner 1968). 2006), South Africa and New Zealand (Dunstan et al. 1998), probably with Pinus Thelephora griseozonata Cooke radiata D. Don. Distribution – SC – Rio Vermelho. Habitat and substrate – Eucalyptus Suillus luteus (L.) Roussel plantations; terricolous. Distribution – PR – Colombo; RS – Literature – Giachini et al. (2000: Canela, Nova Petrópolis, Pelotas. 1169). Habitat and substrate – Pinus plantatio- Comments – Thelephora griseozonata ns, plantations of P. elliottii Engelm.; is indigenous to New Zealand, but also found terricolous. in North America under pines. Literature – de Meijer (2001: 113, 2006: 44), Putzke et al. (1994: 88), Thelephora palmata (Scop.) Fr. Sobestiansky (2005: 454), Watling, de Meijer Distribution – PR – Pontal do Paraná. (1997: 238). Habitat and substrate: „restinga‟; occurring on Comments – Suillus luteus is found in dead wood. the Northern hemisphere. It is common in pine Literature – Meijer (2001: 113, 2006:

79 Mycosphere Doi 10.5943/mycosphere/4/1/5 44). Literature – Putzke (2003: 579); Rick Comments – Thelephora palmata is (1961a: 304); Singer (1953a: 64); Singer et al. common in Europe and North America (1983: 173), Sulzbacher et al. (2007: 146). (Phillips 2006), mainly in coniferous woods. It Comments: Singer et al. (1983) pointed out the is regarded as common in the North temperate possibility of this species being zone. According to Chen et al. (2001), the ectomycorrhizal since it is a member of the species is ectomycorrhizal. section Sericella (Fr.) Quél. (= Sericeocutis Singer), a section known to contain Ehrh. ectomycorrhizal species. Distribution – PR – General Carneiro; SC – Rio Vermelho, Três Barras. vaccinum (Schaeff.) P. Kumm. Habitat and substrate – Eucalyptus Distribution – RS – locality unknown. plantations, Pinus plantations, plantations of P. Habitat and substrate – forest; unknown taeda L.; terricolous, occurring on dead wood. substrate. Literature: de Meijer (2001: 114, 2006: 44), Literature – Rick (1961: 304). Giachini et al. (2000: 1169), Guerrero & Comments – Tricholoma vaccinum is Homrich (1999: 46). mycorrhizal with conifer, especially Picea spp. Comments – Thelephora terrestris is and Pinus spp. It is common in Europe and known from Europe, North America, Japan, widely distributed in the Northern portion of China, Jamaica, Brazil, Uruguay, Australia, the Northern Hemisphere. In North America it New Zealand and South Africa. It occurs either is common in the Rockies and the Pacific on the ground in coniferous woods, Northwest (Trudell & Ammirati 2009). saprophytic, mycorrhizal on Betula, Picea, Pinus and Eucalyptus (Agerer 1987-2008) or Xerocomus basius de Meijer & Watling semi-parasitic (sensu Corner 1968). Distribution – PR – Campina Grande do Thelephora terrestris was introduced to Sul, Colombo, Curitiba, São José dos Pinhais. Australia during the establishment of exotic Habitat and substrate – dense pines (Corner 1968). Malajczuk et al. (1982) ombrophilous forest, mixed ombrophilous reviewed host-sporocarp records and concluded montane forest; terricolous. that several fungal associates of pine, such as Literature – de Meijer (2008: 44, 125, T. terrestris, were absent from eucalypt stands. 373). Comments – the species Tricholoma atrosquamosum Sacc. ectomycorrhizal status is unknown (de Meijer Distribution – PR – Colombo. 2008). According to Tedersoo et al. (2010), Habitat and substrate – Eucalyptus Xerocomus is an ectomycorrhizal genus. plantations; unknown substrate. Literature – de Meijer (2006: 45). Xerocomus brasiliensis (Rick) Singer Comments – Tricholoma atrosquamo- Distribution – RS – Nova Petrópolis, sum is native in Europe and has been also São Leopoldo. collected in North America (Phillips 2006). Habitat and substrate – open places This species can be found in deciduous (Fagus among Gramineae, subtropical and tropical spp., Quercus spp.) and coniferous (Pinus spp., forest, Eucalyptus plantations; terricolous, on Picea spp.) stands, mainly on calcareous soils. rotten wood. The species was also identified from roots of Literature – Gómez (1996: 61), Putzke the heterotrophic plant Pityopus californicus et al. (1994: 92), Singer & Digilio (1957: 260), (Eastwood) H. F. Copel., and from California, Singer et al. (1983: 65), Singer (1953a: 100, as USA (Bidartondo & Bruns 2001). Boletus brasiliensis Rick, 101, as Phylloporus flavipes Rick), Sobestiansky (2005: 443, as Tricholoma sulphurellum Rick Boletus brasiliensis). Distribution – RS – Esmeralda. Comments – Xerocomus brasiliensis Habitat and substrate – Araucaria forms mycorrhiza with Eucalyptus (Singer et forest; terricolous. al. 1983) or is regarded as a putative

80 Mycosphere Doi 10.5943/mycosphere/4/1/5 ectomycorrhizal fungus (Sobestiansky 2005). Habitat and substrate – mixed According to Tedersoo et al. (2010) the genus ombrophilous forest; terricolous. is ectomycorrhizal. Literature – de Meijer (2006: 14, 2008: 372). Doubtful and undescribed taxa cited from Comments – Clavulina rugosa is Southern Brazil putative ectomycorrhizal (de Meijer 2008). This species is frequently found under conifers Amanita cf. crebresulcata Bas and hardwoods. It is presumably mycorrhizal. Distribution – PR – Antonina. Its close relatives in the genus Clavulina were Habitat and substrate – dense confirmed to be mycorrhizal with Abies in ombrophilous forest; terricolous. Europe (Grebenc et al. 2009). The Dicymbe Literature – de Meijer (2006: 11). forest in South America has several Clavulina Comments – Wartchow & Maia (2007) reported species (Henkel et al. 2012). It is recorded this species from the state of widely distributed in North America and Pernambuco in a tropical rain forest. The Europe (Kuo 2007 – mycorrhizal association was not observed (Bas http://www.mushroomexpert.com/clavulina_ru 1978). gosa.html).

Amanita sp. “A” subg. Amanita sect. Ovigerae Cortinarius spp. /records from de Meijer Distribution – PR – locality unknown. (2001, 2006) Habitat and substrate – „restinga‟; Distribution – PR – Cerro Azul, probably terricolous. Cornélio Procópio. Tunas do Paraná. Literature – de Meijer (2001: 112, as Habitat and substrate – dense Amanita sp. “A” subg. Amanita sect. ombrophilous forest, seasonal semi-deciduous Ovigerae). submontane/montane forest, and mixed ombrophilous forest; terricolous, occurring on Cantharellus cf. cinnabarinus (Schwein.) dead wood. Schwein. Literature – de Meijer (2001: 112, as Distribution – PR – Antonina, Cortinarius sp. A (subg. Sericeocybe sect. Morretes, São José dos Pinhais. Pallidoviolaceae), de Meijer (2001: 112, as Habitat and substrate – mixed Cortinarius sp. B (subg. Sericeocybe sect. ombrophilous forest, dense ombrophilous Sericeocybe), de Meijer (2001: 112, as forest, „restinga‟; standing trunks of living Cortinarius sp. C (subg. Telamonia), de Meijer dicotyledonous trees. (2001: 112, as Cortinarius sp. D (subg. Literature – de Meijer (2001: 112, Telamonia), de Meijer (2001: 112, as 2006: 13, 2008: 44, 372). Cortinarius sp. E (subg. Telamonia), de Meijer Comments – Cantharellus cinnabarinus (2006: 16, as Cortinarius sp. A (subg. is native or common in eastern North America, Sericeocybe sect. Sericeocybe), de Meijer Mexico (Pilz et al. 2003), Europe, Africa and (2006: 17, as Cortinarius sp. B (subg. Asia (Watling et al. 2002). Pilz et al. (2003) Sericeocybe sect. Pallidoviolaceae). also noted that C. cibarius is likely to include multiple cryptic species in different regions. Inocybe aff. cingulatipes (Corner & E. Horak) According to Wartchow et al. (2012a), it is Garrido highly unlikely that the material with the Distribution – PR – São José dos epithet „cinnabarinus‟ occurs naturally in Pinhais. tropical South America. For more information Habitat and substrate – mixed see Buyck et al. (2011) and Buyck & Hofstetter ombrophilous forest; unknown substrate. (2011). Literature – de Meijer (2006: 26).

Clavulina aff. rugosa (Bull.) J. Schröt. Inocybe aff. conspicuospora Buyck & Eyssart. Distribution– PR – Curitiba. Distribution – PR – Fênix.

81 Mycosphere Doi 10.5943/mycosphere/4/1/5 Habitat and substrate – seasonal semi- Lactarius aff. necator (Bull.) Pers. deciduous, alluvial forest; unknown substrate. Distribution – RS – locality unknown. Literature – de Meijer (2006: 26). Habitat and substrate – forest; terricolous. Inocybe aff. crassicystidiata Pegler Literature – Rick (1961a: 364, as L. Distribution – PR – Antonina. adustus Rick), Singer (1953a: 83 as L. Habitat and substrate – dense aductus). ombrophilous forest; unknown substrate. Comments – As Singer (1953) noted, Literature – de Meijer (2006: 26). no specimens are preserved in Rick‟s herbaria.

Inocybe aff. incognita (E. Horak) Garrido Lactarius spp./records from de Meijer (2001) Distribution – PR – Morretes. Distribution – PR – locality unknown. Habitat and substrate – dense Habitat and substrate – dense ombrophilous forest; unknown substrate. ombrophilous forest; terricolous. Literature – de Meijer (2006: 26). Literature – de Meijer (2001: 113), as Lactarius sp. “A” (aff. venezuelanus Dennis), Inocybe cf. matrisdei Singer as Lactarius sp. “B”). Distribution – PR – locality unknown. Habitat and substrate: „restinga‟; terricolous. Phaeoclavulina cf. camellia (Corner) Giachini Literature – de Meijer (2001: 112). Distribution – PR – Campina Grande do Comments – Its association with any Sul, Colombo, Piraquara, São José dos Pinhais. particular host tree and the full characteristics Habitat and substrate: dense ombrophilous of the forest type have not been described forest, mixed ombrophilous forest; terricolous, (Singer et al. 1983). occurring on dead wood. Literature – de Meijer (2001: 112, Inocybe aff. pahangi (Corner & E. Horak) 2006: 41, 2008: 373, all as Ramaria cf. Garrido camellia Corner). Distribution – PR – Guaraqueçaba. Habitat and substrate – „restinga‟; Phaeoclavulina aff. eumorpha (P. Karst.) unknown substrate. Giachini Literature – de Meijer (2006: 26). Distribution – PR – Fênix, Morretes, Paranaguá, Rio Branco do Sul, São José dos Inocybe aff. xerophytica Pegler Pinhais. Distribution – PR – Paranaguá. Habitat and substrate – dense Habitat and substrate – „restinga‟; ombrophilous forest, mixed ombrophilous unknown substrate. forest, seasonal semi-deciduous Literature – de Meijer (2006: 26). submontane/montane forest; unknown substrate. Inocybe spp./records from de Meijer (2001) Literature – de Meijer (2006: 41, as Distribution – PR – locality unknown. Ramaria aff. patagonica (Speg.) Corner). Habitat and substrate – dense ombrophilous forest, „restinga‟, seasonal semi- Phaeoclavulina spp. / records from de Meijer deciduous submontane/montane forest; (2006) terricolous. Distribution – PR: São José dos Literature – de Meijer (2001: 112, as Pinhais. Inocybe sp. A (subg. Inocybe sect. Inocybe), as Habitat and substrate – mixed Inocybe sp. B (subg. Inocybe sect. Inocybe), as ombrophilous forest; terricolous. Inocybe sp. C (subg. Inocybe sect. Literature – de Meijer (2006: 41, as Petiginosae), as Inocybe sp. D (subg. Inocybe Ramaria sp. “A” (subgen. Echinoramaria ser. sect. Petiginosae). Grandisporae).

82 Mycosphere Doi 10.5943/mycosphere/4/1/5 Ramaria cf. aureofulva Corner Pinus-plantations, seasonal semi-deciduous Distribution – PR – São José dos alluvial forest; terricolous. Pinhais. Literature – de Meijer (2001: 112-113, as Habitat and substrate – mixed Tricholoma sp. “A” aff. imbricatum (Fr.: Fr.) ombrophilous forest; unknown substrate. Kummer; Tricholoma sp. “B” subg. Literature – de Meijer (2006: 41, 2008: Tricholoma sect. Genuina, and Tricholoma sp. 44, 373). “C”. subg. Tricholoma sect. Tricholoma.”).

Ramaria cf. reticulata (Berk. & Cooke) Corner Tylopilus spp./records from de Meijer (2001, var. macrospora Corner 2006), Watling & de Meijer (1997) Distribution – PR – Cornélio Procópio. Distribution – PR – locality unknown. Habitat and substrate – seasonal semi- Habitat and substrate – dense deciduous submontane/montane forest; ombrophilous forest; terricolous or growing at unknown substrate. the base of a living tree-fern trunk (Watling & Literature – de Meijer (2006: 41). de Meijer 1997: 240). Literature – de Meijer (2001: 112, Russula cf. dennisii Singer ex Buyck 2006: 45, both as cf. Tylopilus sp.), Watling & Distribution – PR – Antonina. de Meijer (1997: 240). Habitat and substrate – dense Comments – Putative ectomycorrhizal, ombrophilous forest; unknown substrate. based on the fact that several species from the Literature: de Meijer (2006: 42). genus are known to be ectomycorrhizal (Halling et al. 2008, Tedersoo et al. 2010). Russula spp./records from de Meijer (2001) Distribution – PR – locality unknown. Xerocomus aff. coccolobae Pegler Habitat and substrate: dense Distribution – PR – Curitiba, Campina ombrophilous forest, gallery forest in area of Grande do Sul, Mandirituba, São José dos savanna, „restinga‟; terricolous. Pinhais. Literature – de Meijer (2001: 113, as Habitat and substrate – dense Russula sp. “A”, as Russula sp. “B”, as Russula ombrophilous forest, mixed ombrophilous sp. “C”, as Russula sp. “D”). forest; terricolous. Literature – de Meijer (2001: 112, Thelephora spp./records from de Meijer 2006: 45, as Xerocomus sp. “A”), Watling & (2001) de Meijer (1997: 239, as Xerocomus cf. Distribution – PR – locality unknown. coccolobae Pegler). Habitat and substrate – mixed ombrophilous forest; occurring on dead wood. Ectomycorrhizal fungi from Southern Brazil Literature – de Meijer (2001: 113, as in perspective of the origin and plant Thelephora sp. “A”). partners From a list of 144 species and affiliated Tricholoma cf. eucalypticum A. Pearson species, over half (80 species) are considered to Distribution – PR – Curitiba. be introduced from other continents. In Habitat and substrate – Eucalyptus particular, from Santa Catarina and Rio Grande plantations; terricolous. do Sul most of the recorded ECM species were Literature – de Meijer (2001: 113, introduced (Fig. 1). In Parana the dominance of 2006: 45). native ECM species was observed mainly due to de Meijer‟s studies on macromycetes (Buyck Tricholoma spp./records from de Meijer & de Meijer 1999, de Meijer 2001, 2006, 2008, (2001) Watling & de Meijer 1997). That author Distribution – PR – locality unknown. described 37 potentialy ECM species from Habitat and substrate – mixed mixed ombrophilous forests (including ombrophilous forest, Eucalyptus-plantations, Araucaria forest).

83 Mycosphere Doi 10.5943/mycosphere/4/1/5

Fig. 1 – Number of introduced or native ectomycorrhizal fungal species recorded from the states of Parana, Rio Grande do Sul and Santa Catarina (Southern Brazil), and the total for all three states.

The differences among states (Fig.1) Most of the ECM fungi introduced to reflect the focus of research in past decades Southern Brazil originate from North America (Fidalgo 1962, 1968, Putzke 1994), in and Europe. In Southern Brazil these fungi particular the mycological studies of were found associated with Pinus spp. or introduced species plantations (Cortez et al. recorded in association with both Pinus and 2008a, Cortez et al. 2011a, Giachini et al. Eucalyptus. As expected, most of the species 2000, Giachini et al. 2004, Sulzbacher et al. originally known from Australia and New 2010, Putzke 1999; in parts also Cortez et al. Zealand were found associated with Eucalyptus 2008, b, 2009, 2011b, Guzmán 1970, Putzke et spp. plantations. Most of the ectomycorrhizal al. 1994, Rick 1961a, b, Sobestiansky 2005) fungi associated with other broadleaf hosts and the fact that plantations in the whole were also native to Europe or North America country cover about 5.98 million ha. These (Fig. 3). As for origin, the rest of the world plantations where Eucalyptus spp. and Pinus (Asia, Africa) did not contribute significantly spp. dominated as plant ECM partners, to the number of ectomycorrhizal species contributed equally to the number of recorded observed in Southern Brazil. introduced ECM species. Most of the introduced ECM species listed were published Conclusion in Giachini et al. (2000), who in particular In total 144 ectomycorrhizal or putatively surveyed the ECM fungal communities in the ectomycorrhizal taxa were recorded in state of Santa Catarina. Their work resulted in publications for the area of Southern Brazil 49 reported taxa, including three new described (states of Rio Grande do Sul, Santa Catarina species to Southern Brazil. Other introduced and Parana). In this region, climate, soil and plantation tree species (Acacia, Castanea, vegetation are very different from the rest of Populus, Salix) are of minor importance and of the country (Dieckow et al. 2009). Thus, these a low influence to the recorded conditions play a particular role in defining the ectomycorrhizal diversity and were only rarely relation between fungi and host. Over half of available in the analysed references (Fig. 2). introduced plant partners (mainly Eucalyptus

84 Mycosphere Doi 10.5943/mycosphere/4/1/5

Fig. 2 – Number of introduced ectomycorrhizal fungal species recorded and potential ectomycorrhizal plant partners. Data based on published scientific papers summarised for three Southern Brazil states. Some species were found associated with more then one category of ectomycorrhizal partner or the host was unknown.

Fig. 3 – Global distribution/origin of the ectomycorrhizal fungi recorded in Southern Brazil. Species associated with hosts grouped either at the genus level, or at a higher level if available data were too general. The origin data is based on the combination of host and fungus origin.

85 Mycosphere Doi 10.5943/mycosphere/4/1/5 spp. and Pinus spp.) are well-known ECM in 23, 25–73. the Northern hemisphere or Australia and were Agerer R, Rambold G. 2004 – 2010 [first introduced into Southern Brazil by means of posted on 2004-06-01; most recent the hand. Still, several of the listed species, are update: 2009-01-26]. DEEMY – An found only in South America (including Information System for Characterization Southern Brazil). For these, the majority has no and Determination of Ectomycorrhizae. definite record as being ectomycorrhizal and www.deemy.de – München, Germany. requires ECM description and/or molecular Agerer R, Waller K. 1993 – Mycorrhizae of confirmation (sensu Agerer 1991). Therefore, Entoloma saepium: parasitism or there is still a need for further investigation in ? Mycorrhiza 3, 145–154. to provide information on their ECM Arora D. 1986 – Mushrooms Demystified: a status, host partners range and ecological and Comprehensive Guide to the Fleshy seasonal distribution. Additionally, a number Fungi. Berkeley, California: Ten Speed of the species still need more detailed Press. taxonomic revision, including modern Arora D. 2008 – California porcini: Three new morphological analysis, application of taxa, observations on their harvest, and molecular markers and phylogenetic species the tragedy of no commons. Economic concept as a tool to better support their Botany 62, 356–375. placement within a taxonomic classification. Ba AM, Thoen D. 1990 – First syntheses of Future work aims to fill the missing gaps in between Afzelia knowledge of the diversity, mycorrhizal status africana Sm. (Caesalpinioideae) and and general ecology of the species listed here. native fungi West Africa. New Phytologist 114, 99–103. Acknowledgements Baltazar JM, Ryvarden L, Gilbertoni, TB. 2010 We are grateful to National Council for – The genus Coltricia in Brazil: new Scientific and Technological Development records and two new species. Mycologia (CNPq - Brazil) and the Coordination for the 102, 1253–1262. Improvement of Higher Education Personnel Bas C. 1978 – Studies in Amanita. I. Some (CAPES - Brazil) for their financial support. Amazonian species. Persoonia 10, 1–22. The work was co-financed by the Brazil – Bas C, de Meijer AAR. 1993 – Amanita Slovenia bilateral project (BI-BR/11-13-005) grallipes, a new species in Amanita and the Research Programme Forest Biology, subsection Vittadiniae from Southern Ecology and Technology (P4-0107) of the Brazil. Persoonia 15, 345–350. Slovenian Research Agency. Special thanks go Baseia IG, Milanez AI. 2000 – First record of to Adriano Afonso Spielmann and Bruno Scleroderma polyrhizum Pers. Tomio Goto for their valuable suggestions to (Gasteromycetes) from Brazil. Acta the manuscript. We thank to Admir José Botanica Brasilica 14, 181–184. Giachini and Felipe Wartchow for constructive Baseia IG, Milanez AI. 2002 – Rhizopogon comments on the earlier version of the (Gasteromycetes): hypogeous fungi in manuscript. exotic forests from the State of São Paulo, Brazil. Acta Botanica Brasilica 16, References 55–60. Baseia IG, Cortez VG, Calonge FD. 2006 – Agerer R. (1987–2006) – Colour atlas of Rick species revision: Mitremyces Ectomycorrhizae. 1st-13th delivery. zanchianus versus Calostoma Einhorn Verlag, Schwäbisch Gmünd. zanchianum. Mycotaxon 95, 113–116. Agerer R. 1991 – Characterization of Baseia IG, Silva BDB, Leite AG, Maia LC. ectomycorrhiza. In: Norris JR, Read DJ, 2007 – O gênero Calostoma (Boletales, Varma AK, eds. Techniques for the study Agaricomycetidae) em áreas de cerrado e of mycorrhiza. Methods in Microbiology semi-árido no Brasil. Acta Botanica Brasilica 21, 277–280.

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